Tethya cranium var. australiensis Tetilla? australiensis Spiretta raphidiophora Tetilla hirsuta Tetilla ternatensis Ternate Tetilla ternatensis Tetilla australiensis Tetilla ternatensis Tetilla ternatensis Tetilla lindgreni Tetilla australiensis Tethya hebes Cinachyra isis Tetilla cinachyroides Cinachyra nuda Cinachyra vaccinata Cinachyra providentiae Tetilla (Cinachyrella) hirsuta Cinachyra australiensis C. australiensis porosa C. australiensis schulzei Cinachyrella anatriaenilla Unravelling the moons: review of the genera Paratetilla and Cinachyrella in the Indo-Pacific (Demospongiae, Tetractinellida, Tetillidae) Santodomingo, Nadiezhda Becking, Leontine E. ZooKeys 2018 791 1 46 VCPP Carter, 1886 Carter 1886 Demospongiae Tetillidae Cinachyrella CoL Animalia Cinachyrella australiensis Tetractinellida 12 13 Porifera species australiensis   Tethya cranium var. australiensisCarter, 1886: 127 (holotype seen).  Tetilla? australiensis; Sollas, 1888: 43.  Spiretta raphidiophoraLendenfeld, 1888: 43 (type seen).  Tetilla hirsutaDendy, 1889: 75 (type seen).  Tetilla ternatensisLindgren, 1898: 329 pl. 17, fig. 14; pl. 19, Fig. 25 a-e, a', b'. TernateNot Tetilla ternatensis; Kieschnick*, 1896: 527.  Tetilla australiensis; Thiele, 1899: 6, pl.1 fig.1; pl. 5, fig.1 a-e. Celebes Sea.  Tetilla ternatensis; Kirkpatrick, 1900: 132 (material seen) Not Tetilla ternatensisKieschnick*, 1896: 527.  Tetilla lindgreniLendenfeld, 1903: 18.  Tetilla australiensis; Lendenfeld, 1903: 20.  Tethya hebesLendenfeld, 1907: 98, pl. XVI, figs 19-38. 19'South NW Australia, 91 m depth (syntype seen).  Cinachyra isisLendenfeld, 1907: 143, pl. XV, figs 54-58, XVI, figs 1-4. Mermaid Strasse (NW Australia) (syntype seen); Dendy, 1922: 16, pl. 10, figs 3a-b.  Tetilla cinachyroidesHentschel, 1911: 281, textfig. 1. NW Australia, Barrow Island.  Cinachyra nudaHentschel, 1912:333, pl. XIII, fig.2; pl. XVIII fig. 13. Aru Island (type seen).  Cinachyra vaccinataDendy, 1922: 14, pl. 1, fig. 4; pl. 11, figs 1a-l. Diego Garcia, Chagos Island (type seen).  Cinachyra providentiaeDendy, 1922: 18, pl.1, figs 5-5a; pl. 10, figs2 a-f. Providence Island (type seen).  Tetilla (Cinachyrella) hirsuta; Wilson, 1925: 365, pl. 39, fig.4.  Cinachyra australiensis; Burton, 1934: 523. In part, not C. australiensisin porosa-group, nor C. australiensisin schulzei-group; de Laubenfels, 1954: 241, text-fig. 166.  Cinachyrella anatriaenillaFernandez, Kelly, Bell, 2017: 83, figs 2-4.  Material examined. Holotype NHMUK 1886.12.15.367, Port Phillip Heads, Southeast Australia (as Tethya cranium var. australiensis). Holotype NHMUK 1886.8.27.634, Port Jackson, Sidney, Australia (as Spiretta raphidiophoraLendenfeld, 1888). NHMUK unreg. type, Gulf of Manaar, Sri Lanka (as Tetilla hirsutaDendy, 1889). NHMUK 1898.12.20.20 Christmas islands (as Tetilla ternatensisKirkpatrick, 1900). Holotype NHMUK 1908.9.24.19-21, 19°17'S116°E, Gazelle Exp., Western Australia, (as Tethya hebesLendenfeld, 1907). Syntype NHMUK 1908.9.24.74, Mermaid Strait, NW Australia (as Cinachyra isisLendenfeld, 1907). RMNH unreg. fragment taken from the type (pers. comm. NJ de Voogd) and available in Naturalis collections, Aru Island, Indonesia, as Cinachyra nudaHentschel, 1912. Holotype NHMUK 1921.11.7.6, Diego Garcia, Chagos Islands (as Cinachyra vaccinataDendy, 