Preliminary study on Acropora - Scientific Journals - Tübitak
Preliminary study on Acropora - Scientific Journals - Tübitak
Preliminary study on Acropora - Scientific Journals - Tübitak
You also want an ePaper? Increase the reach of your titles
YUMPU automatically turns print PDFs into web optimized ePapers that Google loves.
* Corresp<strong>on</strong>dence: rahmani@coe.ac.ir<br />
308<br />
Turkish Journal of Zoology<br />
http://journals.tubitak.gov.tr/zoology/<br />
Research Article<br />
Turk J Zool<br />
(2013) 37: 308-320<br />
© TÜBİTAK<br />
doi:10.3906/zoo-1112-21<br />
<str<strong>on</strong>g>Preliminary</str<strong>on</strong>g> <str<strong>on</strong>g>study</str<strong>on</strong>g> <strong>on</strong> <strong>Acropora</strong> (Scleractinia: Astrocoeniina: Acroporidae)<br />
of the Persian Gulf, with emphasis <strong>on</strong> the north and northeastern areas<br />
Mohammadreza RAHMANI 1,2, *, Hassan RAHIMIAN 1<br />
1 Faculty of Biology, College of Science, University of Tehran, Tehran, Iran<br />
2 University of Envir<strong>on</strong>ment, Standard Square, Karaj, Iran<br />
Received: 16.12.2011 Accepted: 02.10.2012 Published Online: 29.04.2013 Printed: 29.05.2013<br />
Abstract: Due to a lack of adequate tax<strong>on</strong>omic informati<strong>on</strong> <strong>on</strong> <strong>Acropora</strong> Oken, 1815 species in the Persian Gulf, especially those<br />
inhabiting the coasts of the northern and northeastern areas, an appraisal of the diversity of this genus required comprehensive and<br />
careful examinati<strong>on</strong> of a large number of specimens. To satisfy such a need, 10–15 cm pieces of more than 1100 col<strong>on</strong>ies of staghorn<br />
corals were collected from the subtidal z<strong>on</strong>es of different islands. The specimens were studied using both light stereo and scanning<br />
electr<strong>on</strong> microscopes and were identified with regi<strong>on</strong>al and global keys. During the <str<strong>on</strong>g>study</str<strong>on</strong>g>, 9 species of <strong>Acropora</strong> were identified, including<br />
A. tortuosa (Dana, 1846) and A. mossambica Riegl, 1995, which are new to the Persian Gulf. Furthermore, A. horrida (Dana, 1846), A.<br />
nasuta (Dana, 1846), A. aspera (Dana, 1846), and A. muricata (Linnaeus, 1758) were identified, all of which were recorded for the first<br />
time from Iran. By adding these species to those previously reported from the area, the numbers of <strong>Acropora</strong> species in the <str<strong>on</strong>g>study</str<strong>on</strong>g> area<br />
and in the Persian Gulf reach 10 and 15, respectively.<br />
Key words: Khark, Larak, Farur, tax<strong>on</strong>omic key, islands, Iran, coral, <strong>Acropora</strong><br />
1. Introducti<strong>on</strong><br />
The informati<strong>on</strong> available <strong>on</strong> different aspects of<br />
scleractinian diversity and biology is exhaustive (Wallace,<br />
1999; Meyer and Paulay, 2000; Ver<strong>on</strong>, 2000). From a<br />
tax<strong>on</strong>omic point of view, staghorn corals, members<br />
of genus <strong>Acropora</strong> Oken, 1815, are am<strong>on</strong>gst the most<br />
c<strong>on</strong>troversial taxa due to their high intraspecific diversity<br />
and interspecific similarities. Those facts are evident in the<br />
c<strong>on</strong>stant changes of the total diversity estimates given by<br />
several authors (see Wallace, 1999; Ver<strong>on</strong>, 2000). Although<br />
recent molecular studies (van Oppen et al., 2001; Fukami et<br />
al., 2003; Wolstenholme et al., 2003, Wolstenholme, 2004;<br />
Fukami et al., 2007; Chen et al., 2009) raised some hopes in<br />
solving such problems, morphological identificati<strong>on</strong> is still<br />
the most preferred procedure in the tax<strong>on</strong>omical studies<br />
c<strong>on</strong>ducted <strong>on</strong> such a group. Wallace (1999) resolved<br />
most of the tax<strong>on</strong>omical problems of this genus, hence<br />
improving the accuracy of surveys.<br />
Despite the numerous studies carried out <strong>on</strong> <strong>Acropora</strong><br />
worldwide, the data available <strong>on</strong> scleractinian species<br />
diversity in the Persian Gulf in general and <strong>on</strong> the genus<br />
<strong>Acropora</strong> in particular are restricted (Spalding et al., 2001),<br />
even though the corals of the southern part of the Persian<br />
Gulf are better known than those of the northern and<br />
northeastern parts.<br />
Coles (2003) reported 11 species of <strong>Acropora</strong>, including<br />
A. clathrata (Brook, 1891), in the Persian Gulf. In her<br />
previous m<strong>on</strong>ograph, which included the Persian Gulf and<br />
the Gulf of Oman, Wallace (1999) reported 13 <strong>Acropora</strong><br />
species, but excluded A. clathrata. On the northern<br />
coast of the Persian Gulf, however, <strong>on</strong>ly 5 <strong>Acropora</strong><br />
species, including A. clathrata, have been reported so far<br />
(Maghsoudlou, 2008).<br />
