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LEUCALTIS TENUIS HÔZAWA, 1929 (FIGS 11, 12; TABLE 6), Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on page 713, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Hozawa S. 1929. Studies on the calcareous sponges of Japan. Journal of the Faculty of Sciences, Imperial UniVersity of Tokyo 1: 277 - 389."]}

GENUS ROWELLA GEN.NOV. Z o o b a n k r e g i s t r a t i o n: h t t p s: / / z o o b a n k.o r g / urn:lsid:zoobank.org:act: 5163DADD-AD0B-47B9- A595-369DACFA329C. Type species: Leucettusa simplicissima Burton, 1932. Synonym: Leucetta – Poléjaeff, 1883: 28; non Haeckel, 1872: 117. Leucettusa – Dendy & Row, 1913: 738; Burton, 1963: 50; Borojević et al., 1990: 258; 2002: 1148; Hooper & Wiedenmayer, 1994: 481; Voigt et al., 2012: 11; Klautau et al., 2013: 454; Riesgo et al., 2018: 835. Etymology: Named after Dr Harold Row for his efforts on the systematics of calcareous sponges and for being the co-author of Leucaltidae. Diagnosis: Leucettidae with an amorphous or tubular massive body.The well-developed cortical skeleton can be comprised of single or multiple layers of triactines and/ or tetractines. The choanosomal and the atrial skeletons are comprised of pygmy triactines and tetractines. Aquiferous system leuconoid, syconoid or both. Remarks: Most of the species here assigned to RoƜella have a tubular body but several others can grow ramifications (e.g. R. simplicissima), while some have a pyriform or amorphous shape (e.g. R. pyriformis and R. dyctiogaster). More than the external morphology, every member of this genus has a peculiar spicule category, named primarily as pygmy triactines and/ or tetractines by Poléjaeff (1883: 67). These spicules are present in the choanosomal and atrial regions, in different proportions among species. We opt to follow Poléjaeff’s termination and refer to these spicules as pygmy in our descriptions. It is important to highlight that Leucaltis and RoƜella were not recovered as sister-taxa. Hence, pygmy spicules are different from the small choanosomal and atrial spicules of Leucaltis, differing from what was previously thought (e.g. Borojević et al., 1990). Pygmy spicules may have even a reduced actine (assuming a V-shape). As Poléjaeff (1883) just called them pygmy, not defining them, we are defining them by their size range (from 10.0–83.0 μm length to 2.5–13.0 μm width) and location scattered in the choanosomal and atrial skeletons. The presence of these morphological traits should be considered in the diagnosis of the genus. We could not resolve doubts on Leucettusa soyo (Hôzawa, 1933). This species had its position questioned in previous works (e.g. Dendy & Row, 1913; Rapp, 2004; Cavalcanti et al., 2013) and was allocated in different calcinean genera. Hôzawa’s (1933) description is doubtful, because important morphological characters, such as the presence of anastomosed tubes, were not clearly mentioned. He described a massive sponge, with radial choanocyte chambers around the atrium. Yet, he also mentioned the presence of ‘ascon-tubes’ and described the aquiferous system as ‘Dendy’s type D’ (Dendy, 1891), i.e. solenoid (Cavalcanti & Klautau, 2011). This is the only description available for this species and it is not satisfactory for its identification. We consider this a case of species inquirenda and highlight Cavalcanti et al. ’s (2013) statement that its position is uncertain and new material is necessary to make a proper genus assignment. Scope: Ten previously described ‘ Leucettusa ’ species are being transferred to the new genus RoƜella: R. connectens (Brøndsted, 1926); R. dictyogaster (Row & Hôzawa, 1931); R. haeckeliana (Poléjaeff, 1883); R. imperfecta (Poléjaeff, 1883); R. lancifera (Dendy, 1924); R. mariae (Brøndsted, 1926); R. pyriformis (Brønsted, 1926); R. simplicissima (Burton, 1932); R. tubulosa (Dendy, 1924) and R. Ʋera (Poléjaeff, 1883). Leucettusa nuda (Azevedo et al., 2009) is considered a junior synonym of R. simplicissima. Leucettusa soyo (Hôzawa, 1933) is considered species inquirenda.

ROWELLA MARIAE (BRØNDSTED, 1926) COMB.NOV. (FIG. 24) Synonyms: Leucettusa mariae – Brøndsted, 1926: 302; Burton, 1963: 555; Cavalcanti et al., 2013: 276; Leucascus mariae – Rapp, 2004: 124; Lanna et al., 2007: 1560. Type specimen: Possible holotype – NHMD-89631/ CAL-242 (only pictures were analysed). Type locality: Cape Maria van Diemen, New Zealand (34°24ʹ S, 172°30ʹ E; inaccurate coordinates). Three Kings – North Cape MEOW ecoregion. Description (Fig. 24A): Sponge body tubular, widen at the apical region and narrow at the base. External colour beige in ethanol. Consistency rough to the touch. Outer and atrial surfaces smooth. One big osculum is located on a depression at the apical region of the tube, surrounded by a membrane. Atrial cavity wide and spacious. The aquiferous system was not described by Brøndsted (1926), but it is probably leuconoid, as at that time Leucettusa was considered leuconoid. Skeleton: Cortical skeleton well developed, having approximately the same thickness as the choanosome. It is comprised of several layers of tangential triactines. Choanosomal and atrial skeletons reduced, mostly devoid of spicules, but sparse pygmy triactines and tetractines can be found., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on page 732, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Brondsted HV. 1926. Papers from Dr. Th. Mortensen's Pacific Expedition 1914 - 16. XXXV. Sponges from New Zealand. Part II. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 81: 295 - 331.","Burton M. 1963. A reVision of the classification of the calcareous sponges. London: British Museum (Natural History).","Cavalcanti FF, Rapp HT, Klautau M. 2013. Taxonomic revision of Leucascus Dendy, 1892 (Porifera: Calcarea) with revalidation of Ascoleucetta Dendy & Frederick, 1924 and description of three new species. Zootaxa 3619: 275 - 314.","Rapp HT. 2004. The first record of the genus Leucascus Dendy, 1892 from the Atlantic Ocean, with description of Leucascus lobatus sp. nov. (Porifera, Calcarea, Calcinea) from Greenland. Steenstrupia 28: 1 - 9.","Lanna E, Rossi AL, Cavalcanti FF, Hajdu E, Klautau M. 2007. Calcareous sponges from Sao Paulo State, Brazil (Porifera: Calcarea: Calcinea) with the description of two new species. Journal of the Marine Biological Association of the UK 87: 1553 - 1561."]}

FAMILY LEUCALTIDAE DENDY & ROW, 1913 The most recent diagnosis of Leucaltidae is: Clathrinida with a tubular, ramified or even anastomosed cormus with many oscula, or individualized with a large central atrium and a single osculum. The sponge wall is comprised of a distinct cortex sustained by a well-developed skeleton, and a choanosome. The skeleton of the choanosome and the atrial wall may be absent or comprised of small and dispersed triactines and tetractines (Borojević et al., 2002). According to our molecular and morphological results, only Leucaltis should be included in Leucaltidae. The aquiferous system of Leucaltis does not fit into any of the five types accepted up to date. Past works have considered it analogous to the leuconoid (e.g. Lanna et al., 2009) or syconoid (e.g. Wörheide & Hooper, 1999) aquiferous systems, although the choanocyte chambers are not spherical nor do they have a regular radial orientation around the atrium. Pinacocytes are restricted to the external and atrial surfaces. Dendy (1893) described the ‘ type C’ for this organization of the choanocyte chambers, which are mostly elongated and ramified, commonly fusing with each other (Fig. 3). It must not be mistaken with the solenoid aquiferous system because, in Leucaltis, the choanoderm is present inside the ramified chambers, instead of revesting the internal part of the anastomosed choanocyte tubes. Finally, it differs from the sylleibid aquiferous system because choanocyte chambers do not group around a common exhalant chamber. This kind of aquiferous system is present in five out of the six accepted species (following this work). Leucaltis sambucus is said to be leuconoid, but samples of this species were not found, hence we cannot corroborate if it has indeed a leuconoid aquiferous system. Considering that the aquiferous system of Leucaltis has been referred to as leuconoid in past descriptions, it is possible that L. sambucus is not leuconoid, but has the distinct aquiferous system of the other Leucaltis species. Leucaltis corticata is mentioned as having a similar aquiferous system to Leucaltis clathria (Poléjaeff, 1883: 10): ‘with the extension of the flagellated chambers into larger cavities of irregular outline, which come into contact with each other and anastomose, thus forming still larger sinus-like spaces’. For that reason, we draw attention to the presence of the new aquiferous system of Leucaltis in Leucaltis corticata. We suggest the name kladonoid for this new aquiferous system, derived from the Greek kladotós (= ΚΛαδωτός), branched, ramified. For Leucaltidae, we propose to modify the diagnosis to: Clathrinida with a body comprised of large, ramified and anastomosed tubes; never clathroid. Each tube has a distinct cortex, a choanoderm and a central atrium. The skeleton of the choanosome and atrium is comprised of regular and/or sagittal triactines and tetractines much smaller than the cortical ones. Aquiferous system kladonoid, with elongated and ramified choanocyte chambers., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on pages 697-698, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Dendy A, Row H. 1913. The classification and phylogeny of the calcareous sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London 1913: 704 - 813.","Borojevic R, Boury-Esnault N, Manuel M, Vacelet J. 2002. Order Clathrinida Hartman, 1958. In: Hooper JNA, Van Soest RWM, eds. Systema Porifera: a guide to the classification of sponges. New York: Kluwer Academic / Plenum Publishers, 1141 - 1152.","Lanna E, Cavalcanti FF, Cardoso L, Muricy G, Klautau M. 2009. Taxonomy of calcareous sponges (Porifera, Calcarea) from Potiguar Basin, NE Brazil. Zootaxa 1973: 1 - 27.","Worheide G, Hooper JNA. 1999. Calcarea from the Great Barrier Reef. I: cryptic Calcinea from Heron Island and Wistari Reef (Capricorn-Bunker group). Memoirs of the Queensland Museum 43: 859 - 891.","Dendy A. 1893. Studies on the comparative anatomy of sponges. V. Observations on the structure and classification of the Calcarea Heterocoela. Quarterly Journal of Microscopical Science 35: 159 - 257.","Polejaeff N. 1883. Report on the Calcarea dredged by the HMS Challenger during the years 1873 - 1876. Report on the Scientific Results of the Voyage Challenger (Zoology) 8: 1 - 76."]}

FAMILY LEUCETTIDAE DE LAUBENFELS, 1936 Recent molecular data draw attention to the relationship of the genus we are proposing here, RoƜella gen. nov., with other genera within the family Leucettidae de Laubenfels, 1936 (e.g. Voigt et al., 2012, 2017; Klautau et al., 2013; Riesgo et al., 2018) and we confirm this relationship. The current diagnosis of Leucettidae is: Clathrinida with a solid body. The aquiferous system is always leuconoid. The choanoskeleton is well-developed, in the form of a regular network comprised of triactines and/or tetractines. The cortex is thin and comprised of spicules similar to those of the choanoskeleton (Borojević et al., 2002). Several morphological aspects are similar among RoƜella, Leucetta and Pericharax. The three genera have a solid, massive body and a leuconoid aquiferous system (occasionally syconoid in RoƜella); triactines and tetractines, different from the large cortical category, are present in the choanosomal and atrial skeletons. The cortical region is more developed in RoƜella, with several large layers of spicules, whereas it is thin in Leucetta and Pericharax. To include the genus RoƜella in Leucettidae, the diagnosis of this family is being modified to: Clathrinida with a solid massive body, never anastomosed. Aquiferous system leuconoid or syconoid. The choanoskeleton is disorganized and formed by a network of triactines and/or tetractines of equal size or smaller than those found in the cortex. Cortical spicules are often found also in the choanoskeleton., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on pages 714-715, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["de Laubenfels MW. 1936. A discussion of the sponge fauna of the dry Tortugas in particular and the West Indies in general, with material for a revision of the families and orders of the Porifera. Tortugas Laboratory Occasional Papers 467: 1 - 225.","Voigt O, Wulfing E, Worheide G. 2012. Molecular phylogenetic evaluation of classification and scenarios of character evolution in calcareous sponges (Porifera, Class Calcarea). PLoS One 7: e 334171 - e 334116.","Voigt O, Erpenbeck D, Gonzalez-Pech RA, Al-Aidaroos AM, Berumen ML, Worheide G. 2017. Calcinea of the Red Sea: providing a DNA barcode inventory with description of four new species. Marine BiodiVersity 47: 1009 - 1034.","Klautau M, Azevedo F, Condor-Lujan B, Rapp HT, Collins A, Russo CADM. 2013. A molecular phylogeny for the order Clathrinida rekindles and refines Haeckel's taxonomic proposal for calcareous sponges. IntegratiVe and ComparatiVe Biology 53: 447 - 461.","Riesgo A, Cavalcanti FF, Kenny NJ, Rios P, Cristobo J, Lanna E. 2018. Integrative systematics of clathrinid sponges: morphological, reproductive and phylogenetic characterisation of a new species of Leucetta from Antarctica (Porifera, Calcarea, Calcinea) with notes on the occurrence of flagellated sperm. InVertebrate Systematics 32: 827 - 841.","Borojevic R, Boury-Esnault N, Manuel M, Vacelet J. 2002. Order Clathrinida Hartman, 1958. In: Hooper JNA, Van Soest RWM, eds. Systema Porifera: a guide to the classification of sponges. New York: Kluwer Academic / Plenum Publishers, 1141 - 1152.","Hozawa S. 1929. Studies on the calcareous sponges of Japan. Journal of the Faculty of Sciences, Imperial UniVersity of Tokyo 1: 277 - 389."]}

ROWELLA TUBULOSA (DENDY, 1924) COMB.NOV. (FIGS 26, 27; TABLE 14) Synonyms and citations: Leucettusa tubulosa – Dendy, 1924: 476; Burton, 1963: 50; Kelly et al., 2009: 45; Kelly, 2018: 20. Type specimen: One lectotype (BMNH 1923.10.1.2) and two paralectotypes (BMNH 1923.10.1.3-4). Type locality: Near Three Kings Islands, New Zealand (34°24ʹ S, 172°06ʹ E). Three Kings–North Cape MEOW ecoregion. Description: Sponge body tubular, cylindrical to vase-shaped and ramified at the base. The types are dark brown to black in the outer region because of the fixation with osmium tetroxide. Internally, the sponge is beige (Fig. 26A, B). Incompressible and rough to the touch. Outer surface smooth (Fig. 26C, D). Atrial surface hispid due to the apical actines of the tetractines (Fig. 26E). One osculum is present at the end of each tube. They are simple circular apertures or slits without membrane (Fig. 26B). The body wall is thick (Fig. 26C). The atrial cavity is spacious and the excurrent canals are evident. Aquiferous system leuconoid with spherical to subspherical choanocyte chambers (Fig. 26E). Granular cells are distributed all over the body, more frequently in the cortical and atrial surfaces (Fig. 26F). Skeleton: Cortical skeleton well developed, although not as thick as the choanosome (Fig. 26C). It is comprised of several layers of tangential triactines and tetractines (Fig. 26 C-D). Choanosomal skeleton comprised of pygmy triactines and tetractines, mostly present around canals. Large cortical spicules can also be found in the choanosome (Fig. 26C). Pygmy spicules are also present in the atrium, lying tangentially, with tetractines being more abundant and projecting their apical actine into the atrial cavity (Fig. 26E–F).

LEUCALTIS CORTICATA (HAECKEL, 1872) COMB.NOV. (FIG. 6; TABLE 2) Synonyms: Leucetta corticata – Haeckel, 1872: 129; Poléjaeff, 1883: 29; Aphroceras corticata – Haeckel, 1872: 130; Leucettusa corticata – Dendy & Row, 1913: 739; Burton, 1963: 550; Borojević et al., 1990: 259; 2002: 1148. Type specimen: No type material designated. The specimen used in the original description was not found. Type locality: Cuba, Caribbean Sea (21°58ʹ N, 77°23ʹ W; inaccurate coordinates). Greater Antilles MEOW ecoregion. Material examined: Original description. Description: Sponge formed by a mass of anastomosed tubes (Fig. 6A). Tubes are mostly cylindrical, compressed in few parts. Colour brown after fixation. Outer and atrial surfaces smooth. Osculum not apparent. Aquiferous system described as formed by elongated choanocyte chambers, occasionally fusing REVISION OF LEUCALTIS AND LEUCETTUSA 703 a Haeckel (1872) did not mention which category had these measurements. with each other (Haeckel, 1872: 235; Poléjaeff, 1883: 10), which matches the kladonoid type. Skeleton (Fig. 6B): Cortical skeleton well developed, comprised of several layers of tangential triactines. Choanosomal skeleton reduced, with triactines sparse around the canals. These spicules are also present in the atrium, laying tangentially., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on pages 702-703, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Haeckel E. 1872. Die KalkschMamme. Eine monographie, Vols 1 - 3. Berlin: Reimer.","Polejaeff N. 1883. Report on the Calcarea dredged by the HMS Challenger during the years 1873 - 1876. Report on the Scientific Results of the Voyage Challenger (Zoology) 8: 1 - 76.","Dendy A, Row H. 1913. The classification and phylogeny of the calcareous sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London 1913: 704 - 813.","Burton M. 1963. A reVision of the classification of the calcareous sponges. London: British Museum (Natural History).","Borojevic R, Boury-Esnault N, Vacelet J. 1990. A revision of the intraspecific classification of the subclass Calcinea (Porifera, class Calcarea). Bulletin du Museum National d'Histoire Naturelle. Section A, Zoologie, Biologie et Ecologie Animales 12: 243 - 276."]}

GENUS ROWELLA GEN.NOV. Z o o b a n k r e g i s t r a t i o n: h t t p s: / / z o o b a n k.o r g / urn:lsid:zoobank.org:act: 5163DADD-AD0B-47B9- A595-369DACFA329C. Type species: Leucettusa simplicissima Burton, 1932. Synonym: Leucetta – Poléjaeff, 1883: 28; non Haeckel, 1872: 117. Leucettusa – Dendy & Row, 1913: 738; Burton, 1963: 50; Borojević et al., 1990: 258; 2002: 1148; Hooper & Wiedenmayer, 1994: 481; Voigt et al., 2012: 11; Klautau et al., 2013: 454; Riesgo et al., 2018: 835. Etymology: Named after Dr Harold Row for his efforts on the systematics of calcareous sponges and for being the co-author of Leucaltidae. Diagnosis: Leucettidae with an amorphous or tubular massive body.The well-developed cortical skeleton can be comprised of single or multiple layers of triactines and/ or tetractines. The choanosomal and the atrial skeletons are comprised of pygmy triactines and tetractines. Aquiferous system leuconoid, syconoid or both. Remarks: Most of the species here assigned to RoƜella have a tubular body but several others can grow ramifications (e.g. R. simplicissima), while some have a pyriform or amorphous shape (e.g. R. pyriformis and R. dyctiogaster). More than the external morphology, every member of this genus has a peculiar spicule category, named primarily as pygmy triactines and/ or tetractines by Poléjaeff (1883: 67). These spicules are present in the choanosomal and atrial regions, in different proportions among species. We opt to follow Poléjaeff’s termination and refer to these spicules as pygmy in our descriptions. It is important to highlight that Leucaltis and RoƜella were not recovered as sister-taxa. Hence, pygmy spicules are different from the small choanosomal and atrial spicules of Leucaltis, differing from what was previously thought (e.g. Borojević et al., 1990). Pygmy spicules may have even a reduced actine (assuming a V-shape). As Poléjaeff (1883) just called them pygmy, not defining them, we are defining them by their size range (from 10.0–83.0 μm length to 2.5–13.0 μm width) and location scattered in the choanosomal and atrial skeletons. The presence of these morphological traits should be considered in the diagnosis of the genus. We could not resolve doubts on Leucettusa soyo (Hôzawa, 1933). This species had its position questioned in previous works (e.g. Dendy & Row, 1913; Rapp, 2004; Cavalcanti et al., 2013) and was allocated in different calcinean genera. Hôzawa’s (1933) description is doubtful, because important morphological characters, such as the presence of anastomosed tubes, were not clearly mentioned. He described a massive sponge, with radial choanocyte chambers around the atrium. Yet, he also mentioned the presence of ‘ascon-tubes’ and described the aquiferous system as ‘Dendy’s type D’ (Dendy, 1891), i.e. solenoid (Cavalcanti & Klautau, 2011). This is the only description available for this species and it is not satisfactory for its identification. We consider this a case of species inquirenda and highlight Cavalcanti et al. ’s (2013) statement that its position is uncertain and new material is necessary to make a proper genus assignment. Scope: Ten previously described ‘ Leucettusa ’ species are being transferred to the new genus RoƜella: R. connectens (Brøndsted, 1926); R. dictyogaster (Row & Hôzawa, 1931); R. haeckeliana (Poléjaeff, 1883); R. imperfecta (Poléjaeff, 1883); R. lancifera (Dendy, 1924); R. mariae (Brøndsted, 1926); R. pyriformis (Brønsted, 1926); R. simplicissima (Burton, 1932); R. tubulosa (Dendy, 1924) and R. Ʋera (Poléjaeff, 1883). Leucettusa nuda (Azevedo et al., 2009) is considered a junior synonym of R. simplicissima. Leucettusa soyo (Hôzawa, 1933) is considered species inquirenda., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on pages 715-716, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Burton M. 1932. Sponges. DiscoVery Reports 6: 237 - 392.","Polejaeff N. 1883. Report on the Calcarea dredged by the HMS Challenger during the years 1873 - 1876. Report on the Scientific Results of the Voyage Challenger (Zoology) 8: 1 - 76.","Haeckel E. 1872. Die KalkschMamme. Eine monographie, Vols 1 - 3. Berlin: Reimer.","Dendy A, Row H. 1913. The classification and phylogeny of the calcareous sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London 1913: 704 - 813.","Burton M. 1963. A reVision of the classification of the calcareous sponges. London: British Museum (Natural History).","Borojevic R, Boury-Esnault N, Vacelet J. 1990. A revision of the intraspecific classification of the subclass Calcinea (Porifera, class Calcarea). Bulletin du Museum National d'Histoire Naturelle. Section A, Zoologie, Biologie et Ecologie Animales 12: 243 - 276.","Hooper JNA, Wiedenmayer F. 1994. Porifera. In: Wells A, ed. Zoological catalogue of Australia, Vol. 12. Melbourne: CSIRO.","Voigt O, Wulfing E, Worheide G. 2012. Molecular phylogenetic evaluation of classification and scenarios of character evolution in calcareous sponges (Porifera, Class Calcarea). PLoS One 7: e 334171 - e 334116.","Klautau M, Azevedo F, Condor-Lujan B, Rapp HT, Collins A, Russo CADM. 2013. A molecular phylogeny for the order Clathrinida rekindles and refines Haeckel's taxonomic proposal for calcareous sponges. IntegratiVe and ComparatiVe Biology 53: 447 - 461.","Riesgo A, Cavalcanti FF, Kenny NJ, Rios P, Cristobo J, Lanna E. 2018. Integrative systematics of clathrinid sponges: morphological, reproductive and phylogenetic characterisation of a new species of Leucetta from Antarctica (Porifera, Calcarea, Calcinea) with notes on the occurrence of flagellated sperm. InVertebrate Systematics 32: 827 - 841.","Hozawa S. 1929. Studies on the calcareous sponges of Japan. Journal of the Faculty of Sciences, Imperial UniVersity of Tokyo 1: 277 - 389.","Hozawa S. 1933. Report on the calcareous sponges obtained by the survey of the Continental Shelf bordering on Japan. Science Reports of the Tohoku Imperial UniVersity 8: 1 - 20.","Rapp HT. 2004. The first record of the genus Leucascus Dendy, 1892 from the Atlantic Ocean, with description of Leucascus lobatus sp. nov. (Porifera, Calcarea, Calcinea) from Greenland. Steenstrupia 28: 1 - 9.","Cavalcanti FF, Rapp HT, Klautau M. 2013. Taxonomic revision of Leucascus Dendy, 1892 (Porifera: Calcarea) with revalidation of Ascoleucetta Dendy & Frederick, 1924 and description of three new species. Zootaxa 3619: 275 - 314.","Dendy A. 1891. A monograph of the Victorian sponges. I. - The organisation and classification of the Calcarea Homocoela, with description of the Victorian species. Transactions of the Royal Society of Victoria 3: 1 - 81.","Cavalcanti FF, Klautau M. 2011. Solenoid: a new aquiferous system to Porifera. Zoomorphology 130: 255 - 260.","Brondsted HV. 1926. Papers from Dr. Th. Mortensen's Pacific Expedition 1914 - 16. XXXV. Sponges from New Zealand. Part II. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 81: 295 - 331.","Row RWH, Hozawa S. 1931. Report on the Calcarea obtained by the Hamburg South-West Australian Expedition of 1905. Science Reports of the Tohoku Imperial UniVersity (4) 6: 727 - 809, pls XIX - XXI.","Dendy A. 1924. Porifera. Part I. Non-Antarctic sponges. Natural History Report. British Antarctic (Terra NoVa) Expedition, 1910 (Zoology) 6: 269 - 392, pls I - XV.","Azevedo F, Hajdu E, Willenz P, Klautau M. 2009. New records of Calcareous sponges (Porifera, Calcarea) from the Chilean coast. Zootaxa 2072: 1 - 30."]}

