Research Article |
Nadiezhda Santodomingo‡§, Leontine E. Becking§|¶
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Corresponding author: Nadiezhda Santodomingo ( n.santodomingo@nhm.ac.uk ) Corresponding author: Leontine E. Becking ( lebecking@gmail.com ) Academic editor: Martin Pfannkuchen
© 2018 Nadiezhda Santodomingo, Leontine E. Becking.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Santodomingo N, Becking LE (2018) Unravelling the moons: review of the genera Paratetilla and Cinachyrella in the Indo-Pacific (Demospongiae, Tetractinellida, Tetillidae). ZooKeys 791: 1-46. https://doi.org/10.3897/zookeys.791.27546
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Abstract
Paratetilla bacca (Selenka, 1867) and Cinachyrella australiensis (Carter, 1886) occur in a broad range of marine environments and are allegedly widely distributed species in the Indo-Pacific. We coin the term ‘moon sponges’ for these species as they are spherical in shape with numerous porocalices resembling the lunar surface. Both species have a complex taxonomic history with high synonymization, in particular by
Keywords
anchialine systems, coral reef, mangrove, marine lake, Porifera
Introduction
Moon sponges include two good examples of allegedly widely distributed species in the Indo-Pacific: Paratetilla bacca (Selenka, 1867) and Cinachyrella australiensis (Carter, 1886). They are conspicuous dwellers of a broad range of marine environments, including coral reefs, rocky shores, and coastal mangroves, as well as landlocked marine systems called marine lakes (e.g.
The genera Paratetilla and Cinachyrella, belong to the family Tetillidae, suborder Spirophorina, order Tetractinellida, class Demospongiae. As spirophorids, they are characterized by the presence of rugose sigmaspires (
Valid genera of Tetillidae Sollas, 1888 and principal characteristics used to distinguish them. (+) present, (-) absent. (AN) Antarctic, (AT) Atlantic, (CA) Caribbean, (IP) Indo-Pacific. Modified from
| Genus | Cortex (reinforced by) | Porocalices (shape) | Accessory spicules | Valid species | Distribution |
|---|---|---|---|---|---|
| Tetilla Schmidt, 1868 | – | – | – | 54 | AT, CA, IP |
| Craniella Schmidt, 1870 | + (minute smooth oxea) | – | – | 42 | AN, AT, CA, IP |
| Cinachyra Sollas, 1886 | + (minute smooth oxea) | + (flask) | – | 3 | AN, AT |
| Paratetilla Dendy, 1905 | – | + (hemi-spherical or narrow) | + (calthrop-like) | 5 | IP |
| Cinachyrella Wilson, 1925 | – | + (hemi-spherical) | – | 42 | AT, CA, IP |
| Amphitethya Lendenfeld, 1907 | + (amphiclads) | – | + (amphiclads) | 2 | IP |
| Fangophilina Schmidt, 1880 | – | + (differentiated, narrow) | – | 4 | AT, CA, IP |
| Acanthotetilla Burton, 1959 | + (megacanthoxea) | + (narrow) | + (megacanthoxea) | 7 | AT, CA, IP |
| Antarctotetilla Carella et al., 2016 | pseudocortex (oxeas loosely arranged) | – | – | 4 | AN |
| Levantiniella Carella et al., 2016 | – | + (small, rounded) | – | 1 | AN |
The species P. bacca and C. australiensis share an obscure taxonomic history, including incomplete descriptions, intermingled identifications, and tens of different species synonymized (see synonyms of C. australiensis in
Materials and methods
Taxonomic revision
Literature from 1867 to date was reviewed in order to compile the descriptions of the 11 nominal species for the genus Paratetilla Dendy, 1905. The Cinachyrella species revision was based on the literature cited by
Sampling
Individuals of Cinachyrella spp. and Paratetilla spp. were collected by snorkelling and SCUBA diving during expeditions to Bali (2003), Bunaken (Sulawesi, 2006), Pulau Seribu (Java, 2005), Raja Ampat (Papua, 2007), Berau (East Kalimantan, 2008), and Ternate (Moluccas, 2009). Sampling was systematically achieved in marine habitats such as coral reefs and mangroves, and within marine lakes (Raja Ampat and Berau). Specimens were photographed in situ and notes made on morphological and ecological features such as color, size, depth, and substrate. A total of 237 specimens were collected and preserved in ethanol 70%; an additional 11 specimens from the Naturalis Biodiversity Center collection from Indonesia and elsewhere were reviewed as well as 20 type specimens. Table
Number of samples reviewed per taxon. The column “Indonesia” refers to all samples recently collected in Indonesia (years 2006–2011), “other material” to older specimens in museum collections from Indonesia or other countries; “types” refer to type specimens of valid species and junior synonyms.
| Species | Indonesia | Other material | Types | Total |
|---|---|---|---|---|
| Paratetilla bacca | 38 | 4 | 4 | 46 |
| Paratetilla arcifera | 21 | 4 | 1 | 26 |
| Cinachyrella australiensis | 117 | 3 | 9 | 129 |
| Cinachyrella porosa | 47 | – | 5 | 52 |
| Cinachyrella paterifera | 14 | – | 1 | 15 |
| Total | 237 | 11 | 20 | 268 |
Morphology
Radial and superficial histological sections of sponges were hand cut with a surgical blade; tissue sections were dried on a heat-plate more than 1 hour, mounted in Durcupan ACR resin and examined using light microscopy. Spicule preparations were made by dissociation of a fragment of sponge in sodium hypochlorite and consecutive washing steps, three times in distilled water, twice in 70% ethanol, and suspending in 95% ethanol. The dissociated spicules were dropped onto glass microscope slides, dried and mounted in Durcupan for light microscopy. Spicule preparations for Scanning Electron Microscopy (SEM) were made after two extra washing steps with 95% ethanol. Spicule dimensions and character definitions follow
Results and discussion
Systematic descriptions
Order Astrophorida
Family Tetillidae Sollas, 1888
Genus Paratetilla Dendy, 1905
The genus Paratetilla was established by
Historic milestones in the taxonomy of the genus Paratetilla Dendy, 1905. Asterisk (*) indicates misidentification of Cinachyrella specimens as Paratetilla.