Due to the lack of adequate tax<strong>on</strong>omical informati<strong>on</strong><br />
about the <strong>Acropora</strong> species of the Iranian parts of the<br />
Persian Gulf, and taking into account the importance<br />
of the genus <strong>Acropora</strong> from an endemicity point of view<br />
and its use as a model for <str<strong>on</strong>g>study</str<strong>on</strong>g>ing the effects of global<br />
warming <strong>on</strong> marine taxa (Wallace and Muir, 2005), the<br />
present <str<strong>on</strong>g>study</str<strong>on</strong>g> was c<strong>on</strong>ducted to ascertain the distributi<strong>on</strong><br />
and the diversity of <strong>Acropora</strong> species in the area.<br />
2. Materials and methods<br />
On the basis of the studies performed <strong>on</strong> the scale of<br />
envir<strong>on</strong>mental stresses (see Kämpf and Sadrinasab, 2006;<br />
Nezlin et al., 2007), the Persian Gulf was divided into 3<br />
distinct parts: the eastern, central, and western parts.<br />
Taking this fact into c<strong>on</strong>siderati<strong>on</strong>, in each part, a coralrich<br />
island was selected for sampling: these were Larak<br />
Island (26°49′N–26°53′N, 56°19′E–56°25′E), Farur Island
(26°14′N–26°19′N, 54°29′E–54°32′E), and Khark Island<br />
(29°12′N–29°16′N, 50°16′E–50°20′E), from the eastern,<br />
central, and western parts of the Persian Gulf, respectively.<br />
Al<strong>on</strong>g the coasts of these 3 islands, 40 locati<strong>on</strong>s<br />
were selected (Figure 1) for sampling. The geographic<br />
coordinates of each stati<strong>on</strong> were obtained using a Garmin<br />
Etrex GPS receiver. From the 1169 col<strong>on</strong>ies, pieces of 10<br />
to 15 cm were collected from October 2008 to December<br />
2009. The first author carried out the underwater<br />
photography. At each stati<strong>on</strong>, the specimens were collected<br />
by scuba diving to the depths of 3, 6, and 9 m, al<strong>on</strong>g a line<br />
at least 120 m l<strong>on</strong>g.<br />
c b<br />
30°<br />
28°<br />
26°<br />
2 km 2 km<br />
IRAQ<br />
KUWAIT<br />
Khark<br />
SAUDI ARABIA<br />
N<br />
W E<br />
S<br />
Khark<br />
Farsi<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
BAHRAIN<br />
Bushehr<br />
Farur<br />
Persian Gulf<br />
The specimens were bleached in 5 ppt sodium<br />
hypochlorite and transferred to the laboratory where<br />
further studies took place. To <str<strong>on</strong>g>study</str<strong>on</strong>g> the gross and fine<br />
morphologies of the specimens, a Nik<strong>on</strong> SMZ 1000<br />
stereomicroscope and a Zeiss DSM960A scanning electr<strong>on</strong><br />
microscope were used, respectively. Based <strong>on</strong> even minute<br />
morphological differences, 55 specimens representing<br />
9 species (excluding <strong>Acropora</strong> aspera) were selected<br />
and shipped to the Museum of Tropical Queensland for<br />
further studies and identificati<strong>on</strong>s. Of these specimens,<br />
C.C. Wallace identified 45. Mostly due to their small size,<br />
10 specimens remained unidentified. The morphological<br />
N<br />
W E<br />
S<br />
IRAN<br />
Larak<br />
0 200km<br />
QATAR<br />
U.A.E.<br />
48° 50° 52° 54° 56°<br />
TIR<br />
N<br />
W E<br />
S<br />
Bandar Abbas<br />
Hormuz<br />
Lavan<br />
Larak<br />
Hendourabi<br />
Hengam<br />
Kish<br />
FarurT<strong>on</strong>be-e<br />
Bozorg<br />
T<strong>on</strong>be-e Kouchak<br />
Farurgan<br />
SirriAbu Musa<br />
Qeshm<br />
OMAN<br />
Figure 1. Sampling sites in the Persian Gulf al<strong>on</strong>g the coasts of a) Larak Island, b) Farur Island, and c) Khark Island.<br />
a<br />
N<br />
W E<br />
S<br />
2 km<br />
309
descripti<strong>on</strong>s as well as the terminology and measuring<br />
procedures used in this <str<strong>on</strong>g>study</str<strong>on</strong>g> were performed <strong>on</strong> the basis<br />
of the descripti<strong>on</strong>s and the nomenclature suggested by<br />
Wallace (1978), Ver<strong>on</strong> and Wallace (1984), Riegl (1995),<br />
Wallace (1999), Ver<strong>on</strong> (2000), and Wolstenholme et al.<br />
(2003).<br />
In the present paper, a regi<strong>on</strong>al tax<strong>on</strong>omic key <strong>on</strong><br />
the <strong>Acropora</strong> species is also suggested, in which the<br />
characteristics of the specimens found in the present <str<strong>on</strong>g>study</str<strong>on</strong>g><br />
were used for species of the northern and northeastern<br />
parts of the Persian Gulf. For the species of the southern<br />
part, the descripti<strong>on</strong>s given by Wallace (1999) were used.<br />
3. Results<br />
All 1169 specimens were examined, but from those, for<br />
each species a few specimens c<strong>on</strong>taining most of the<br />
plasticity range of the characteristics of that species were<br />
selected for descripti<strong>on</strong> purposes. Throughout this article,<br />
the specimens’ data are presented in the following order:<br />