1922). Holotype NHMUK 1921.11.7.8, Providence Island, Seychelles (as Cinachyra providentiaeDendy, 1922). INDONESIA. East Kalimantan, Berau reef, RMNH.POR.11101, RMNH.POR.11102, RMNH.POR.11103, RMNH.POR.11104, RMNH.POR.11105, RMNH.POR.11106, RMNH.POR.11107, RMNH.POR.11108, RMNH.POR.11109, RMNH.POR.11110, RMNH.POR.11111, RMNH.POR11112, RMNH.POR.11113, RMNH.POR.11114, RMNH.POR.11115, RMNH.POR.11116, RMNH.POR.11117, RMNH.POR.11210, RMNH.POR.11124, RMNH.POR.11125, RMNH.POR.11126, RMNH.POR.11127, RMNH.POR.11128, RMNH.POR.11129, RMNH.POR.11130, RMNH.POR.11118, RMNH.POR.11119, RMNH.POR.11120, RMNH.POR.11121, RMNH.POR.11122, RMNH.POR.11123; RMNH.POR.11132; RMNH.POR.11133, RMNH.POR.11134, RMNH.POR.11135, RMNH.POR.11136; Pea Bay, RMNH.POR.11162; Haji Buang Lake, RMNH.POR.11137, RMNH.POR.3511, RMNH. POR.3512, RMNH.POR.3513, RMNH.POR.3516, RMNH.POR.3517; Kakaban Lake, RMNH.POR.11161, RMNH.POR.11138, RMNH.POR.11139, RMNH.POR.11140, RMNH.POR.11141, RMNH.POR.11142, RMNH.POR.11143, RMNH.POR.11144, RMNH.POR.11145, RMNH.POR.11146, RMNH.POR.11147, RMNH.POR.11148, RMNH.POR.11149, RMNH.POR.11150, RMNH.POR.11151, RMNH.POR.11152, RMNH.POR.11153, RMNH.POR.11154, RMNH.POR.11155, RMNH.POR.11156, RMNH.POR.11157, RMNH.POR.11158, RMNH.POR.11159, RMNH.POR.11160. Java, Thousand Islands, RMNH.POR.1969. Ternate, Ternate reef, RMNH.POR.11308. Sulawesi, Bunaken, RMNH.POR.3108, RMNH.POR.3112, RMNH.POR.3119, RMNH.POR.3122. West Papua, Sawaundarek Lake, RMNH.POR.11163, RMNH.POR.11164, RMNH.POR.11165, RMNH.POR.11166, RMNH.POR.11167; Gam Island, Wallace Lake, RMNH.POR.11168, RMNH.POR.11169 Outside Wallace Lake, RMNH.POR.11170, RMNH.POR.11171, RMNH.POR.11172, RMNH.POR.11173; Gam Island, Blue Water Mangrove, RMNH.POR.11174, RMNH.POR.11175, RMNH.POR.11176, RMNH.POR.11177, RMNH.POR.11178, RMNH.POR.11179, RMNH.POR.11180, RMNH.POR.11181, RMNH.POR.11182, RMNH.POR.11183, RMNH.POR.11184, RMNH.POR.11185, RMNH.POR.11186, RMNH.POR.11187, RMNH.POR.11188, RMNH.POR.11189, RMNH.POR.11190, RMNH.POR.11191, RMNH.POR.11192; Ctenophore Lake, RMNH.POR.11193, RMNH.POR.11194, RMNH.POR.11195, RMNH.POR.11196, RMNH.POR.11197; Outside Ctenophore Lake, RMNH.POR.11198, RMNH.POR.11199, RMNH.POR.11200, RMNH.POR.11201; Big Caulerpa Lake, RMNH.POR.11202, RMNH.POR.11203; Outside Big Caulerpa lake, RMNH.POR.11204; Gam Island, RMNH.POR.11205, RMNH.POR.11206. Other material: Singapore, RMNH.POR.3520, RMNH.POR.2440, RMNH.POR. 2505. Other types and material examined (not included as synonyms of C. australiensis): NHMUK 1892.8.8.8. Macclesfield Bank, South China Sea Cinachyra schulzei(unpublished material). Holotype NHMUK 1908.9.24.75 Red Sea, Cinachyra trochiformisKeller, 1891. Holotype NHMUK 1907.2.1.14, Gulf of Manaar, Sri Lanka, Tetilla poculiferaDendy, 1905. Holotype NHMUK 1912.2.1.35, Tella Tella Kebira, Red Sea, Chrotella ibisRow, 1911. RMNH unreg. fragment taken from the type (pers. comm. NJ de Voogd) available in Naturalis collections, Kei Island, Indonesia, Cinachyra mertoniHentschel, 1912.  