Research Article |
Corresponding author: Renata Manconi ( rmanconi@uniss.it ) Academic editor: Martin Pfannkuchen
© 2016 Renata Manconi, Erica Perino, Roberto Pronzato.
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:
Manconi R, Pronzato R, Perino E (2016) A new species of Agelas from the Zanzibar Archipelago, western Indian Ocean (Porifera, Demospongiae). ZooKeys 553: 1-31. https://doi.org/10.3897/zookeys.553.5999
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A new sponge species (Demospongiae: Agelasida: Agelasidae) is described from the eastern coast of Unguja Island in the Zanzibar Archipelago. Agelas sansibarica sp. n. is compared to all other Agelas species described so far. The new species differs from its congeners mainly in its three categories of verticillate spicules (acanthostyles, acanthostrongyles, and acanthoxeas) and their sizes. Acanthostrongyles, well represented in the spicular complement, are an exclusive trait of the new species widening the morphological range of the genus. Summarizing on spicular complement and spicular morphotraits of 36 species belonging to the genus Agelas: i) 32 species show only acanthostyles from Indo-Pacific (n = 14), Atlantic (n = 17), and Mediterranean (n = 1); ii) three Indo-Pacific species show acanthostyles and acanthoxeas; iii) one species A. sansibarica sp. n. from the western Indian Ocean is characterised by the unique trait of three categories of verticillate spicules (acanthostyles, acanthostrongyles and acanthoxeas). A key for the Indo-Pacific species is supplied together with short descriptions, illustrations, and geographic range; literature on chemical bioprospecting of the genus Agelas is also provided.
Biodiversity, sponges, morpho-taxonomy, diagnostic key, geographic range, Unguja Island
The sponge fauna of the Zanzibar Archipelago is poorly studied and data are reported almost exclusively in very old papers (
The widespread genus Agelas, including until now 35 valid species, was established by Duchassaing and Michelotti (1864: 76) describing the type species Agelas dispar from the Caribbean Sea. Agelas oroides is the only Mediterranean species, and is endemic. The western Atlantic (Gulf of Mexico, Caribbean, and Brazil) hosts 17 species. The majority of the latter (13) were recently revised while the remaining four species were considered dubious or suggested as synonyms (
The Indo-Pacific species of Agelas number 18, including the new species here described. The most widespread species is Agelas mauritiana (including its oxeata variety) recorded in the Australian western Pacific, and the Indian Ocean from the Mascarenes Archipelago (type locality), Seychelles Archipelago, Madagascar, and Mozambique Channel to the southern Red Sea and east to Sri Lanka.
Several species (14) are reported only once from the type locality i.e. Agelas axifera Hentschel, 1911; A. bispiculata Vacelet, Vasseur & Lévi, 1976; A. braekmani Thomas, 1998 (1997); A. carpenteri (Gray, 1867); A. cavernosa Thiele, 1903; A. citrina Gotera & Alcolado, 1987; A. dendromorpha Lévi, 1993; A. inaequalis Pulitzer-Finali, 1986; A. linnaei de Voogd, Parra-Velandia & Van Soest, 2008; A. nakamurai Hoshino, 1985; A. nemoechinata Hoshino, 1985; A. novaecaledoniae Lévi & Lévi, 1983; A. robusta,
In the framework of sponges, applied research on bioactive compounds at a global level (e.g.
The present paper aims to report the discovery of a new species of Agelas from the Zanzibar Archipelago comparing it to all species belonging to the genus. To support global sharing of information on faunistics and taxonomy of Porifera from not widely accessible data sources an updated overview on the morphology and geographic distribution of Agelas species from the Indo-Pacific area is also provided together with a brief description and exhaustive iconography, as well as a dichotomous key to Indo-Pacific species.
Representative fragments of six sponge specimens from the Unguja Island were studied. Growth form, surface traits, skeletal architecture, shape and size of the spicules are considered diagnostic morphotraits. Spicule dimensions are given as maximum, minimum, and medium lengths and widths of ca. 400 spicules.
The skeletal architecture was investigated by hand-cut sections of the ectosome and choanosome. To study the three-dimensional arrangements of fibres and spicules in the skeleton, fragments ca. 10 × 10 × 2 mm were cut, cleaned with 5% sodium hypochlorite (NaClO) for 24 h in a warm temperature (35–40 °C), then washed and stirred five times in abundant double distilled water, washed and stirred twice in ethanol 95%, and finally allowed to air dry and gold-sputtered or mounted in Eukitt. The skeletal samples were than studied by light microscope (LM) and scanning electron microscope (SEM). Spicule preparations were made by dissolving a small fragment of the specimen in 65% boiling nitric acid (HNO3)and cleaned spicules were rinsed four times with water, once with 95% ethanol. The spicules were air-dried on slides, mounted in Eukitt, and observed by a Leitz Dialux 20 EB (LM), as well as on aluminium stubs and coated with gold for the observation with a Vega3 TESCAN type LMU (SEM).