| Year | Author | Descriptions / Statements |
|---|---|---|
| 1867 | Selenka | Description of Stelletta bacca. Selenka’s material was collected in Samoa Island and due to the presence of triaenes this species was associated to the family Corticatae (now Astrophorida: Ancorinidae). The description is brief but the sketches included are illustrative, including “Vierstrahler” (=calthrop-like) spicules. Sigma-like spicules are neither mentioned in the description nor drawn in the figures. Currently, type specimen could not be located. |
| 1883 | Carter | Description of Tethya merguiensis, including sigmaspires, calthrop-like spicules, oxeas and triaenes and their respective measurements and sketches. |
| 1884 | Ridley | In his monograph, Ridley kept Stelletta bacca in the genus Stelletta. The diagnostic characteristic for Stelletta for his decision was the absence of bacillar or acerate flesh-spicules. He also noticed that the Samoan Stelletta “is probably a Tethya, as its stellate agrees with the large stellate of that genus, and its forks are rare and probably foreign to the sponge” (see footnote in |
| 1887 | Vosmaer | Statement about Stelletta bacca mentioning that it can hardly belong to Stelletta genus without further argumentation. |
| 1888 | Sollas | Establishment of Family Tetillidae, type genus Tetilla Schmidt, 1868. Sponges in this family have sigmaspires (microscleres) and slender protriaenes (megascleres) as diagnostic characters. In this family Sollas included the species Craniella (Alcyonium) cranium |
| 1896* | Kieschnick | Description of Tetilla ternatensis based on material from Ternate Island (Indonesia); he mentioned “Vierstrahler” (=calthrops). |
| 1897 | Lindgren |
Tethya merguiensis Carter, 1873 as junior synonym to Stelletta bacca, based on a comment by |
| 1898* | Lindgren | Redescription of Tetilla bacca, with Tetilla merguiensis as junior synonym, including material of Torres Straits (North Australia), two localities at Java (Indonesia) and Carter’s specimens from Mergui Archipelago. Size range for each station is shown for oxeas and triaenes, arguing that larger spicules are found to the west while smaller sizes to the east. Redescription of Tetilla ternatensis based on Java material. It is remarkable that he mentioned the presence of numerous microxeas (240 × 4 µm) and sigmaspires 24 µm. |
| 1898 | Kieschnick | Description of Tetilla amboinensis, Tetilla violacea and Tetilla rubra from Amboina Island, all of them with “Vierstrahler” (=calthrop-like) spicules. T. amboinensis and T. violacea with calthrops in a layer below the surface of the sponge; while the former is characterized by smaller number of triaenes and bundles of oxeas up to the surface of the sponge, the latter by very abundant triaenes, bundles of oxeas projected over the surface of the sponge, and a typical violet color. T. rubra separated from the other two by its brick-red color and with calthrops mainly on the basal part of the sponge. |
| 1900 | Kieschnick | Extensive description of the same three new species. |
| 1900* | Thiele | Redescription of Tetilla ternatensis Kieschnick, 1896. Thiele drew attention on the misidentification of T. ternatensis by |
| 1900* | Kirkpatrick | Extension of the geographical range of T. bacca and T. ternatensis to Christmas Island. T. bacca specimens were described with identical spicules to |
| 1903* | Lendenfeld | Designation of a new species Tetilla lindgreni based on Lindgren’s specimens (1898) from Java and Kirkpatrick’s specimens (1900) from Christmas islands, both identified as T. ternatensis without calthrops and with small microxeas. Thus, Lendenfeld concluded that those specimens belong to a new species (T. lindgreni) because they did not show calthrops as in the original description of |
| 1905 | Dendy | The genus Paratetilla was erected within the family Tetillidae, based on the presence of calthrop-like spicules. Thus, Tetilla bacca is transferred to Paratetilla genus, including their junior synonyms T. merguiensis, as well as the three Kieschnick’s species T. ternatensis, T. amboinensis and T. violacea based on Thiele’s annotation (1903). Description of Paratetilla cineriformis based on material from Gulf of Manaar (Sri Lanka). Although the spicules shown by P. cineriformis resembled T. merguiensis, |
| 1907 | Lendenfeld | The genus Amphytethya was created based on its characteristic amphitriaenes. Many other species under the genus Cinachyra, Fangophilina and Tetilla were described. |
| 1911 | Row | Description of Paratetilla eccentrica from the Red Sea. Cortical triaenes (= calthrop-like) with high modifications, in some cases even becoming into “walking-sticks”. |
| 1912 | Hentschel | Description of Paratetilla aruensis from Aru- and Kei- Islands (Indonesia), with characteristic amphitriaenes. Relocation of the genus Amphitethya Lendenfeld, 1907 as a junior synonym of Paratetilla. |
| 1922 | Dendy | All nominal species with calthrop-like spicules were synonymized to Paratetilla bacca, except for P. aruensis Hentschel, 1912. Two varieties were identified: P. bacca var. violacea based on T. violacea characteristics, and the new variety P. bacca var. corrugata from Diego Garcia in the Indian Ocean. |
| 1925 | Wilson | Description of Paratetilla arcifera from Philippines. Wilson recognized as valid four additional species: P. bacca (Selenka, 1867), P. amboinensis (Kieschnick, 1898), P. cineriformis (Dendy, 1905) and P. eccentrica (Row, 1911). However, he also commented that P. bacca is a comprehensive variable species, as previously proposed by |
| 1954 | de Laubenfels | Description of Paratetilla lipotriaena from Micronesia (West-Central Pacific), characterized by variable calthrop-like spicules and the absence of triaenes, and relatively similar to P. eccentrica Row, 1911. |
| 1959 | Burton | All nominal species described within the genus Paratetilla were included as synonyms of P. bacca, except for P. lipotriaena. |
| 1987 | Rützler | Review of Family Tetillidae, including seven genera (all except for Fangophilina). Nomination of Paratetilla cineriformis as type species of genus Paratetilla. |
| 1994 | Hooper and Wiedenmayer | Review of all Paratetilla bacca synonyms based on |
| 2002 | van Soest and Rützler | Review of the eight genera included within family Tetillidae. Although Paratetilla characters were a combination of two descriptions, a paragraph in the discussion included the size differences between both Selenka’s and Carter’s material (Stelletta bacca and Tethya merguiensis, respectively). The origin of calthrop-like spicules was also discussed as probably modified plagiotriaenes resembling some Cinachyrella species, arguing the possibility of the inclusion of the widespread species Paratetilla bacca within Cinachyrella genus. |
| 2008 | van Soest and Beglinger | Redescription of Paratetilla corrugata based on material from the Gulf of Oman, and giving validity to the variety P. bacca var. corrugata by |
| 2018 | van Soest et al. (WPD) | Junior synonyms for Paratetilla bacca (Selenka, 1867): Tetilla bacca (Selenka, 1867), Stelletta bacca bacca Selenka, 1867, Tethya merguiensis Carter, 1883, Stelletta bacca Selenka, 1887, Tetilla violacea Kieschnick, 1896, Tetilla ternatensis Kieschnick, 1896, Tetilla rubra Kieschnick, 1898, Paratetilla cineriformis Dendy, 1905, Paratetilla eccentrica Row, 1911, Paratetilla arcifera Wilson, 1925. Other accepted Paratetilla species in WPD: Paratetilla amboinensis (Kieschnick, 1898), Paratetilla aruensis Hentschel, 1912, Paratetilla corrugata Dendy, 1922, Paratetilla lipotriaena de Laubenfels, 1954. |
| 2018 | This study | Paratetilla species from Indonesia: Paratetilla bacca (Selenka, 1867), Paratetilla arcifera Wilson, 1925, and Paratetilla corrugata Dendy, 1922 (not observed in our Indonesian material), Paratetilla aruensis Hentschel, 1912 with amphitriaenes, it is suggested to be transferred to Amphitethya. |
Recent checklists and biodiversity studies in the Indo-Pacific have only recorded P. bacca, following Burton’s taxonomic decision in 1959 to synonymize all nominal Paratetilla species except P. lipotriaena. Two exceptions were found in the literature, the review by
Paratetilla bacca
Stelletta bacca Selenka, 1867: 569, pl. xxxv, figs 14, 15 (type not found, material from type locality seen).
Tethya merguiensis Carter, 1883: 366, pl. xv, figs 6–8; Carter, 1887: 80 (type seen).
Tetilla merguiensis ; Sollas, 1888: 14; Topsent, 1897: 441, pl. xviii, fig. 4–5, pl. xxi figs 34.