ROWELLA SIMPLICISSIMA (BURTON, 1932) COMB.NOV. (FIGS 13, 14; TABLE 7) Synonyms: Leucettusa simplicissima – Burton, 1932: 261; Burton, 1963: 51. Leucettusa simplissima – Borojević et al., 1990: 257 (misspelling). Leucaltis nuda – Azevedo et al., 2009: 9; Leucettusa nuda – Klautau et al., 2013: 449; Fontana et al., 2018: 336 Lopes et al., 2018a: 59; Riesgo et al., 2018: 830. Type specimen: Holotype (BMNH 1928.2.15.35). Type locality: Off Sea Lion Island, East Falkland Island, South Atlantic (52°33ʹ S, 59°04ʹ W). Malvinas / Falklands and Araucanian MEOW ecoregion. Description: Sponge tubular or ramified with cylindrical tubes (Fig. 13A). It grows parallel to the substrate and raises the tubes that bear oscula. Colour white in life and creamy-white to beige in ethanol (Fig. 13A, B). Consistency slightly compressible and friable to the touch. Outer surface smooth, while the atrial surface is hispid due to the apical actines of tetractines. Single, circular and naked oscula are present in each tube. Body wall overall thin. Atrial cavity wide and spacious, many excurrent canals are evident (Fig. 13B). Aquiferous system syconoid, with elongated choanocyte chambers (Fig. 13C). Reproductive elements (oocytes) are present in the MNRJ specimens. Skeleton: Oscular margin has a transitional skeleton comprised of sagittal triactines and rare tetractines, gradually becoming regular as the body wall thickens. Cortical skeleton well developed and comprised of several layers of tangential triactines and fewer tetractines (Fig. 13D, E). Choanosomal skeleton with sparse pygmy triactines and tetractines, mostly present around the exhalant canals. The atrial skeleton is not well developed, often with pygmy tangential triactines and tetractines (Fig. 13F). Overall, the pygmy triactines are less abundant than the pygmy tetractines. Spicules (Table 7): Cortical triactines (Fig. 14A). Regular. Variable sizes. Actines are mostly cylindrical, straight, with blunt to sharp tips. Sometimes there is a constriction near the tip. Size – 274.0 (± 69.0) μm length/19.0 (± 5.0) μm width. Cortical tetractines (Fig. 14B). Regular. Variable sizes. Basal actines are cylindrical to conical, straight, with blunt to sharp tips. The apical actine is long, cylindrical to slightly conical, straight, smooth, with blunt tips. Its tip can be straight, spiral or curved. Tetractines are less abundant and larger than the 718 M. V. LOPES and M. KLAUTAU triactines. Size – basal actines: 315.0 (± 137.0) μm length/35.0 (± 10.0) μm width; apical actine: 454.0 (± 132.0) μm length/36.0 (± 10.8) μm width. Choanosomal and atrial triactines (Fig. 14C). Regular. Pygmy. Actines are conical, straight with blunt to sharp tips. Sometimes they become cylindrical and are thicker near the tip. They are less abundant than the tetractines. Size – 35.5 (± 5.2) μm length/7.5 (± 1.7) μm width. Choanosomal and atrial tetractines (Fig. 14D). Regular. Pygmy. Basal actines are conical, straight with blunt tips. Sometimes they become cylindrical and are thicker near the tip. The apical actine is long, thick, slightly conical, smooth, with sharp tips and slightly curved at the distal part. Size – basal actines: 33.8 (± 4.9) μm length/6.9 (± 1.1) μm width; apical actine: 63.8 (± 9.3) μm length/8.1 (± 1.1) μm width. Ecology: Coarse sand, shell and stone grounds. The specimens from Chile were found in vertical substrate. Depth range 22 to 75 m. Geographical distribution (MEOW ecoregion): Malvinas /Falklands and Araucanian (Sea Lion Island, Falkland Islands, South Atlantic – Burton, 1932), Chiloense (Reñihue Fjord, Central-South Chile – Azevedo et al., 2009). Remarks: Burton described Leucettusa simplicissima in 1932 based on a couple of specimens collected in the Falklands during the Discovery Expedition. He stated that the diagnostic character of this species was the underdeveloped atrial skeleton, with occasional pygmy triactines. Indeed, the atrial skeleton has scattered pygmy spicules, nonetheless, pygmy tetractines are also present and they are even more abundant than the pygmy triactines. Despite this, the pygmy tetractines were overlooked in previous works (Burton, 1932, 1963; Borojević et al., 1990, 2002). The authors also overlooked the cortical tetractines, which are numerous, although less abundant than the cortical triactines. aTaken from Azevedo et al. (2009). H, holotype. The main character that separates R. simplicissima from other species of RoƜella is the syconoid aquiferous system, although two other species currently classified as Leucettusa present this kind of aquiferous system: L. Ʋera (now R. Ʋera) from Kerguelen and L. nuda from the Chilean fjords. The aquiferous system of R. Ʋera can be easely differentiated from that of R. simplicissima because its elongated chambers are restricted only to the region underneath the cortex, while in the lower parts of the choanosome, closer to the atrium, chambers are spherical (Poléjaeff, 1883; Borojević & Grua, 1965). For Leucettusa nuda the story is different. That species was originally described as Leucaltis because of the organisation of the aquiferous system (Azevedo et al., 2009) and, later, with molecular data, it was transferred to Leucettusa (Klautau et al., 2013). Leucettusa nuda and R. simplicissima occur in the Magellanic Province (Spalding et al., 2007) and both have the same morphological characters: syconoid aquiferous system, triactines and tetractines in the cortical skeleton and pygmy triactines and tetractines in the choanosome and atrium. Moreover, the shape and size of the spicules are similar in both species (Table 7). Taking this into consideration, we conclude that L. nuda is a junior synonym of R. simplicissima. In the original description of R. simplicissima, it was stated that the only difference between this species and R. haeckeliana is the presence of tetractines in the cortex of the latter (Burton, 1932). However, after re-analysing the holotype skeleton of R. simplicissima, we found cortical tetractines in it. In fact, the two species are morphologically similar, but they differ from each other by the aquiferous system, which is syconoid in R. simplicissima and leuconoid in R. haeckeliana, and by the shape of the pygmy spicules, with basal actines always straight in R. simplicissima and casually curved in the osculum in R. haeckeliana. Moreover, the pygmy spicules of R. haeckeliana are broadened at the distal portion of the actine. The cortical tetractines are abundant in R. simplicissima and rare in R. haeckeliana. Therefore, we consider them distinct species, although we have found that specimens from the Falkland Islands identified as ‘ Leucettusa ’ haeckeliana (Burton, 1932) are probably R. simplicissima (see Remarks under R. haeckeliana)., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on pages 716-719, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Burton M. 1932. Sponges. DiscoVery Reports 6: 237 - 392.","Burton M. 1963. A reVision of the classification of the calcareous sponges. London: British Museum (Natural History).","Borojevic R, Boury-Esnault N, Vacelet J. 1990. A revision of the intraspecific classification of the subclass Calcinea (Porifera, class Calcarea). Bulletin du Museum National d'Histoire Naturelle. Section A, Zoologie, Biologie et Ecologie Animales 12: 243 - 276.","Azevedo F, Hajdu E, Willenz P, Klautau M. 2009. New records of Calcareous sponges (Porifera, Calcarea) from the Chilean coast. Zootaxa 2072: 1 - 30.","Klautau M, Azevedo F, Condor-Lujan B, Rapp HT, Collins A, Russo CADM. 2013. A molecular phylogeny for the order Clathrinida rekindles and refines Haeckel's taxonomic proposal for calcareous sponges. IntegratiVe and ComparatiVe Biology 53: 447 - 461.","Fontana T, Condor-Lujan B, Azevedo F, Perez T, Klautau M. 2018. Diversity and distribution of calcareous sponges (subclass Calcinea) from Martinique. Zootaxa 4410: 331 - 369.","Lopes MV, Condor-Lujan B, Azevedo F, Perez T, Klautau M. 2018 a. A new genus of calcareous sponge discovered in the Caribbean Sea: Bidderia gen. nov. (Porifera, Calcarea, Calcinea). Zootaxa 4526: 56 - 70.","Riesgo A, Cavalcanti FF, Kenny NJ, Rios P, Cristobo J, Lanna E. 2018. Integrative systematics of clathrinid sponges: morphological, reproductive and phylogenetic characterisation of a new species of Leucetta from Antarctica (Porifera, Calcarea, Calcinea) with notes on the occurrence of flagellated sperm. InVertebrate Systematics 32: 827 - 841.","Polejaeff N. 1883. Report on the Calcarea dredged by the HMS Challenger during the years 1873 - 1876. Report on the Scientific Results of the Voyage Challenger (Zoology) 8: 1 - 76.","Borojevic R, Grua P. 1965. Eponges calcaires de Kerguelen. Systematique et ecologie. ArchiVes de Zoologie Experimentale et Generale 105: 1 - 29.","Spalding MD, Foz HE, Allen GR, Davidson N, Ferdana ZA, Finlayson M, Halpern BS, Jorge MA, Lombana A, Lourie SA, Martin KD, McManus E, Molnar J, Recchia CA, Robertson J. 2007. Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. Bioscience 57: 573 - 583."]}

ROWELLA VERA (POLÉJAEFF, 1883) COMB.NOV. (FIG. 28; TABLE 15) Synonyms: Leucetta Ʋera – Poléjaeff, 1883: 68; Leucettusa Ʋera – Dendy & Row, 1913: 739; Burton, 1963: 557; Borojević & Grua, 1965: 11. Leucilla Ʋera – Fell, 1950: 10. Type specimen: Holotype (BMNH 1884.4.22.60). Type locality: Off Kerguelen Island, South Indian Ocean (49°30ʹ S, 70°30ʹ E). Kerguelen Islands MEOW ecoregion. Description: The holotype is only a small fragment (Fig. 28A, B). Colour light beige (surface) and brown (mesohyl) in ethanol. Compressible and soft to the touch. Outer surface smooth. Atrial surface hispid due to the apical actines of tetractines. Osculum could not be seen. Body wall thick. Atrial cavity spacious with conspicuous excurrent canals. Aquiferous system peculiar: near the cortex (one-third of the body wall) the choanocyte chambers are elongated and cylindrical (syconoid), while near the atrial surface (other twothirds) they are subspherical (leuconoid) (Fig. 28C). Large granular cells found surrounding the canals. Few young oocytes present in the holotype. Skeleton: Oscular margin with sagittal triactines that gradually become regular as the body wall thickens. Cortical skeleton well developed, although not as thick as the choanosome. It is comprised of several layers of tangential triactines and tetractines (Fig. 28C). The apical actine of the cortical tetractines crosses the choanosome, sometimes penetrating the atrium. The choanosomal skeleton is comprised of pygmy triactines and tetractines, mostly present around canals. These spicules are also present in the atrium, laying tangentially, with pygmy tetractines being more abundant and projecting their apical actine into the atrial cavity., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on page 738, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Polejaeff N. 1883. Report on the Calcarea dredged by the HMS Challenger during the years 1873 - 1876. Report on the Scientific Results of the Voyage Challenger (Zoology) 8: 1 - 76.","Dendy A, Row H. 1913. The classification and phylogeny of the calcareous sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London 1913: 704 - 813.","Burton M. 1963. A reVision of the classification of the calcareous sponges. London: British Museum (Natural History).","Borojevic R, Grua P. 1965. Eponges calcaires de Kerguelen. Systematique et ecologie. ArchiVes de Zoologie Experimentale et Generale 105: 1 - 29.","Fell HB. 1950. The Kirk collection of sponges (Porifera) in the Zoology Museum, Victoria University College. Zoology Publications from Victoria UniVersity of Wellington 4: 1 - 12."]}

ROWELLA DICTYOGASTER (ROW & HÔZAWA, 1931) COMB.NOV. (FIG. 17; TABLE 9) Synonyms: Leucettusa dictyogaster – Row & Hôzawa, 1931: 751; Burton, 1963: 576; non Leucettusa dictyogaster – Dendy & Row, 1913: 739; Dendy, 1924: 280 (nomen nudum). Type specimen: Holotype (Tokyo Science Faculty Spec. No. BC) possibly lost. Type locality: Koombana Bay, Bunbury, Western Australia (22°29ʹ S, 113°57ʹ E). Leeuwin MEOW ecoregion. Material examined: Original description. Description: Sponge massive and amorphous with a single osculum atop (Fig. 17A). Colour greyish in ethanol. Consistency hard outside and soft inside. Outer surface smooth and uneven with few depressions. Osculum oval with a membrane. Body wall thick, tapering towards the top and the base of the REVISION OF LEUCALTIS AND LEUCETTUSA 723 sponge. Atrial cavity narrow and slightly hispid due to the apical actine of the atrial tetractines. Many visible excurrent canals reach the atrial surface. Aquiferous system leuconoid with oval to spherical choanocyte chambers and lacunar canals. Skeleton: The oscular margin has sagittal triactines, which possibly derive from the regular cortical triactines (Fig. 17B). Cortical skeleton well developed, comprised of several layers of tangential triactines and microdiactines. No spicules were mentioned for the choanosomal region. Atrial skeleton comprised of pygmy tetractines. Spicules (Table 9): Cortical microdiactines (Fig. 17C). Cylindrical, sinuous and curved. Both tips are sharp; however, one is fusiform, while the other is arrow-shaped. Size – 30.0–130.0 μm length/6.0–10.0 μm width. Cortical triactines (Fig. 17D). Regular. Actines are cylindrical, with sharp tips. Size – 380.0–580.0 μm length/30.0–50.0 μm width. Atrial tetractines (Fig. 17E). Sagittal. Pygmy. Basal actines are conical, with sharp tips. The paired actines are slightly shorter than the unpaired one. The apical actine is long, conical, smooth, slightly curved, with sharp tips. Size – paired actine: 20.0–40.0 μm length/6.0 μm width; unpaired actine: 30.0–50.0 μm length/8.0 μm width; apical actine: 100.0–120.0 μm length/8.0 μm width. Ecology: No information available. Geographical distribution (MEOW ecoregion): Leeuwin (Koombana Bay – originally Bunbury Bay – Bunbury, Western Australia – Row & Hôzawa, 1931). Remarks: Row & Hôzawa (1931) described Leucettusa dictyogaster after five specimens collected in Bunbury Bay, Western Australia. Curiously, prior to the species description, this species was mentioned in the systematic review of Dendy & Row (1913) without any description other than the aquiferous system. The specimens used for the species description are lost but, since it possesses a massive body and pygmy spicules, we propose to transfer it to the new genus RoƜella. One unique character of R. dictyogaster is the presence of microdiactines in the cortical skeleton. These spicules are commonly found in the subclass Calcaronea and rare in calcinean species (e.g. Ascandra minchini Borojević, 1966). Although they could be considered a contamination, they were mentioned by Row & Hôzawa (1931) as being present in all the five specimens analysed. Therefore, we consider microdiactines part of this species. Disregarding these spicules, the skeleton of R. dictyogaster is simple. The only categories found are the cortical triactines and the atrial tetractines. Oscular triactines are considered modified cortical triactines. We have confirmed that some categories, mostly pygmy spicules, are often overlooked in this group of species possibly because of the size and, sometimes, low abundancy. It is also possible that R. dictyogaster has rare tetractines in the cortex and scattered pygmy spicules in the choanosome. It is necessary to re-analyse specimens from Bunbury to check these spicules and properly perform taxonomic comparisons with other RoƜella species., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on pages 722-724, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Row RWH, Hozawa S. 1931. Report on the Calcarea obtained by the Hamburg South-West Australian Expedition of 1905. Science Reports of the Tohoku Imperial UniVersity (4) 6: 727 - 809, pls XIX - XXI.","Burton M. 1963. A reVision of the classification of the calcareous sponges. London: British Museum (Natural History).","Dendy A, Row H. 1913. The classification and phylogeny of the calcareous sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London 1913: 704 - 813.","Dendy A. 1924. Porifera. Part I. Non-Antarctic sponges. Natural History Report. British Antarctic (Terra NoVa) Expedition, 1910 (Zoology) 6: 269 - 392, pls I - XV.","Borojevic R. 1966. Eponges calcaires des cotes de France. II. Le genre Ascandra Haeckel emend. ArchiVes de Zoologie Experimentale et Generale 107: 357 - 367."]}

LEUCALTIS NODUSGORDII (POLÉJAEFF, 1883) (FIGS 9, 10; TABLE 4), Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on page 708, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Polejaeff N. 1883. Report on the Calcarea dredged by the HMS Challenger during the years 1873 - 1876. Report on the Scientific Results of the Voyage Challenger (Zoology) 8: 1 - 76."]}