sampling locati<strong>on</strong>; coordinates of the sampling locati<strong>on</strong>;<br />
locati<strong>on</strong> depth, sampling date, and the museum reference<br />
number. The order followed for the descripti<strong>on</strong> of 2 or<br />
more specimens was also the same.<br />
Whenever branch diameters are provided, they are<br />
presented in the following order: branch diameter at the<br />
base, at the midpoint, and at 5 mm below the tip of the<br />
branch.<br />
From the northern and northeastern parts of the<br />
Persian Gulf, 9 species of <strong>Acropora</strong> were identified during<br />
the <str<strong>on</strong>g>study</str<strong>on</strong>g>. Am<strong>on</strong>gst those species, 6 are reported for the<br />
first time from the area of the <str<strong>on</strong>g>study</str<strong>on</strong>g>, and 2 other species<br />
are new to the Persian Gulf. Although the ecological and<br />
populati<strong>on</strong> data collected during the sampling are not<br />
analyzed in the present paper in general, am<strong>on</strong>gst the<br />
species found in this <str<strong>on</strong>g>study</str<strong>on</strong>g>, A. downingi and A. arabensis<br />
were the most comm<strong>on</strong> species, with A. valida next in<br />
abundance and the rest of the species being rare.<br />
The specimens described in this article are deposited<br />
in the Zoological Museum of the University of Tehran<br />
(ZUTC) and the Nati<strong>on</strong>al Museum of Natural History<br />
(MMTT).<br />
In the descripti<strong>on</strong> of each species, we chose not to<br />
menti<strong>on</strong> the syn<strong>on</strong>yms, as they are available in detail from<br />
Wallace (1999).<br />
<strong>Acropora</strong> valida (Dana, 1846); Figures 2a–2g<br />
Material examined: Iran, Larak Island: 26°53′10.1″N,<br />
56°23′58.3″E; depth 6 m; 7 October 2008; MMTT Cnid.<br />
1024. Larak Island: 26°52′45.8″N, 56°20′26.8″E; depth<br />
6 m; 14 October 2008; MMTT Cnid. 1319. Larak Island:<br />
26°53′12.1″N, 56°23′32.1″E; depth 3 m; 8 October 2008;<br />
ZUTC Cnid. 1115.<br />
Descripti<strong>on</strong>: Branching pattern: Caespitose (usually<br />
for small col<strong>on</strong>ies) to corymbose (usually for large<br />
310<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
col<strong>on</strong>ies); main branches of large col<strong>on</strong>ies sometimes<br />
grow horiz<strong>on</strong>tally; col<strong>on</strong>y size up to 45 cm (rarely more<br />
than 55 cm) (Figure 2a); branches cylindrical to slightly<br />
tapering (Figure 2b); branch diameters: 6.5–11.9 mm,<br />
7.3–11.7 mm, and 7.2–9.8 mm; branch length up to 29.4<br />
mm; growth determinate or semideterminate depending<br />
<strong>on</strong> the corallum.<br />
Axial corallites: C<strong>on</strong>spicuous, cylindrical, twice the<br />
size of radial corallites; outer diameter 2.0–2.6 mm; calice<br />
diameter and thickness of axial corallites walls similar<br />
(0.8–1.2 mm); round opening; most axial corallites c<strong>on</strong>tain<br />
2 septa cycles; primary septa up to 2/3R, but l<strong>on</strong>ger in the<br />
deeper parts of the calice; sec<strong>on</strong>dary septa up to 1/3R,<br />
sometimes dentate; tertiary septa rarely present (Figure 2f).<br />
Radial corallites: Appressed tubular with oval opening;<br />
more or less equal in size; profile length 2.0–3.3 mm; some<br />
tubular or tubo-nariform corallites (usually having thick<br />
lips) scattered al<strong>on</strong>g branches (Figures 2c and 2d); primary<br />
septa present up to 2/3R; directive septa prominent;<br />
sec<strong>on</strong>dary septa development variable in different<br />
corallites: up to 1/5R in those near the branch base, absent<br />
or partially developed in those near the branch tip.<br />
Coenosteum: Broken-costate to costate <strong>on</strong> outer walls<br />
of radial corallites (Figure 2e); densely arranged lines of<br />
laterally flattened spinules <strong>on</strong> the outer edge of radial<br />
corallite walls; reticulate with simple to slightly elaborated<br />
spinules between radials (Figure 2g).<br />
Color: Pale to cream brown.<br />
Remarks: Comparis<strong>on</strong> of the specimens of A. valida<br />
from this <str<strong>on</strong>g>study</str<strong>on</strong>g> with those described by Wallace (1999)<br />
showed that the branches were relatively thinner and<br />
shorter, and axial corallites were smaller and had thinner<br />
walls in the specimens from the <str<strong>on</strong>g>study</str<strong>on</strong>g> area.<br />
<strong>Acropora</strong> aspera (Dana, 1846); Figures 3a–3g<br />
Material examined: Iran, Farur Island: 26°17′44.1″N,<br />
54°32′22.4″E; depth 6 m; 28 October 2009; MMTT Cnid.<br />
1347. Khark Island: 29°16′33.8″N, 50°18′21.5″E; depth 3<br />
m; 19 December 2009; MMTT Cnid. 1699.<br />
Descripti<strong>on</strong>: Branching pattern: Arborescent (Figure<br />
3a); branch length up to 71.7 mm, tapering slightly towards<br />
the tip, and tapering abruptly at the tip (Figure 3b); branch<br />
diameters: 11.3–13.0 mm, 9.5–10.8 mm, and 6.8–8.4 mm;<br />