Description. External morphology. Globular sponges, size from 4 to 10 cm in diameter (Figure 6A, B). Surface hispid due to the projecting spicules; covered by numerous porocalices. Porocalices are abundant bowl-shape with open oval apertures, up to 10 x5 mm and 5 mm deep, or bottle-shape, up to 18 x6.5 mm, with minuscule apertures (2-3 mm diameter), size of porocalices can vary between habitats; a cloaca, defined as a central exhalant cavity (Boury-Esnault and Ruetzler, 1997), is distinguishable at the top of some specimens (Figure 6A); in preserved material some porocalices are open. Color generally bright yellow when alive, which turns paler or even white in ethanol. In the field, the sponge can appear brownish due to sediment or greenish due to association with algae.  Figure 6. Cinachyrella australiensis. A, C, E-H, L RMNH.POR.11139, Kakaban lake, Indonesia (left side) B, D, I-K, M holotype NHMUK 1886.12.15.367, Port Phillip Heads, Australia (right side) A In situ photograph showing porocalices B dry specimen, lateral view C skeleton showing acanthose microxeas (am) and radial bundles with oxeas D spicule montage showing acanthose microxeas (am), and oxeas (ox) E protriaene F anatriaene G Acanthose microxea, full lenght H acanthose microxea, detail I acanthose microxea, full length J acanthose microxea, detail K oxea, end detail L, M sigmaspires. Scale bars: 1 cm(B); 500 μm(C, D); 20 μm( E-G, I); 5 μm(H, J, L, M); 50 μm(K). Skeleton. No cortex. Skeleton composed by bundles of oxeas and triaenes radiating from a central core. Megascleres. Holotype and Indonesian specimens'measurements are shown in Table 6. Holotype, oxeas 3375 -4135.5-5500 mm x15 -24.7-37.5 mm (Figure 6D, K); no triaenes were observed in the type specimen; in Carter'sdescription, protriaenes are described (135 mm long) and the absence of anatriaenes was explained as their heads broke off when collected; Indonesian specimens have a wide size range of oxea 1000-5500 mm (Figure 6C), abundant anatriaenes (Figure 6F), with rhabd 2250 -3224.4-4250 mm x2.5 -5.7-10 mm, cladi thin, mainly with obtuse angles 30 -70.6-100 mm x20 -51.7-80 mm x2.5 -4.9-10 mm; protriaenes (Figure 6E), with thin and long cladi (20 -57.1-80 mm x25 -86.9-170 mm x2.5 -7.5-12.5 mm), rhabd up to 5800 x20 mm, tapering to dimensions of <1 mm; few prodiaenes also observed, having smaller cladi (20-30 mm x20-30 mm); no calthrop-like triaenes.  Table 6. Spicule measurements of eight specimens Cinachyrella australiensis, five specimens of C. anomala, four specimens of C. pateriferafrom different regions (n = 10 per spicule type and dimension with minimum-mean-maximum). Asterisk (*) indicate that rhabd of spicules were broken and no measurement was possible. Double asterisk (**) indicate that a particular spicule type was not observed.     Cinachyrella australiensis  Cinachyrella porosa  Cinachyrella paterifera  BMNH86.12.15.367 (Holotype) RMNH.POR.11120 RMNH.POR.11146 RMNH.POR.11123 RMNH.POR.11139 RMNH.POR.11118 RMNH.POR.11308 RMNH.POR.11192 BMNH86.8.27.632-3 (Holotype) RMNH.POR.11226 RMNH.POR.11244 RMNH.POR.11262 RMNH.POR.11309 USNM21314 (Holotype) RMNH.POR.11207 RMNH.POR.11213 RMNH.POR.11208  Locality  Habitat  Oxeas 4135.5 3332 2912 3822.8 3066 4315 2676 2658.3 2553.2 2138.1 2702.5 2304.2 2710 3011.5 3580.6 3060 2748.2  24.7 40 33.9 41.7 26.4 59.1 30.3 31.3 29.4 28.2 29.5 29.4 32.6 34.5 46.1 35.6 34.2  Anatriaene 3271.9 3317.9 3083.3 4250  6 5.3 4.9 5 5 8.4 5.3 5.8 12 7.8 5 6.1 7.5 6.6 8.8 5  63.5 74.2 58 55 81 79.4 83 67.6 71 65 62.9 62.5 24.2 37.5 90 79.3  49.5 60.4 42.5 45 49.5 49.1 66 42 56 52 51.4 50.4 6.5 25.7 65 58.2  4.5 3.5 5.6 5 5.1 5.6- 5 