Specimens were deposited at the Museo civico di Storia Naturale “Giacomo Doria” of Genoa, Italy (acronym
Phylum Porifera Grant, 1835
Class Demospongiae Sollas, 1885
Order Agelasida Hartman, 1980
Family Agelasidae Verrill, 1907
Genus Agelas Duchassaing & Michelotti, 1864
Chalinopsis Schmidt, 1870 (junior synonym)
Ectyon Gray, 1867 (junior synonym)
Oroidea Gray, 1867 (junior synonym)
Pachychalinopsis Schmidt, 1880 (nomen nudum)
Siphonochalinopsis Schmidt, 1880 (nomen nudum)
Diagnosis (emended from
Holotype:
Agelas with unique spicular complement composed of three spicular categories, oxeas, styles and strongyles with spines arranged in a variable number of verticilles.
The speciphic epithet refers to the Zanzibar Archipelago.
Coral reef, quite common at 7–12 m of depth. Water temperature 28–31 °C. Salinity 20–36‰ (
Western Indian Ocean, but only recorded from the type locality to date.
Growth form massive, thick, rounded, 6–10 cm in diameter. Colour in life purple to orange and light brown. Consistency firm and elastic. Surface rough to the touch, finely hispid, finely conulose for tips of ascending fibres supporting the dermal membrane, with regularly scattered circular and convoluted depressions (0.5 cm in diameter) very similar to those of A. cerebrum. Oscules few, small, irregularly scattered. Choanosomal skeleton as an irregularly reticulate network of spongin fibres. Primary fibres 50–110 (71.67 ±17.63) μm in diameter, strongly echinate by single, scattered spicules to groups of diverging spicules; ascending primary fibres cored by spicules also present. Secondary fibres 20–50 (35 ± 9) μm in diameter notably echinate and cored by spicules. Tertiary network not observed.
Megascleres as three categories of monaxons mostly with acute spines. Acanthostyles 90–250 (180.72 ± 28.66) × 7.5–20 (13.46 ± 2.59) μm ornate by verticillate spines arranged as 11–27 (17.8 ± 2.86) whorls. Acanthoxeas 130–295 (195 ± 43.09) × 7.5–15 (12.17 ± 1.89) μm ornate by verticillate spines arranged as 14–26 (19.24 ± 3.47) whorls. Acanthostrongyles 80–245 (148.18 ± 36.82) × 4–17 (11.09 ± 4.24) μm ornate by verticillate spines arranged as 9–26 (15.76 ± 3.85) whorls. Annulate spicules apparently young.
The new species is characterized by the co-presence of three categories of spicules never recorded in other Agelas species. Acanthostrongyles are abundant, ca. 20–30 % of the total number of spicules.
Madagascar, Mozambique Channel, Seychelles and Mascarene archipelagos (Western Indian Ocean province) harbour four species, whereas Japan (Ryukyu Archipelago) and New Caledonia enumerates two species each. Only one species each is recorded from Philippines, Papua New Guinea, and Funafuti. Only one species each is harboured in the Red Sea/Gulf of Aden, Sri Lanka, Moluccas, Sunda Shelf/Java Sea (Indonesia), Hong Kong, Funafuti, and Australia (Fig.
To discriminate between all 36 Agelas species by diverging diagnostic morphotraits is notably difficult, as highlighted in the previous section. Morphotraits of the genus are extremely conservative and different species appear very similar. Focusing on the Indo-Pacific species our attempt was not as completely successful as is also the case for the Atlantic species by
Atlanto-Mediterranean Agelas species (18) seems to possess only acanthostrongyles, including the uncertain A. fascicularis, A. flabelliformis, A. inaequalis, and A. rudis not redescribed by
Among the 17 previously known Indo-Pacific Agelas species, the spicular complement of 14 species is composed of acanthostyles in a single or two-dimensional classes (see
Agelas ceylonica. a very low quality image of the specimen studied by
Agelas mauritiana. a drawing of a massive specimen b skeleton fragment c–g spicular complement b–dA. mauritiana var. oxeataa modified from
The most common and studied Indo-Pacific species, i.e. A. mauritiana, are characterized by a single spicular type acanthostyles, which are extremely variable in morphology, abundance of spines, and dimensional range (sometime more than three times in length) (see Table
References | Acanthostyles μm | Whorls nº | Colour | Habitus size (cm) | Consistency |
---|---|---|---|---|---|
|
132 | 15–18 | - | - | - |
|
200 × 14–15 | 16 | - | - | - |
|
176 × 16 | – | dark brown | tubular 3.1 × 1.6 length × diameter |
firm resilient |
|
170–180 × 10–14 | 12–18 | - | - | - |
|
150–160 × 8–12 | 16–20 | brown | massive 6–10 × 4 |
firm resilient soft |
|
275 × 12–13 | 15–17 | - | - | - |
|
140–230 × 8–10 | 10–15 | - | - | - |
|
135–250 | 13–20 | - | - | - |
|
80–180 × 7.5–12.5 | 18–23 | brown | encrusting 1.6–1.5 thickness |
firm resilient coriaceus |
|
112–212 × 6–8 | 6–8 | pale yellow | encrusting cavernous |
firm resilient compressible |
The new species A. sansibarica is characterized by the unique morphotrait of three categories of megascleres, i.e. acanthostyles, acanthoxeas, and acanthostrongyles with spines arranged in verticilles regularly scattered along the entire spicule. No other Agelas species exhibit this spicular combination. Acanthostrongyles, well identifiable and abundant, represent an exclusive diagnostic trait of the new species. The functional role of acanthostrongyles is doubled since echinanting spicules arm both the fibres surface and the core of the axial part of fibres.