Tetilla ternatensis Kieschnick, 1896: 527. Thiele, 1900: 39, pl. ii, fig 13; Not Tetilla ternatensis Lindgren, 1898: 329 pl. 17, fig. 14; pl. 19, Fig. 25 a-e, a’, b’.
Tetilla bacca ; Lindgren, 1897: 485; Lindgren, 1898: 328; Thiele, 1900: 39, pl. ii, fig 13; Kirkpatrick, 1900: 132 (material seen); Lendenfeld, 1903: 19.
Tetilla amboinensis Kieschnick, 1898: 10.
Tetilla violacea Kieschnick, 1898: 15.
Tetilla rubra Kieschnick, 1898: 18.
Paratetilla cineriformis Dendy, 1905: 97, pl. iii, fig. 7 (type seen).
Paratetilla eccentrica Row, 1911: 306, pl. xxxv, fig. 1, pI. xxxvi, fig. 8 (type seen).
Cinachyra amboinensis ; Hentschel, 1912: 331.
Paratetilla bacca ; Dendy, 1922: 21 (material seen).
Paratetilla bacca var. violacea ; Dendy, 1922: 22, pl. 1, fig. 6 (material seen).
Paratetilla lipotriaena de Laubenfels, 1954: 244, text figure no. 168 (type seen).
Material examined
Neotype ZMA.POR.13029, Tutuila Island, American Samoa. Holotype of first junior synonym Tethya merguiensis Carter, 1883 (?)
Other material: East Kalimantan, Makassar Straits, ZMA.POR.1735, Siboga Expedition, St. 81. Singapore,
Description
External morphology. Globular sponges, size between 1 and 5 cm in diameter. Surface hispid due to the projecting spicules, covered by numerous porocalices (Figure
Paratetilla bacca. A,B, G-M
Skeleton. No cortex. Choanosomal skeleton composed by bundles of oxeas and triaenes radiating from a central core, ⅕–⅓ of the diameter of the sponge.
Megascleres. The material from Indonesia and the type of P. merguiensis have oxeas 850–3085.3–4500 mm × 5–41.5–65 mm (Table
Spicule measurements of six specimens of Paratetilla bacca and five specimens of P. arcifera from different regions (n = 10 per spicule type and dimension with minimum-mean-maximum). Asterisk (*) indicate that rhabds of spicules were broken and no measurement was possible.
| Measurements | Paratetilla bacca | Paratetilla arcifera | ||||||||||
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| NHMUK94. 11.16.17/16 |
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USNM21278 (Holotype) |
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| Locality | Mergui Archipelago | Kakaban Lake | Haji Buang Lake | Berau | Raja Ampat | Ternate | Philippines | Berau | Berau | Ternate | Manado | |
| Habitat | Reef | Marine Lake | Marine Lake | Reef | Mangrove | Reef | Reef | Reef | Reef | Reef | Reef | |
| Oxeas | Length | 3114.36–3114.6–3115 | 850–2340.8–3150 | 1000–2922–3850 | 2520–3324.6–3850 | 3100–3270–3500 | 1250–3540–4500 | 1650–2435–3125 | 1650–3093–4500 | 1600–3041–4175 | 840–1996–3100 | 3100–3600–4000 |
| Width | 40.8–42.5–51 | 5–29.9–40 | 12.5–36–50 | 30–48.8–60 | 30–42–55 | 25–49.6–65 | 20–39.5–65 | 25–42.2–55 | 20–33.7–50 | 10–25.4–50 | 27.5–43–52.5 | |
| Anatriaenes | Rhabd length | * | 3000–3677.8–4600 | 3900–4741.7–5300 | 4250–5057.1–6000 | * | * | * | * | * | * | * |
| Rhabd width | 15–16–20 | 5–12.9–20 | 12.5–14.8–17.5 | 7.5–10.7–15 | 7.5–11.3–15 | 5–7.5–10 | * | * | * | 5–5.7–7.5 | 5–6.3–7.5 | |
| Cladi total | 20–27.9–40 | 37.5–58.7–75 | 80–91.5–100 | 40–73.6–90 | 60–74.5–80 | 22.5–49.5–75 | 40–68–80 | 60–65.6–80 | 22.5–39.8–60 | 40–48.2–50 | 50–65–75 | |
| Cladi length | 40.8–52–71.4 | 25–44.4–65 | 50–63.9–75 | 20–40.5–50 | 40–57.3–75 | 12.5–31.8–50 | 25–39.4–45 | 30–38.7–47.5 | 10–21.3–30 | 15–20.9–25 | 35–40.6–50 | |
| Cladi width | 10–12–15 | 7.5–12.1–20 | 7.5–10.9–15 | 7.5–10.7–15 | 7.5–9.5–10 | 5–7-7.5 | 5–8.2–10 | 5–6.6–10 | 2.5–5-7.5 | 5–5-5 | 5–6.3–7.5 | |
| Protriaenes | Rhabd length | * | 3900–3900–3900 | 3000–4434.6–5850 | 3100–3800–4500 | * | * | * | * | * | * | * |
| Rhabd width | * | 7.5–10–12.5 | 10–13.3–17.5 | 5–8.6–15 | 2.5–4.8–5 | 5–5-5 | * | * | * | * | 5–6-7.5 | |
| Cladi total | * | 27.5–46.9–70 | 40–70.7–100 | 50–67.9–100 | 30–54–70 | 30–30–30 | * | 30–38.3–40 | 40–40–40 | * | 30–42.8–50 | |
| Cladi length | * | 47.5–84.4–100 | 85–140.5–185 | 110–141.4–200 | 50–133–170 | 37.5–37.5–37.5 | * | 50–61.7–70 | 75–75–75 | * | 25–40.3–60 | |
| Cladi width | * | 5–7.5–10 | 7.5–9.5–12.5 | 5–7.9–10 | 2.5–2.5–2.5 | 5–5-5 | * | 2.5–5.4–7.5 | 5–5-5 | * | 2.5–4.3–5 | |
| Calthrops | C1 | 42.5–168.1–255 | 110–266–475 | 270–369.8–510 | 140–296.7–360 | 220–301–350 | 250–375.7–600 | 320–362.5–430 | 150–253.9–375 | 110–154–220 | 150–192.5–250 | 120–179–220 |
| C2 | 22.5–92.9–183.6 | 90–225–325 | 220–346.4–460 | 140–281–350 | 210–284–350 | 240–291.4–350 | 230–287.5–320 | 75–239.4–390 | 90–134.4–160 | 70–125–230 | 90–129–200 | |
| C3 | 20–106–234.6 | 40–203.7–325 | 50–292.5–400 | 25–218.7–345 | 180–254–310 | 200–272.9–350 | 120–195–300 | 140–245.5–355 | 60–110–150 | 50–93.3–160 | 90–129–200 | |
| Width | 3.5–12.2–20.4 | 7.5–18.3–35 | 15–31.7–45 | 10–18.3–27.5 | 10–14.5–17.5 | 17.5–20.4–25 | 15–18.8–22.5 | 10–18.3–25 | 10–12.5–15 | 10–13.8–17.5 | 12.5–15–20 | |
| Microxea | 173.4–195.3–224.4 | 105–136.2–212.5 | 170–213.6–250 | 250–316.8–385 | 210–264–300 | 250–323.6–380 | 180–308.4–380 | 270–323.2–400 | 200–342–500 | 340–370–410 | 250–367–450 | |
| Sigmaspires | 12.5–14.4–17.5 | 10–13–17.5 | 12.5–15.4–25 | 12.5–14.2–17.5 | 12.5–13.8–15 | 12.5–14–17.5 | 7.5–12.5–17.5 | 12.5–15.4–17.5 | 15–16.3–20 | 12.5–15.3–17.5 | 12.5–14–17.5 | |
Microscleres. Thin microxeas are common, 105–241.6–380 mm, ‘hair-like’. Sigmaspires, 10–14.1–25 mm, C-S shape (Figure
Ecology
Inhabiting all studied environments in Indonesia, including coral reefs, mangroves, and marine lakes. Specimens more common in mangroves and marine lakes, and shallow reef flats where they are usually found on dead coral skeletons or coral rubble, typically ranging in depth from 0–5m. No specimens collected from deeper coral reefs in Indonesia.
