BenzoniFrancescaEchinophyllia tarae sp. n. (Cnidaria, Anthozoa, Scleractinia), a new reef coral species from the Gambier Islands, French PolynesiaZookeys24720132013318597910.3897/zookeys.318.5351 Echinophyllia tarae urn:lsid:zoobank.org:act:9C41F53B-3FEE-47BA-8017-48C252F65F5E http://species-id.net/wiki/Echinophyllia_tarae sp. n.Figures 2–8, 9a, b, 10b, dMaterial examined.

Holotype: MNHN-IK.2012-8000 (Figures 24). Type Locality: Taravai Island, Gambier, French Polynesia (MV Tara, Tara Oceans Expedition, Site 9), 23°9.404'N, 135°1.769'E, 10 m, 30 June 2011, coll. F. Benzoni.

Holotype of Echinophyllia tarae sp. n. (MNHN-IK.2012–8000) a the colony in situ before collection, and b the corallum after removal of the animal tissues. C1 to 4 indicate the position of same corallite (C) in the two images. Numbers are assigned in decreasing order of corallite size, C1 being the largest. Stippled line on the specimen in b shows the boundary of living tissue at the time of collection.

Details of corallites, septa, and costae in the holotype of Echinophyllia tarae sp. n. (MNHN-IK.2012–8000) a top view of the largest corallite in the colony, b lateral view of the same corallite shown in a, c top view of the second largest corallite, and d of the third e top and f side view of the costae. C1 to 3 indicate the corallites as shown in Figure 2. White arrows in e indicate the position of exothecal alveoli at the insertion of costae.

Specimens of Echinophyllia tarae sp. n. collected for this study a side view of the holotype (MNHN-IK.2012–8000) b specimen UNIMIB TO-GA028 c specimen UNIMIB TO-GA186 d specimen UNIMIB TO-GA099 e specimen UNIMIB TO-GA071. C1 to 3 indicate the holotype corallites as shown in Figures 1 and 2.

Corallum: The holotype is a knob-shaped, plocoid, encrusting colony attached to a fragment of a dead tabular Acropora coral (Figure 2). The specimen is 9.2 cm high, and 8.5 x 5 cm wide at the base in its original growth position.

Corallites: The 12 corallites are oval in shape and variable in size (Figure 2, 3), ranging from 3.3 cm in diameter (C1 in Figure 2, 3a) to 1.0 cm. Corallites are organically united (see terminology in Budd et al. 2012). The central position of C1 (the largest corallite) is less obvious in the holotypes due to its knob-shaped growth form. Corallites protrude up to 1 cm and are directed in different directions (Figure 2). Corallite wall is septothecal.

Costosepta: Variable in number depending on the size of the corallite (Figure 3a–d), exsert and thickened over the theca. The largest corallite contains 26 septa arranged in five orders (Figure 3a). Septa of the first three orders are thicker than the others. Septal teeth are elliptical in outline, large and high (> 0.6 mm) according to the parameters defined by Budd et al. (2012) (Figure 3a–d), and their tips are irregular bulbous. Tooth spacing is very high (> 2 mm). Septal side granulation is weak. Paliform lobes (see Benzoni et al. 2011 for a definition) thick and well developed, forming an obvious crown around the columella, which was also visible in vivo. Paliform lobes always present and of similar size at the proximal margin of the first two orders of septa (Figures 3a–d). In larger corallites, in which more than four orders are present, they can also form in front of the third order but then they are of smaller dimensions (Figure 3b). The costal parts of the costosepta are thick and unequal. They are strongly ornamented by triangular shaped teeth (Figure 3e, f), which bear fine granules on the tip (Figure 3f). Exothecal alveoli are present at the insertion of lower order costae (Figure 3e). Costae cover the whole surface of the coenosteum between corallites.

Columella: Well developed, deep in the fossa (Figure 3b) made by a mesh of twisted intermingled processes derived from the inner end of the higher order septa: the first two in smaller corallites (Figure 3a–d), and up to the fourth order in larger corallites (Figure 10b–d).

Colour: The living colony was mottled brown. Tips of septa and costae ornamentation varied from light beige to white.