growth indeterminate.<br />
Axial corallites: C<strong>on</strong>spicuous; larger than radial<br />
corallites; dome-shaped; outer diameter 2.7–4.0 mm;<br />
calice diameter 1.3–1.6 mm; both primary and sec<strong>on</strong>dary<br />
septa present, primary septa up to 2/3R, sec<strong>on</strong>dary septa<br />
up to 1/2R (Figure 3f).<br />
Radial corallites: Labellate; horiz<strong>on</strong>tal extensi<strong>on</strong> of the<br />
outer walls and absence of inner walls give a gutter-shaped<br />
appearance (Figures 3c and 3d); profile length 1.2–1.7<br />
mm; subimmersed radial corallites scattered between<br />
labellate corallites; crowded; radial corallites in 2 sizes,
Figure 2. <strong>Acropora</strong> valida: a) live col<strong>on</strong>y; b) part of col<strong>on</strong>y; c)<br />
porti<strong>on</strong> of branch; d) SEM micrograph showing micrograph<br />
showing a few radial corallites; e) SEM micrograph showing<br />
coenosteum <strong>on</strong> a radial corallite; f) SEM micrograph showing<br />
top view of an axial corallite; g) SEM micrograph showing<br />
coenosteum between radial corallites.<br />
both with a round opening; development of septal cycles<br />
varies depending <strong>on</strong> the positi<strong>on</strong> of the radial corallites<br />
<strong>on</strong> the branch. In those located from the branch base to<br />
near its tip 2 complete cycles are present, but these are<br />
incomplete and poorly developed in the radial corallites<br />
located at the branch tips; primary septa up to 1/2R, except<br />
for the directives that are well developed; sec<strong>on</strong>dary septa,<br />
if present, a low ridge maximum 1/3R.<br />
Coenosteum: Costate to broken-costate <strong>on</strong> radial<br />
corallites proper (Figure 3e); sometimes densely arranged<br />
lines of laterally flattened spinules <strong>on</strong> radial corallites;<br />
coenosteum between radial corallites reticulate with<br />
dispersed, laterally flattened, and slightly elaborated<br />
spinules (Figure 3g).<br />
Color: Brown to cream-brown or greenish brown.<br />
<strong>Acropora</strong> horrida (Dana, 1846); Figures 4a–4g<br />
Material examined: Iran, Farur Island: 26°17′44.1″N,<br />
54°32′22.4″E; depth 6 m; 28 October 2009; MMTT Cnid.<br />
1348. Larak Island: 26°52′45.8″N, 56°20′26.8″E; depth 6<br />
m; 14 October 2008; ZUTC Cnid. 1305.<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
Figure 3. <strong>Acropora</strong> aspera: a) porti<strong>on</strong> of col<strong>on</strong>y; b) porti<strong>on</strong> of<br />
branch; c) SEM micrograph showing terminal part of branch; d)<br />
SEM micrograph from lateral view of radial corallites, note the<br />
gutter-shaped appearances formed by the outer walls; e) SEM<br />
micrograph showing coenosteum <strong>on</strong> a radial corallite; f) top view<br />
of axial corallite showing extent of primary and sec<strong>on</strong>dary septa; g)<br />
SEM micrograph showing coenosteum between radial corallites.<br />
Descripti<strong>on</strong>: Branching pattern: Arborescent; slightly<br />
tapering branches (Figures 4a and 4b); branch diameters:<br />
7.9–11.3 mm, 7.2–10.5 mm, 6.0–9.4 mm; branch length up<br />
to 26.1 mm; growth indeterminate.<br />
Axial corallites: C<strong>on</strong>spicuous; outer diameter 2.0–2.6<br />
mm; calice diameter 1.2–1.6 mm; primary septa present,<br />
maximum 3/4R; sometimes dentate; sec<strong>on</strong>dary septa<br />
absent (Figure 4f).<br />
Radial corallites: Extremely fragile; with different sizes;<br />
ragged surface (Figure 4c); profile length 2.0–4.5 mm;<br />
round opening; septa poorly developed, particularly the<br />
inner septa; primary septa up to 1/3R; outer directive<br />
septum more developed than the inner directive;<br />
sec<strong>on</strong>dary septa absent.<br />
Coenosteum: Broken-costate to reticulate <strong>on</strong> radial<br />
corallites proper (Figure 4d); reticulate between radial<br />
corallites; simple, laterally flattened spinules scattered <strong>on</strong><br />
and between radials (Figures 4e and 4g).<br />
Color: Brown.<br />
311
Figure 4. <strong>Acropora</strong> horrida: a) porti<strong>on</strong> of col<strong>on</strong>y; b) porti<strong>on</strong> of<br />
branch; c) SEM micrograph showing terminal part of branch;<br />
d) SEM micrograph showing coenosteum <strong>on</strong> radial corallite; e)<br />
SEM micrograph showing 2 simple, laterally flattened spinules<br />
between radial corallites; f) top view of axial corallite, note the<br />
absence of sec<strong>on</strong>dary septal cycle; g) high-magnificati<strong>on</strong> SEM<br />
micrograph showing ornamentati<strong>on</strong>s <strong>on</strong> coenosteum between<br />
radial corallites.<br />
Remarks: Of this species, 1 specimen was sent to the<br />
Museum of Tropical Queensland, where C.C. Wallace<br />
identified it as <strong>Acropora</strong> cf. horrida. Detailed and careful<br />
examinati<strong>on</strong> of other specimens from the same dive and<br />
