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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."]}

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 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."]}

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."]}

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."]}