Other material

(Gambier, French Polynesia, Tara Oceans Expedition): UNIMIB TO-GA028, Akamaru Island (Site 2), 23°11.082'N, 134°54.331'E, 26 June 2011, coll. F. Benzoni; UNIMIB TO-GA071 Makaroa Island (Site 6), 23°12.960'N, 134°57.991'E, 28 June 2011, coll. F. Benzoni; UNIMIB TO-GA099 Taravai Island (Site 11), 23°9.540'N, 135°3.055'E, 1 July 2011, coll. F. Benzoni; UNIMIB TO-GA186 Taravai Island (Site 9), 23°9.404'N, 135°1.769'E, 30 June 2011, coll F. Benzoni.

Variation of skeletal structures: Colony size is relatively small (Figures 28), the largest colony is 20 cm wide (Figure 6f). Corallum generally encrusting, its plane following the surface of the underlying substrate (Figures 4b, d, e) but also knob-like (Figures 4a, c) with foliose margins where they become detached from the substrate. The number of corallites per colony is low, ranging from 1 to 15. A large, central primary corallite is always present (Figures 2, 6a, b). Secondary corallites are often inclined in various directions and show variable diameter sizes within the same colony (Figures 5a–e, Table 1). The largest corallite observed (specimen UNIMIB TO-GA099, Figure 4d) is 3.5 cm in diameter, the smallest, in the same specimen, 0.9 mm (Figure 5e). The numbers of septa, orders of septa, and paliform lobes vary between and within colonies. First order septa always thicker than the others and in some cases up to 4 mm thick (Figure 5f). Columella always present, large and oval in the largest corallite, less developed is smaller corallites. Costae typically thick, alternating in size (Figure 5c, d) and strongly dentate, although variably so between specimens.

Variation in shape and size of the corallites of Echinophyllia tarae sp. n. a and b two lateral views of specimen UNIMIB TO-GA186 c specimen UNIMIB TO-GA071 d UNIMIB TO-GA028 e specimen UNIMIB TO-GA099 f close up-of a corallite of the same specimen as in a and b. White arrows in e indicate the position of secondary corallites.

Variation of corallum shape, costosepta ornamentation, and colouration in smaller colonies of Echinophyllia tarae sp. n. observed in situ a young one polyp brown coloured specimen settled on crustose coralline algae, Mangareva Island (Site 15) b a dark brown two polyp colony with green oral discs, Taravai Island (Site 21) c one polyp bright green specimen with a large raised central corallite in which the crown of pali is clearly visible and spiky and well developed costae, Makaroa Island (Site 6) d a light green colony with two raised polyps in central position, well developed crown of pali and costae ornamentation smoothening towards the colony periphery, northern lagoon pinnacles (Site 16) e brown colony with one raised polyps in central position and markedly teethed costae, Mangareva Island (Site 13) f in situ image of specimen UNIMIB TO-GA028 (Figures 4b, 5d) showing the irregularly shaped colony with a very rugged and almost inflated appearance, Akamaru Island (Site 2). Sites are indicated in Figure 1.

Dimensions of the examined specimens of Echinophyllia tarae sp. n. For each specimen the total number (n.) of corallites, the maximum and minimum diameter of the largest and smallest corallite in the colony, the number of septa, and of paliform lobes is listed. LC = largest corallite; SC smallest corallite.

Specimen coden. of corallitesDiam. LC max-min: (cm)Diam. SC max-min: (cm)n. of septa LCn. of paliform lobes LC
MNHN-IK.2012–8000123.3 – 2.01.0 – 0.92611
UNIMIB TO-GA02843.1 – 2.41.1 – 0.93914
UNIMIB TO-GA07142.3 – 2.11.0 – 0.9348
UNIMIB TO-GA08443.2 – 2.41.9 – 1.82911
UNIMIB TO-GA09933.5 – 3.01.2 – 1.05224

Field characteristics and colouration: The colouration is showing much variation, ranging from dark brown (Figures 7a–c, e), to mottled brown (Figures 2a, 7f), beige (Figures 6e, f), and bright green (Figures 6c, 7d). The tissue on the tips of septal teeth and costal spines teeth can be lighter in colour (Figure 6c) or white (Figure 7c), possibly as a result of tissue being less thick above these structures. In very mottled colonies (Figure 7f) or with lighter colouration of the tissue over costoseptal teeth (Figure 7b) the size and shape of the corallites may be hard to detect. The crown of paliform lobes is always prominent (Figure 5) and often obvious, especially in the largest corallite (Figures 6, 7b).