5.6 10.5 6.6 4.6 5.4 5.8 6.4 7.5 4.3  Protriaene 4550 4262.5 3522.5 2375 4210 4689.3  13.3 14.7 12.5 5.9 8.3 5 7.3 8.8 6.5 3.6 5 10 9.5 7.8 15.2  46.9 58.8 60.8 55 50.4 70.5 44.4 51.25 59 44.3 53.8 32.5 53.9 70 37.3  66.3 93.8 95.8 79.5 77.9 108- 73 77.5 100 67.1 78.3 31.3 74.1 82.5 51.6  7.9 11.6 10.8 4.8 5.8 4 5.1 7.5 5.5 2.5 4 7.5 6.8 6.6 11.6  Strongyle 4000 3041.7 2650 2650 3350 2800 2975 1862.5  50 43.3 39.4 35 45 45 43.1 45.3 37.5  Acanthose microxea 166.9 197.3 165 230 183.5 189.5 191.4 154  Sigmaspires 14.4 15 12.3 14.8 12 16.5 15.7 14.8 8.6 8.5 8.9 8 8 13.2 15.3 14.5 16.2  Protriaene (hair-like) Rhabd length 698.9  Width 2.5 2.5 2.5 2.2  Cladi total 12.5 10.9 13.6 11.7  Cladi length 15.5 21.6 14.5 14.4  Cladi width 2.5 2.1 2.5 1.7 Microscleres. Numerous acanthose microxeas, holotype, 117 -166.9-260 mm (Figure 6I, J), slightly larger in the Indonesian material 137.5 -184.7-270 mm (Figure 6G, H); sigmaspires vary within the same range in both, holotype and Indonesian specimens, 10 -14.4-20 mm, C-S shape (Figure 6L, M).  Ecology.  Cinachyrella australiensisoccurs in reefs, mangroves, and marine lakes, ranging in depths from 0 to at least 30 m, possibly deeper. Specimens can be covered by sand and mud; or in symbiosis with algae, resulting in green external color. This species produces 1-2 mm sized buds (Figure 8) and buds are extensively observed in specimens collected from marine lake habitats.  Distribution.  Cinachyrella australiensishas a wide distribution in Indonesia, including Berau, Bunaken, Raja Ampat, Ternate, and Java. Previous Indonesian records are from Spermonde Archipelago in Sulawesi (de Voogd and Cleary 2005, Becking et al. 2006, de Voogd et al. 2006), North Sulawesi ( Calcinai et al. 2017), Berau ( de Voogd et al. 2009, Becking et al. 2013), Thousand Islands in Java ( de Voogd and Cleary 2008), and Raja Ampat ( Becking 2008). In addition, this species has also been found in Gulf of Oman ( van Soest and Beglinger 2008), Seychelles Islands ( Thomas 1973) Southwest Madagascar ( Vacelet et al. 1976), Zanzibar ( Pulitzer-Finali 1993), Thailand ( Kritsanapuntu et al. 2001a- b, Putchakarn 2007), Singapore ( Lim et al. 2008), Vietnam ( Azzini et al. 2007), Philippines ( Longakit et al. 2005), Northern Territory of Australia ( McDonald et al. 2002), and the Great Barrier Reef in Australia ( Burton 1934).  Figure 7. Distribution of Cinachyrella australiensis. Red dot: type locality, Tethya cranium var. australiensisCarter, 1886, Port Phillip Heads, Southeast Australia. Green dots: Indonesian localities where the species was collected recently. Yellow triangles: Non-Indonesian localities, Seychelles Islands, Southwest Madagascar, Zanzibar, Thailand, Singapore, Vietnam, Philippines, Northern Territory of Australia, and the Great Barrier Reef in Australia. Circled numbers: type localities of synonymized species 1 Spiretta raphidiophoraLendenfeld, 1888, Port Jackson, Sidney, Australia 2 Tetilla hirsutaDendy, 1889, Gulf of Manaar, Sri Lanka 3 Tetilla lindgreniLendenfeld, 1903, Christmas Island 4 Tetilla poculiferaDendy, 1905, Gulf of Manaar, Sri Lanka 5 Tethya hebes, 1907, at 19° South on the NW coast of Australia 6 Cinachyra isisLendenfeld, 1907, Mermaid Strait, NW Australia 7 Tetilla cinachyroidesHentschel, 1911, Barrow Island, NW Australia 8 Cinachyra nudaHentschel, 1912, Aru Island, Indonesia 9 Cinachyra vaccinataDendy, 1922, Diego Garcia, Chagos Islands 10 Cinachyra providentiaeDendy, 1922, Providence Island, Seychelles 11 Cinachyrella anatriaenillaFernandez, Kelly, Bell, 2017, American Samoa.  