of additi<strong>on</strong>al specimens collected later <strong>on</strong> allowed us to<br />
c<strong>on</strong>firm the identificati<strong>on</strong>.<br />
Comparis<strong>on</strong> between the measurements of the<br />
specimen found in the present <str<strong>on</strong>g>study</str<strong>on</strong>g> with those described<br />
by Wallace (1999) indicated that branches of almost the<br />
same diameter were much shorter in our specimens. The<br />
other measurements more or less overlapped.<br />
<strong>Acropora</strong> downingi Wallace, 1999; Figures 5a–5f<br />
Material examined: Iran, Larak Island: 26°53′07.8″N,<br />
56°23′59.0″E; depth 3 m; 7 October 2008; MMTT Cnid.<br />
1022. Larak Island: 26°53′16.3″N, 56°21′01.3″E; depth 6<br />
m, 11 October 2008; MMTT Cnid. 1172. Khark Island:<br />
29°12′36.6″N, 50°19′56.5″E; depth 3 m; 14 December<br />
2009; MMTT Cnid. 1404. Farur Island: 26°17′44.1″N,<br />
312<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
54°32′22.4″E; depth 6 m; 28 October 2009; MMTT Cnid.<br />
1382. Khark Island: 29°16′34.4″N, 50°18′31.7″E; depth 9<br />
m; 20 December 2009; ZUTC Cnid. 1708.<br />
Descripti<strong>on</strong>: Branching pattern: Arborescent table,<br />
large, some up to 3 m in diameter (Figure 5a), both single<br />
and multilayered table forms present; some branches<br />
grow upwards; branches tapering slightly towards the tip<br />
(Figures 5b and 5c), anastomosed branches forming solid<br />
central plate; branch diameters: 5.4–14.8 mm, 4.8–13.6<br />
mm, 4.2–9.1 mm; branch length up to 62.0 mm; growth<br />
determinate.<br />
Axial corallites: Small cylindrical; outer diameter<br />
0.9–2.5 mm; calice diameter 0.6–1.6 mm; round to oval<br />
opening; 1 or 2 septal cycles present (Figure 5e), primary<br />
septa 1/2R, directive septa sometimes touching; sec<strong>on</strong>dary<br />
septa absent or inc<strong>on</strong>spicuous; septal length increasing<br />
downwards.<br />
Radial corallites: Crowded and fragile; various sizes<br />
(Figure 5c); tubular, with oblique opening and upward<br />
sharp edges (Figure 5d); profile length 1–4 mm; some<br />
corallites immersed, especially those at the base of<br />
branches; primary septa seldom present (in most cases<br />
<strong>on</strong>ly directive septa present), when present maximum<br />
1/4R.<br />
Coenosteum: Coenosteum <strong>on</strong> radial corallites proper<br />
costate (Figure 5d), broken-costate, or linearly arranged,<br />
laterally flattened spinules; coenosteum between radials<br />
reticulate with simple spinules (Figure 5f).<br />
Color: Brown to dark green or gray; branch tips pale<br />
or white.<br />
Remarks: Our preliminary studies indicated that<br />
<strong>Acropora</strong> downingi has 2 forms in the <str<strong>on</strong>g>study</str<strong>on</strong>g> area. In <strong>on</strong>e<br />
form, some branches grow upwards, unlike in the sec<strong>on</strong>d<br />
form, which has no upright branches <strong>on</strong> the table. The<br />
latter form might have been misidentified as A. clathrata.<br />
Further tax<strong>on</strong>omic work <strong>on</strong> these growth forms is<br />
currently underway. A. clathrata was not found during<br />
the present <str<strong>on</strong>g>study</str<strong>on</strong>g>. C<strong>on</strong>sequently, it is likely that either the<br />
reports of A. clathrata from the southern (and western)<br />
Persian Gulf (Sheppard and Sheppard, 1991; Hodgs<strong>on</strong><br />
and Carpenter, 1996; Riegl, 1999; Abdel-Moati, 2011) are<br />
a misidentificati<strong>on</strong> of A. downingi (see Wallace, 1999) or<br />
that A. clathrata is restricted to the southern (and western)<br />
part of the Persian Gulf, or it is an extremely rare species in<br />
this part of the Persian Gulf (see discussi<strong>on</strong>).<br />
<strong>Acropora</strong> arabensis Hodgs<strong>on</strong> and Carpenter, 1996;<br />
Figures 6a–6g<br />
Material examined: Iran, Larak Island: 26°53′16.3″N,<br />
56°21′01.3″E; depth 6 m; 11 October 2008; MMTT<br />
Cnid. 1153. Larak Island: 26°53′16.9″N, 56°21′00.4″E;<br />
depth 9 m; 11 October 2008; MMTT Cnid. 1239. Larak<br />
Island: 26°53′12.1″N, 56°23′32.1″E; depth 3 m; 8 October<br />
2008; MMTT Cnid. 1125. Farur Island: 26°16′52.4″N,
Figure 5. <strong>Acropora</strong> downingi: a) live col<strong>on</strong>y; b) porti<strong>on</strong> of col<strong>on</strong>y;<br />
c) porti<strong>on</strong> of branch, note different sizes of radial corallites; d)<br />
SEM micrograph showing coenosteum <strong>on</strong> a radial corallite<br />
proper; e) top view of axial corallite; f) SEM micrograph showing<br />
coenosteum between radial corallites.<br />
54°29′36.4″E; depth 9 m; 30 October 2009; MMTT Cnid.<br />
1377. Khark Island: 29°13′53.5″N, 50°17′53.5″E; depth 3<br />
m; 15 December 2009; ZUTC Cnid. 1503.<br />
Descripti<strong>on</strong>: Branching pattern: Arborescent,<br />
sometimes caespitose; col<strong>on</strong>ies mostly small, up to 35 cm<br />
in diameter (Figure 6a), seldom larger col<strong>on</strong>ies, up to 75<br />