Variation of shape, spikiness of septa and costae, and colouration oflarge colonies Echinophyllia tarae sp. n. observed in situ a brown encrusting colony with free margins, bright green oral discs and raised corallites, Akamaru Island (Site 2) b brown encrusting colony with white oral discs, raised corallites (larger one in the stippled circle), and very spiky costae, Taravai Island (Site 9 – type locality), the prominent crown of paliform lobes of the largest corallite is indicated by the white arrow c brown knob shaped colony with bright green oral discs and raised corallites, note the white colouration of the tips of costae teeth, Taravai Island (Site 9 – type locality) d a bright green knob shaped colony, Taravai Island (Site 9 – type locality) e brown encrusting colony with bright green oral discs and relatively low-lying corallites, note the uniform colouration of the costae, Taravai Island (Site 9 – type locality) f mottled brown encrusting colony with free margins and relatively low-lying corallites, note the uniform colouration of the costae, Taravai Island (Site 9 – type locality). Sites are indicated in Figure 1.

Frequently observed patterns of partial death and re-growth of Echinophyllia tarae sp. n. in the fielda concave colony with a large central corallite showing a peripheral rim of skeleton encrusted by pink coralline algae and surrounded by zoanthids and corallimorpharians, Mangareva Island (Site 19) b a similar situation as in a but with re-growth occurring over previously dead colonies, note the variation of colouration in adjacent corals, Taravai Island (Site 11). White arrows in b indicate the position of larger central corallites. Sites are indicated in Figure 1.

Ecology: Echinophyllia tarae sp. n. inhabits protected reef habitats and was observed between 5 and 20 m depth. It commonly grows on dead coral fragments, usually parts of branching or tabular Acropora colonies, which are covered by crustose coralline algae and fleshy macroalgae (Figures 67). This species can grow on well-illuminated surfaces but also encrusts shaded underhangs. In well-lit conditions the appearance is typically corrugated (Figures 6a–b, e–f, 7a–d). However, in some cases a certain degree of inflation of the soft tissues was observed (Figures 6a–b, f), although this generally depends on the very developed ornamentation of the underlying costosepta, which is most obvious when a live colony (Figure 6f) is compared with the clean skeleton (Figures 4b, 5d). In poorly lit conditions the overall appearance is smoother and the colouration more uniform (e.g. Figures 6d, 7e–f) although the oral discs remain generally brightly coloured and different from the rest of the tissues. Re-growth of partially dead colonies, especially at the margins, is common (Figure 8). Such patters of partial death and recovery could result from competition with other benthic invertebrates, like soft-bodied corallimorpharians and zoanthids which can co-occurr with this species (Figure 8a). The observed patterns of partial death may also be caused by deposition of sediment on the living corals. In fact, Echinophyllia tarae sp. n. is most commonly found at sheltered sites characterized by calm water conditions and muddy sediment which could be stirred up and deposit on benthic organisms suffocating them (Erftemeijer et al. 2012). The dead parts of the corallum are generally encrusted by coralline algae over which the coral can re-grow or re-settle (Figure 8b).

Occurrence:This species was commonly encountered on the fringing reefs off Mangareva, Aukena, Tekava, Akamaru, Kamaka, Makaroa, Agakauitai, and Taravai islands as well as at the base of the lagoon pinnacles found in the lagoon north of Mangareva Island (Figure 1, Table 2). Its distribution outside the Gambier archipelago is unknown although it could also occur in the Austral Islands (see Discussion section). No record is known from other localities.

Affinities:In its encrusting growth form, and in the presence of a central larger and protruding corallite this species is similar to Echinophyllia echinata and Echinomorpha nishihirai (see Discussion). Ongoing molecular analyses will reveal the phylogenetic relationships of this species with its congeners.

Occurrence of Echinophyllia tarae sp. n. encountered during the Tara Oceans Expedition in the Gambier Islands. For each site the site code (as in Figure 1), island name and reef type, coordinates, sampling date, and recorded presence or absence of the species are listed.