Distribution
Paratetilla bacca has a wide distribution in Indonesia, including Berau, Bunaken, Raja Ampat, Ternate, and Java. Previous Indonesian records are from Spermonde Archipelago (
Distribution of Paratetilla bacca. Red dot: type locality, Stelletta bacca Selenka, 1867, American Samoa. Green dots: Indonesian localities where the species was collected recently. Yellow triangles: Records from localities outside Indonesia, Zanzibar, Southwest Madagascar, Seychelles, Thailand, Singapore, Christmas Island, and Philippines. Circled numbers: type localities of synonymized species, 1 Tethya merguiensis Carter, 1873, Mergui Archipelago 2 Tetilla ternatensis Kieschnick, 1896, Ternate Island 3 Tetilla amboinensis Kieschnick, 1898, Ambon Island 4 Tetilla violacea Kieschnick, 1898, Ambon Island 5 Tetilla rubra Kieschnick, 1898, Ambon Island 6 Paratetilla cineriformis Dendy, 1905, Gulf of Manaar, Sri Lanka. 7 Paratetilla eccentrica Row, 1911, Tella Tella Kabira, Red Sea 8 Paratetilla lipotriaena de Laubenfels, 1954, Matalanim, Eastern Pohnpei, Micronesia.
Remarks
We did not succeed in locating the holotype of Paratetilla bacca, despite concerted effort. At this time, we assume that the type is no longer available. The description by Selenka of the type specimen does not mention the occurrence of any type of sigmaspires. It is a matter of speculation whether
According to the WPD (
In a molecular phylogenetic study, which was based in part on specimens that we review in the current study (see Suppl. material
Paratetilla arcifera
Paratetilla arcifera Wilson, 1925: 380; plate 40, fig. 2; plate 48, fig. 6 (type seen).
Material examined
Holotype USNM 21278, Albatross Stn. 5400, Malapascua Island, Cebu, Philippines, 46 m, 16 Mar 1909. INDONESIA. East Kalimantan, Berau reef,
Description
External morphology. Globular sponges, size from 3 to 6 cm in diameter (Figs
Paratetilla arcifera. Holotype USNM 21278, Malapascua Island, Cebu, Philippines A preserved specimen showing large porocalices B Labels of holotype C skeleton, showing calthrops and radial bundles D skeleton, showing oxeas, calthrops, and anatriaenes E oxea, end detail F-I different calthrop shapes and sizes J anatriaene K, L protriaene, different types M thin microxea, detail N thin microxea, full length O sigmaspires. Scale bars: 1 cm (A); 500 μm (C, D); 100 μm (E); 50 μm (F–I, N); 20 μm (J); 40 μm (K, L); 5 μm (M, O).
Paratetilla arcifera from Indonesia
Skeleton. No cortex. Skeleton composed by bundles of oxeas and triaenes radiating from a central core, and spaced between each other, giving a softer consistency (Figs
Megascleres. Holotype and Indonesian specimen size ranges are summarized in Table
Microscleres. Thin microxeas are common, 180–308.4–380 mm, ‘hair-like’ (Figs
Ecology
Coral reef habitats at depths from 1- 20/30 m. Absent from marine lakes, mangroves and other localities with higher sedimentation and/or variable salinity.
Distribution
Occur in coral reefs of Berau, Bunaken, Ternate, and Raja Ampat. An additional record from its type locality, Philippines (Wilson, 1925) could be inferred from the literature (see
Remarks
Spicule sizes for most Indonesian specimens vary within the holotype ranges, except for the Ternate population, which exhibits smaller sizes and lack of protriaenes (Table
Genus Cinachyrella Wilson, 1925
Currently, 42 species are valid within the genus Cinachyrella according to the WPD (
Historic milestones in the taxonomy of Cinachyrella australiensis and other Cinachyrella species from Indonesia. Asterisk (*) indicates misidentification of Cinachyrella specimens as Paratetilla.
| Year | Author | Descriptions / Statements |
|---|---|---|
| 1873 | Gray | Description of the monotypic genera Psetalia and Labaria, with the species P. globulosa and L. hemisphaerica, respectively. |
| 1886 | Carter | Description of Tethya cranium var. australiensis from Port Phillip Heads (South Australia) collected at 36 m depth. This species was characterized by the presence of minutely spined (= acanthose) microxea (210 μm). |
| 1888 | Sollas | Establishment of Family Tetillidae. Tethya cranium var. australiensis was redescribed as Tetilla (?) australiensis. In addition, Sollas noted that the characteristic microxeas of T. australiensis were also present in T. merguiensis as well, but were more abundant in T. australiensis. |
| 1888 | Lendenfeld | Description of genus Spiretta within Family Tetillidae, including two new species S. raphidiophora and S. porosa, from Port Jackson (SE Australia) and Port Denison (NE Australia), respectively. The former with microxea (240 × 2 µm) and the latter without them. |
| 1891 | Keller | Description of Cinachyra schulzei from the Red Sea and Mozambique, with microxea 250 × 5 µm. |
| 1896* | Kieschnick | Description of Tetilla ternatensis based on material from Ternate Island (Indonesia). He mentioned “Vierstrahler” (= calthrops). |
| 1898* | Lindgren | Redescription of Tetilla ternatensis based on Java material. It is remarkable that he mentioned the presence of numerous microxea (240 × 4 µm) and sigmaspires 24 µm. |
| 1898 | Kieschnick | Description of Tetilla schulzei from material of NE Australia to Ambon Island, with microxea (198-220 µm × 4 µm). T. schulzei has ‘oscula’ that we interpret as porocalices. Although Kieschnick entitled T. schulzei as new species, it is not clear if he was aware of Cinachyra schulzei described by |
| 1899 | Thiele | Record of Tetilla australiensis from Sulawesi (Indonesia). Specimens with acanthose microxea (180-200 µm × 2.5 µm). |
| 1900* | Thiele | With the redescription of Tetilla ternatensis Kieschnick, 1896, Thiele noticed the misidentification of T. ternatensis by |
| 1900 | Kieschnick | Additional record of Tetilla schulzei from Ambon Islands, including description of the specimens, with microxea from 198 to 220 µm × 4 µm. |
| 1900* | Kirkpatrick | Extension of geographical range of T. bacca and T. ternatensis to Christmas Island. T. bacca specimens were described with identical spicules to Lindgren’s material from Java. T. ternatensis also similar to Lindgren’s material of T. ternatensis, this is having microxeas and missing calthrops. |
| 1902 | Sollas | Description of Cinachyra malaccensis from Malaysia. Cup-shaped porocalices are described together with different spicules, except for microxea. In the available figures, no microxeas are shown. |
| 1903* | Thiele | Redescription of Tetilla ternatensis Kieschnick, 1898. He drew attention on the misidentification of T. ternatensis by |
| 1903* | Lendenfeld | Designation of a new species Tetilla lindgreni based on T. ternatensis material described by |
| 1905 | Dendy | Monograph on sponges from Sri Lanka. Description of Tetilla anomala, showing remarkable siliceous micro-spherules (4 µm) and no microxeas. Description of Tetilla poculifera with smooth microxeas (230 × 5 µm). Description of Tetilla limicola, pink-color and root tuft; neither porocalices nor microxea are described. The genus Paratetilla was established. |