Figure 8. Budding and sediment capture of Cinachyrellaspecies A Three individuals of C. porosain Haji Buang lake, East Kalimantan, Indonesia, showing distribution of buds beyond the individuals and sediment capture B Close up of C. porosawith detail of buds. Each individual is approximately 4 cm in diameter.  Remarks. In the type description of C. australiensis Carter (1886), the author did not observe anatriaenes as it can be interpreted from his statement: "I saw no anchors (smaller tetractinellids with recurved arms); but as their heads when exposed are generally broken off (for they catch in everything that they touch), it does not follow that they do not form part of the spiculation, particularly as they are present in most of the other species that I have been described (sic)". We examined the holotype kept at the Natural History Museum (NHMUK 1886.12.15.367) and found neither anatriaenes nor protriaenes. In addition, most of the oxeas were broken in the type specimen. Within all the examined material there is a high variability in the presence or absence oftriaenes without a distinct geographic pattern. This variation may be related to where the sponge was cut, as it seems that triaenes are particularly abundant around the porocalices compared to other parts of the sponge. These fragile spicules are also easily broken off. We still assign our specimens to the species C. australiensisdue to the characteristic presence of acanthose microscleres. It is furthermore one of the most common names used in the literature since its description and without further evidence we do not want to cause more confusion. Further examination of Cinachyrellaspecimens from Australia, in particular from the type locality of C. australiensis, will shed more light in this situation. It is quite possible that after a review of specimens from Southern Australia, it will be evident that the Indonesian specimens that we assign to C. australiensisshould in fact be assigned to another species. In that case one of the junior synonyms should be used, e.g. C. raphidiophoraor C. hirsuta. Although our focus was on Indonesian species, it was unavoidable to attempt, for the first time after Burton'sreview (1934), check the status of his large list of junior synonyms, because some of them were described or later found in Indonesian localities. We gathered as many type specimens as possible, most of them repositories of the NHMUK(London) and NMNH (Washington DC). The main criteria we used to suggest a species as junior synonym of C. australiensiswere the presence of acanthose microxea and that the mega- and micro-scleres have the same size range of the species. Therefore, here we include as junior synonyms the following species from Burton'slist: Spiretta raphidiophoraLendenfeld, 1888; Tetilla hirsutaDendy, 1889; Cinachyra isisLenfenfeld, 1907; Tetilla cinachyroidesHentschel, 1911; Cinachyra nudaHentschel, 1912; Cinachyra vaccinataDendy, 1922; Cinachyra providentiaeDendy, 1922. They all fulfill the C. australiensisdescription.