cm in diameter; most branches upright, tapering slightly<br />
towards the tip and terete (Figures 6b and 6c); branch<br />
diameters: 5.5–14.0 mm, 5.3–13.5 mm, and 4.9–11.9 mm;<br />
branch length up to 47.0 mm; growth semideterminate.<br />
Axial corallites: C<strong>on</strong>spicuous, outer diameter 2.3–3.1<br />
mm; calice diameter 0.7–1.6 mm; at least 1 complete septal<br />
cycle present; primary septa maximum 3/4R; directive<br />
septa well developed; sec<strong>on</strong>dary usually absent, when<br />
present up to 1/4R (Figure 6f).<br />
Radial corallites: Regular and sturdy; appressed tubular<br />
to nariform (Figure 6d), gradually changing from branch<br />
tip towards its base; calice mostly oblique, sometimes<br />
dimidiate; 2 sizes; profile length 1.2–5.4 mm; outer walls<br />
extended al<strong>on</strong>g the branches; a few subimmersed corallites<br />
scattered <strong>on</strong> branches; dense immersed corallites at the base<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
Figure 6. <strong>Acropora</strong> arabensis: a) live col<strong>on</strong>y, note the distinct<br />
white branch tips; b) porti<strong>on</strong> of the col<strong>on</strong>y; c) porti<strong>on</strong> of branch;<br />
d) SEM micrograph showing close up view of radial corallites;<br />
e) SEM micrograph showing broken costae ornamentati<strong>on</strong> <strong>on</strong><br />
radial corallite; f) top view of axial corallite; g) SEM micrograph<br />
showing coenosteum between radial corallites.<br />
and between branches; at the base of branches first cycle<br />
complete, maximum septal development 1/3R, directive<br />
septa prominent, sometimes touching, particularly<br />
towards the bottom of calice; near the branches tip primary<br />
cycle mostly incomplete, directive septa present maximum<br />
1/4R; sec<strong>on</strong>dary cycle usually absent (especially in radials<br />
near the branch tip) or very short.<br />
Coenosteum: Broken-costate <strong>on</strong> radial corallites<br />
(Figure 6e); reticulate with simple, dense, laterally flattened<br />
spinules between radials (Figure 6g).<br />
Color: Dark brown to pale brown <strong>on</strong> branches, pale or<br />
white <strong>on</strong> branch tips (Figure 6a).<br />
Remarks: In general, most col<strong>on</strong>ies of A. arabensis<br />
are arborescent in the <str<strong>on</strong>g>study</str<strong>on</strong>g> area, while those described<br />
in Wallace (1999) were caespitose to low arborescent. The<br />
branches were also relatively thinner and much shorter<br />
in our specimens. C<strong>on</strong>versely, axial corallites of the<br />
specimens found in these parts of the Persian Gulf were<br />
larger than those described by Wallace (1999). Differences<br />
were also observed in the coenosteum: while that of the<br />
specimens in the present <str<strong>on</strong>g>study</str<strong>on</strong>g> is broken-costate <strong>on</strong> the<br />
313
adial corallites and reticulate between the corallites, it is<br />
uniformly reticulate in the specimens of Wallace (1999).<br />
<strong>Acropora</strong> mossambica Riegl, 1995; Figures 7a–7h<br />
Material examined: Iran, Larak Island: 26°52′45.8″N,<br />
56°20′26.8″E; depth 6 m; 14 October 2008; MMTT Cnid.<br />
1323. Larak Island: 26°53′12.1″N, 56°23′32.1″E; depth 3<br />
m; 8 October 2008; MMTT Cnid. 1118.<br />
Descripti<strong>on</strong>: Branching pattern: Corymbose; branches<br />
very thick (Figure 7a); some sec<strong>on</strong>dary branches fuse<br />
to main branches at branching area, forming flattened<br />
branches; branch diameters: 10.9–13.2 mm, 8.8–13.5 mm,<br />
and 7.8–11.3 mm; branch length up to 24.2 mm.<br />
Axial corallites: Size more than twice that of radials;<br />
walls thick; round to slightly oval opening; outer diameter<br />
3.1–4.3 mm; calice diameter 1.3–1.8 mm; 1 or 2 septal<br />
cycles, poorly developed; primary septa maximum 1/3R;<br />
directive septa prominent, often touching at bottom of<br />
calice; sec<strong>on</strong>dary septa mostly absent, when present up to<br />
1/5R (Figure 7g).<br />
Radial corallites: Tubo-nariform, different sizes<br />
(Figures 7d and 7e); some corallites appressed, scattered<br />
<strong>on</strong> branches (Figure 7c); subimmersed and immersed<br />
radials more numerous at the base and between branches<br />
(Figure 7b); oval opening; outer walls thick, roughly 7<br />
times thicker than the inner walls; profile length 2–3 mm;<br />
septa poorly developed; primary septa maximum 1/5R;<br />
sec<strong>on</strong>dary septa absent or minute.<br />
Coenosteum: Costate to broken-costate <strong>on</strong> radial<br />
corallites (in most cases, costate at the tips and brokencostate<br />
at the bases of branches), sometimes densely<br />
arranged lines of laterally flattened spinules <strong>on</strong> radial<br />
corallites (Figure 7f); simple spinules between radials<br />
(Figure 7h).<br />
Color: Pale to dark brown.<br />
Remarks: So far, A. mossambica has <strong>on</strong>ly been reported<br />