| 1906 | Baer | Description of Tethya armata from Zanzibar (Africa, Indian Ocean). It is characterized by a dermal cortex formed by microxea (166-296 µm × 1-2 µm). |
| 1907 | Lendenfeld | Description of Cinachyra isis and Tethya hebes from NW Australia, the first one exhibiting smaller microxea (130-160 µm × 2-5.5 µm), and the second one larger rough microxea (= acanthose microxea, 250-275 µm × 4-6 µm). Description of Cinachyra alba-tridens, C. alba-obtusa, and C. alba-bidens species, slightly differentiated by the geometry and abundance of triaenes. He kept the three species because they were collected in three distant localities, Chagos Archipelago, Papua New Guinea, and Tonga Islands, respectively; “alba-group” species do not contain microxeas, and sigmaspires are small (<10 µm). |
| 1911 | Row | Description of Chrotella ibis from the Red Sea. Species with smooth microxea (150 × 2.1 µm), sharing this character with Tetilla poculifera, and Paratetilla species P. merguiensis, P. eccentrica and P. cineriformis. In his description, Row clearly differentiated his species from T. australiensis due to the latter having acanthose microxea. |
| 1911 | Hentschel | Description of Tetilla cinachyroides from South Australia. Species with acanthose microxea (112-168 µm × 2.5 µm), sigmaspires (10-12 µm) and spherules (5 µm). |
| 1912 | Hentschel | Description of Cinachyra mertoni and Cinachyra nuda from Aru- and Kei- Islands (Indonesia). Both species contain microxea, the first one smooth 250 µm, whereas in the second one they are acanthose, from 200-280 µm, and no anatriaenes were found. A third species, Tethya clavigera, with oscula (similar to porocalices) and no microxea was also described. |
| 1922 | Dendy | Description of Cinachyra vaccinata and C. providentiae from the Indian Ocean. Both of them with microxea (no mention whether acanthose or not), being 200 × 4 µm in the former, and 220 × 5.5 µm in the latter one. C. vaccinata characterized by small hair-like protri- and prodiaenes, terminating in an elongated oval swelling tip unique to this species. C. providentiae with bottle-shaped porocalices. |
| 1925 | Wilson | Establishment of Cinachyrella as a subgenus of Tetilla, with type species Tetilla hirsuta Dendy, 1889. The characters used to distinguish Cinachyrella species from the other were special depressions (=porocalices) and no specialization of a cortical zone. Wilson included the following species within Cinachyrella: Cinachyra malaccensis Sollas, 1902; Tetilla limicola Dendy, 1905; Tetilla anomala Dendy, 1905; Cinachyra isis Lendenfeld, 1907; C. hamata Lendenfeld, 1907; C. alba-tridens Lendenfeld, 1907; C. alba-bidens Lendenfeld, 1907; C. alba-obtusa Lendenfeld, 1907; C. vertex Lendenfeld, 1907; Tetilla cinachyroides Hentschel, 1911; Cinachyra phacoides Hentschel, 1911; Tethya clavigera Hentschel, 1912; Cinachyra mertoni Hentschel, 1912; Cinachyra nuda Hentschel, 1912; Cinachyra vaccinata Dendy, 1922; Cinachyra providentiae Dendy, 1922. In addition, Cinachyrella crustata and Cinachyrella paterifera were described from Philippines. C. crustata with distinctive long and stout promonoenes, no microxea. C. paterifera with a characteristic cloaca (= large osculum) on top and root-like structure to attach to sediments, microxea (250 × 2 µm) observed in two specimens although almost absent in the third one of the type series, pointing out a high variability in the presence of microxea within the same individual. |
| 1934 | Burton | Taxonomic revision of Cinachyra australiensis. In his compilation, Burton grouped 16 nominal species described in 32 references and designated them as junior synonyms of the widespread species C. australiensis. Three different groups were recognized: the australiensis-group characterized by the presence of acanthose microxea; the schulzei-group with smooth microxea; and the porosa-group without microxea. Description of genus Raphidotethya. |
| 1954 | de Laubenfels | Identification of Cinachyra porosa and Cinachyra australiensis from Micronesia (West-Central Pacific). |
| 1973 | Thomas | Records of Cinachyra cavernosa (Lamarck, 1815) from the Seychelles Islands, having, microxea (126 × 2 µm) sometimes granulated (= acanthose). Among the junior synonyms of C. cavernosa, Thomas included Tethya cranium var. australiensis Carter, 1886, Chrotella australiensis Burton, 1937, and Chrotella cavernosa Burton, 1959. However, in the WPD ( |
| 1982 | Pulitzer-Finali | Description of Cinachyra tenuiviolacea from the Great Barrier Reef (Australia), characterized by a light violet color, small oxeas (up to 2500 µm × 13-25 µm), atrophic anatriaenes, no microxeas, and no protriaenes in the choanosome. |
| 1987 | Rützler | Review of Family Tetillidae, including seven genera (all except for Fangophilina). Subgenus Cinachyrella was elevated to the hierarchy of genus. |
| 1992 | Rützler and Smith | Review of four species of Cinachyrella for the Caribbean region, mainly described by Uliczka (1929). Geometry and size ranges of all spicule types were shown. According to their descriptions, Cinachyrella kuekenthali is the most similar species to C. australiensis, since both of them have acanthose microxea. |
| 1994 | Hooper and Wiedenmayer | Compilation of Cinachyra australiensis synonyms based on |
| 2002 | van Soest and Rützler | Review of the eight genera of tetillids, including Cinachyrella. Cinachyra australiensis was transferred into the genus Cinachyrella. The authors considered that all junior synonyms proposed for C. australiensis by |
| 2018 | van Soest et al. (WPD) | Accepted synonyms of Cinachyrella australiensis (Carter, 1886): Tethya australiensis Carter, 1886; Spiretta porosa Lendenfeld, 1888; Cinachyra malaccensis Sollas, 1902; Tetilla lindgreni Lendenfeld, 1903; Tethya armata Baer, 1906; Cinachyra isis Lendenfeld, 1907; Tetilla cinachyroides Hentschel, 1911; and Cinachyra providentiae Dendy, 1922. Valid Cinachyrella spp. from the Indo-pacific (excluding species only found in the Red Sea) comprise 6 species |
| 2018 | This study | From our detailed examination of Indonesian material and type material, we conclude that in Indonesia there are three species: Cinachyrella australiensis (Carter, 1886), Cinachyrella porosa (Lendenfeld, 1888), and Cinachyrella paterifera Wilson, 1922. Further investigations will reveal if the five species from the C. schulzei- group can be synonymized or belong to separate and distinctive species. |
Cinachyrella australiensis has been recorded from a wide geographic area from the Gulf of Oman (
Ecological studies on the morphological plasticity of C. australiensis from North Australia (
Cinachyrella australiensis
Tethya cranium var. australiensis Carter, 1886: 127 (holotype seen).
