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Zookeys
2015 Aug 12;517:1-37. doi: 10.3897/zookeys.517.9308.
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Scleractinian corals (Fungiidae, Agariciidae and Euphylliidae) of Pulau Layang-Layang, Spratly Islands, with a note on Pavonamaldivensis (Gardiner, 1905).
Waheed Z
,
Benzoni F
,
van der Meij SE
,
Terraneo TI
,
Hoeksema BW
.
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Layang-Layang is a small island part of an oceanic atoll in the Spratly Islands off Sabah, Malaysia. As the reef coral fauna in this part of the South China Sea is poorly known, a survey was carried out in 2013 to study the species composition of the scleractinian coral families Fungiidae, Agariciidae and Euphylliidae. A total of 56 species was recorded. The addition of three previously reported coral species brings the total to 59, consisting of 32 Fungiidae, 22 Agariciidae, and five Euphylliidae. Of these, 32 species are new records for Layang-Layang, which include five rarely reported species, i.e., the fungiids Lithophyllonranjithi, Podabaciasinai, Sandalolithaboucheti, and the agariciids Leptoseriskalayaanensis and Leptoseristroglodyta. The coral fauna of Layang-Layang is poor compared to other areas in Sabah, which may be related to its recovery from a crown-of-thorns seastar outbreak in 2010, and its low habitat diversity, which is dominated by reef slopes consisting of steep outer walls. Based on integrative molecular and morphological analyses, a Pavona variety with small and extremely thin coralla was revealed as Pavonamaldivensis. Since specimens from Sabah previously identified as Pavonamaldivensis were found to belong to Pavonaexplanulata, the affinities and distinctions of Pavonamaldivensis and Pavonaexplanulata are discussed.
Figure 1. Research area at Pulau Layang-Layang, Spratly Islands. Survey sites are numbered 1-18, and HR for the House Reef. Depth contours are indicated in metres (smaller font). For a complete georeferenced list of localities and depths of survey sites, see Table 1.
Figure 2. Fungiidae recorded at Pulau Layang-Layang in this study. a
Ctenactis
albitentaculata
b
Cycloseris
boschmai
c
Cycloseris
costulata
d
Cycloseris
explanulata
e
Cycloseris
mokai
f
Cycloseris
tenuis
g
Danafungia
horrida
h
Danafungia
scruposa.
Figure 3. Fungiidae recorded at Pulau Layang-Layang in this study. a
Fungia
fungites b Halomitra
pileus
c
Herpolitha
limax
d
Lithophyllon
ranjithi
e
Lithophyllon
repanda
f
Lithophyllon
scabra
g
Lithophyllon
undulatum
h
Lobactis
scutaria.
Figure 4. Fungiidae recorded at Pulau Layang-Layang in this study. a
Pleuractis
granulosa
b
Pleuractis
gravis
c
Pleuractis
moluccensis
d
Podabacia
sinai
e
Polyphyllia
talpina
f
Sandalolitha
boucheti
g
Sandalolitha
dentata
h
Sandalolitha
robusta.
Figure 5. Agariciidae recorded at Pulau Layang-Layang in this study. a
Coeloseris
mayeri
b
Gardineroseris
planulata
c
Leptoseris
foliosa
d
Leptoseris
glabra
e
Leptoseris
hawaiiensis
f
Leptoseris
incrustans
g
Leptoseris
kalayaanensis
h
Leptoseris
mycetoseroides.
Figure 6. Agariciidae recorded at Pulau Layang-Layang in this study. a
Leptoseris
scabra
b
Leptoseris
solida
c
Leptoseris
troglodyta
d
Leptoseris
yabei
e
Pachyseris
rugosa
f
Pacyhseris
speciosa
g
Pavona
bipartita
h
Pavona
clavus.
Figure 7. Agariciidae recorded at Pulau Layang-Layang in this study. a
Pavona
duerdeni
b
Pavona
explanulata
c
Pavona
maldivensis (registration no. IPMB-C 13.00007) d
Pavona
minuta
e
Pavona
varians
f
Pavona
venosa.
Figure 8. Euphylliidae recorded at Pulau Layang-Layang in this study. a
Euphyllia
glabrescens
b
Euphyllia
paraancora
c
Physogyra
lichtensteini.
Figure 9. Undetermined Leptoseris coral species recorded at Pulau Layang-Layang in this study a
Leptoseris sp. 1 (registration no. IPMB-C 13.00009) b
Leptoseris sp. 2 c
Leptoseris sp. 3.
Figure 10. Maximum likelihood phylogram of Pavona
maldivensis and Pavona
explanulata based on combined mitochondrial intergenic spacer between CO1 and 16S-rRna and nuclear marker ITS1-5.8S-ITS2 sequences. Support values for maximum likelihood, maximum parsimony (>70) and bayesian posterior probabilities (>80) are given at the nodes. Dashes (-) indicate nodes without statistical support. Letters in parentheses correspond to images a–l in rows L and R. L: coral colonies in situ, R: SEM images of calices. Locality for the coral samples are: LAC = Layang-Layang, LOY = Loyalty Islands, MAL = the Maldives, BAN = Banggi, East Malaysia, TER = Ternate, Indonesia, RED = Redang, Peninsular Malaysia. a–b
Pavona
maldivensis from Layang-Layang c–d
Pavona
maldivensis from Loyalty Islands e–f
Pavona
maldivensis from the Maldives g–h
Pavona
explanulata from Banggi, Sabah i–j
Pavona
explanulata from Banggi, Sabah k–l
Pavona
explanulata from Ternate, Indonesia. Additional samples KF437780
Pavona
maldivensis and KF437762
Pavona
cf.
explanulata are from Pearl and Hermes Atoll, the northwest Hawaiian Islands and O’ahu, Hawaii, respectively (Luck et al. 2013). Outgroup LK934507, HE978501
Leptoseris
foliosa is from Prony Bay, New Caledonia (Benzoni et al. 2012b, Terraneo et al. 2014).
Figure 11. Specimen RED05 Pavona
cf.
explanulata from Pulau Redang, Peninsular Malaysia. a coral colony in situ at 13 m depth b corallites detail at the plate margin c coral fragment preserved in ethanol d, e SEM images showing calices.
Figure 12. Variation of radial elements in Pavona
maldivensis (a, c) and Pavona
explanulata (b, d). a specimen LAC23 from Layang-Layang, b specimen BAN02 from Banggi, North Borneo, c specimen LOY12 from Loyalty Islands, and d specimen TER28 from Ternate, Indonesia. Lateral faces of septa are covered with granules (g) as indicated in a and d. Aligned granulations along the lateral faces of septa (menianae) are indicated by white arrows in b, c, d.
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