in the literature from high-energy intertidal habitats in<br />
SE Africa (Riegl, 1995). Thus, finding this species in the<br />
Persian Gulf is noteworthy. For c<strong>on</strong>firmati<strong>on</strong>, a specimen<br />
of this species was sent to the Museum of Tropical<br />
Queensland, where it was identified by C.C. Wallace as<br />
<strong>Acropora</strong> cf. mossambica. Characteristics supporting the<br />
identificati<strong>on</strong> included the branch shape, size of axial<br />
corallites, and shape of radial corallites. C<strong>on</strong>sequently, this<br />
is the first report of A. mossambica from the Persian Gulf.<br />
Misidentificati<strong>on</strong> of A. mossambica as similar species, e.g.,<br />
A. selago and A. tenuis, both present in the area (Riegl,<br />
1999; Wallace, 1999), was possible. However, the larger size<br />
of axial corallites together with the differences observed in<br />
the forms of radial corallites supported the identificati<strong>on</strong><br />
as A. mossambica.<br />
Comparis<strong>on</strong> between measurements of the specimen<br />
found in the present <str<strong>on</strong>g>study</str<strong>on</strong>g> with those described by Riegl<br />
(1995) suggests that the axial corallites are narrower and<br />
314<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
Figure 7. <strong>Acropora</strong> mossambica: a) porti<strong>on</strong> of col<strong>on</strong>y; b)<br />
top view of plate; c) porti<strong>on</strong> of branch; d) SEM micrograph<br />
showing terminal part of branch; e) SEM micrograph showing<br />
close up view of radial corallites; f) SEM micrograph showing<br />
coenosteum <strong>on</strong> radial corallite; g) top view of axial corallite; h)<br />
SEM micrograph showing coenosteum between radial corallites.<br />
smaller in our specimens, but very limited details for other<br />
measurements were provided by Riegl (1995).<br />
<strong>Acropora</strong> muricata (Linnaeus, 1758); Figures 8a–8i<br />
Material examined: Iran, Larak Island: 26°53′15.9″N,<br />
56°21′02.3″E; depth 6 m; 11 October 2008; MMTT Cnid.<br />
1182. Larak Island: 26°53′15.9″N, 56°21′02.3″E; depth 3<br />
m; 11 October 2008; MMTT Cnid. 1185.<br />
Descripti<strong>on</strong>: Branching pattern: Arborescent (Figure<br />
8a) with l<strong>on</strong>g branches (maximum 160 mm), slightly<br />
tapering (Figures 8b and 8c); branches relatively thick;<br />
branch diameters: 10.0–16.6 mm, 9.7–15.4 mm, and 8.8–<br />
14.1 mm; growth determinate.<br />
Axial corallites: Axial corallites thicker and substantially<br />
larger than radials; outer diameter 2.5–3.2 mm; calice<br />
diameter 1.1–1.5 mm; calice usually not perfectly round;<br />
2 complete septal cycles present (Figure 8g); primary septa<br />
up to 3/4R; sec<strong>on</strong>dary septa up to 2/3R.<br />
Radial corallites: Two sizes, arranged spirally <strong>on</strong><br />
branches; mostly appressed tubular, some nariform, with<br />
round to oval openings (Figures 8d and 8e); subimmersed
and immersed corallites at base of branches; profile length<br />
up to 3.6 mm; 2 septal cycles developed (Figure 8g); inner<br />
and outer septa similar in size; primary septa maximum<br />
3/4R; sec<strong>on</strong>dary septa up to 2/3R.<br />
Coenosteum: Similar both <strong>on</strong> and between radial<br />
corallites (Figures 8f and 8h), linear arrangement of<br />
elaborated spinules <strong>on</strong> coenosteum, some spinules forked<br />
(Figures 8h and 8i).<br />
Color: Pale brown, brick red, or pale yellow.<br />
Remarks: A. muricata has not been previously reported<br />
in this part of the Persian Gulf. In the beginning, that<br />
caused some doubts as to whether our specimens really<br />
bel<strong>on</strong>ged to A. muricata, as in some areas of the Persian<br />
Gulf A. phara<strong>on</strong>is forms open arborescent col<strong>on</strong>ies similar<br />
to those formed by A. muricata. Nevertheless, other<br />
characteristics, including the absence of branchlets in A.<br />
muricata and the presence of a stalk in A. phara<strong>on</strong>is, help<br />
to distinguish these 2 species. A specimen was sent to<br />
the Museum of Tropical Queensland, where C.C. Wallace<br />
identified it as <strong>Acropora</strong> cf. muricata. Although there are<br />
Figure 8. <strong>Acropora</strong> muricata: a) live col<strong>on</strong>y; b) porti<strong>on</strong> of col<strong>on</strong>y;<br />
c) porti<strong>on</strong> of a branch; d) SEM micrograph showing branch tip;<br />
e) SEM micrograph showing close up view of radial corallites; f)<br />
SEM micrograph showing coenosteum <strong>on</strong> radial corallite; g) top<br />
view of axial corallite; h) SEM micrograph showing coenosteum<br />
between radial corallites; i) SEM micrograph showing close up<br />
view of coenosteum between radial corallites.<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
literature records of A. phara<strong>on</strong>is from the Persian Gulf<br />
(Coles and Fadlallah, 1991; Sheppard and Sheppard,<br />