LEUCALTIS CLATHRIA HAECKEL, 1872 (FIGS 4, 5; TABLE 1) Synonyms: Artynas clathria – Haeckel, 1872: 159. Leucaltis clathria – Haeckel, 1872: 159; Arndt, 1941: 46; Burton, 1963: 598; Borojević & Peixinho, 1976: 1002; Borojević et al., 2002: 1148; Dohrmann et al., 2006: 832; Muricy et al., 2008: 131; Lanna et al., 2009: 13; Muricy et al., 2011: 35; Klautau et al., 2013: 449; Van Soest & De Voogd, 2015: 43; Cóndor-Luján & Klautau, 2016: 232; Van Soest, 2017: 198, Cóndor-Luján et al., 2018: 61; Fontana et al., 2018: 354; Lopes et al., 2018a: 59; 2018b: 139; non Dendy, 1913: 16; Leucetta clathria – Poléjaeff, 1883: 29; Heteropegma nodus gordii – Poléjaeff, 1883: 45 (in part, Bermuda material only); Heteropegma nodus-gordii – Hanitsch, 1895: 209; Amphoriscus nodusgordii – Breitfuss, 1898: 90; Leucaltis sp. – Thacker et al., 2013: 385. Type specimen: Fragment (MNHN. LBIM. C 1968.667) and slides (BMNH 1956.4.26.42) of Haeckel’s specimen used in the original description (holotype). Type locality: Florida, United States of America (27°54ʹ N, 80°01ʹ W; inaccurate coordinates). Floridian ecoregion. Description: Sponge body tubular and repent with some anastomosis (Fig. 4A, B). The anastomosis is variable; it can be loose and sparse or dense, with tubes closely attached to each other. Tubes are mostly cylindrical, but commonly compressed in few parts, giving the body a folded appearance. Consistency harsh to the touch, relatively firm and friable. Colour white or pink in life and white, beige to light brown or pink after fixation (Fig. 4A, B). Outer surface smooth. Body wall thin (Fig. 4C). Atrial cavity wide and spacious. Atrial surface slightly hispid due to the apical actines of cortical and atrial tetractines. Oscula are few, simple circular apertures without ornamentation (Fig. 5A). The aquiferous system represents the new type, kladonoid, with choanocyte chambers mostly elongated and ramified, commonly these ramifications fuse with each other (Fig. 4C). Skeleton: The oscular margin has a transitional skeleton comprised of large sagittal triactines and tetractines, gradually becoming regular as the body wall thickens. Cortical skeleton well developed, with several layers of tangential triactines and tetractines (Fig. 4D). Choanosomal skeleton reduced, with many regular triactines and tetractines sparse around the canals (Fig. 4E). Excurrent canals and atrial skeleton comprised of sagittal triactines and tetractines, lying tangentially, with the apical actines pointing into the atrial cavity (Fig. 4F)., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on page 699, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Haeckel E. 1872. Die KalkschMamme. Eine monographie, Vols 1 - 3. Berlin: Reimer.","Arndt W. 1941. Eine neuere Ausbeute von Meeresschwammen der West und Sudkuste Portugals. Memorias do Museu de Zoologia da UniVersidade de Coimbra 116: 1 - 75.","Burton M. 1963. A reVision of the classification of the calcareous sponges. London: British Museum (Natural History).","Borojevic R, Peixinho S. 1976. Eponges calcaires du nordnord-est du Bresil. Bulletin du Museum National d´Histoire Naturelle, 3 402: 988 - 1036.","Borojevic R, Boury-Esnault N, Manuel M, Vacelet J. 2002. Order Clathrinida Hartman, 1958. In: Hooper JNA, Van Soest RWM, eds. Systema Porifera: a guide to the classification of sponges. New York: Kluwer Academic / Plenum Publishers, 1141 - 1152.","Dohrmann M, Voigt O, Erpenbeck D, Worheide G. 2006. Non-monophyly of most supraspecific taxa of calcareous sponges (Porifera, Calcarea) revealed by increased taxon sampling and partitioned Bayesian analysis of ribosomal DNA. Molecular Phylogenetics and EVolution 40: 830 - 843.","Muricy G, Esteves EL, Moraes FC, Santos JP, da Silva SM, Klautau M, Lanna E. 2008. BiodiVersidade Marinha da Bacia Potiguar. Rio de Janeiro: Museu Nacional.","Lanna E, Cavalcanti FF, Cardoso L, Muricy G, Klautau M. 2009. Taxonomy of calcareous sponges (Porifera, Calcarea) from Potiguar Basin, NE Brazil. Zootaxa 1973: 1 - 27.","Muricy G, Lopes DA, Hajdu E, Carvalho MS, Moraes FC, Klautau M, Menegola C, Pinheiro U. 2011. Catalogue of Brazilian Porifera. Rio de Janeiro: Museu Nacional.","Klautau M, Azevedo F, Condor-Lujan B, Rapp HT, Collins A, Russo CADM. 2013. A molecular phylogeny for the order Clathrinida rekindles and refines Haeckel's taxonomic proposal for calcareous sponges. IntegratiVe and ComparatiVe Biology 53: 447 - 461.","Van Soest RWM, De Voogd NJ. 2015. Calcareous sponges of Indonesia. Zootaxa 3951: 1 - 105.","Condor-Lujan B, Klautau M. 2016. Nicola gen. nov. with redescription of Nicola tetela (Borojevic & Peixinho, 1976) (Porifera: Calcarea: Calcinea: Clathrinida). Zootaxa 4103: 230 - 238.","Van Soest RWM. 2017. Sponges of the Guyana Shelf. Zootaxa 4217: 1 - 40.","Condor-Lujan B, Louzada TS, Hajdu E, Klautau M. 2018. Morphological and molecular taxonomy of calcareous sponges (Porifera: Calcarea) from Curacao, Caribbean Sea. Zoological Journal of the Linnean Society 182: 1 - 67.","Fontana T, Condor-Lujan B, Azevedo F, Perez T, Klautau M. 2018. Diversity and distribution of calcareous sponges (subclass Calcinea) from Martinique. Zootaxa 4410: 331 - 369.","Lopes MV, Condor-Lujan B, Azevedo F, Perez T, Klautau M. 2018 a. A new genus of calcareous sponge discovered in the Caribbean Sea: Bidderia gen. nov. (Porifera, Calcarea, Calcinea). Zootaxa 4526: 56 - 70.","Lopes MV, Padua A, Condor-Lujan B, Klautau M. 2018 b. Calcareous sponges (Porifera, Calcarea) from Florida: new species, new records and biogeographical affinities. Zootaxa 4526: 127 - 150.","Polejaeff N. 1883. Report on the Calcarea dredged by the HMS Challenger during the years 1873 - 1876. Report on the Scientific Results of the Voyage Challenger (Zoology) 8: 1 - 76.","Hanitsch R. 1895. Notes on a collection of sponges from the west coast of Portugal. Transactions of the LiVerpool Biological Society 9: 205 - 219.","Breitfuss LL. 1898. Kalkschwammfauna der Westkuste Portugals. Zoologische Jahrburger Abteilung fur Systematik 11: 89 - 102.","Thacker RW, Hill AL, Hill MS, Redmond NE, Collins AG, Morrow CC, Spicer L, Carmack CA, Zappe ME, Pohlmann D, Hall C, Diaz M-C, Bangalore PV. 2013. Nearly complete 28 S rRNA gene sequences confirm new hypotheses of sponge evolution. IntegratiVe and ComparatiVe Biology 53: 373 - 387."]}

ROWELLA MARIAE (BRØNDSTED, 1926) COMB.NOV. (FIG. 24) Synonyms: Leucettusa mariae – Brøndsted, 1926: 302; Burton, 1963: 555; Cavalcanti et al., 2013: 276; Leucascus mariae – Rapp, 2004: 124; Lanna et al., 2007: 1560. Type specimen: Possible holotype – NHMD-89631/ CAL-242 (only pictures were analysed). Type locality: Cape Maria van Diemen, New Zealand (34°24ʹ S, 172°30ʹ E; inaccurate coordinates). Three Kings – North Cape MEOW ecoregion. Description (Fig. 24A): Sponge body tubular, widen at the apical region and narrow at the base. External colour beige in ethanol. Consistency rough to the touch. Outer and atrial surfaces smooth. One big osculum is located on a depression at the apical region of the tube, surrounded by a membrane. Atrial cavity wide and spacious. The aquiferous system was not described by Brøndsted (1926), but it is probably leuconoid, as at that time Leucettusa was considered leuconoid. Skeleton: Cortical skeleton well developed, having approximately the same thickness as the choanosome. It is comprised of several layers of tangential triactines. Choanosomal and atrial skeletons reduced, mostly devoid of spicules, but sparse pygmy triactines and tetractines can be found.

ROWELLA LANCIFERA (DENDY, 1924) COMB.NOV. (FIGS 22, 23; TABLE 12) Synonyms: Leucettusa lancifer – Dendy, 1924: 278; Brøndsted, 1926: 301; Burton, 1929: 402; Burton, 1963: 555; Pritchard et al., 1984: 128; Kelly et al., 2009: 45; Leucettusa imperfecta – Burton, 1963: 598; Leucettusa lancifera – Kelly, 2018: 19. T y p e s p e c i m e n: O n e l e c t o t y p e (B M N H 1 9 2 3. 1 9. 1. 5) a n d o n e p a r a l e c t o t y p e (B M N H 1923.10.1.6). L, lectotype; PL, paralectotype. Type locality: Three Kings Islands, New Zealand (34°24ʹ S, 172°06ʹ E; inaccurate coordinates). Three Kings–North Cape MEOW ecoregion. Description: Sponge tubular with vase-shaped form, strongly oval at the top, tapering to the base. Colour white to beige in ethanol (Fig. 22A). Consistency compressible to the touch. Outer surface smooth. Folds of choanosomal tissue delimitate the atrial cavity (Fig. 22B). One apical osculum, elevated and surrounded by a membrane. Aquiferous system leuconoid with subspherical to oval choanocyte chambers (Fig. 22C). Skeleton: Cortical skeleton well developed, having approximately the same thickness as the choanosome; comprised of several layers of tangential triactines (Fig. 22D). Choanosomal and atrial skeletons reduced and comprised of pygmy triactines and tetractines mostly present around canals (Fig. 22E, F). Pygmy tetractines are also abundant near the osculum., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on pages 729-732, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Dendy A. 1924. Porifera. Part I. Non-Antarctic sponges. Natural History Report. British Antarctic (Terra NoVa) Expedition, 1910 (Zoology) 6: 269 - 392, pls I - XV.","Brondsted HV. 1926. Papers from Dr. Th. Mortensen's Pacific Expedition 1914 - 16. XXXV. Sponges from New Zealand. Part II. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 81: 295 - 331.","Burton M. 1929. Porifera. Part II. Antarctic sponges. British Antarctic (' Terra Nova') Expedition, 1910. Natural History Report, London, Zoology 6: 393 - 458.","Pritchard K, Ward V, Battershill C, Bergquist PR. 1984. Marine sponges: forty-six sponges of northern New Zealand. Leigh Laboratory Bulletin 14: 1 - 149.","Kelly M, Edwards AR, Wilkinson MR, Alvarez B, Cook S de C, Bergquist PR, Buckeridge JS, Campbell HJ, Reiswig HM, Valentine C, Vacelet J. 2009. Phylum Porifera: sponges. In: Gordon DP, ed. NeM Zealand inVentory of biodiVersity, Vol 1. Kingdom Animalia: Radiata, Lophotrochozoa, Deuterostomia. Christchurch: Canterbury University Press, 23 - 46.","Burton M. 1963. A reVision of the classification of the calcareous sponges. London: British Museum (Natural History).","Kelly M. 2018. Splendid sponges: a guide to the sponges of NeM Zealand, V. 3. Auckland: National Institute of Water and Atmospheric Research (NIWA) Taihoro Nukurangi."]}

ROWELLA CONNECTENS (BRØNDSTED, 1926) COMB. NOV. (FIGS 15, 16; TABLE 8) Synonyms: Leucandra connectens – Brøndsted, 1926: 308; Burton, 1963: 552. Type specimen: Five syntypes under the same voucher (NHMD-633253). Type locality: Cape Maria van Diemen, Northland, New Zealand (34°24ʹ S, 172°30ʹ E; inaccurate coordinates). Three Kings–North Cape MEOW ecoregion. Description: Sponge body tubular and digitate (Fig. 15A). Colour cream in ethanol. Consistency incompressible but slightly soft to the touch. Outer surface smooth. Cortex with multiple small inhalant apertures, giving it a porous appearance. Oscula present at the apical region of the tubes. They are simple slits or circular and surrounded by membrane with specific skeleton (Fig. 15B). Body wall thick, tapering towards the top and base of the sponge. Atrial cavity narrow and slightly hispid due to the apical actine of tetractines. Aquiferous system leuconoid with spherical choanocyte chambers (Fig. 15C). Skeleton: The oscular margin has sagittal triactines and tetractines, which gradually become regular as the body wall thickens (Fig. 15B). The cortical skeleton is well developed, although it is not as thick as the choanosome (Fig. 15C). The cortical skeleton has several layers of tangential triactines and few tetractines, which project their apical actine into the choanosome, but never reaching the atrium (Fig. 15D). The choanosomal skeleton is comprised of large triactines and pygmy triactines and tetractines, mostly present around the canals. The pygmy spicules are also present in the atrium, laying tangentially (Fig. 15E), with pygmy tetractines being more abundant and projecting their apical actine into the atrial cavity. Spicules (Table 8): Cortical triactines (Fig. 16A). Regular. Variable sizes. Actines are cylindrical to slightly conical, straight or slightly undulated in the middle portion. Tips are blunt. Size – 313.5 (± 77.5) μm length/24.5 (± 7.7) μm width. Cortical tetractines (Fig. 16B). Regular. Basal actines are conical, slightly undulated, with blunt to sharp tips. The apical actine is slightly conical, straight and smooth, with blunt to sharp tips. It is frequently longer than the basal actines. These spicules are less abundant and bigger than the cortical triactines. Size – basal actines: 376.7 (± 83.7) μm length/38.3 (± 3.6) μm width; apical actine: 1,042.6 (± 172.2) μm length/47.1 (± 7.0) μm width. Choanosomal and atrial triactines (Fig. 16C). Regular. Pygmy. Actines are conical and straight, with blunt to rounded tips. They are less abundant than the tetractines in the atrium. Size – 44.5 (± 9.1) μm length/6.6 (± 1.7) μm width. Choanosomal and atrial tetractines (Fig. 16D). Regular. Pygmy. Basal actines are conical and straight, with blunt to rounded tips. The apical actine is conical, smooth, blunt and straight or slightly curved at the distal part. Size – basal actines: 46.8 (± 6.8) μm length/6.7 (± 6.7 μm) width; apical actine: 49.6 (± 7.4) μm length/6.6 (± 1.1) μm width. Ecology: Two specimens were found attached to rock boulders with colonial bryozoans, barnacles, calcareous tubes of polychaetes and algae. They were collected by dredging at 92 m depth. Geographical distribution (MEOW ecoregion): Three K i n g s–N o r t h C a p e (C a p e M a r i a v a n D i e m e n, Northland, New Zealand – Brøndsted, 1926). Remarks: Brøndsted (1926) stated having difficulties assigning this species to any other genus of Calcarea. He mentioned that this species had a classification between Leucandra Haeckel, 1872 and Leucilla Haeckel, 1872 (Calcaronea). The species was then placed in Leucandra due to the absence of the typical inarticulate skeleton of Leucilla and for having a disorganized choanosomal skeleton (Brøndsted, 1926). If Brøndsted had considered the synapomorphies of Calcinea and Calcaronea (subclasses not accepted by some authors at the beginning of the 20 th century), he would have noticed that Leucandra connectens is, in fact, a calcinean sponge. In addition, considering the pygmy spicules present in the choanosome, the well-developed cortical skeleton and the body shape of this sponge, he would see that his species should be placed in the genus Leucettusa sensu Dendy & Row, 1913. Burton (1963) was the only one who analysed this species after its original description. Even though Burton equivocally proposed its synonym with Leucettusa corticata, he drew attention to the fact that Leucandra connectens is not a Leucandra and that it is related with Leucaltidae sensu Dendy & Row, 1913. Following Burton’s idea, and after the re-analysis of the type specimens of Leucandra connectens, we propose the new combination RoƜella connectens. In this species, in addition to the pygmy spicules, large cortical triactines are also present in the choanosome. This trait is also observed in R. imperfecta and R. tubulosa, but they have cortical tetractines often composing the choanosomal skeleton, instead of triactines. Moreover, these large cortical spicules are abundant in the choanosome of R. connectens, opposing to their occasional presence in R. imperfecta and R. tubulosa., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on pages 719-722, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Brondsted HV. 1926. Papers from Dr. Th. Mortensen's Pacific Expedition 1914 - 16. XXXV. Sponges from New Zealand. Part II. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 81: 295 - 331.","Burton M. 1963. A reVision of the classification of the calcareous sponges. London: British Museum (Natural History).","Haeckel E. 1872. Die KalkschMamme. Eine monographie, Vols 1 - 3. Berlin: Reimer.","Dendy A, Row H. 1913. The classification and phylogeny of the calcareous sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London 1913: 704 - 813."]}

ROWELLA TUBULOSA (DENDY, 1924) COMB.NOV. (FIGS 26, 27; TABLE 14) Synonyms and citations: Leucettusa tubulosa – Dendy, 1924: 476; Burton, 1963: 50; Kelly et al., 2009: 45; Kelly, 2018: 20. Type specimen: One lectotype (BMNH 1923.10.1.2) and two paralectotypes (BMNH 1923.10.1.3-4). Type locality: Near Three Kings Islands, New Zealand (34°24ʹ S, 172°06ʹ E). Three Kings–North Cape MEOW ecoregion. Description: Sponge body tubular, cylindrical to vase-shaped and ramified at the base. The types are dark brown to black in the outer region because of the fixation with osmium tetroxide. Internally, the sponge is beige (Fig. 26A, B). Incompressible and rough to the touch. Outer surface smooth (Fig. 26C, D). Atrial surface hispid due to the apical actines of the tetractines (Fig. 26E). One osculum is present at the end of each tube. They are simple circular apertures or slits without membrane (Fig. 26B). The body wall is thick (Fig. 26C). The atrial cavity is spacious and the excurrent canals are evident. Aquiferous system leuconoid with spherical to subspherical choanocyte chambers (Fig. 26E). Granular cells are distributed all over the body, more frequently in the cortical and atrial surfaces (Fig. 26F). Skeleton: Cortical skeleton well developed, although not as thick as the choanosome (Fig. 26C). It is comprised of several layers of tangential triactines and tetractines (Fig. 26 C-D). Choanosomal skeleton comprised of pygmy triactines and tetractines, mostly present around canals. Large cortical spicules can also be found in the choanosome (Fig. 26C). Pygmy spicules are also present in the atrium, lying tangentially, with tetractines being more abundant and projecting their apical actine into the atrial cavity (Fig. 26E–F)., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on pages 735-736, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Dendy A. 1924. Porifera. Part I. Non-Antarctic sponges. Natural History Report. British Antarctic (Terra NoVa) Expedition, 1910 (Zoology) 6: 269 - 392, pls I - XV.","Burton M. 1963. A reVision of the classification of the calcareous sponges. London: British Museum (Natural History).","Kelly M, Edwards AR, Wilkinson MR, Alvarez B, Cook S de C, Bergquist PR, Buckeridge JS, Campbell HJ, Reiswig HM, Valentine C, Vacelet J. 2009. Phylum Porifera: sponges. In: Gordon DP, ed. NeM Zealand inVentory of biodiVersity, Vol 1. Kingdom Animalia: Radiata, Lophotrochozoa, Deuterostomia. Christchurch: Canterbury University Press, 23 - 46.","Kelly M. 2018. Splendid sponges: a guide to the sponges of NeM Zealand, V. 3. Auckland: National Institute of Water and Atmospheric Research (NIWA) Taihoro Nukurangi."]}

GENUS LEUCALTIS HAECKEL, 1872 Type species: Leucaltis clathria Haeckel, 1872 (subsequent designation by Dendy & Row, 1913). Diagnosis: The same as that of the family. Synonyms: Artynas – Haeckel, 1872: 405; Heteropegma – Poléjaeff, 1883: 25; Dendy, 1913: 17; Dendy & Row, 1913: 737; Leucaltis – Haeckel, 1872: 142; Dendy & Row, 1913: 737; Borojević & Peixinho, 1976: 1002; Borojević et al., 1990: 258; 2002: 1147; Lévi, 1998: 75; Wörheide & Hooper, 1999: 877; Borojević & Klautau, 2000: 190; Klautau et al., 2013: 454; Leucaltusa – Haeckel, 1872: 143; Leucetta – Poléjaeff, 1883: 28; Leucettusa – present study. Remarks: The body of Leucaltis species is formed by tubes that anastomose in most parts. Each tube is covered with cortex and atrium with endopinacoderm. Even though the body wall of Leucaltis includes cortical, choanosomal and atrial skeletons, it is thin. It is important to highlight that the choanosomal and atrial spicules of Leucaltis are much smaller than the cortical ones. They are mostly cylindrical, thin and frequently become sagittal at the excurrent canals and at the atrial surface. Scope: Three species of Leucaltis are currently accepted: L. clathria Haeckel, 1872; L. nodusgordii (Poléjaeff, 1883) and L. tenuis Hôzawa, 1929, but we propose the transference of three other species to this genus: Clathrina latitubulata Carter, 1886, Leucettusa corticata (Haeckel, 1872) and Leucettusa sambucus (Preiwisch, 1904)., Published as part of Lopes, Matheus Vieira & Klautau, Michelle, 2023, Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system, pp. 691-746 in Zoological Journal of the Linnean Society 198 on pages 698-699, DOI: 10.1093/zoolinnean/zlad008, http://zenodo.org/record/7894159, {"references":["Haeckel E. 1872. Die KalkschMamme. Eine monographie, Vols 1 - 3. Berlin: Reimer.","Dendy A, Row H. 1913. The classification and phylogeny of the calcareous sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London 1913: 704 - 813.","Polejaeff N. 1883. Report on the Calcarea dredged by the HMS Challenger during the years 1873 - 1876. Report on the Scientific Results of the Voyage Challenger (Zoology) 8: 1 - 76.","Borojevic R, Peixinho S. 1976. Eponges calcaires du nordnord-est du Bresil. Bulletin du Museum National d´Histoire Naturelle, 3 402: 988 - 1036.","Borojevic R, Boury-Esnault N, Vacelet J. 1990. A revision of the intraspecific classification of the subclass Calcinea (Porifera, class Calcarea). Bulletin du Museum National d'Histoire Naturelle. Section A, Zoologie, Biologie et Ecologie Animales 12: 243 - 276.","Levi C. 1998. Sponges of the NeM Caledonian lagoon. Paris: Editions ORSTOM.","Worheide G, Hooper JNA. 1999. Calcarea from the Great Barrier Reef. I: cryptic Calcinea from Heron Island and Wistari Reef (Capricorn-Bunker group). Memoirs of the Queensland Museum 43: 859 - 891.","Borojevic R, Klautau M. 2000. Calcareous sponges from New Caledonia. Zoosystema 22: 187 - 201.","Klautau M, Azevedo F, Condor-Lujan B, Rapp HT, Collins A, Russo CADM. 2013. A molecular phylogeny for the order Clathrinida rekindles and refines Haeckel's taxonomic proposal for calcareous sponges. IntegratiVe and ComparatiVe Biology 53: 447 - 461.","Dendy A. 1893. Studies on the comparative anatomy of sponges. V. Observations on the structure and classification of the Calcarea Heterocoela. Quarterly Journal of Microscopical Science 35: 159 - 257.","Hozawa S. 1929. Studies on the calcareous sponges of Japan. Journal of the Faculty of Sciences, Imperial UniVersity of Tokyo 1: 277 - 389.","Carter HJ. 1886. Descriptions of the sponges from the neighbourhood of Port Phillip Heads, South Australia, continued. Annals and Magazine of Natural History 15 - 18: 431 - 441.","Preiwisch J. 1904. Kalkschwamme aus dem Pacific. Ergebnisse einer Reise nach dem Pacific, Schauinsland 1896 - 97. Zoologische Jahrbucher Abteilung Systematik, Geographie und Biologie der Thiere 19: 9 - 26."]}