Summarizing spicular complements and spicular morphotraits of 36 Agelas species: i) 32 species show only acanthostyles from Atlantic (17), Mediterranean (1), and the majority (14) of the Indo-Pacific areas; ii) three Indo-Pacific species show acanthostyles and acanthoxeas; iii) only one species A. sansibarica sp. n. from the western Indian Ocean show a spicular component composed of acanthostyles, acanthoxeas, and acanthostrongyles.
The present key is an attempt to discriminate between the Indo-Pacific species, but the scenario appears very intricate mainly because morphotraits from many descriptions and illustrations are overlapping (see Table
1 | Spicular complement composed by 1 or 2 spicular types (acanthoxeas, acanthostyles) | 2 |
– | Spicular complement composed by 3 spicular types (acanthoxeas, acanthostyles, acanthostrongyles) | A. sansibarica sp. n. |
2 | Spicular complement composed by 2 spicular types (acanthoxeas and acanthostyles) | 3 |
– | Spicular complement composed by 1 spicular type (acanthostyles) | 4 |
3 | Sponge body cup-shaped | A. axifera |
– | Sponge body blade-shaped | A. novaecaledoniae |
– | Sponge body lobed | A. mauritiana oxeata |
– | Sponge body digitate | A. dendromorpha |
– | Sponge body as slim cylindrical erected axis (branched or unbranched) | A. gracilis |
4 | Acanthostyles of 2-dimensional categories | 5 |
– | Acanthostyles of 1-dimensional category | 7 |
5 | Long acanthostyles spiny only at the tips | A. semiglabra |
– | Long and short acanthostyles almost entirely spiny | A. bispiculata |
7 | Primary and secondary fibres uncored | 8 |
– | Primary and/or secondary fibres cored | 9 |
8 | Acanthostyles longer than 300 μm | A. ceylonica |
– | Acanthostyles length no more than 200 μm | A. cavernosa |
9 | Primary and secondary fibres cored | A. nemoechinata |
– | Primary fibres cored and secondary uncored | 10 |
10 | Acanthostyles (185–265 × 8–15 μm) with 15–23 whorls | A. nakamurai |
– | Acanthostyles (130–220 × 4–21 μm) with 8–18 whorls | A. braekmani |
– | Acanthostyles (80–370 × 5–24 μm) with 11–33 whorls | A. linnaei |
We acknowledge financial support from EU–7FP, BAMMBO (Sustainable Production of Biologically Active Molecules of Marine Based Origin contract n. 265896), Regione Autonoma Sardegna (RAS2012-CRP60215 “Conservazione e valorizzazione delle grotte sarde: biodiversità e ruolo socio-economico-culturale”), Ministero dell’Istruzione, dell’Università e della Ricerca (MIUR–PRIN), and Fondazione Banco di Sardegna. We are grateful to the Swiss Marine NGO for specimen supply. We thank Nicole de Voogd and Zoölogische Mededelingen Leiden for Fig.
Indo-Pacific species belonging to the genus Agelas reported from the literature.
Fig.
Description (modified from
Fig.
Description (modified from
Fig.
Description (modified from
Fig.
Halichondria (?) Carpenter, 1856 in
Ectyon carpenteri Gray, 1867
Description (modified from
Fig.
Description (modified from
Fig.
Description (modified from
Fig.
Description (modified from
Fig.
Description (modified from
Fig.
Description (modified from
Fig.
Description (modified from
Fig.
Ectyon mauritiana Carter, 1883
Description (modified from
Fig.
Description (modified from
Fig.
Description (modified from
Fig.
Description (modified from
Consistency spongy, resilient when wet, hard and difficult to compress when dry. Colour madder brown at the surface to rufous-orange in the interior when dry. Oscules from 1–3 to 5–8 mm in diameter. Ectosomal skeleton not reported. Choanosomal skeleton reticulate network with elliptical meshes (50–250 μm in diameter) of spongin fibres. Primary and secondary fibres almost indistinguishable (20–50 μm in diameter) and cored by 1–3 acanthostyles, occasionally not cored, only slightly echinated. Megascleres verticillate of a single category. Acanthostyles 170–(189)–210 × 9–(11)–13 μm, straight to gently curved, ornate by 16–23 regular whorls, each with eight spines. Habitat. Shallow water, 20 m depth. Geographic distribution. North Pacific Ocean. Recorded from south Kuroshio (east Japan). Type locality: Zamami Island, Ryukyu Archipelago. Remarks. Also recorded from Indonesia (
Fig.
Description (modified from
Fig.
Description (modified from
Fig.
Agelas semiglaber Pulitzer-Finali, 1996
Description (modified from
Several findings from the Seychelles Archipelago (