Tetilla ? australiensis ; Sollas, 1888: 43.
Spiretta raphidiophora Lendenfeld, 1888: 43 (type seen).
Tetilla hirsuta Dendy, 1889: 75 (type seen).
Tetilla ternatensis Lindgren, 1898: 329 pl. 17, fig. 14; pl. 19, Fig. 25 a-e, a’, b’. Ternate Not 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 ternatensis Kieschnick*, 1896: 527.
Tetilla lindgreni Lendenfeld, 1903: 18.
Tetilla australiensis ; Lendenfeld, 1903: 20.
Tethya hebes Lendenfeld, 1907: 98, pl. XVI, figs 19–38. 19`South NW Australia, 91 m depth (syntype seen).
Cinachyra isis Lendenfeld, 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 cinachyroides Hentschel, 1911: 281, textfig. 1. NW Australia, Barrow Island.
Cinachyra nuda Hentschel, 1912:333, pl. XIII, fig.2; pl. XVIII fig. 13. Aru Island (type seen).
Cinachyra vaccinata Dendy, 1922: 14, pl. 1, fig. 4; pl. 11, figs 1a-l. Diego Garcia, Chagos Island (type seen).
Cinachyra providentiae Dendy, 1922: 18, pl.1, figs 5–5a; pl. 10, figs2a–f. Providence Island (type seen).
Tetilla (Cinachyrella) hirsuta ; Wilson, 1925: 365, pl. 39, fig.4.
Cinachyra australiensis ; Burton, 1934: 523. In part, not C. australiensis in porosa-group, nor C. australiensis in schulzei-group; de Laubenfels, 1954: 241, text-fig. 166.
Cinachyrella anatriaenilla Fernandez, Kelly, Bell, 2017: 83, figs 2–4.
Material examined
Holotype
Other material: Singapore,
Other types and material examined (not included as synonyms of C. australiensis):
Description
External morphology. Globular sponges, size from 4 to 10 cm in diameter (Figure
Cinachyrella australiensis. A, C, E-H, L
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
Spicule measurements of eight specimens Cinachyrella australiensis, five specimens of C. anomala, four specimens of C. paterifera from 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 | Port Phillip Heads | Berau | Kakaban | Berau | Kakaban | Berau | Ternate | Raja Ampat | Port Denison | Tanah Banban | Teluk Pea | Raja Ampat | Ternate | Philippines | Berau | Raja Ampat | Berau | |
| Habitat | Reef? | Reef | Marine Lake | Reef | Marine Lake | Reef | Reef | Mangrove | Reef | Marine Lake | Reef | Reef | Reef | Reef | Reef | Reef | Reef | |
| Oxeas | Length | 3375-4135.5-5500 | 1000-3332-4500 | 1375-2912-4000 | 2425-3822.8-5500 | 2250-3066-4250 | 2300-4315-5750 | 1500-2676-3800 | 2000-2658.3-3750 | 820-2553.2-3750 | 550-2138.1-3750 | 1350-2702.5-4000 | 1250-2304.2-3150 | 1500-2710-3500 | 1400-3011.5-4750 | 2850-3580.6-4500 | 1850-3060-5000 | 800-2748.2-4500 |
| Width | 15-24.7-37.5 | 10-40-52.5 | 17.5-33.9-55 | 25-41.7-60 | 17.5-26.4-35 | 37.5-59.1-77.5 | 12.5-30.3-60 | 25-31.3-37.5 | 7.5-29.4-47.5 | 5-28.2-60 | 7.5-29.5-47.5 | 12.5-29.4-40 | 7.5-32.6-45 | 10-34.5-62.5 | 30-46.1-62.5 | 12.5-35.6-55 | 5-34.2-75 | |
| Anatriaene | Rhabd | ** | 2750-3271.9-3650 | 2250-3317.9-4250 | 2700-3083.3-3300 | * | * | * | * | * | * | * | * | * | * | * | * | 4250-4250-4250 |
| Rhabd width | 5-6-7.5 | 2.5-5.3-7.5 | 4-4.9-5 | 5-5-5 | 5-5-5 | 7.5-8.4-10 | 5-5.3-7.5 | 2.5-5.8-7.5 | 10-12-15 | 5-7.8-10 | 5-5-5 | 5-6.1-7.5 | 5-7.5-10 | 5-6.6-10 | 7.5-8.8-10 | 5-5-5 | ||
| Cladi total | 50-63.5-85 | 30-74.2-100 | 45-58-70 | 40-55-70 | 60-81-100 | 55-79.4-100 | 70-83-100 | 50-67.6-100 | 65-71-80 | 60-65-70 | 50-62.9-70 | 50-62.5-75 | 17.5-24.2-30 | 17.5-37.5-75 | 70-90-110 | 65-79.3-110 | ||
| Cladi length | 40-49.5-65 | 20-60.4-80 | 30-42.5-50 | 35-45-60 | 32.5-49.5-57.5 | 30-49.1-67.5 | 50-66-80 | 30-42-60 | 45-56-65 | 40-52-70 | 40-51.4-60 | 30-50.4-62.5 | 2-6.5-10 | 7.5-25.7-80 | 50-65-80 | 42.5-58.2-90 | ||
| Cladi width | 2.5-4.5-5 | 2.5-3.5-5 | 4-5.6-10 | 5-5-5 | 5-5.1-6 | 5-5.6-7.5 | 5-5-5 | 2.5-5.6-7.5 | 10-10.5-12.5 | 2.5-6.6-7.5 | 2.5-4.6-5 | 5-5.4-7.5 | 5-5.8-7.5 | 5-6.4-10 | 7.5-7.5-7.5 | 2.5-4.3-5 | ||
| Protriaene | Rhabd length | ** | 3900-4550-5800 | 3700-4262.5-4750 | 840-3522.5-5000 | 2250-2375-2500 | ** | * | * | * | * | * | * | * | * | 3500-4210-5350 | * | 4300-4689.3-5100 |
| Rhabd width | 10-13.3-20 | 12.5-14.7-15 | 10-12.5-15 | 2.5-5.9-7.5 | 7.5-8.3-10 | 2.5-5-7.5 | 5-7.3-12.5 | 5-8.8-15 | 2.5-6.5-10 | 2.5-3.6-5 | 2.5-5-7.5 | 10-10-10 | 5-9.5-12.5 | 5-7.8-10 | 10-15.2-17.5 | |||
| Cladi total | 20-46.9-70 | 20-58.8-80 | 40-60.8-80 | 40-55-80 | 45-50.4-55 | 25-70.5-100 | 25-44.4-65 | 20-51.25-80 | 40-59-80 | 30-44.3-60 | 50-53.8-67.5 | 30-32.5-35 | 35-53.9-75 | 40-70-100 | 30-37.3-60 | |||
| Cladi length | 35-66.3-110 | 25-93.8-170 | 75-95.8-130 | 50-79.5-110 | 40-77.9-120 | 30-108-150 | 35-73-110 | 30-77.5-125 | 60-100-160 | 40-67.1-100 | 50-78.3-137.5 | 22.5-31.3-40 | 20-74.1-130 | 30-82.5-140 | 40-51.6-80 | |||
| Cladi width | 5-7.9-10 | 10-11.6-12.5 | 10-10.8-12.5 | 2.5-4.8-5 | 5-5.8-7.5 | 2.5-4.3-5 | 5-5.1-7.5 | 5-7.5-15 | 2.5-5.5-10 | 2.5-2.5-2.5 | 2.5-4-5 | 7.5-7.5-7.5 | 2.5-6.8-7.5 | 5-6.6-7.5 | 10-11.6-15 | |||
| Strongyle | Length | ** | -- | 4000-4000-4000 | 2450-3041.7-4200 | 2200-2650-2850 | 2650-2650-2650 | ** | 3350-3350-3350 | ** | ** | * | 2450-2800-3250 | 2100-2975-3700 | 1800-1862.5-1925 | |||
| Width | 50-50-50 | 35-43.3-60 | 32.5-39.4-50 | 35-35-35 | 45-45-45 | 45-45-45 | 40-43.1-50 | 35-45.3-62.5 | 35-37.5-40 | |||||||||
| Acanthose microxea | 117-166.9-260 | 160-197.3-230 | 150-165-200 | 200-230-270 | 150-183.5-240 | 157.5-189.5-225 | 170-191.4-225 | 137.5-154-175 | ||||||||||