FAMILY LEUCALTIDAE DENDY & ROW, 1913 The most recent diagnosis of Leucaltidae is: Clathrinida with a tubular, ramified or even anastomosed cormus with many oscula, or individualized with a large central atrium and a single osculum. The sponge wall is comprised of a distinct cortex sustained by a well-developed skeleton, and a choanosome. The skeleton of the choanosome and the atrial wall may be absent or comprised of small and dispersed triactines and tetractines (Borojević et al., 2002). According to our molecular and morphological results, only Leucaltis should be included in Leucaltidae. The aquiferous system of Leucaltis does not fit into any of the five types accepted up to date. Past works have considered it analogous to the leuconoid (e.g. Lanna et al., 2009) or syconoid (e.g. Wörheide & Hooper, 1999) aquiferous systems, although the choanocyte chambers are not spherical nor do they have a regular radial orientation around the atrium. Pinacocytes are restricted to the external and atrial surfaces. Dendy (1893) described the ‘ type C’ for this organization of the choanocyte chambers, which are mostly elongated and ramified, commonly fusing with each other (Fig. 3). It must not be mistaken with the solenoid aquiferous system because, in Leucaltis, the choanoderm is present inside the ramified chambers, instead of revesting the internal part of the anastomosed choanocyte tubes. Finally, it differs from the sylleibid aquiferous system because choanocyte chambers do not group around a common exhalant chamber. This kind of aquiferous system is present in five out of the six accepted species (following this work). Leucaltis sambucus is said to be leuconoid, but samples of this species were not found, hence we cannot corroborate if it has indeed a leuconoid aquiferous system. Considering that the aquiferous system of Leucaltis has been referred to as leuconoid in past descriptions, it is possible that L. sambucus is not leuconoid, but has the distinct aquiferous system of the other Leucaltis species. Leucaltis corticata is mentioned as having a similar aquiferous system to Leucaltis clathria (Poléjaeff, 1883: 10): ‘with the extension of the flagellated chambers into larger cavities of irregular outline, which come into contact with each other and anastomose, thus forming still larger sinus-like spaces’. For that reason, we draw attention to the presence of the new aquiferous system of Leucaltis in Leucaltis corticata. We suggest the name kladonoid for this new aquiferous system, derived from the Greek kladotós (= ΚΛαδωτός), branched, ramified. For Leucaltidae, we propose to modify the diagnosis to: Clathrinida with a body comprised of large, ramified and anastomosed tubes; never clathroid. Each tube has a distinct cortex, a choanoderm and a central atrium. The skeleton of the choanosome and atrium is comprised of regular and/or sagittal triactines and tetractines much smaller than the cortical ones. Aquiferous system kladonoid, with elongated and ramified choanocyte chambers.

Turner, Thomas L., Lonhart, Steve I. (2023): The Sponges of the Carmel Pinnacles Marine Protected Area. Zootaxa 5318 (2): 151-194, DOI: 10.11646/zootaxa.5318.2.1, URL: http://dx.doi.org/10.11646/zootaxa.5318.2.1

Leucilla nuttingi (Urban, 1902) Figures 1I & 24A, Published as part of Turner, Thomas L. & Lonhart, Steve I., 2023, The Sponges of the Carmel Pinnacles Marine Protected Area, pp. 151-194 in Zootaxa 5318 (2) on page 190, DOI: 10.11646/zootaxa.5318.2.1, http://zenodo.org/record/8162357, {"references":["Urban, F. (1902) Rhabdodermella nuttingi, nov. gen. et nov. spec. Zeitschrift fur Wissenschaftliche Zoologie, 71, 268 - 275."]}

Amphoriscus semoni Breitfuss, 1896 Amphoriscus semoni Breitfuss, 1896 Citations: Amphoriscus semoni Breitfuss 1896: 435; 1898: 221; Dendy & Row 1913: 782; Burton 1963: 542; Van Soest & De Voogd 2015: 94; Klautau, Cavalcanti & Borojevic 2017: 105; Van Soest & De Voogd 2018: 130; Cóndor-Luján et al. 2019: 1825. Type material: ZMB 2698 (Holotype; Ambon Island, Maluku, Indonesia). Not analysed in the present work. Type locality: Ambon Island, Maluku, Indonesia. Analysed material: BMNH 1886.6.7.32 (two slides containing sections of the skeleton). Port Jackson, Australia; previously identified as A. cylindrus in Burton, 1963: 634. Morphology: The holotype of A. semoni was not available for redescription, and the material analysed here (deposited in the NHM) was comprised of two microscope slides. Aquiferous system is syconoid. Anatomy: The material evaluated in the present study (BMNH 1886.6.7.32) was deposited at the Porifera collection of the NHM under the name A. cylindrus (Fig. 9). It is formed by giant cortical tetractines, with paired and unpaired actines laying on the cortex (Figs. 9A, B). The organisation of the skeleton is inarticulate due to the unpaired actine of the subatrial triactines and the apical actine of the cortical tetractines (Figs. 9A–C). The atrium is perforated by the apical actine of the small atrial tetractines (Fig. 9D). Spicules (Tables 5 and 6): A figure containing only the spicule categories could not be prepared due to the lack of a slide of dissociated spicules. Cortical tetractines: Irregular, conical, and sharp. The basal actines (paired and unpaired) are slightly curved from the base to the tips. The apical actine is straight and long. Subatrial triactines: Irregular, cylindrical to slightly conical, sharp. The paired actines are straight. The unpaired actine is longer or the same size as the paired actines. Atrial tetractines: Irregular, cylindrical, small, thin, and sharp. The paired actines are curved. The unpaired actine is similar to the paired ones. The apical actine is straight and short, always very thin. Remarks: BMNH 1886.6.7.32 was deposited in the NHM as A. cylindrus (Burton 1963), but its subatrial skeleton is comprised of triactines and not tetractines, as in A. cylindrus. The only species of Amphoriscus whose organisation is consistent with this specimen—cortical tetractines, subatrial triactines, and atrial tetractines—is A. semoni. The specimen is from Port Jackson, Australia, in the same biogeographical region as the type locality of A. semoni (Ambon Island, Indonesia) and isolated from the type locality of A. cylindrus (the Adriatic Sea). Therefore, we suggest the identification of BMNH 1886.6.7.32 as A. semoni. Figure 9 shows its skeletal organisation, in which the subatrial and atrial layers are clearly evident. In our opinion, these illustrations are important for further recognition of specimens belonging to A. semoni. Among the characters described by Breitfuss (1896), A. semoni is bright white when preserved, and the atrial surface is perforated by the apical actine of the colossal cortical tetractines. The specimens described as A. semoni by Van Soest & De Voogd (2015, 2018) present minor differences in the colour and length of these apical actines. The sample ZMAPOR 08073, collected in Sumbawa, Indonesia, was green alive and beige after fixation. The apical actine of its cortical tetractines does not exceed 600 μm (see Tables 8 and 9; Van Soest & De Voogd 2015), while in the holotype, they vary from 520 to 790 μm (Breitfuss 1896). In their most recent study, Van Soest & De Voogd (2018) described a new set of specimens of A. semoni (ZMAPOR 10527), sampled in Seychelles, Western Indian Ocean, which were white alive and beige after fixation. The apical actines of the cortical tetractines do not perforate the atrium, although they measure 239–882 μm (Van Soest & De Voogd 2018). The differences between the holotype and the other specimens described in both works by Van Soest & De Voogd provide a better understanding of the intraspecific variation of A. semoni. These are precious data, usually rare for Amphoriscus species since the original description, based on one or few specimens, is all that is available for many species of the genus. Amphoriscus semoni resembles two species of the Adriatic Sea, namely A. gregorii and A. bucchichii. However, A. semoni differs in having a cortical skeleton comprised exclusively of tetractines. Distribution: Ambon Island, Indonesia (Breitfuss 1896); Seychelles (Van Soest & De Voogd 2018); Port Jackson, Australia (present study). Corresponding MEOW: Banda Sea, Seychelles, and Manning-Hawkesbury (Spalding et al. 2007).

Amphoriscus pedunculatus Klautau, Cavalcanti & Borojevic, 2017 Amphoriscus pedunculatus Klautau, Cavalcanti & Borojevic, 2017 Citations: Amphoriscus pedunculatus Klautau, Cavalcanti & Borojevic 2017: 105; C��ndor-Luj��n et al. 2019: 1825. Type material: UFRJPOR 5802 (Holotype). Saco do Po��o, S��o Sebasti��o, S��o Paulo State, Brazil (23�� 45��� 40.3��� S ��� 45�� 14��� 53.5��� W); 13 m depth; F. F. Cavalcanti, V. Padula & L. Kremer; 03 December 2008. UFRJPOR 5803 (Paratype). Saco da Ponta Grossa, S��o Sebasti��o, S��o Paulo State, Brazil (23�� 46��� 31.8��� S ��� 45�� 13��� 54.8��� W); 6 m depth; F. F. Cavalcanti & V. Padula, 03 December 2008. Type locality: Saco do Po��o, S��o Sebasti��o, S��o Paulo State, Brazil. Analysed material: UFRJPOR 5802 (holotype; specimen and slides containing sections of the skeleton and dissociated spicules). UFRJPOR 5803 (paratype; specimen and slides containing sections of the skeleton and dissociated spicules). UFRJPOR 5801 (specimen and slides containing sections of the skeleton and dissociated spicules), Serraria Island, S��o Sebasti��o, S��o Paulo, Brazil (23�� 48��� 47.9���S ��� 45�� 13��� 44.0���W); 9 m depth; F. F. Cavalcanti & V. Padula, 04 December 2008. MNRJ 5818 (specimen and slides containing sections of the skeleton and dissociated spicules). Alcatrazes Archipelago, S��o Sebasti��o, S��o Paulo, Brazil (24�� 06��� 00.0��� S ��� 45�� 40��� 59.9��� W); 12 m depth; M. Cust��dio & C. Santos, 03 May 2002. UFRJPOR 5776 (specimen and slides containing sections of the skeleton and dissociated spicules), Saco da Saia, Arraial do Cabo, Rio de Janeiro, Brazil (23�� 00��� 23.0��� S ��� 42�� 00��� 36.0��� W); 1 m depth; G. Muricy, 17 March 1988. Morphology: The specimens are similar in size and shape, with tubular shape and apical osculum surrounded by a delicate fringe of trichoxeas (Fig. 7A). The peduncle is white in live specimens and brownish/orange in ethanolpreserved specimens, but the peduncle is brownish/orange. The surface is slightly hispid due to the trichoxeas of the cortical region. The atrial cavity is hispid and fills the entire body of the specimens. The aquiferous system is syconoid. Anatomy: The cortical region has several broken spicules. It is mostly formed by giant tetractines with long apical actines. Triactines are also present, but they are less abundant (Fig. 7B and C). Trichoxeas are abundant throughout the entire cortex (Fig. 7D) and may reach the choanosome (Fig. 7E). The subatrial region is comprised exclusively of triactines. They vary in size, but the unpaired actine is always long (Fig. 7C). The atrial skeleton is comprised of tetractines (Fig. 7F). Spicules (Table 4; Fig. 8): Trichoxeas: Very thin, fusiform with sharp tips. Straight or slightly curved. Mostly broken (Fig. 7D). Cortical triactines (Fig. 8A): Conical with blunt tips. The paired actines are slightly curved. The unpaired actine is straight. Cortical tetractines (Fig. 8B): Conical with blunt tips. The paired actines are slightly curved, as well as the unpaired one. The apical actine is straight and long. Subatrial triactines (Fig. 8C): Conical to slightly conical and blunt. Paired actines are short and straight. The unpaired actine is straight. Atrial tetractines: Slightly conical and sharp. Paired actines are slightly curved and larger or the same size as the apical actine. The unpaired actine is straight and longer than the other actines. The apical actine is straight (Fig. 8D). Remarks: Amphoriscus pedunculatus was recently described (Klautau et al. 2017), with an original description in which important taxonomic characters were fully described and illustrated. The redescription provided in the present study differs only with respect to the tips of the spicules: cortical and subatrial spicules are described here as having blunt tips while they were reported as being sharp in the original description. In our opinion, the differentiation between blunt and sharp tips may not always be obvious, but the availability of images (as in Figure 8) helps resolve any doubts. Slight differences were also found in the measurements of the cortical tetractines of the holotype: according to Klautau et al. (2017), the range observed in the length of the unpaired actine is larger (120.0��� 238.5 ��58.2���350.0 ��m) than that obtained here (177.8��� 182.9 ��7.2���188.0 ��m). These differences may be related to the existence of several microscopical slides containing dissociated spicules, and the largest spicules may not be evenly distributed. Among the species with peduncle, Amphoriscus pedunculatus is the only one with a subatrial skeleton formed exclusively by triactines. The remaining species, A. chrysalis, A. cyathiscus, and A. testiparus, have both triactines and tetractines or only tetractines. Distribution: The species seems to be restricted to the southeastern Brazilian coast in S��o Sebasti��o (S��o Paulo state), and Arraial do Cabo (Rio de Janeiro state) (Klautau et al. 2017). Distribution: S��o Sebasti��o (S��o Paulo state) and Arraial do Cabo (Rio de Janeiro state). Corresponding MEOW: Southeastern Brazil (Spalding et al. 2007)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on pages 51-53, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287, {"references":["Klautau, M., Cavalcanti, F. F. & Borojevic, R. (2017) The new sponge species Amphoriscus pedunculatus (Porifera, Calcarea). Zootaxa, 4341 (1), 105 - 112. https: // doi. org / 10.11646 / zootaxa. 4341.1.9","Condor-Lujan, B., Azevedo, F., Hajdu, E., Hooker, Y., Willenz, P. & Klautau, M. (2019) Tropical Eastern Pacific Amphoriscidae Dendy, 1892 (Porifera: Calcarea: Calcaronea: Leucosolenida) from the Peruvian coast. Marine Biodiversity, 49 (4), 1 - 18. https: // doi. org / 10.1007 / s 12526 - 019 - 00946 - y","Spalding, M. D., Fox, H. E., Allen, G. R., Davidson, N., Ferdana, Z. A., Finlayson, M., Halpern, B. S., Jorge, M. A., Lombana, A., Lourie, S. A., Martin, K. D., McManus, E., Molnar, J., Recchia, C. A. & Robertson, J. (2007) Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. BioScience, 57 (7), 573 - 583. https: // doi. org / 10.1641 / B 570707"]}

Amphoriscus elongatus Pol��jaeff, 1883 Citations and synonymies: Amphoriscus elongatus Pol��jaeff 1883: 48, pl. iv, fig. 5, pl. v, fig. 4; Dendy & Row 1913: 782; Burton 1956: 117; Burton 1963: 538; Klautau et al. 2017: 106; C��ndor-Luj��n et al. 2019: 1825. Type material: BMNH 1884.4.22.27 (holotype). Station 145, off Prince Edward Islands (46�� 40��� S ��� 37�� 50��� E), 275 to 567 m depth, 27 December 1873. Type locality: off Prince Edward Islands, Indian Ocean. Analysed material: BMNH 1884.4.22.27 (holotype; specimen and one slide containing sections of the skeleton). BMNH 1955.12.13.10 (one slide containing sections of the skeleton) and BMNH 1955.12.13.11 (one slide containing sections of the skeleton), both from Plymouth; R.W.H. Row collection. Morphology: The colour of the holotype in ethanol is white (Fig. 4A). It is a fragment with tubular shape and apical osculum, measuring 4 cm x 0.2 cm (length x width). The surface is slightly hispid. Syconoid aquiferous system (Fig. 5). Anatomy: The skeleton is typical of the genus Amphoriscus, and the inarticulation is evident (Figs. 4D and 5D, E). The cortical region is formed by giant tetractines and small sagittal triactines (Figs. 4B, C). The apical actine of these giant tetractines may reach the atrial cavity. All the analysed samples exhibited trichoxeas perforating the cortex without forming tufts (Fig. 5C). The subatrial region is formed by abundant triactines with long unpaired actine and rare tetractines (Fig. 5D). The latter are more frequent in the specimens BMNH 1955.12.13.10 and BMNH 1955.12.13.11 but also occur in the holotype. Additionally, in these specimens from Plymouth, we also observed a few modified subatrial triactines in which the actines that surround the atrial wall have different lengths and shapes (Fig. 5E). These spicules were found in the sections. Due to the occurrence of other spicules overlapping them, it was not clear if they were pseudosagittal. All the subatrial spicules point their unpaired actines to the cortex, in opposition to the apical actine of the cortical tetractines (Figs. 4D and 5D, E). The atrial region is comprised exclusively of tetractines (Fig. 5F). Spicules (Table 1): A figure containing only the spicule categories, commonly shown in studies of the taxonomy of calcareous sponges, could not be prepared since only slides containing sections of the skeleton were available. Cortical tetractines: Giant, conical with blunt tips. Paired actines are curved and long. The unpaired actine is curved and smaller or the same size as the paired actines. The apical actine is straight and long. Cortical triactines: Slightly conical and sharp. Paired actines are straight to slightly curved. Unpaired actine is shorter and straight. Subatrial triactines: Cylindrical to slightly conical with sharp tips. Paired actines are straight. Unpaired actine is straight and long, sometimes reaching the cortical region. Subatrial tetractines: Rare, with cylindrical actines and sharp tips. Paired actines are slightly curved. Unpaired actine is straight and larger than the paired ones. Apical actine is short and curved. Atrial tetractines: Vary in size. Cylindrical actines with sharp tips. Paired actines are long, thin, and straight. Unpaired actine is slightly curved. Apical actine is straight. Remarks: The original description of Amphoriscus elongatus was based on a single specimen sampled during the Challenger expedition (Pol��jaeff 1883). It was tubular and elongated (hence the species name elongatus). Close to the osculum, it was divided into two tubes, a feature that can no longer be recognised in the type (Fig. 4A). According to Pol��jaeff (1883), A. elongatus has ���an important anatomical peculiarity���the tendency of the radial tubes to meet in threes, fours, or in larger numbers around the same shallow invagination of the gastric cavity���. We could not clearly understand this organisation described by Pol��jaeff (1883), even after analysing the holotype and additional specimens. Nevertheless, we are convinced that the aquiferous system of A. elongatus is not a typical syconoid in which choanocyte chambers run in parallel from the atrium to the cortex. Instead, choanocyte chambers of A. elongatus seem to branch, possibly causing, when sectioned, the holes observed in Figure 5 along the chambers��� length. More studies are needed to better describe this organisation, which depends on the discovery of fresh specimens for histological and electron microscopy analyses. Our results indicate an important difference between our description of the holotype and that provided by Pol��jaeff (1883), namely the occurrence of rare subatrial tetractines. We first observed this characteristic in the slides of the specimens from Plymouth and then, after a careful reanalysis, also in the holotype. With respect to the geographical distribution, BMNH 1955.12.13.10 and BMNH 1955.12.13.11 are from Plymouth, United Kingdom (Northern Atlantic Ocean), while the holotype BMNH 1884.4.22.27 was sampled ���off Prince Edward Islands��� (Southern Indian Ocean). In general, representatives of a species with such a disjunct distribution are viewed as questionable in the literature, mainly due to the low dispersal capability of the larvae of calcareous sponges (Maldonado 2006). The only morphological difference between the three analysed samples is the putative presence of pseudosagittal triactines in the subatrial region of those from Plymouth. These spicules were not found in the holotype, and we assumed that the absence could be the result of plasticity. The analysis of a larger set of specimens, preferably using an integrative approach, is needed to elucidate this question. The species that most closely resembles A. elongatus in its skeletal composition is A. pedunculatus (cortical triactines, tetractines and trichoxeas, subatrial triactines, and atrial tetractines). These species differ mainly in the presence of rare subatrial tetractines in the former and the size of the cortical tetractines, which are larger in A. elongatus (paired actines: 443.6��59.0/ 48.7��7.9��m; unpaired actine: 397.2��42.1/ 52.7��4.9 ��m; apical actine: 453.8��60.3/48.2��6.4��m) than in A. pedunculatus (paired actines: 225.5��39.3/ 27.2 ��4.3 ��m; unpaired actine: 182.9��7.2/ 27.2��3.8 ��m; apical actine: 303.8��50.7/ 29.4��5.4 ��m). Distribution: Off Prince Edward Islands, South Africa (Pol��jaeff 1883) and Plymouth, United Kingdom (present study). Corresponding MEOW: Prince Edward Islands and Celtic Seas (Spalding et al. 2007)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on pages 45-48, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287, {"references":["Polejaeff, N. (1883) Report on the Calcarea dredged by H. M. S. ' Challenger', during the years 1873 - 1876. Report on the Scientific Results of the Voyage of H. M. S. ' Challenger', 1873 - 1876. Zoology, 8 (2), 1 - 76.","Dendy, A. & Row, R. W. H. (1913) The Classification and Phylogeny of the Calcareous Sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London, 3, 704 - 813. https: // doi. org / 10.1111 / j. 1469 - 7998.1913. tb 06152. x","Burton, M. (1956) The sponges of West Africa. Atlantide Report (Scientific Results of the Danish Expedition to the Coasts of Tropical West Africa, 1945 - 1946, Copenhagen, 4, 111 - 147.","Burton, M. (1963) A revision of the Classification of the Calcareous Sponges: with a Catalogue of the specimens in the British Museum. Order of the trustees of the British Museum (Natural History), London, 693 pp.","Klautau, M., Cavalcanti, F. F. & Borojevic, R. (2017) The new sponge species Amphoriscus pedunculatus (Porifera, Calcarea). Zootaxa, 4341 (1), 105 - 112. https: // doi. org / 10.11646 / zootaxa. 4341.1.9","Condor-Lujan, B., Azevedo, F., Hajdu, E., Hooker, Y., Willenz, P. & Klautau, M. (2019) Tropical Eastern Pacific Amphoriscidae Dendy, 1892 (Porifera: Calcarea: Calcaronea: Leucosolenida) from the Peruvian coast. Marine Biodiversity, 49 (4), 1 - 18. https: // doi. org / 10.1007 / s 12526 - 019 - 00946 - y","Maldonado, M. (2006) The ecology of the sponge larva. Canadian Journal of Zoology, 84 (2), 175 - 194. https: // doi. org / 10.1139 / z 05 - 177","Spalding, M. D., Fox, H. E., Allen, G. R., Davidson, N., Ferdana, Z. A., Finlayson, M., Halpern, B. S., Jorge, M. A., Lombana, A., Lourie, S. A., Martin, K. D., McManus, E., Molnar, J., Recchia, C. A. & Robertson, J. (2007) Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. BioScience, 57 (7), 573 - 583. https: // doi. org / 10.1641 / B 570707"]}