SiteIsland, reef typeLatitude, LongitudeDateEchinophyllia tarae sp. n.
1Akamaru, fringing reef23°10.61'S, 134°54.37'W26/06/11recorded
2Akamaru, fringing reef23°11.08'S, 134°55.33'W26/06/11recorded
3Aukena, fringing reef23°08.44'S, 134°55.18'W27/06/11recorded
4Aukena, fringing reef23°08.56'S, 134°54.74'W27/06/11recorded
5Kamaka, fringing reef23°14.19'S, 134°57.47'W28/06/11recorded
6Makaroa, fringing reef23°12.96'S, 134°57.99'W28/06/11recorded
7Kamaka, fringing reef23°14.98'S, 134°57.80'W29/06/11not recorded
8Makaroa, fringing reef23°13.32'S, 134°58.34'W29/06/11recorded
9Taravai, fringing reef23°09.40'S, 135°01.77'W30/06/11recorded
10Agakauitai, fringing reef23°10.17'S, 135°02.52'W30/06/11recorded
11Taravai, fringing reef23°09.54'S, 135°03.05'W01/07/11recorded
12Agakauitai, fringing reef23°10.35'S, 135°01.99'W01/07/11recorded
13Mangareva, fringing reef23°08.25'S, 134°57.11'W02/07/11recorded
14Mangareva/Totegie23°04.78'S, 134°54.99'W04/07/11not recorded
15Mangareva, fringing reef23°05.45'S, 134°55.69'W04/07/11recorded
16Lagoon pinnacles23°01.55'S, 134°55.69'W05/07/11recorded
17Lagoon pinnacles23°04.12'S, 134°55.83'W05/07/11not recorded
18Mangareva, fringing reef23°05.57'S, 134°59.16'W07/07/11not recorded
19Mangareva, fringing reef23°06.14'S, 134°59.23'W07/07/11recorded
20Taravai, fringing reef23°08.72'S, 135°03.09'W08/07/11recorded
21Taravai, fringing reef23°08.03'S, 135°02.14'W08/07/11recorded
22outer barrier north23°04.21'S, 135°00.85'W09/07/11not recorded
23outer barrier north23°03.79'S, 135°00.49'W09/07/11not recorded
24Tekava, fringing reef23°10.13'S, 134°51.51'W10/07/11recorded
Etymology.

This species is named after MV Tara, which allowed the exploration of coral reefs in Gambier. Moreover, the name “tara” in the Polynesian language may refer to a spiny, pointed object, which applies well to the new species typically featuring pointed skeletal structures. In the same language, Tara is also the name of a sea goddess.

Map of a the Pacific Ocean, showing the position of French Polynesia b the island groups belonging to French Polynesia, and c the study area in the Gambier Archipelago showing the 24 sampling sites of the Tara Oceans leg and the sites where Echinophyllia tarae sp. n. was found. Stippled rectangle in a shows the position of the map shown in b. Arrow in b shows the position of the Gambier Archipelago shown in c.

Comparison of in situ morphology between specimens of Echinophyllia tarae sp. n., Echinophyllia echinata, and Echinomorpha nishihirai and corallum morphology of the latter two a Echinophyllia tarae sp. n. resembling Echinophyllia echinata, Agakauitai Island (Site 10) b another colony of the new species resembling Echinomorpha nishihirai, Taravai Island (Site 11) c Echinophyllia echinata from Cap Bocage, New Caledonia (IRD HS 3171) d Echinomorpha nishihirai, Ryukyu Islands, Japan, picture by K. Yanagiya e same specimen as in c, f holotype of Echinomorpha nishihirai (MTQ G 32483), Okinawa Island, Japan. Images c and e from the IRD LagPlon database (http://lagplon.ird.nc/consultv2_5/rechSimple.faces). Sites are indicated in Figure 1.

Comparison ofcentralcorallitemorphology between a Echinophyllia echinata b and d Echinophyllia tarae sp. n., and c Echinomorpha nishihirai: a side view of the central corallite of thesame Echinophyllia echinata specimen as in Figure 8c (IRD HS 3171) b side view of the central protocorallite of Echinophyllia tarae sp. n. (UNIMIB TO-GA099), c top view of the central corallite of Echinomorpha nishihirai holotype (MTQ G 32483) d top view of the same corallite as in b.

BuddAFFukamiHSmithNDKnowltonN (2012) Taxonomic classification of the reef coral family Mussidae (Cnidaria: Anthozoa: Scleractinia).Zoological Journal of the Linnean Society166: 465-529. doi: 10.1111/j.1096-3642.2012.00855.xBenzoniFArrigoniRStefaniFPichonM (2011) Phylogeny of the coral genus Plesiastrea (Cnidaria, Scleractinia).Contributions to Zoology80: 231-249.ErftemeijerPLARieglBHoeksemaBWToddPA (2012) Environmental impacts of dredging and other sediment disturbances on corals: A review.Marine Pollution Bulletin64: 173765. doi: 10.1016/j.marpolbul.2012.05.008