| Sigmaspires | 10-14.4-17.5 | 12.5-15-17.5 | 10-12.3-15 | 12.5-14.8-17.5 | 10-12-15 | 15-16.5-20 | 12.5-15.7-20 | 12.5-14.8-17.5 | 5-8.6-12.5 | 7.5-8.5-10 | 5-8.9-12.5 | 5-8-10 | 5-8-10 | 10-13.2-17.5 | 12.5-15.3-17.5 | 12.5-14.5-17.5 | 12.5-16.2-20 | |
| Protriaene (hair-like) | Rhabd length | * | * | * | 550-698.9-820 | |||||||||||||
| Width | 2.5-2.5-2.5 | 2.5-2.5-2.5 | 2.5-2.5-2.5 | 2-2.2-2.5 | ||||||||||||||
| Cladi total | 7.5-12.5-17.5 | 7.5-10.9-17.5 | 12.5-13.6-15 | 7.5-11.7-20 | ||||||||||||||
| Cladi length | 12-15.5-20 | 17.5-21.6-25 | 12-14.5-17.5 | 10-14.4-25 | ||||||||||||||
| Cladi width | 2.5-2.5-2.5 | 2-2.1-2.5 | 2.5-2.5-2.5 | 1-1.7-2.5 | ||||||||||||||
Microscleres. Numerous acanthose microxeas, holotype, 117–166.9–260 mm (Figure
Ecology
Cinachyrella australiensis occurs 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
Distribution
Cinachyrella australiensis has 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,
Distribution of Cinachyrella australiensis. Red dot: type locality, Tethya cranium var. australiensis Carter, 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 raphidiophora Lendenfeld, 1888, Port Jackson, Sidney, Australia 2 Tetilla hirsuta Dendy, 1889, Gulf of Manaar, Sri Lanka 3 Tetilla lindgreni Lendenfeld, 1903, Christmas Island 4 Tetilla poculifera Dendy, 1905, Gulf of Manaar, Sri Lanka 5 Tethya hebes, 1907, at 19° South on the NW coast of Australia 6 Cinachyra isis Lendenfeld, 1907, Mermaid Strait, NW Australia 7 Tetilla cinachyroides Hentschel, 1911, Barrow Island, NW Australia 8 Cinachyra nuda Hentschel, 1912, Aru Island, Indonesia 9 Cinachyra vaccinata Dendy, 1922, Diego Garcia, Chagos Islands 10 Cinachyra providentiae Dendy, 1922, Providence Island, Seychelles 11 Cinachyrella anatriaenilla Fernandez, Kelly, Bell, 2017, American Samoa.
Budding and sediment capture of Cinachyrella species A Three individuals of C. porosa in Haji Buang lake, East Kalimantan, Indonesia, showing distribution of buds beyond the individuals and sediment capture B Close up of C. porosa with detail of buds. Each individual is approximately 4 cm in diameter.
Remarks
In the type description of C. australiensis
Although our focus was on Indonesian species, it was unavoidable to attempt, for the first time after Burton’s review (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
Here we provide further remarks on the following species, in chronologic order
Tetilla lindgreni Lendenfeld, 1903 was described as a new species to separate it from T. ternatensis Kieschnick, 1896, as T. ternatensisis is a Paratetilla based on the presence of calthrop-like spicules. Lendenfeld noticed that both,
Tethya hebes Lendenfeld, 1907 has acanthose microxea and it has most of C. australiensis characters, yet it was excluded from Lendenfeld’s Cinachyrinae-group (with porocalices) because he did not observe porocalices. The type specimens of T. hebes examined at the NHM (
We exclude from C. australiensis some junior synonyms that are part of the schulzei-group species proposed by
We also exclude from C. australiensis two of the junior synonyms still present in the WPD (
In our view, the recently erected species of Cinachyrella anatriaenilla is junior synonym of C. australiensis, because the oxea and the microscleres fall within the size range of the type species of C. australiensis as well as the specimens we have included in this review. The authors distinguish their species from C. australiensis on the basis of having only one category of oxeas versus two categories in C. anatriaenilla. However, we do not recognize size classes in oxea in any of the Cinachyrella specimens and types, but rather a continuos range in size (1000–5500 mm for C. australiensis). The oxea of C. anatriaenilla fall within the size range of the type specimen of C. australiensis as well as the other reviewed material of C. australiensis. In addition, the authors based their statements on the revision of the type specimen of C. kuekenthali, which is from the west Atlantic, but they did not review the type specimen of C. australiensis nor any of the other species with acanthose microxea from the Indo-Pacific.
Recent molecular studies (
Cinachyrella porosa
Spiretta porosa Lendenfeld, 1888: 43 (type seen).
Cinachyra malaccensis Sollas, 1902: 219, pl. XIV, fig. 2; pl. XV, fig. 5. Malacca Strait.
Tetilla porosa ; Lendenfeld, 1903: 22.
Tetilla anomala Dendy, 1905: 91, pl. III, fig.5 (type seen).
Cinachyra albatridens Lendenfeld, 1907: 149, pl. XV, figs 7–9 (type seen).
Cinachyra albaobtusa Lendenfeld, 1907: 154, pl. XVI, figs 45–52 (type seen).
Cinachyra albabidens Lendenfeld, 1907: 151, pl. XVI, figs 39–44 (type seen).
Tethya clavigera Hentschel, 1912: 327, pl. XVI, fig.1, pl. XVIII, fig. 10 In Aru Island, Beach Ngaiboor Trangan.
Cinachyra anomala ; Dendy, 1922: 20, pl. 1, fig. 3 (material seen).
Cinachyra porosa ; de Laubenfels, 1954: 240, pl. XI, fig. b (material seen).
Material examined
Holotype
Description
External morphology. Globular sponges, size from 3 to 5 cm in diameter (Figs
Cinachyrella porosa. Holotype NHMUK1886.8.29.632-633, Port Denison, Australia. A preserved material showing porocalices and internal structure B Labels of the type specimen C skeleton D electron micrograph showing oxea fragments and triaenes rhabds E oxea, end detail F protriaene G prodiaene H, I anatriaenes J sigmaspires. Scale bars: 1 cm (A, C); 500 μm (D); 50 μm (E); 40 μm (F–I); 5 μm (J).