Order BAERIDA Borojević, Boury-Esnault & Vacelet, 2000 ���Leuconoid Calcaronea with the skeleton either composed exclusively of microdiactines or in which microdiactines constitute exclusively or predominantly a specific sector of the skeleton, such as choanoskeleton or atrial skeleton. Large or giant spicules are frequently present in the cortical skeleton, from which they can partially or fully invade the choanoderm. In sponges with a reinforced cortex, the inhalant pores can be restricted to a sieve-like ostia-bearing region. Dagger-shaped small tetractines (pugioles) are frequently the sole skeleton of the exhalant aquiferous system. Although the skeleton may be highly reinforced by the presence of dense layers of microdiactines in a specific region, an aspicular calcareous skeleton is not present��� (Borojevic et al. 2002)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on page 63, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287, {"references":["Borojevic, R., Boury-Esnault, N. & Vacelet, J. (2000) A revision of the supraspecific classification of the subclass Calcaronea (Porifera, class Calcarea). Zoosystema, 22 (2), 203 - 263.","Borojevic, R., Boury-Esnault, N., Manuel, M. & Vacelet, J. (2002) Order Leucosolenida Hartman, 1958. In: Hooper, J. N. A. & Van Soest, R. W. M. (Eds.), Systema Porifera: a guide to the classification of sponges. Vol. 2. Kluwer Academic / Plenum Publishers, New York, Boston, Dordrecht, London and Moscow, pp. 1157 - 1184. https: // doi. org / 10.1007 / 978 - 1 - 4615 - 0747 - 5 _ 120"]}

Comparison of Amphoriscus species In order to facilitate the comparison between species of Amphoriscus and, consequently, their taxonomic identification, the skeletal composition and type locality of all Amphoriscus species is summarised below (Tab. 8).

Amphoriscus decennis sp. nov. Etymology: From the Latin decem + annus (meaning decades). The name is related to the time span the specimens remained in the collection before being recognised as a new species. Diagnosis: Amphoriscus with cortical skeleton comprised of trichoxeas and one type of tetractine variable in size, subatrial skeleton comprised of triactines and atrial tetractines. Type material: UFRJPOR 9031 holotype (��le Riou, Marseille, France; 05/VI/1967). UFRJPOR 9032, UFRJPOR 9033, UFRJPOR 9034, and UFRJPOR 9035��� paratypes (��le Riou, Marseille, France; 05/VI/1967). Type locality: ��le Riou, Marseille, France. Additional material: UFRJPOR 5822 (��le Riou, Marseille, France; 05/VI/1967). Several fragments from one or more specimens. BMNH 1955.12.13.9 (Plymouth; R. W. H. Row). Morphology: The specimens are cylindrical, with differences in thickness along the body, as they are thicker at the base. Smooth surface, colour white in spirit. The osculum is apical and naked. The holotype (UFRJPOR 9031) measures 2.0 x 1.0 cm (height x width), while the largest specimen, the paratype UFRJPOR 9035, measures 3.0 x 1.0 cm (Fig. 10A). The atrial cavity occupies the entire body of the specimen. The aquiferous system is syconoid and seems slightly disorganised in the paratype UFRJPOR 9032, possibly due to the presence of embryos and amphiblastulae. Anatomy: The organisation is typical of Amphoriscidae, with an inarticulate skeleton formed mainly by giant cortical tetractines (Fig. 10B). Smaller tetractines also occur in the cortex and are less abundant than the giant ones. Trichoxeas are present (Fig. 10C). The inarticulation of the skeleton is due to the apical actine of the cortical tetractines and the unpaired actines of subatrial triactines (Fig. 10D). The atrial region is comprised exclusively of tetractines with short apical actines (Fig. 10E, F). These short apical actines and those of the giant cortical tetractines perforate the atrial cavity, making the atrial surface hispid. Spicules (Table 7; Fig. 11): Cortical tetractines: Actines are conical and blunt. The paired actines are curved, while the unpaired one is straight. Apical actine is long, straight, and usually extends up to the atrial cavity (Figs. 11A, A���). Subatrial triactines: Actines are cylindrical to slightly conical, with blunt tips. The paired actines are curved from the base to the tips and are always smaller than the unpaired one (Fig. 11B). Atrial tetractines: Actines are cylindrical and sharp. The paired actines are long and curved from the base to the tips. Unpaired actine is curved and usually smaller than the paired actines. The apical actine is short and straight or slightly curved (Figs. 11C, C���). Remarks: Amphoriscus decennis sp. nov. does not present anchoring spicules or peduncle, which differs from A. ancora, A. chrysalis, A. cyathiscus, A. pedunculatus, A. synapta, and A. testiparus. Among the remaining species, A. semoni most closely resembles the new species due to its skeletal organisation: cortical tetractines, subatrial triactines, and atrial tetractines. Nevertheless, the atrial tetractines of A. decennis sp. nov. have smaller apical actines [holotype: 47.0��� 61.2 ��9.3���73.4/ 9.4��� 10.7 ��1.1���13.2 ��m] than the type of A. semoni [according to the original description, the apical actine is 100���130/ 9 ��m; Breitfuss (1896)]. Among the proposed paratypes, the largest apical actine of the new species was 97.2 ��m (maximum value, obtained for the specimen UFRJPOR 9033). Although close to the minimum value of the size range described by Breitfuss (1896), the measurements found here for A. decennis sp. nov. for this actine are, in general, smaller than in A. semoni. Additionally, cortical spicules are considerably thicker in A. decennis sp. nov. (holotype ���paired: 31.9���58.5 ��m, unpaired 30.1���41.4 ��m, apical 35.0���52.1 ��m versus paired: 10���20 ��m, unpaired 10���20 ��m, apical 19���27 ��m in A. semoni). Finally, only A. decennis sp. nov. has trichoxeas. The presence of small cortical tetractines in A. decennis sp. nov. may also be useful to differentiate it from A. semoni, although we considered them to belong to the same category as the giant cortical tetractines. The geographical distribution of both species is also different. Type specimens of A. decennis sp. nov. were sampled in the Mediterranean Sea (more specifically at ��le Riou, Marseille, France), while A. semoni is known to occur across the central Indo-Pacific region (type locality��� Ambon Island, Indonesia). A specimen of A. decennis sp. nov. from Plymouth was also recognised (as discussed along with the remarks of A. chrysalis), which remains consistent with a separate species distribution from that of A. semoni. A wide distributional range is not common in calcareous sponges and has been associated with erroneous taxonomic identification [such as the specimen BMNH 1886.6.7.32, from Australia, previously identified as A. cylindrus by Burton (1963)] or with the occurrence of introduced species. Since there is no reason to suspect that A. semoni could have invaded regions outside the Indo-Pacific, we consider the distribution additional evidence to support the new species. Distribution: Mediterranean Sea and Plymouth, United Kingdom (BMNH 1955.12.13.9; see remarks of A. chrysalis). Corresponding MEOW: Western Mediterranean and Celtic Seas (Spalding et al. 2007)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on pages 57-58, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287, {"references":["Breitfuss, L. (1896) Amphoriscus semoni, eine neue Art heterocoeler Kalkschwamme. Zoologischer Anzeiger, 19 (515), 435 - 436.","Burton, M. (1963) A revision of the Classification of the Calcareous Sponges: with a Catalogue of the specimens in the British Museum. Order of the trustees of the British Museum (Natural History), London, 693 pp.","Spalding, M. D., Fox, H. E., Allen, G. R., Davidson, N., Ferdana, Z. A., Finlayson, M., Halpern, B. S., Jorge, M. A., Lombana, A., Lourie, S. A., Martin, K. D., McManus, E., Molnar, J., Recchia, C. A. & Robertson, J. (2007) Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. BioScience, 57 (7), 573 - 583. https: // doi. org / 10.1641 / B 570707"]}

Family AMPHORISCIDAE Dendy, 1892 “ Leucosolenida with syconoid, sylleibid or leuconoid organisation, and a distinct cortex supported by tangential tetractines whose centripetal apical actines cross the outer part or the whole of the choanosome. Tangential triactines and small tetractines may also be present in the cortex. The choanoskeleton is typically inarticulated, composed of apical actines of cortical tetractines and the unpaired actines of subatrial spicules. In species with a thick wall, scattered triactines and/or tetractines may also be present, either among the spicules of the inarticulated choanoskeleton or forming a distinct subatrial layer. An atrial skeleton is always present” (Borojevic et al. 2002).

Amphoriscus chrysalis (Schmidt, 1864) Ute chrysalis Schmidt, 1864 (type species by subsequent designation; Dendy & Row,1913) Citations and synonymies: Ute chrysalis Schmidt 1864: 23 (original description); Amphoriscus chrysalis Haeckel 1870: 177; Sycilla chrysalis Haeckel 1872: 256; Sycurus chrysalis Haeckel 1872: 256 (variation of Sycilla chrysalis); Amphoriscus chrysalis Pol��jaeff 1883: 7; Dendy & Row 1913: 782; Breitfuss 1935: 29; Burton 1963: 535; Vacelet 1981: 165; Radolović et al. 2015: 305; Klautau, Cavalcanti & Borojevic 2017: 105; C��ndor-Luj��n et al. 2019: 1825. Type material: Unknown. Type locality: Lesina, Lissa, Adriatic Sea. Analysed material: LBIM 1968-557 L (Glenans, France; four slides containing sections of the skeleton). LBIM 1968-556 L (Marseille, France; three slides containing sections of the skeleton). LBIM 1968-327 (Roscoff, France; four slides containing sections of the skeleton). LBIM 1968-355 (Roscoff, France; two slides containing sections of the skeleton). LBIM 1968-326 (Roscoff, France; seven slides containing sections of the skeleton and dissociated spicules). LBIM 1968-280 (Roscoff, France; one slide containing dissociated spicules). BMNH 1954.8.12.195 (Plymouth; M.B. L. collection; one slide containing sections of the skeleton). BMNH 195512.13.9 (Plymouth; R.W.H. collection; one slide containing sections of the skeleton; not A. chrysalis ��� see below). SME-500 (Biscay Bay, France; one specimen and one slide containing sections of the skeleton). Morphology: Colour in spirit varies from yellowish-white to yellowish-brown (Burton 1963). The specimen SME-500 has a tubular shape and measures ca. 4 mm x 1 mm (Fig. 1A). According to Schmidt (1864), the type had a peduncle, which is not visible in SME-500. Smooth surface and a single osculum without fringe of trichoxeas. Syconoid aquiferous system. Abundant larvae were observed in the choanosome of LBIM 1968-556 L (Figs. 1B, D, E). Anatomy: Skeleton typical of the family Amphoriscidae (Fig. 1B). The cortical region is formed mainly by giant and abundant tetractines and a few tangential triactines, pierced by tufts of trichoxeas (Fig. 1C). Abundant subatrial tetractines and triactines point their unpaired actine towards the cortex (Fig. 1E). The atrial skeleton is formed exclusively by tetractines, and these have a very long apical actine (Fig. 1F). Some lacunae occur along the subcortical and subatrial regions, mainly in the specimen LBIM 1968-556 L, possibly due to fixation and/or the occurrence of tissue retraction over time (Fig. 1D). Fusiform diactines were found in the spicules slides of LBIM 1968-326, but these were not observed in the sections of the skeleton of the same specimen and were not found in any other analysed material. In addition, BMNH 1955.12.13.9 differs from the remaining analysed samples in its skeletal composition: cortical tetractines, subatrial triactines, and atrial tetractines, and thus cannot be classified as A. chrysalis (see Remarks). Spicules: Among the analysed material, slides containing dissociated spicules were available only for LBIM 1968-326. The values shown here therefore only reflect this sample. Cortical tetractines (Fig. 2A): Conical with blunt tips. The paired actines are curved, and one of the tips is broken in most of the spicules (118.8��� 298.3 ��90.5���520.2/ 23.8��� 38.9 ��7.9���53.1 ��m). The unpaired actine is smaller or the same size as the paired actines (106.1��� 203.6 ��107.4���485.1/ 22.2��� 41.5 ��8.0���58.6 ��m). The apical actine is straight and longer than the other actines (281.5��� 421.6 ��87.2���636.2/ 23.8��� 39.8 ��7.7���53.7 ��m). Cortical triactines (Fig. 2B): Conical with blunt tips, less abundant than the cortical tetractines. Paired actines are slightly curved (246.2��� 360.5 ��66.8���466.0/ 22.3��� 35.6 ��6.3���45.2 ��m). The unpaired actine is straight, sometimes slightly lanceolate (241.7��� 468.8 ��140.7���673.9/ 23.1��� 37.2 ��8.1���52.6 ��m). Subatrial triactines and tetractines (Figs. 2C, D): Cylindrical to slightly conical and blunt. The paired actines are short and straight (triactines: 102.1��� 145.0 ��23.3���185.3/ 6.2��� 10.2 ��1.9���13.6 ��m; tetractines: 92.9��� 139.7 ��45.5���253.1/ 6.3��� 8.5 ��2.0���11.4 ��m). The unpaired actines are straight and long (triactines: 232.5��� 340.4 ��49.6���435.3/ 8.5��� 13.0 ��2.4���17.4 ��m; tetractines: 142.6��� 271.3 ��67.6���350.8/ 8.0��� 9.6 ��1.2���11.1 ��m). The apical actine of the tetractines is short and curved (20.3��� 31.0 ��14.3���68.6/ 5.3��� 7.6 ��1.7���10.1 ��m). Atrial tetractines(Fig.2E):Cylindrical with sharp tips.The paired actines are long and curved (100.3��� 173.0 ��49.3��� 315.4/ 6.0��� 14.1 ��15.9���97.1 ��m). The unpaired actine is straight (62.5��� 169.3 ��95.9���445.6/ 4.9��� 12.3 ��3.7���21.2 ��m). The apical actine is straight and long. It is generally twice as large as the paired actines (102.2��� 271.7 ��126.4���527.2/ 7.9��� 11.9 ��2.6���17.8 ��m). Remarks: According to the original description (Schmidt 1864), Amphoriscus chrysalis is comprised only of tetractines (the presence of only this spicule category and the external morphology are the only characters mentioned by the author). The material analysed here has cortical and subatrial triactines, differing from the original description. Haeckel (1872) also described A. chrysalis as having only tetractines, though subatrial (and possibly cortical) triactines similar to those we observed were illustrated in his work (Plate 43, figures 2 and 3). It is not clear if Haeckel analysed the same material described by Schmidt (1864), but the short original description lacking either spicule measurements or illustrations of the spicule types and skeleton show the importance of Haeckel���s (1872) work for the recognition of A. chrysalis. The difficulty in recognising this species is also caused by a lack of type material. Our results and the figure provided by Haeckel (1872) thus allow us to state that A. chrysalis has triactines. We were also able to recognise the main feature of A. chrysalis reported by Haeckel (1872) in the samples analysed here: the long apical actine of the atrial tetractines (>100 ��m). This feature was important for supporting our decision to identify some samples as A. chrysalis sensu Haeckel (1872). Among the analysed samples, BMNH 1954.8.12.195 and BMNH 1955.12.13.9 had been listed by Burton (1963: 634) under the name A. chrysalis. Our results indicate that the latter does not belong to A. chrysalis due to its different skeletal composition. Here, we allocate it in A. decennis sp. nov., while BMNH 1954.8.12.195 remains A. chrysalis. Some of the samples analysed in this study had previously been identified as A. chrysalis by Borojevic et al. (1968), namely: LBIM 1968-557, LBIM 1968-556 L, LBIM 1968-327, LBIM 1968-355, LBIM 1968-326, and LBIM 1968-280. The list of species from Roscoff was provided without morphological details, meaning that these specimens are described for the first time in this study. Compared to the other species of the genus, the most similar to A. chrysalis is A. elongatus. However, the main difference between them is the abundance of tetractines: the former has abundant tetractines in the subatrial region, while in A. elongatus tetractines were observed to be rare. Additionally, the actines of the subatrial triactines of A. chrysalis are more conical and bear blunt tips, unlike A. elongatus, which has more cylindrical actines and with sharp tips. Distribution: Amphoriscus chrysalis is one of the few species of the genus that has several records besides its type locality. It has been described in the Adriatic and Celtic Seas (Schmidt 1864; Borojevic et al. 1968) and is reported here in the Mediterranean part of France and the Bay of Biscay (Fig. 3). The corresponding marine ecoregions of the world (MEOW) are the Celtic Seas, South European Atlantic Shelf, Adriatic Sea, and Western Mediterranean (Spalding et al. 2007)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on pages 41-45, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287, {"references":["Schmidt, O. (1864) Supplement der Spongien des adriatischen Meeres. Enthaltend die Histologie und systematische Erganzungen. Wilhelm Engelmann, Leipzig, 48 pp.","Dendy, A. & Row, R. W. H. (1913) The Classification and Phylogeny of the Calcareous Sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London, 3, 704 - 813. https: // doi. org / 10.1111 / j. 1469 - 7998.1913. tb 06152. x","Haeckel, E. (1870) Prodromus of a system of the calcareous sponges. Annals and Magazine of Natural History, 4 (5), 176 - 191. https: // doi. org / 10.1080 / 00222937008696137","Haeckel, E. (1872) Die Kalkschwamme. Eine Monographie in zwei Banden Text und einem Atlas mit 60 Tafeln Abbildungen. Vols. 1 - 3. Reimer, Berlin, 484 pp., 418 pp. & 60 pp.","Polejaeff, N. (1883) Report on the Calcarea dredged by H. M. S. ' Challenger', during the years 1873 - 1876. Report on the Scientific Results of the Voyage of H. M. S. ' Challenger', 1873 - 1876. Zoology, 8 (2), 1 - 76.","Breitfuss, L. (1935) La spugne calcarea dell'Adriatico con riflesso a tutto il Mediterraneo. Memorie Reale Comitato Talassographico Italiano, Venezia, 2223, 1 - 45.","Burton, M. (1963) A revision of the Classification of the Calcareous Sponges: with a Catalogue of the specimens in the British Museum. Order of the trustees of the British Museum (Natural History), London, 693 pp.","Vacelet, J. (1981) Etude qualitative et quantitative des salissures biologiques de plaques experimentales immergees en pleine eau. Les eponges. Tethys, 10 (2), 165 - 172.","Radolovic, M., Bakran-Petricioli, T., Petricioli, D., Suric, M. & Perica, D. (2015) Biological response to geochemical and hydrological processes in a shallow submarine cave. Mediterranean Marine Science, 16 (2), 305 - 324. https: // doi. org / 10.12681 / mms. 1146","Klautau, M., Cavalcanti, F. F. & Borojevic, R. (2017) The new sponge species Amphoriscus pedunculatus (Porifera, Calcarea). Zootaxa, 4341 (1), 105 - 112. https: // doi. org / 10.11646 / zootaxa. 4341.1.9","Condor-Lujan, B., Azevedo, F., Hajdu, E., Hooker, Y., Willenz, P. & Klautau, M. (2019) Tropical Eastern Pacific Amphoriscidae Dendy, 1892 (Porifera: Calcarea: Calcaronea: Leucosolenida) from the Peruvian coast. Marine Biodiversity, 49 (4), 1 - 18. https: // doi. org / 10.1007 / s 12526 - 019 - 00946 - y","Borojevic, R., Cabioch, L. & Levi, C. (1968) Inventaire de la faune marine de Roscoff. Spongiaires. Cahiers de Biologie Marine, 9 (1), 1 - 44.","Spalding, M. D., Fox, H. E., Allen, G. R., Davidson, N., Ferdana, Z. A., Finlayson, M., Halpern, B. S., Jorge, M. A., Lombana, A., Lourie, S. A., Martin, K. D., McManus, E., Molnar, J., Recchia, C. A. & Robertson, J. (2007) Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. BioScience, 57 (7), 573 - 583. https: // doi. org / 10.1641 / B 570707"]}

Family BAERIIDAE Borojević, Boury-Esnault & Vacelet, 2000 “ Baerida with a choanoskeleton consisting of giant triactines, and/or of tetractines in no particular order, and/or of very numerous microdiactines. No traces of radial organisation can be seen in the choanoskeleton. The cortical skeleton consists of triactines, giant diactines, and/or numerous microdiactines, and occasionally the basal actines of cortical giant tetractines. The choanoskeleton consists of scattered spicules similar to those observed in the cortex, to which numerous microdiactines can be added or which can be entirely replaced by microdiactines. The exhalant aquiferous system is formed by ramified canals that have no tangential skeleton, being loosely or densely covered by harpoon-shaped pugioles and/or microdiactines” (Borojevic et al. 2002).

Comparison of Amphoriscus species In order to facilitate the comparison between species of Amphoriscus and, consequently, their taxonomic identification, the skeletal composition and type locality of all Amphoriscus species is summarised below (Tab. 8)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on page 64, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287

Family BAERIIDAE Borojević, Boury-Esnault & Vacelet, 2000 ��� Baerida with a choanoskeleton consisting of giant triactines, and/or of tetractines in no particular order, and/or of very numerous microdiactines. No traces of radial organisation can be seen in the choanoskeleton. The cortical skeleton consists of triactines, giant diactines, and/or numerous microdiactines, and occasionally the basal actines of cortical giant tetractines. The choanoskeleton consists of scattered spicules similar to those observed in the cortex, to which numerous microdiactines can be added or which can be entirely replaced by microdiactines. The exhalant aquiferous system is formed by ramified canals that have no tangential skeleton, being loosely or densely covered by harpoon-shaped pugioles and/or microdiactines��� (Borojevic et al. 2002)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on page 63, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287, {"references":["Borojevic, R., Boury-Esnault, N. & Vacelet, J. (2000) A revision of the supraspecific classification of the subclass Calcaronea (Porifera, class Calcarea). Zoosystema, 22 (2), 203 - 263.","Borojevic, R., Boury-Esnault, N., Manuel, M. & Vacelet, J. (2002) Order Leucosolenida Hartman, 1958. In: Hooper, J. N. A. & Van Soest, R. W. M. (Eds.), Systema Porifera: a guide to the classification of sponges. Vol. 2. Kluwer Academic / Plenum Publishers, New York, Boston, Dordrecht, London and Moscow, pp. 1157 - 1184. https: // doi. org / 10.1007 / 978 - 1 - 4615 - 0747 - 5 _ 120"]}

Genus Leuconia Grant, 1833 ��� Baeriidae in which the choanoskeleton consists of giant triactines and/or tetractines, lying without apparent order, and of very numerous microdiactines. A cavity equivalent to the atrium, localised only under the oscula, has a skeleton supported by tangential triactines. All the other exhalant canals have a skeleton composed of harpoon-shaped pugioles��� (Borojevic et al. 2002)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on page 63, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287, {"references":["Borojevic, R., Boury-Esnault, N., Manuel, M. & Vacelet, J. (2002) Order Leucosolenida Hartman, 1958. In: Hooper, J. N. A. & Van Soest, R. W. M. (Eds.), Systema Porifera: a guide to the classification of sponges. Vol. 2. Kluwer Academic / Plenum Publishers, New York, Boston, Dordrecht, London and Moscow, pp. 1157 - 1184. https: // doi. org / 10.1007 / 978 - 1 - 4615 - 0747 - 5 _ 120"]}

Amphoriscus semoni Breitfuss, 1896 Amphoriscus semoni Breitfuss, 1896 Citations: Amphoriscus semoni Breitfuss 1896: 435; 1898: 221; Dendy & Row 1913: 782; Burton 1963: 542; Van Soest & De Voogd 2015: 94; Klautau, Cavalcanti & Borojevic 2017: 105; Van Soest & De Voogd 2018: 130; C��ndor-Luj��n et al. 2019: 1825. Type material: ZMB 2698 (Holotype; Ambon Island, Maluku, Indonesia). Not analysed in the present work. Type locality: Ambon Island, Maluku, Indonesia. Analysed material: BMNH 1886.6.7.32 (two slides containing sections of the skeleton). Port Jackson, Australia; previously identified as A. cylindrus in Burton, 1963: 634. Morphology: The holotype of A. semoni was not available for redescription, and the material analysed here (deposited in the NHM) was comprised of two microscope slides. Aquiferous system is syconoid. Anatomy: The material evaluated in the present study (BMNH 1886.6.7.32) was deposited at the Porifera collection of the NHM under the name A. cylindrus (Fig. 9). It is formed by giant cortical tetractines, with paired and unpaired actines laying on the cortex (Figs. 9A, B). The organisation of the skeleton is inarticulate due to the unpaired actine of the subatrial triactines and the apical actine of the cortical tetractines (Figs. 9A���C). The atrium is perforated by the apical actine of the small atrial tetractines (Fig. 9D). Spicules (Tables 5 and 6): A figure containing only the spicule categories could not be prepared due to the lack of a slide of dissociated spicules. Cortical tetractines: Irregular, conical, and sharp. The basal actines (paired and unpaired) are slightly curved from the base to the tips. The apical actine is straight and long. Subatrial triactines: Irregular, cylindrical to slightly conical, sharp. The paired actines are straight. The unpaired actine is longer or the same size as the paired actines. Atrial tetractines: Irregular, cylindrical, small, thin, and sharp. The paired actines are curved. The unpaired actine is similar to the paired ones. The apical actine is straight and short, always very thin. Remarks: BMNH 1886.6.7.32 was deposited in the NHM as A. cylindrus (Burton 1963), but its subatrial skeleton is comprised of triactines and not tetractines, as in A. cylindrus. The only species of Amphoriscus whose organisation is consistent with this specimen���cortical tetractines, subatrial triactines, and atrial tetractines���is A. semoni. The specimen is from Port Jackson, Australia, in the same biogeographical region as the type locality of A. semoni (Ambon Island, Indonesia) and isolated from the type locality of A. cylindrus (the Adriatic Sea). Therefore, we suggest the identification of BMNH 1886.6.7.32 as A. semoni. Figure 9 shows its skeletal organisation, in which the subatrial and atrial layers are clearly evident. In our opinion, these illustrations are important for further recognition of specimens belonging to A. semoni. Among the characters described by Breitfuss (1896), A. semoni is bright white when preserved, and the atrial surface is perforated by the apical actine of the colossal cortical tetractines. The specimens described as A. semoni by Van Soest & De Voogd (2015, 2018) present minor differences in the colour and length of these apical actines. The sample ZMAPOR 08073, collected in Sumbawa, Indonesia, was green alive and beige after fixation. The apical actine of its cortical tetractines does not exceed 600 ��m (see Tables 8 and 9; Van Soest & De Voogd 2015), while in the holotype, they vary from 520 to 790 ��m (Breitfuss 1896). In their most recent study, Van Soest & De Voogd (2018) described a new set of specimens of A. semoni (ZMAPOR 10527), sampled in Seychelles, Western Indian Ocean, which were white alive and beige after fixation. The apical actines of the cortical tetractines do not perforate the atrium, although they measure 239���882 ��m (Van Soest & De Voogd 2018). The differences between the holotype and the other specimens described in both works by Van Soest & De Voogd provide a better understanding of the intraspecific variation of A. semoni. These are precious data, usually rare for Amphoriscus species since the original description, based on one or few specimens, is all that is available for many species of the genus. Amphoriscus semoni resembles two species of the Adriatic Sea, namely A. gregorii and A. bucchichii. However, A. semoni differs in having a cortical skeleton comprised exclusively of tetractines. Distribution: Ambon Island, Indonesia (Breitfuss 1896); Seychelles (Van Soest & De Voogd 2018); Port Jackson, Australia (present study). Corresponding MEOW: Banda Sea, Seychelles, and Manning-Hawkesbury (Spalding et al. 2007)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on pages 54-57, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287, {"references":["Breitfuss, L. (1896) Amphoriscus semoni, eine neue Art heterocoeler Kalkschwamme. Zoologischer Anzeiger, 19 (515), 435 - 436.","Dendy, A. & Row, R. W. H. (1913) The Classification and Phylogeny of the Calcareous Sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London, 3, 704 - 813. https: // doi. org / 10.1111 / j. 1469 - 7998.1913. tb 06152. x","Burton, M. (1963) A revision of the Classification of the Calcareous Sponges: with a Catalogue of the specimens in the British Museum. Order of the trustees of the British Museum (Natural History), London, 693 pp.","Van Soest, R. W. M. & De Voogd, N. J. (2015) Calcareous sponges of Indonesia. Zootaxa, 3951 (1), 1 - 105. https: // doi. org / 10.11646 / zootaxa. 3951.1.1","Klautau, M., Cavalcanti, F. F. & Borojevic, R. (2017) The new sponge species Amphoriscus pedunculatus (Porifera, Calcarea). Zootaxa, 4341 (1), 105 - 112. https: // doi. org / 10.11646 / zootaxa. 4341.1.9","Van Soest, R. W. M. & De Voogd, N. J. (2018) Calcareous sponges of the Western Indian Ocean and Red Sea. Zootaxa, 4426 (1), 1 - 160. https: // doi. org / 10.11646 / zootaxa. 4426.1.1","Condor-Lujan, B., Azevedo, F., Hajdu, E., Hooker, Y., Willenz, P. & Klautau, M. (2019) Tropical Eastern Pacific Amphoriscidae Dendy, 1892 (Porifera: Calcarea: Calcaronea: Leucosolenida) from the Peruvian coast. Marine Biodiversity, 49 (4), 1 - 18. https: // doi. org / 10.1007 / s 12526 - 019 - 00946 - y","Spalding, M. D., Fox, H. E., Allen, G. R., Davidson, N., Ferdana, Z. A., Finlayson, M., Halpern, B. S., Jorge, M. A., Lombana, A., Lourie, S. A., Martin, K. D., McManus, E., Molnar, J., Recchia, C. A. & Robertson, J. (2007) Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. BioScience, 57 (7), 573 - 583. https: // doi. org / 10.1641 / B 570707"]}

Amphoriscus chrysalis (Schmidt, 1864) Ute chrysalis Schmidt, 1864 (type species by subsequent designation; Dendy & Row,1913) Citations and synonymies: Ute chrysalis Schmidt 1864: 23 (original description); Amphoriscus chrysalis Haeckel 1870: 177; Sycilla chrysalis Haeckel 1872: 256; Sycurus chrysalis Haeckel 1872: 256 (variation of Sycilla chrysalis); Amphoriscus chrysalis Poléjaeff 1883: 7; Dendy & Row 1913: 782; Breitfuss 1935: 29; Burton 1963: 535; Vacelet 1981: 165; Radolović et al. 2015: 305; Klautau, Cavalcanti & Borojevic 2017: 105; Cóndor-Luján et al. 2019: 1825. Type material: Unknown. Type locality: Lesina, Lissa, Adriatic Sea. Analysed material: LBIM 1968-557 L (Glenans, France; four slides containing sections of the skeleton). LBIM 1968-556 L (Marseille, France; three slides containing sections of the skeleton). LBIM 1968-327 (Roscoff, France; four slides containing sections of the skeleton). LBIM 1968-355 (Roscoff, France; two slides containing sections of the skeleton). LBIM 1968-326 (Roscoff, France; seven slides containing sections of the skeleton and dissociated spicules). LBIM 1968-280 (Roscoff, France; one slide containing dissociated spicules). BMNH 1954.8.12.195 (Plymouth; M.B. L. collection; one slide containing sections of the skeleton). BMNH 195512.13.9 (Plymouth; R.W.H. collection; one slide containing sections of the skeleton; not A. chrysalis — see below). SME-500 (Biscay Bay, France; one specimen and one slide containing sections of the skeleton). Morphology: Colour in spirit varies from yellowish-white to yellowish-brown (Burton 1963). The specimen SME-500 has a tubular shape and measures ca. 4 mm x 1 mm (Fig. 1A). According to Schmidt (1864), the type had a peduncle, which is not visible in SME-500. Smooth surface and a single osculum without fringe of trichoxeas. Syconoid aquiferous system. Abundant larvae were observed in the choanosome of LBIM 1968-556 L (Figs. 1B, D, E). Anatomy: Skeleton typical of the family Amphoriscidae (Fig. 1B). The cortical region is formed mainly by giant and abundant tetractines and a few tangential triactines, pierced by tufts of trichoxeas (Fig. 1C). Abundant subatrial tetractines and triactines point their unpaired actine towards the cortex (Fig. 1E). The atrial skeleton is formed exclusively by tetractines, and these have a very long apical actine (Fig. 1F). Some lacunae occur along the subcortical and subatrial regions, mainly in the specimen LBIM 1968-556 L, possibly due to fixation and/or the occurrence of tissue retraction over time (Fig. 1D). Fusiform diactines were found in the spicules slides of LBIM 1968-326, but these were not observed in the sections of the skeleton of the same specimen and were not found in any other analysed material. In addition, BMNH 1955.12.13.9 differs from the remaining analysed samples in its skeletal composition: cortical tetractines, subatrial triactines, and atrial tetractines, and thus cannot be classified as A. chrysalis (see Remarks). Spicules: Among the analysed material, slides containing dissociated spicules were available only for LBIM 1968-326. The values shown here therefore only reflect this sample. Cortical tetractines (Fig. 2A): Conical with blunt tips. The paired actines are curved, and one of the tips is broken in most of the spicules (118.8– 298.3 ±90.5–520.2/ 23.8– 38.9 ±7.9–53.1 μm). The unpaired actine is smaller or the same size as the paired actines (106.1– 203.6 ±107.4–485.1/ 22.2– 41.5 ±8.0–58.6 μm). The apical actine is straight and longer than the other actines (281.5– 421.6 ±87.2–636.2/ 23.8– 39.8 ±7.7–53.7 μm). Cortical triactines (Fig. 2B): Conical with blunt tips, less abundant than the cortical tetractines. Paired actines are slightly curved (246.2– 360.5 ±66.8–466.0/ 22.3– 35.6 ±6.3–45.2 μm). The unpaired actine is straight, sometimes slightly lanceolate (241.7– 468.8 ±140.7–673.9/ 23.1– 37.2 ±8.1–52.6 μm). Subatrial triactines and tetractines (Figs. 2C, D): Cylindrical to slightly conical and blunt. The paired actines are short and straight (triactines: 102.1– 145.0 ±23.3–185.3/ 6.2– 10.2 ±1.9–13.6 μm; tetractines: 92.9– 139.7 ±45.5–253.1/ 6.3– 8.5 ±2.0–11.4 μm). The unpaired actines are straight and long (triactines: 232.5– 340.4 ±49.6–435.3/ 8.5– 13.0 ±2.4–17.4 μm; tetractines: 142.6– 271.3 ±67.6–350.8/ 8.0– 9.6 ±1.2–11.1 μm). The apical actine of the tetractines is short and curved (20.3– 31.0 ±14.3–68.6/ 5.3– 7.6 ±1.7–10.1 μm). Atrial tetractines(Fig.2E):Cylindrical with sharp tips.The paired actines are long and curved (100.3– 173.0 ±49.3– 315.4/ 6.0– 14.1 ±15.9–97.1 μm). The unpaired actine is straight (62.5– 169.3 ±95.9–445.6/ 4.9– 12.3 ±3.7–21.2 μm). The apical actine is straight and long. It is generally twice as large as the paired actines (102.2– 271.7 ±126.4–527.2/ 7.9– 11.9 ±2.6–17.8 μm). Remarks: According to the original description (Schmidt 1864), Amphoriscus chrysalis is comprised only of tetractines (the presence of only this spicule category and the external morphology are the only characters mentioned by the author). The material analysed here has cortical and subatrial triactines, differing from the original description. Haeckel (1872) also described A. chrysalis as having only tetractines, though subatrial (and possibly cortical) triactines similar to those we observed were illustrated in his work (Plate 43, figures 2 and 3). It is not clear if Haeckel analysed the same material described by Schmidt (1864), but the short original description lacking either spicule measurements or illustrations of the spicule types and skeleton show the importance of Haeckel’s (1872) work for the recognition of A. chrysalis. The difficulty in recognising this species is also caused by a lack of type material. Our results and the figure provided by Haeckel (1872) thus allow us to state that A. chrysalis has triactines. We were also able to recognise the main feature of A. chrysalis reported by Haeckel (1872) in the samples analysed here: the long apical actine of the atrial tetractines (>100 μm). This feature was important for supporting our decision to identify some samples as A. chrysalis sensu Haeckel (1872). Among the analysed samples, BMNH 1954.8.12.195 and BMNH 1955.12.13.9 had been listed by Burton (1963: 634) under the name A. chrysalis. Our results indicate that the latter does not belong to A. chrysalis due to its different skeletal composition. Here, we allocate it in A. decennis sp. nov., while BMNH 1954.8.12.195 remains A. chrysalis. Some of the samples analysed in this study had previously been identified as A. chrysalis by Borojevic et al. (1968), namely: LBIM 1968-557, LBIM 1968-556 L, LBIM 1968-327, LBIM 1968-355, LBIM 1968-326, and LBIM 1968-280. The list of species from Roscoff was provided without morphological details, meaning that these specimens are described for the first time in this study. Compared to the other species of the genus, the most similar to A. chrysalis is A. elongatus. However, the main difference between them is the abundance of tetractines: the former has abundant tetractines in the subatrial region, while in A. elongatus tetractines were observed to be rare. Additionally, the actines of the subatrial triactines of A. chrysalis are more conical and bear blunt tips, unlike A. elongatus, which has more cylindrical actines and with sharp tips. Distribution: Amphoriscus chrysalis is one of the few species of the genus that has several records besides its type locality. It has been described in the Adriatic and Celtic Seas (Schmidt 1864; Borojevic et al. 1968) and is reported here in the Mediterranean part of France and the Bay of Biscay (Fig. 3). The corresponding marine ecoregions of the world (MEOW) are the Celtic Seas, South European Atlantic Shelf, Adriatic Sea, and Western Mediterranean (Spalding et al. 2007).

Order BAERIDA Borojević, Boury-Esnault & Vacelet, 2000 “Leuconoid Calcaronea with the skeleton either composed exclusively of microdiactines or in which microdiactines constitute exclusively or predominantly a specific sector of the skeleton, such as choanoskeleton or atrial skeleton. Large or giant spicules are frequently present in the cortical skeleton, from which they can partially or fully invade the choanoderm. In sponges with a reinforced cortex, the inhalant pores can be restricted to a sieve-like ostia-bearing region. Dagger-shaped small tetractines (pugioles) are frequently the sole skeleton of the exhalant aquiferous system. Although the skeleton may be highly reinforced by the presence of dense layers of microdiactines in a specific region, an aspicular calcareous skeleton is not present” (Borojevic et al. 2002).