Cinachyrella porosa from Indonesia. A, C, E-I, N,
Skeleton. No cortex. Skeleton composed by bundles of oxeas and triaenes radiating from a central core (Figs
Megascleres. Measurements are shown in Table
Microscleres. No microxeas. Sigmaspires 5–8.6–12.5 mm in the holotype (Figure
Ecology
Occurs in reefs, mangroves, and marine lakes. Predominantly in shallow areas. Notably, a large population inhabit the marine lake of Tanah Bambam, where C. porosa was the dominant representative of moon sponges. This species produces 1–2 mm sized buds (Figure
Distribution
According to the material examined in this revision, we observed that this species is widely distributed in the Indo-Pacific, from the Chagos archipelago, Sri Lanka, Australia, and Tonga Islands. In Indonesia, C. porosa has been collected in East Kalimantan, Java, Ternate, and West Papua.
Distribution of Cinachyrella porosa. Red dot: type locality, Spiretta porosa Lendenfeld, 1888, Port Denison, Queensland, Australia. Green dots: Indonesian localities where the species was collected recently. Circled numbers: type localities of synonymized species 1 Cinachyra malaccensis Sollas, 1902, Malacca Strait, Malaysia 2 Tetilla anomala Dendy, 1905, Chilaw, Sri Lanka 3 Cinachyra albatridens Lendenfeld, 1907, Diego Garcia, Chagos Archipelago 4 Cinachyra albaobtusa Lendenfeld, 1907, Anachoreten (=Keniet) Islands, Papua New Guinea 5 Cinachyra albabidens Lendenfeld, 1907, Tonga Islands 6 Tethya clavigera Hentschel, 1912, Aru Island, Indonesia.
Remarks
Cinachyrella porosa is distinguished from C. australiensis by the absence of acanthose microxea and smaller size of sigmaspires. The first species described with these two diagnostic characteristics was Spiretta porosa Lendenfeld, 1888, subsequently transferred to the genus Tetilla (
Cinachyrella paterifera
Tetilla (Cinachyrella) paterifera Wilson, 1925: 375; plate 39, figs 6, 8; plate 48, fig. 4 (type seen).
Material examined
Holotype USNM21314, South of Tumindao Reef, Tibutu Island, Sibutu Group, Sulu Archipelago, Philippines, 18 m, 27 Feb 1908. INDONESIA. East Kalimantan, Berau reef,
Description
External morphology. Globular sponges, size from 5 to 7 cm in diameter attached to the substrate by a large peduncle/shaft 3 × 2.5 cm (Figure
Cinachyrella paterifera. A, C, E-H, M
Megascleres. The holotype and Indonesian measurements are shown in Table
Microscleres. No microxeas; sigmaspires 10–13.2–17.5 mm in the holotype (Figure
Ecology
The species occurs mainly in reefs, and it is rare in marine lakes and mangroves. It usually inhabits sand bottoms, in which the penduncle serves as a support structure.
Distribution
Indonesia, including East Kalimantan and West Papua. It is also known from Sibutu Island in Philippines (
Remarks
Cinachyrella paterifera has a characteristic elongated peduncle, it is pink to violet colored, and it contains abnormal anatriaenes. Interestingly,
Identification key for Indonesian Paratetilla and Cinachyrella species
| 1 | Porocalices present; calthrops | 2 |
| – | Porocalices present; no calthrops, all triaenes –if present– are long-shafted | 4 |
| 2 | Trichodragmata present | Paratetilla corrugata |
| – | Trichodragmata not present | 3 |
| 3 | High numbers of porocalices, small size (up to 5 mm), brown color | Paratetilla bacca |
| – | Few porocalices, large size (7–15 mm), orange color, fleshy consistency | Paratetilla arcifera |
| 4 | Microxea present | 5 |
| – | Microxea not present | 6 |
| 5 | Acanthose microxea present (115–270 μm); sigmaspires 10–20 μm | Cinachyrella australiensis |
| – | Smooth microxea | Cinachyrella schulzei-group |
| 6 | Small sigmaspires (5–10, few up to 12.5 μm), generally yellow color and ball-shape | Cinachyrella porosa |
| – | Large sigmaspires (10–20 μm), generally pink color, sometimes with peduncle to attach it to the substrate, pear-shape; protriaenes in two different classes; few anatriaenes with reduced and deformed cladi | Cinachyrella paterifera |
Final remarks
Our results contribute to the understanding of the taxonomy and systematics of the Indo-Pacific tetillids. A review of the taxonomic history of the genus Paratetilla and the species Cinachyrella australiensis, showed some cases of misinterpreted synonyms, misidentifications and lack of detailed descriptions for some species. The concept of a single widespread species is refuted for Paratetilla bacca (
The species of Paratetilla and Cinachyrella are clearly highly adaptable and widely distributed sponges. All species in the current study are distributed across Indonesia. It is remarkable that they are all sympatric, some species occuring together in the same marine lake. We have reviewed specimens from East Kalimantan, North Sulawesi, and West Papua. It is highly likely that there are more species in Indonesia in regions that have not been sampled as extensively. Further investigations into Paratetilla and Cinachyrella from the Molluccas, Nusa Tenggara, South Kalimantan, Eastern Papua, and also the virtually unexplored deep sea of Indonesia, will likely lead to the discovery of more species within these genera. Most species occur in all studied habitats (marine lakes, mangroves, and reefs) with a high degree of tolerance for high temperature and sedimentation, as has been observed in other families of sponges (Schönberg 2015). The exceptions to this high tolerance were P. arcifera and C. paterifera, which were only seen in reefs with little sedimentation or sediment resuspension. High budding was observed in specimens of Cinachyrella australiensis and C. porosa residing in marine lakes, while no budding was observed in the same species in the reefs.
Previous molecular phylogenetic studies indicate that P. bacca, P. arcifera, C. porosa, and C. paterifera are distinct monophyletic species, while Cinachyrella australiensis may consist of a species complex with morphologically cryptic species (
Acknowledgements
We greatly appreciate the valuable help and discussions with Dr. NJ de Voogd and Dr. RWM van Soest. We credit Dr. RWM van Soest for thinking of the common name ‘Moon sponges’. Ms. Estradivari and Mr. Bahruddin were indispensable in our quest for Cinderellas.We would like to thank the following people for their help in field logistics and/or lab assistence: Dr. B Hoeksema, Dr. Suharsono, Dr. Y Tuti, Dr. Abdunnur, Mr. M Ammer, Dr. M Erdmann, Ms. E Dondorp, Dr. W Renema, Ms. E Oberhauser, Dr. E Gittenberger, Mr. J van Oyen, Mr. E van Egmond, Mr. R Suhr, Dr. H Breeuwer, and the staff of Papua Diving, TNC/WWF Berau Office, Nabucco Island Dive Resort, Derawan Dive Resort, and at the molecular labs of IBED UvA and Leiden University. Mrs. C Valentine, Mr. A Cabrinovic, Ms. E Sherlock and Dr. T White are acknowledged for making available type specimens at the NHM London, and Dr. K Rützler at the
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Supplementary materials
Table S1. Full collection details of each sample
Figure S1. Type material of Tethya merguiensis