Amphoriscus elongatus Poléjaeff, 1883 Citations and synonymies: Amphoriscus elongatus Poléjaeff 1883: 48, pl. iv, fig. 5, pl. v, fig. 4; Dendy & Row 1913: 782; Burton 1956: 117; Burton 1963: 538; Klautau et al. 2017: 106; Cóndor-Luján et al. 2019: 1825. Type material: BMNH 1884.4.22.27 (holotype). Station 145, off Prince Edward Islands (46° 40’ S – 37° 50’ E), 275 to 567 m depth, 27 December 1873. Type locality: off Prince Edward Islands, Indian Ocean. Analysed material: BMNH 1884.4.22.27 (holotype; specimen and one slide containing sections of the skeleton). BMNH 1955.12.13.10 (one slide containing sections of the skeleton) and BMNH 1955.12.13.11 (one slide containing sections of the skeleton), both from Plymouth; R.W.H. Row collection. Morphology: The colour of the holotype in ethanol is white (Fig. 4A). It is a fragment with tubular shape and apical osculum, measuring 4 cm x 0.2 cm (length x width). The surface is slightly hispid. Syconoid aquiferous system (Fig. 5). Anatomy: The skeleton is typical of the genus Amphoriscus, and the inarticulation is evident (Figs. 4D and 5D, E). The cortical region is formed by giant tetractines and small sagittal triactines (Figs. 4B, C). The apical actine of these giant tetractines may reach the atrial cavity. All the analysed samples exhibited trichoxeas perforating the cortex without forming tufts (Fig. 5C). The subatrial region is formed by abundant triactines with long unpaired actine and rare tetractines (Fig. 5D). The latter are more frequent in the specimens BMNH 1955.12.13.10 and BMNH 1955.12.13.11 but also occur in the holotype. Additionally, in these specimens from Plymouth, we also observed a few modified subatrial triactines in which the actines that surround the atrial wall have different lengths and shapes (Fig. 5E). These spicules were found in the sections. Due to the occurrence of other spicules overlapping them, it was not clear if they were pseudosagittal. All the subatrial spicules point their unpaired actines to the cortex, in opposition to the apical actine of the cortical tetractines (Figs. 4D and 5D, E). The atrial region is comprised exclusively of tetractines (Fig. 5F). Spicules (Table 1): A figure containing only the spicule categories, commonly shown in studies of the taxonomy of calcareous sponges, could not be prepared since only slides containing sections of the skeleton were available. Cortical tetractines: Giant, conical with blunt tips. Paired actines are curved and long. The unpaired actine is curved and smaller or the same size as the paired actines. The apical actine is straight and long. Cortical triactines: Slightly conical and sharp. Paired actines are straight to slightly curved. Unpaired actine is shorter and straight. Subatrial triactines: Cylindrical to slightly conical with sharp tips. Paired actines are straight. Unpaired actine is straight and long, sometimes reaching the cortical region. Subatrial tetractines: Rare, with cylindrical actines and sharp tips. Paired actines are slightly curved. Unpaired actine is straight and larger than the paired ones. Apical actine is short and curved. Atrial tetractines: Vary in size. Cylindrical actines with sharp tips. Paired actines are long, thin, and straight. Unpaired actine is slightly curved. Apical actine is straight. Remarks: The original description of Amphoriscus elongatus was based on a single specimen sampled during the Challenger expedition (Poléjaeff 1883). It was tubular and elongated (hence the species name elongatus). Close to the osculum, it was divided into two tubes, a feature that can no longer be recognised in the type (Fig. 4A). According to Poléjaeff (1883), A. elongatus has “an important anatomical peculiarity—the tendency of the radial tubes to meet in threes, fours, or in larger numbers around the same shallow invagination of the gastric cavity”. We could not clearly understand this organisation described by Poléjaeff (1883), even after analysing the holotype and additional specimens. Nevertheless, we are convinced that the aquiferous system of A. elongatus is not a typical syconoid in which choanocyte chambers run in parallel from the atrium to the cortex. Instead, choanocyte chambers of A. elongatus seem to branch, possibly causing, when sectioned, the holes observed in Figure 5 along the chambers’ length. More studies are needed to better describe this organisation, which depends on the discovery of fresh specimens for histological and electron microscopy analyses. Our results indicate an important difference between our description of the holotype and that provided by Poléjaeff (1883), namely the occurrence of rare subatrial tetractines. We first observed this characteristic in the slides of the specimens from Plymouth and then, after a careful reanalysis, also in the holotype. With respect to the geographical distribution, BMNH 1955.12.13.10 and BMNH 1955.12.13.11 are from Plymouth, United Kingdom (Northern Atlantic Ocean), while the holotype BMNH 1884.4.22.27 was sampled “off Prince Edward Islands” (Southern Indian Ocean). In general, representatives of a species with such a disjunct distribution are viewed as questionable in the literature, mainly due to the low dispersal capability of the larvae of calcareous sponges (Maldonado 2006). The only morphological difference between the three analysed samples is the putative presence of pseudosagittal triactines in the subatrial region of those from Plymouth. These spicules were not found in the holotype, and we assumed that the absence could be the result of plasticity. The analysis of a larger set of specimens, preferably using an integrative approach, is needed to elucidate this question. The species that most closely resembles A. elongatus in its skeletal composition is A. pedunculatus (cortical triactines, tetractines and trichoxeas, subatrial triactines, and atrial tetractines). These species differ mainly in the presence of rare subatrial tetractines in the former and the size of the cortical tetractines, which are larger in A. elongatus (paired actines: 443.6±59.0/ 48.7±7.9μm; unpaired actine: 397.2±42.1/ 52.7±4.9 μm; apical actine: 453.8±60.3/48.2±6.4μm) than in A. pedunculatus (paired actines: 225.5±39.3/ 27.2 ±4.3 μm; unpaired actine: 182.9±7.2/ 27.2±3.8 μm; apical actine: 303.8±50.7/ 29.4±5.4 μm). Distribution: Off Prince Edward Islands, South Africa (Poléjaeff 1883) and Plymouth, United Kingdom (present study). Corresponding MEOW: Prince Edward Islands and Celtic Seas (Spalding et al. 2007).

Family AMPHORISCIDAE Dendy, 1892 ��� Leucosolenida with syconoid, sylleibid or leuconoid organisation, and a distinct cortex supported by tangential tetractines whose centripetal apical actines cross the outer part or the whole of the choanosome. Tangential triactines and small tetractines may also be present in the cortex. The choanoskeleton is typically inarticulated, composed of apical actines of cortical tetractines and the unpaired actines of subatrial spicules. In species with a thick wall, scattered triactines and/or tetractines may also be present, either among the spicules of the inarticulated choanoskeleton or forming a distinct subatrial layer. An atrial skeleton is always present��� (Borojevic et al. 2002)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on page 41, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287, {"references":["Dendy, A. (1892) Synopsis of the Australian Calcarea Heterocoela; with a proposed Classification of the group and descriptions of some new genera and species. Proceedings of the Royal Society of Victoria, New Series, 5, 69 - 116.","Borojevic, R., Boury-Esnault, N., Manuel, M. & Vacelet, J. (2002) Order Leucosolenida Hartman, 1958. In: Hooper, J. N. A. & Van Soest, R. W. M. (Eds.), Systema Porifera: a guide to the classification of sponges. Vol. 2. Kluwer Academic / Plenum Publishers, New York, Boston, Dordrecht, London and Moscow, pp. 1157 - 1184. https: // doi. org / 10.1007 / 978 - 1 - 4615 - 0747 - 5 _ 120"]}

Amphoriscus gastrorhabdifera (Burton, 1932) Leucaltis gastrorhabdifera Burton, 1932 Citations and Synonymies: Leucaltis gastrorhabdifera Burton, 1932: 259, figs. 4-5; Amphoriscus gastrorhabdifera Burton, 1963: 136, 147, 548; Klautau et al. 2017: 105-106; C��ndor-Luj��n et al. 2019: 1825. Type material: BMNH 1928.2.15.833 (Holotype; St. 6, Tristan da Cunha, South Atlantic; 80���140 m deep). Type locality: Tristan da Cunha, South Atlantic. Morphology: Colour is beige after fixation. The holotype is a tubular fragment (Fig. 12A), and a fringe of trichoxeas was present, according to Burton (1932). The type of aquiferous system is unclear. Anatomy: The skeleton has no similarity with the other species of Amphoriscus. The cortical region is formed by triactines of varying sizes and subcortical giant tetractines (which are possibly the only typical character of Amphoriscidae). However, there is no typical inarticulation (Fig. 12B). The presence of several broken spicules makes the visualisation of the triactines difficult (Fig. 12C). The apical actines of the subcortical tetractines cross the entire choanosome, often perforating the atrium (Figs. 12D, E). The subatrial/atrial region is comprised exclusively of large tangential diactines. They are abundant and line the atrial cavity (Figs. 12E, F). A specific atrial skeleton comprised of triactines or tetractines is absent. According to Burton (1932), large diactines similar to those found in the inner part of the sponge were found projecting from the cortex, but they were not observed here. The aquiferous system could not be recognised and was not mentioned along the original description. Spicules: Cortical triactines: Regular, actines are slightly conical and sharp. The actines are straight (paired actines: 123.1��� 150.7 ��21.6���204.4/ 8.4��� 12.2 �� 1.7���15.3 ��m; unpaired actines: 81.5��� 133.6 ��23.2���180.0/ 8.9��� 122.3 ��2.0���18.3 ��m) (Fig. 13A). Subcortical tetractines: Giant, actines are slightly conical to conical and blunt. The paired actines are slightly curved (111.4��� 146.6 ��21.8���192.1/ 10.8��� 14.5 �� 2.0���19.2 ��m). The unpaired actine is curved from the base to the tip (93.9��� 116.7 ��21.5���156.6/ 14.1��� 15.2 ��1.1���16.9 ��m). The apical actine is long and conical (174.3��� 228.3 ��29.1���285.4/ 16.9��� 20.8 ��2.2���26.3 ��m) (Fig. 13B). Diactines: Large, sinuous along their length. Most of them have one of the tips blunt while the other is sharp (330.0��� 464.8 ��89.6���632.6/ 9.0��� 18.9 ��4.0���25.7 ��m) (Fig. 13C). Remarks: This species was originally described as Leucaltis gastrorhabdifera (subclass Calcinea) and was later transferred to Amphoriscus (subclass Calcaronea) (Burton 1932, 1963). Such a reallocation would, by the standards of modern-day research on Calcarea systematics, be considered out of the ordinary and require a detailed explanation, yet at that time Burton did not note the reasons supporting his decision in any detail. In his discussion of the genus Leucaltis, Burton (1963) argued that ��� L. gastrorhabdifera is aberrant and is here doubtfully assigned to Amphoriscus ���, suggesting that the only certainty the author had was that it was not Leucaltis. He named the species ��� Amphoriscus ? gastrorhabdifera ���, and although we suspect that he was influenced by the presence of the giant cortical tetractines, this cannot be confirmed.Also, there is no information or illustration in the literature about the aquiferous system of A. gastrorhabdifera, which would be essential to support its allocation in Amphoriscus. Whether or not it was one of the characters used by Burton to assign the species in Amphoriscus is a question that remains unanswered. We analysed the holotype (BMNH 1928.2.15.833) in this study. The microscopical slides contain sections of the skeleton that were probably not stained since it was not possible to unequivocally confirm whether A.gastrorhabdifera has the syconoid aquiferous system typical of the genus. We observed the main morphological characters reported in the original description, such as the presence of cortical triactines, subcortical giant tetractines with a long apical actine, and diactines. Burton (1932) mentioned diactines protruding through the cortex, but we did not observe this characteristic. The absence of a subatrial layer of triactines/tetractines was also confirmed and, consequently, the absence of inarticulate skeletal organisation. Therefore, the most important diagnostic characters typical of Amphoriscus either could not be confirmed (the syconoid aquiferous system) or are absent (the inarticulation formed by the apical actines of giant cortical tetractines and the unpaired actine of subatrial spicules). In order to assess the possibility of assigning A. gastrorhabdifera in another genus, a diagnosis of all genera under the order Leucosolenida was carried out. The presence of triactines and tetractines and of an atrial layer of diactines is a remarkable character found only in Sycodorus, although members of this genus are syconoid. Tangential triactines and tetractines also occur along the atrial cavity, suggesting that, even if A. gastrorhabdifera is syconoid, it could not be allocated in Sycodorus. An alternative decision would be to propose a new genus. The reasons why we did not choose this option to solve the problem of A. gastrorhabdifera were as follows: (i) the holotype is tiny, and no reports of additional specimens exist that could enrich our understanding of the morphology of the new genus; (ii) the lack of data on the aquiferous system could raise doubts on the family in which the genus should be inserted and also makes the description of a robust diagnosis difficult; (iii) the species is represented only by the holotype, found at Tristan da Cunha, Southern Atlantic Ocean, at a depth of 80 to 140 m. The lack of perspective in finding fresh samples to elucidate the questions mentioned earlier suggests that the problem with the species would persist (not as Amphoriscus but as a newly named genus); finally, (iv) we cannot undoubtedly rule out A. gastrorhabdifera actually belonging to Amphoriscus as the absence of an inarticulate organisation could be a secondary character caused by the loss of subatrial spicules along the species��� evolutionary history. The latter question will be resolved after the species is tested in phylogenetic analyses, though, as highlighted above, there is no fresh material available. Therefore, we decided to be conservative and avoid potentially increasing the number of open questions on the classification of this species. Amphoriscus gastrorhabdifera is thus maintained in the genus, but we indicate that it should be considered as incertae sedis until the discovery of additional samples., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on pages 60-62, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287, {"references":["Burton, M. (1932) Sponges. Discovery Reports, 6, 237 - 392. https: // doi. org / 10.5962 / bhl. part. 24379","Burton, M. (1963) A revision of the Classification of the Calcareous Sponges: with a Catalogue of the specimens in the British Museum. Order of the trustees of the British Museum (Natural History), London, 693 pp.","Klautau, M., Cavalcanti, F. F. & Borojevic, R. (2017) The new sponge species Amphoriscus pedunculatus (Porifera, Calcarea). Zootaxa, 4341 (1), 105 - 112. https: // doi. org / 10.11646 / zootaxa. 4341.1.9","Condor-Lujan, B., Azevedo, F., Hajdu, E., Hooker, Y., Willenz, P. & Klautau, M. (2019) Tropical Eastern Pacific Amphoriscidae Dendy, 1892 (Porifera: Calcarea: Calcaronea: Leucosolenida) from the Peruvian coast. Marine Biodiversity, 49 (4), 1 - 18. https: // doi. org / 10.1007 / s 12526 - 019 - 00946 - y"]}

Amphoriscus decennis sp. nov. Etymology: From the Latin decem + annus (meaning decades). The name is related to the time span the specimens remained in the collection before being recognised as a new species. Diagnosis: Amphoriscus with cortical skeleton comprised of trichoxeas and one type of tetractine variable in size, subatrial skeleton comprised of triactines and atrial tetractines. Type material: UFRJPOR 9031 holotype (Île Riou, Marseille, France; 05/VI/1967). UFRJPOR 9032, UFRJPOR 9033, UFRJPOR 9034, and UFRJPOR 9035— paratypes (Île Riou, Marseille, France; 05/VI/1967). Type locality: Île Riou, Marseille, France. Additional material: UFRJPOR 5822 (Île Riou, Marseille, France; 05/VI/1967). Several fragments from one or more specimens. BMNH 1955.12.13.9 (Plymouth; R. W. H. Row). Morphology: The specimens are cylindrical, with differences in thickness along the body, as they are thicker at the base. Smooth surface, colour white in spirit. The osculum is apical and naked. The holotype (UFRJPOR 9031) measures 2.0 x 1.0 cm (height x width), while the largest specimen, the paratype UFRJPOR 9035, measures 3.0 x 1.0 cm (Fig. 10A). The atrial cavity occupies the entire body of the specimen. The aquiferous system is syconoid and seems slightly disorganised in the paratype UFRJPOR 9032, possibly due to the presence of embryos and amphiblastulae. Anatomy: The organisation is typical of Amphoriscidae, with an inarticulate skeleton formed mainly by giant cortical tetractines (Fig. 10B). Smaller tetractines also occur in the cortex and are less abundant than the giant ones. Trichoxeas are present (Fig. 10C). The inarticulation of the skeleton is due to the apical actine of the cortical tetractines and the unpaired actines of subatrial triactines (Fig. 10D). The atrial region is comprised exclusively of tetractines with short apical actines (Fig. 10E, F). These short apical actines and those of the giant cortical tetractines perforate the atrial cavity, making the atrial surface hispid. Spicules (Table 7; Fig. 11): Cortical tetractines: Actines are conical and blunt. The paired actines are curved, while the unpaired one is straight. Apical actine is long, straight, and usually extends up to the atrial cavity (Figs. 11A, A’). Subatrial triactines: Actines are cylindrical to slightly conical, with blunt tips. The paired actines are curved from the base to the tips and are always smaller than the unpaired one (Fig. 11B). Atrial tetractines: Actines are cylindrical and sharp. The paired actines are long and curved from the base to the tips. Unpaired actine is curved and usually smaller than the paired actines. The apical actine is short and straight or slightly curved (Figs. 11C, C’). Remarks: Amphoriscus decennis sp. nov. does not present anchoring spicules or peduncle, which differs from A. ancora, A. chrysalis, A. cyathiscus, A. pedunculatus, A. synapta, and A. testiparus. Among the remaining species, A. semoni most closely resembles the new species due to its skeletal organisation: cortical tetractines, subatrial triactines, and atrial tetractines. Nevertheless, the atrial tetractines of A. decennis sp. nov. have smaller apical actines [holotype: 47.0– 61.2 ±9.3–73.4/ 9.4– 10.7 ±1.1–13.2 μm] than the type of A. semoni [according to the original description, the apical actine is 100–130/ 9 μm; Breitfuss (1896)]. Among the proposed paratypes, the largest apical actine of the new species was 97.2 μm (maximum value, obtained for the specimen UFRJPOR 9033). Although close to the minimum value of the size range described by Breitfuss (1896), the measurements found here for A. decennis sp. nov. for this actine are, in general, smaller than in A. semoni. Additionally, cortical spicules are considerably thicker in A. decennis sp. nov. (holotype —paired: 31.9–58.5 μm, unpaired 30.1–41.4 μm, apical 35.0–52.1 μm versus paired: 10–20 μm, unpaired 10–20 μm, apical 19–27 μm in A. semoni). Finally, only A. decennis sp. nov. has trichoxeas. The presence of small cortical tetractines in A. decennis sp. nov. may also be useful to differentiate it from A. semoni, although we considered them to belong to the same category as the giant cortical tetractines. The geographical distribution of both species is also different. Type specimens of A. decennis sp. nov. were sampled in the Mediterranean Sea (more specifically at Île Riou, Marseille, France), while A. semoni is known to occur across the central Indo-Pacific region (type locality— Ambon Island, Indonesia). A specimen of A. decennis sp. nov. from Plymouth was also recognised (as discussed along with the remarks of A. chrysalis), which remains consistent with a separate species distribution from that of A. semoni. A wide distributional range is not common in calcareous sponges and has been associated with erroneous taxonomic identification [such as the specimen BMNH 1886.6.7.32, from Australia, previously identified as A. cylindrus by Burton (1963)] or with the occurrence of introduced species. Since there is no reason to suspect that A. semoni could have invaded regions outside the Indo-Pacific, we consider the distribution additional evidence to support the new species. Distribution: Mediterranean Sea and Plymouth, United Kingdom (BMNH 1955.12.13.9; see remarks of A. chrysalis). Corresponding MEOW: Western Mediterranean and Celtic Seas (Spalding et al. 2007).

Amphoriscus gastrorhabdifera (Burton, 1932) Leucaltis gastrorhabdifera Burton, 1932 Citations and Synonymies: Leucaltis gastrorhabdifera Burton, 1932: 259, figs. 4-5; Amphoriscus gastrorhabdifera Burton, 1963: 136, 147, 548; Klautau et al. 2017: 105-106; Cóndor-Luján et al. 2019: 1825. Type material: BMNH 1928.2.15.833 (Holotype; St. 6, Tristan da Cunha, South Atlantic; 80–140 m deep). Type locality: Tristan da Cunha, South Atlantic. Morphology: Colour is beige after fixation. The holotype is a tubular fragment (Fig. 12A), and a fringe of trichoxeas was present, according to Burton (1932). The type of aquiferous system is unclear. Anatomy: The skeleton has no similarity with the other species of Amphoriscus. The cortical region is formed by triactines of varying sizes and subcortical giant tetractines (which are possibly the only typical character of Amphoriscidae). However, there is no typical inarticulation (Fig. 12B). The presence of several broken spicules makes the visualisation of the triactines difficult (Fig. 12C). The apical actines of the subcortical tetractines cross the entire choanosome, often perforating the atrium (Figs. 12D, E). The subatrial/atrial region is comprised exclusively of large tangential diactines. They are abundant and line the atrial cavity (Figs. 12E, F). A specific atrial skeleton comprised of triactines or tetractines is absent. According to Burton (1932), large diactines similar to those found in the inner part of the sponge were found projecting from the cortex, but they were not observed here. The aquiferous system could not be recognised and was not mentioned along the original description. Spicules: Cortical triactines: Regular, actines are slightly conical and sharp. The actines are straight (paired actines: 123.1– 150.7 ±21.6–204.4/ 8.4– 12.2 ± 1.7–15.3 μm; unpaired actines: 81.5– 133.6 ±23.2–180.0/ 8.9– 122.3 ±2.0–18.3 μm) (Fig. 13A). Subcortical tetractines: Giant, actines are slightly conical to conical and blunt. The paired actines are slightly curved (111.4– 146.6 ±21.8–192.1/ 10.8– 14.5 ± 2.0–19.2 μm). The unpaired actine is curved from the base to the tip (93.9– 116.7 ±21.5–156.6/ 14.1– 15.2 ±1.1–16.9 μm). The apical actine is long and conical (174.3– 228.3 ±29.1–285.4/ 16.9– 20.8 ±2.2–26.3 μm) (Fig. 13B). Diactines: Large, sinuous along their length. Most of them have one of the tips blunt while the other is sharp (330.0– 464.8 ±89.6–632.6/ 9.0– 18.9 ±4.0–25.7 μm) (Fig. 13C). Remarks: This species was originally described as Leucaltis gastrorhabdifera (subclass Calcinea) and was later transferred to Amphoriscus (subclass Calcaronea) (Burton 1932, 1963). Such a reallocation would, by the standards of modern-day research on Calcarea systematics, be considered out of the ordinary and require a detailed explanation, yet at that time Burton did not note the reasons supporting his decision in any detail. In his discussion of the genus Leucaltis, Burton (1963) argued that “ L. gastrorhabdifera is aberrant and is here doubtfully assigned to Amphoriscus ”, suggesting that the only certainty the author had was that it was not Leucaltis. He named the species “ Amphoriscus ? gastrorhabdifera ”, and although we suspect that he was influenced by the presence of the giant cortical tetractines, this cannot be confirmed.Also, there is no information or illustration in the literature about the aquiferous system of A. gastrorhabdifera, which would be essential to support its allocation in Amphoriscus. Whether or not it was one of the characters used by Burton to assign the species in Amphoriscus is a question that remains unanswered. We analysed the holotype (BMNH 1928.2.15.833) in this study. The microscopical slides contain sections of the skeleton that were probably not stained since it was not possible to unequivocally confirm whether A.gastrorhabdifera has the syconoid aquiferous system typical of the genus. We observed the main morphological characters reported in the original description, such as the presence of cortical triactines, subcortical giant tetractines with a long apical actine, and diactines. Burton (1932) mentioned diactines protruding through the cortex, but we did not observe this characteristic. The absence of a subatrial layer of triactines/tetractines was also confirmed and, consequently, the absence of inarticulate skeletal organisation. Therefore, the most important diagnostic characters typical of Amphoriscus either could not be confirmed (the syconoid aquiferous system) or are absent (the inarticulation formed by the apical actines of giant cortical tetractines and the unpaired actine of subatrial spicules). In order to assess the possibility of assigning A. gastrorhabdifera in another genus, a diagnosis of all genera under the order Leucosolenida was carried out. The presence of triactines and tetractines and of an atrial layer of diactines is a remarkable character found only in Sycodorus, although members of this genus are syconoid. Tangential triactines and tetractines also occur along the atrial cavity, suggesting that, even if A. gastrorhabdifera is syconoid, it could not be allocated in Sycodorus. An alternative decision would be to propose a new genus. The reasons why we did not choose this option to solve the problem of A. gastrorhabdifera were as follows: (i) the holotype is tiny, and no reports of additional specimens exist that could enrich our understanding of the morphology of the new genus; (ii) the lack of data on the aquiferous system could raise doubts on the family in which the genus should be inserted and also makes the description of a robust diagnosis difficult; (iii) the species is represented only by the holotype, found at Tristan da Cunha, Southern Atlantic Ocean, at a depth of 80 to 140 m. The lack of perspective in finding fresh samples to elucidate the questions mentioned earlier suggests that the problem with the species would persist (not as Amphoriscus but as a newly named genus); finally, (iv) we cannot undoubtedly rule out A. gastrorhabdifera actually belonging to Amphoriscus as the absence of an inarticulate organisation could be a secondary character caused by the loss of subatrial spicules along the species’ evolutionary history. The latter question will be resolved after the species is tested in phylogenetic analyses, though, as highlighted above, there is no fresh material available. Therefore, we decided to be conservative and avoid potentially increasing the number of open questions on the classification of this species. Amphoriscus gastrorhabdifera is thus maintained in the genus, but we indicate that it should be considered as incertae sedis until the discovery of additional samples.

Genus Leuconia Grant, 1833 “ Baeriidae in which the choanoskeleton consists of giant triactines and/or tetractines, lying without apparent order, and of very numerous microdiactines. A cavity equivalent to the atrium, localised only under the oscula, has a skeleton supported by tangential triactines. All the other exhalant canals have a skeleton composed of harpoon-shaped pugioles” (Borojevic et al. 2002).

Genus Amphoriscus Haeckel, 1870 “ Amphoriscidae with syconoid organisation of the aquiferous system. Scattered spicules in the choanosome are always absent” (Borojevic et al. 2002).

Amphoriscus gregorii (Lendenfeld, 1891) Ebnerella gregorii Lendenfeld, 1891 Citations and synonymies: Ebnerella gregorii Lendenfeld 1891: 290, pl. xi, fig. 66, xiv, figs. 117-123. Amphoriscus gregorii Dendy & Row 1913: 782; Topsent 1934: 11; Breitfuss 1935: 30; Burton 1963: 133; Klautau et al. 2017: 105; C��ndor-Luj��n et al. 2019: 1825. Type material: BMNH 1896.11.5.18 (paratype). Lesina, Adriatic Sea (The holotype had not been located). Type locality: Lesina, Adriatic Sea. Analysed material: BMNH 1896.11.5.18 (paratype; two slides containing sections of the skeleton). Morphology: The type specimen was not available. However, according to the original description, the sponge has a branched body formed by several tubes with apical osculum and unified at the base (Lendenfeld 1891). The aquiferous system is syconoid. Amphiblastulae are easily observed in some of the histological sections (Fig. 6A). Anatomy: The cortical skeleton is formed by giant tetractines and has trichoxeas perpendicularly disposed. The cortical tetractines have a long apical actine, which often protrude through the atrial cavity (Figs. 6B, C, E). The subatrial region is formed exclusively by triactines (Fig. 6E). The atrium presents tetractines with short apical actine and few triactines (Fig. 6F). Spicules (Tables 2 and 3): A figure containing only the spicule categories, in line with common practice, could not be prepared since only slides containing sections of the skeleton were available. Cortical tetractines: Slightly conical with blunt tips. The paired actines are long and slightly curved, being usually broken for unknown reasons. The unpaired actine is curved and smaller than the paired ones. The apical actine is long and thick, commonly perforating the atrial wall. Subatrial triactines: Slightly conical to cylindrical, sharp tips. The paired actines are straight or have a slight curvature when touching the atrial wall. The unpaired actine is straight and long, extending up to the cortical region. Atrial tetractines: Cylindrical and sharp. Paired actines are long and curved. The unpaired actine is straight and smaller or the same size as the paired ones. The apical actine is short and curved. Atrial triactines: Not abundant, similar in shape to the atrial tetractines. Remarks: Lendenfeld (1891) described a set of specimens, with the presence of embryos reported for at least one. Sections of this specimen were represented by the author in plate IV, figure 123b (Lendenfeld 1891), though we cannot unequivocally confirm that it corresponds to the paratype analysed here based solely on the illustration. Our suspicion was raised by the presence of embryos in BMNH 1896.11.5.18, which can be easily observed along the walls of the choanocyte chambers (Fig. 6). Ebnerella was erected to allocate species of Amphoriscus with diactines (Lendenfeld 1891). Ebnerella gregorii was the only species originally described to this genus, which was soon abandoned (Dendy & Row 1913). After analysing the paratype, we figured out that the diactines mentioned by Lendenfeld (1891) correspond to trichoxeas. Burton (1963) questioned the presence of atrial triactines in A. gregorii. These spicules were confirmed in this study (Fig. 6F). They are not abundant and usually have a broken unpaired actine, which makes it difficult to recognise them immediately. Compared to other species belonging to Amphoriscus, A. gregorii most closely resembles A. bucchichii. They differ mainly by the presence of atrial triactines in the former, while the latter only has atrial tetractines.Additionally, only one category of tetractines can be seen in the cortical region in A. gregorii, while reports on A. bucchichii by Ebner (1887) mention the presence of subcortical tetractines in addition to the cortical tetractines. Amphoriscus chrysalis, A. cyathiscus, A. cylindrus, A. kryptoraphis, A. oviparus, A. salfii, A. synapta, A. testiparus, and A. urna have tetractines in the subatrial region, which are absent in A. gregorii. Amphoriscus elongatus has triactines in the cortical region while A. gregorii has only cortical tetractines. Finally, the recently described species, A. ancora and A. pedunculatus, have anchoring spicules and peduncle, absent in A. gregorii. Distribution: Lesina, Adriatic Sea (Lendenfeld 1891). Corresponding MEOW: Adriatic Sea (Spalding et al. 2007)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on pages 49-51, DOI: 10.11646/zootaxa.5061.1.2, http://zenodo.org/record/5642287, {"references":["Lendenfeld, R. V. (1891) Die Spongien der Adria, I. Die Kalkschwamme. Zeitschrift fur wissenschaftliche Zoologie, 53 (2), 185 - 321.","Dendy, A. & Row, R. W. H. (1913) The Classification and Phylogeny of the Calcareous Sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London, 3, 704 - 813. https: // doi. org / 10.1111 / j. 1469 - 7998.1913. tb 06152. x","Topsent, E. (1934) Apercu de la faune des Eponges calcaires de la Mediterranee. Bulletin de l'Institut Oceanographique de Monaco, 659, 1 - 20.","Breitfuss, L. (1935) La spugne calcarea dell'Adriatico con riflesso a tutto il Mediterraneo. Memorie Reale Comitato Talassographico Italiano, Venezia, 2223, 1 - 45.","Burton, M. (1963) A revision of the Classification of the Calcareous Sponges: with a Catalogue of the specimens in the British Museum. Order of the trustees of the British Museum (Natural History), London, 693 pp.","Klautau, M., Cavalcanti, F. F. & Borojevic, R. (2017) The new sponge species Amphoriscus pedunculatus (Porifera, Calcarea). Zootaxa, 4341 (1), 105 - 112. https: // doi. org / 10.11646 / zootaxa. 4341.1.9","Condor-Lujan, B., Azevedo, F., Hajdu, E., Hooker, Y., Willenz, P. & Klautau, M. (2019) Tropical Eastern Pacific Amphoriscidae Dendy, 1892 (Porifera: Calcarea: Calcaronea: Leucosolenida) from the Peruvian coast. Marine Biodiversity, 49 (4), 1 - 18. https: // doi. org / 10.1007 / s 12526 - 019 - 00946 - y","Ebner, V. V. (1887) Amphoriscus bucchichii n. sp. Zoologische Jahrbucher, 2, 981 - 982.","Spalding, M. D., Fox, H. E., Allen, G. R., Davidson, N., Ferdana, Z. A., Finlayson, M., Halpern, B. S., Jorge, M. A., Lombana, A., Lourie, S. A., Martin, K. D., McManus, E., Molnar, J., Recchia, C. A. & Robertson, J. (2007) Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. BioScience, 57 (7), 573 - 583. https: // doi. org / 10.1641 / B 570707"]}

Leuconia dohrni (Sar��, 1960) comb. nov. Amphoriscus dohrni Sar��, 1960 Citations and synonymies: Amphoriscus dohrni Sar��, 1960: 26; Klautau, Cavalcanti & Borojevic 2017: 109; C��ndor-Luj��n et al. 2019: 1825. Type material: SZN POR207, holotype ���not analysed by the present work. Ischia Island, Gulf of Naples, Tyrrhenian Sea; depth 40 m. Type locality: Ischia Island, Gulf of Naples, Tyrrhenian Sea. Morphology: As explained above, the type material (SZN POR207) could not be analysed, and additional samples were not available or could not be located. According to the original description (Sar�� 1960), the specimen has an ellipsoidal shape and apical osculum without fringe of trichoxeas. No data about the aquiferous system is provided, and the lack of illustrations of the skeletal or histological organisations prevented any inferences from being made. Anatomy: The cortical skeleton is formed by triactines and giant tetractines, which point the apical actines to the middle of the sponge body (Sar�� 1960). Microdiactines are present along the whole skeleton, but Sar�� (1960) emphasised they are more abundant at the atrial skeleton, which is also formed by small and particular tetractines (see remarks below). Spicules (according to the original description): Cortical tetractines: Giant, actines have similar length (700���1200/70���120 ��m). Cortical triactines: Smaller than the cortical tetractines (170���350/15���35 ��m). Atrial tetractines (see remarks below): They have a particular (harpoon) shape. Apical (70���80/ 7���9 ��m) and unpaired (55���70 ��m) actines are considerably longer than the paired ones (20���35 ��m). Microdiactines: Lanceolate (60���70/4 ��m). Remarks: Due to the small size of the type specimen (Amphoriscus were not reported along the description of this species, such as the syconoid aquiferous system and the inarticulate skeleton formed by the apical actines of the cortical tetractines and the unpaired actine of the subatrial spicules; and (2) the shape of the small atrial tetractines, of which ���la forma �� molto caratteristica��� (Sar�� 1960), suggesting that they are pugioles. These spicules correspond to small harpoon���(or dagger-) shaped spicules present in the order Baerida (Borojevic et al. 2002; Alvizu et al. 2018, 2019). Besides the presence of pugioles, the description provided by Sar�� (1960) contains other characters found in the definition of Baerida: the presence of cortical large and giant spicules (triactines and tetractines, respectively, in the case of A. dohrni), and the presence of microdiactines abundant in the atrial skeleton (although it is not clear if they are more abundant than the atrial pugioles). Therefore, based on these features, there is no support for keeping the species inside the genus Amphoriscus, and we propose to transfer it to Baerida. According to Van Soest et al. (2021), Baerida is formed by four families: Baeriidae Borojevic, Boury-Esnault & Vacelet, 2000, Petrobionidae Borojevic, 1979, Trichogypsiidae Borojevic, Boury-Esnault & Vacelet, 2000, and Lepidoleuconidae Vacelet, 1967. Among the families mentioned above, only Baeriidae and Petrobionidae include pugioles in their composition (Borojevic et al. 2002; Alvizu et al. 2019). Petrobionidae has a basal calcareous skeleton of calcite. This family is monotypic, and Petrobiona massiliana Vacelet & L��vi, 1958 does not resemble ��� A ���. dohrni, in which the basal skeleton is absent (Borojevic et al. 2002). Baeriidae includes four genera: Leuconia Grant, 1833, Eilhardia Pol��jaeff, 1883, Lamontia Kirk, 1895, and Leucopsila Dendy & Row, 1913. However, only Leuconia and possibly Lamontia are pugiole-bearing genera. The latter is characterised by having diactines perforating the cortex and microdiactines throughout the choanoskeleton, characters that were not reported for ��� Amphoriscus ��� dohrni (Sar�� 1960; Borojevic et al. 2002). Since this species does have atrial pugioles, we decided to allocate it within Leuconia�� despite the absence of sagittal triactines in the atrium. We therefore highlight the need for a taxonomic revision of Leuconia and the entire order Baerida, echoing recent suggestions from Alvizu et al. (2018, 2019)., Published as part of Chagas, Cl��slei & Cavalcanti, Fernanda F., 2021, Partial taxonomic revision of Amphoriscus Haeckel, 1870 (Porifera: Calcarea) with description of A. decennis sp. nov., pp. 39-68 in Zootaxa 5061 (1) on pages 63-64, DOI: 10.11646/zootaxa.5061.1.2, {"references":["Sara, M. (1960) Poriferi del litorale dell'isola d'lschia e loro ripartizione ambienti. Pubblicazioni della Stazione zoologica di Napoli, 31 (3), 421 - 472.","Klautau, M., Cavalcanti, F. F. & Borojevic, R. (2017) The new sponge species Amphoriscus pedunculatus (Porifera, Calcarea). Zootaxa, 4341 (1), 105 - 112. https: // doi. org / 10.11646 / zootaxa. 4341.1.9","Condor-Lujan, B., Azevedo, F., Hajdu, E., Hooker, Y., Willenz, P. & Klautau, M. (2019) Tropical Eastern Pacific Amphoriscidae Dendy, 1892 (Porifera: Calcarea: Calcaronea: Leucosolenida) from the Peruvian coast. Marine Biodiversity, 49 (4), 1 - 18. https: // doi. org / 10.1007 / s 12526 - 019 - 00946 - y","Borojevic, R., Boury-Esnault, N., Manuel, M. & Vacelet, J. (2002) Order Leucosolenida Hartman, 1958. In: Hooper, J. N. A. & Van Soest, R. W. M. (Eds.), Systema Porifera: a guide to the classification of sponges. Vol. 2. Kluwer Academic / Plenum Publishers, New York, Boston, Dordrecht, London and Moscow, pp. 1157 - 1184. https: // doi. org / 10.1007 / 978 - 1 - 4615 - 0747 - 5 _ 120","Alvizu, A., Eilertsen, M. H., Xavier, J. R. & Rapp, H. T. (2018) Increased taxon sampling provides new insights into the phylogeny and evolution of the subclass Calcaronea (Porifera, Calcarea). Organisms Diversity & Evolution, 18 (3), 279 - 290. https: // doi. org / 10.1007 / s 13127 - 018 - 0368 - 4","Alvizu, A., Xavier, J. R. & Rapp, H. T. (2019) Description of new chiactine-bearing sponges provides insights into the higher classification of Calcaronea (Porifera: Calcarea). Zootaxa, 4615 (2), 201 - 251. https: // doi. org / 10.11646 / zootaxa. 4615.2.1","Borojevic, R., Boury-Esnault, N. & Vacelet, J. (2000) A revision of the supraspecific classification of the subclass Calcaronea (Porifera, class Calcarea). Zoosystema, 22 (2), 203 - 263.","Polejaeff, N. (1883) Report on the Calcarea dredged by H. M. S. ' Challenger', during the years 1873 - 1876. Report on the Scientific Results of the Voyage of H. M. S. ' Challenger', 1873 - 1876. Zoology, 8 (2), 1 - 76.","Dendy, A. & Row, R. W. H. (1913) The Classification and Phylogeny of the Calcareous Sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London, 3, 704 - 813. https: // doi. org / 10.1111 / j. 1469 - 7998.1913. tb 06152. x"]}

Amphoriscus is the type genus of the family Amphoriscidae. Since its nomination in the 19th century, its diagnosis has undergone significant changes. Most of the species currently assigned to Amphoriscus have only been reported once, when they were first described. Furthermore, unlike other Amphoriscidae genera, new Amphoriscus species are not commonly described. Therefore, the understanding of the diversity, distribution, and morphology of the genus remains fragmented, a lacuna that is being filled slowly. In this study, several species were revisited by the redescription of type and/or additional specimens. This results in considerable advances, including changes in the geographical distribution of A. cylindrus and A. chrysalis, a proposal of reallocation of A. dohrni in Leuconia, recognition of A. gastrorhabdifera as incertae sedis, a detailed description of the type species based on a set of specimens and, finally, the description of a new species, A. decennis sp. nov.

Family Amphoriscidae Dendy 1893 Genus Leucilla Haeckel, 1872 Diagnosis. Amphoriscidae with sylleibid or leuconoid organisation. The choanoskeleton is formed primarily by the apical actines of giant cortical tractines and the unpaired actines of subatrial triactines or tetractines. It may contain dispersed spicules, but a typical articulate choanoskeleton is always absent (from Borojevic et al. 2002)., Published as part of Sim-Smith, Carina, Hickman, Cleveland & Kelly, Michelle, 2021, New shallow-water sponges (Porifera) from the Galápagos Islands, pp. 1-71 in Zootaxa 5012 (1) on page 60, DOI: 10.11646/zootaxa.5012.1.1, http://zenodo.org/record/5158062, {"references":["Dendy, A. O. (1893) Synopsis of the Australian Calcarea Heterocoela; with a proposed classification of the group and descriptions of some new genera and species. Proceedings of the Royal Society of Victoria, New Series, 5, 69 - 116.","Haeckel, E. (1872) Die Kalkschwamme. Eine Monographie in zwei Banden Text und einem Atlas mit 60 Tafeln Abbildungen. G. Reimer, Berlin, 418 pp. [in German]","Borojevic, R., Boury-Esnault, N., Manuel, M. & Vacelet, J. (2002) Order Leucosolenida Hartman, 1958. In: Hooper, J. N. A. & Van Soest, R. W. M. (Eds.), Systema Porifera: A Guide to the Classification of Sponges. Vol. 1. Klewer Academic / Plenum Publishers, New York, New York, pp. 1157 - 1184. https: // doi. org / 10.1007 / 978 - 1 - 4615 - 0747 - 5 _ 120"]}

Family Clathrinidae Minchin, 1900 Genus Clathrina Gray, 1867 Diagnosis. Calcinea in which the cormus comprises anastomosed tubes. A stalk may be present. The skeleton contains regular (equiangular and equiradiate) and/or parasagittal triactines, to which diactines and tripods may be added. Asconoid aquiferous system (from Azevedo et al. 2015)., Published as part of Sim-Smith, Carina, Hickman, Cleveland & Kelly, Michelle, 2021, New shallow-water sponges (Porifera) from the Galápagos Islands, pp. 1-71 in Zootaxa 5012 (1) on page 56, DOI: 10.11646/zootaxa.5012.1.1, http://zenodo.org/record/5158062, {"references":["Minchin, E. A. (1900) Chapter III. Sponges. In: Lankester, E. R. (Ed.), A Treatise on Zoology. Part II. The Porifera and Coelenterata. Vol. 2. Adam & Charles Black, London, pp. 1 - 178.","Gray, J. E. (1867) Notes on the arrangement of sponges, with the descriptions of some new genera. Proceedings of the Zoological Society of London, 1867 (2), 492 - 558.","Azevedo, F., Condor-Lujan, B., Willenz, P., Hajdu, E., Hooker, Y. & Klautau, M. (2015) Integrative taxonomy of calcareous sponges (subclass Calcinea) from the Peruvian coast: morphology, molecules, and biogeography. Zoological Journal of the Linnean Society, 173, 787 - 817. https: // doi. org / 10.1111 / zoj. 12213"]}

GENUS LEUCETTA HAECKEL, 1872 Type species: Leucetta primigenia Haeckel, 1872. Diagnosis: ��� Leucettidae with a homogeneous organisation of the wall and a typical leuconoid aquiferous system. There is neither a clear distinction between the cortex and the choanoskeleton, nor the presence of a distinct layer of subcortical inhalant cavities. The atrium is frequently reduced to a system of exhalant canals that open directly into the osculum��� (Borojević et al., 2002)., Published as part of Klautau, Michelle, Lopes, Matheus Vieira, Tavares, Gabriela & P��rez, Thierry, 2022, Integrative taxonomy of calcareous sponges (Porifera: Calcarea) from R��union Island, Indian Ocean, pp. 671-725 in Zoological Journal of the Linnean Society 194 on page 699, DOI: 10.1093/zoolinnean/zlab014, http://zenodo.org/record/6354284, {"references":["Haeckel E. 1872. Die Kalkschwamme. Eine monographie, Vols 1 - 3. Berlin: Reimer.","Borojevic R, Boury-Esnault N, Manuel M, Vacelet J. 2002. Order Clathrinida Hartman, 1958. In: Hooper JNA, Van Soest RWM, eds. Systema Porifera: a guide to the classification of sponges. New York: Kluwer Academic / Plenum Publishers, 1141 - 1152."]}

GENUS LELAPIELLA VACELET, 1977 Type species: Lelapiella incrustans Vacelet, 1977 (by monotypy). Diagnosis: Lelapiellidae encrusting and without fused spicules. Radial growing. Cortical skeleton composed chiefly of a layer of large tripods, diactines may be present. Choanosome with triactines and tetractines. Tracts of diactines may be present (modified from Borojević et al., 2002)., Published as part of Klautau, Michelle, Lopes, Matheus Vieira, Tavares, Gabriela & P��rez, Thierry, 2022, Integrative taxonomy of calcareous sponges (Porifera: Calcarea) from R��union Island, Indian Ocean, pp. 671-725 in Zoological Journal of the Linnean Society 194 on page 702, DOI: 10.1093/zoolinnean/zlab014, http://zenodo.org/record/6354284, {"references":["Vacelet J. 1977. Eponges pharetronides actuelles et sclerosponges de Polynesie francaise, de Madagascar et de la Reunion. Bulletin du Museum National d'Histoire Naturelle de Paris 444: 345 - 368.","Borojevic R, Boury-Esnault N, Manuel M, Vacelet J. 2002. Order Clathrinida Hartman, 1958. In: Hooper JNA, Van Soest RWM, eds. Systema Porifera: a guide to the classification of sponges. New York: Kluwer Academic / Plenum Publishers, 1141 - 1152."]}