1991; Mostafavi et al., 2010), there is actually no formal<br />
c<strong>on</strong>firmed record based <strong>on</strong> voucher skeletal specimens of<br />
A. phara<strong>on</strong>is in this area. That, by itself, does not exclude<br />
the possibility of its presence in the area; however, our<br />
extensive and careful sampling did not reveal any A.<br />
phara<strong>on</strong>is, either.<br />
From a morphological point of view, almost all<br />
the characteristics of our specimens overlapped those<br />
described by Wallace (1999) except in the morphology of<br />
coenosteum, which is costate <strong>on</strong> and reticulated between<br />
the radial corallites of the specimens examined by Wallace.<br />
<strong>Acropora</strong> nasuta (Dana, 1846); Figures 9a–9g<br />
Material examined: Iran, Larak Island: 26°53′07.8″N,<br />
56°23′59.0″E; depth 3 m; 7 October 2008; MMTT Cnid.<br />
1018. Larak Island: 26°52′45.8″N, 56°20′26.8″E; depth<br />
6 m; 14 October 2008; MMTT Cnid. 1322. Larak Island:<br />
26°53′15.0″N, 56°20′59.3″E; depth 6 m; 13 October 2008;<br />
ZUTC Cnid. 1260.<br />
Descripti<strong>on</strong>: Branching pattern: Corymbose, but<br />
tending to form small tables; branches cylindrical (Figure<br />
9a), tapering abruptly at the tip (Figure 9b), slender in<br />
attachment point to the col<strong>on</strong>y; col<strong>on</strong>ies small; irregular<br />
branching; branch diameters: 8.9–15.2 mm, 8.0–14.2 mm,<br />
and 7.7–10.5 mm; branch length up to 54.9 mm; growth<br />
determinate.<br />
Axial corallites: Cylindrical, larger than radial corallites;<br />
outer diameter 2.8–3.5 mm; calice diameter 1.1–1.3 mm;<br />
2 complete septal cycles; primary septa maximum up to<br />
2/3R; sec<strong>on</strong>dary septa 1/3R; directive septa slightly l<strong>on</strong>ger<br />
than the others (Figure 9f).<br />
Radial corallites: Nariform to tubo-nariform (Figure<br />
9c); round to oblique opening (Figure 9e), 2 sizes; profile<br />
length 2.2–3.0 mm; immersed corallite number increasing<br />
towards the base and between branches (Figure 9a); 1<br />
complete septal cycle always present; primary septa up to<br />
1/2R; lines of laterally flattened spinules usually present;<br />
sec<strong>on</strong>dary septa absent, or sometimes forming lines of<br />
small spinules.<br />
Coenosteum: Broken costae or laterally flattened<br />
spinules in rows <strong>on</strong> radials, with spinules mostly joined by<br />
short lamellae (Figure 9d); between radials reticulate with<br />
simple spinules (Figure 9g).<br />
Color: Pale brown to brown.<br />
Remarks: By comparing the measurements of<br />
specimens found in the present <str<strong>on</strong>g>study</str<strong>on</strong>g> with those described<br />
in Wallace (1999), it was evident that the branches are<br />
both thicker and shorter, and axial corallites are larger in<br />
specimens of present <str<strong>on</strong>g>study</str<strong>on</strong>g>.<br />
<strong>Acropora</strong> tortuosa (Dana, 1846); Figures 10a–10h<br />
Material examined: Iran, Farur Island: 26°17′44.1″N,<br />
54°32′22.4″E; depth 6 m, 28 October 2009; MMTT Cnid.<br />
1387.<br />
315
Figure 9. <strong>Acropora</strong> nasuta: a) porti<strong>on</strong> of col<strong>on</strong>y; b) porti<strong>on</strong> of<br />
branch; c) SEM micrograph showing terminal part of branch; d)<br />
SEM micrograph showing coenosteum <strong>on</strong> a radial corallite; e)<br />
fr<strong>on</strong>t view of few radial corallites; f) top view of an axial corallite;<br />
g) SEM micrograph showing coenosteum between radial<br />
corallites.<br />
Descripti<strong>on</strong>: Branching pattern: Arborescent (Figure<br />
10a); branches thin and l<strong>on</strong>g (maximum 110 mm),<br />
slightly tapering (Figures 10b and 10c); branch diameters:<br />
8.0–12.2 mm, 7.5–11.2 mm, and 5.0–6.6 mm; growth<br />
indeterminate.<br />
Axial corallites: Small but slightly larger than large radial<br />
corallites; outer diameter 2.0–2.8 mm; calice diameter<br />
1.0–1.4 mm; wall thickness about half the calice diameter;<br />
oval opening; 1 complete septal cycle present up to 3/4R;<br />
directive septa larger than the others, sometimes touching<br />
at bottom of calice; sec<strong>on</strong>dary septa usually absent, but if<br />
present, <strong>on</strong>ly developed to maximum 1/4R (Figure 10g).<br />
Radial corallites: Arranged in rows <strong>on</strong> branches, 2 sizes,<br />
tubular, round opening; calice large compared to radial<br />
size (Figures 10b and 10d); profile length 2.05–2.83 mm;<br />
appressed and subimmersed corallites scattered <strong>on</strong> the<br />
bases of main branches; septa poorly developed, primary<br />
septa up to 1/4R, some septa dentate, sometimes <strong>on</strong>ly<br />
directive septa present; sec<strong>on</strong>dary septa absent.<br />
316<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
Coenosteum: Covered with elaborate, laterally flattened<br />
spinules, both <strong>on</strong> and between radial corallites (Figures<br />
10e, 10f, and 10h).<br />
Color: Pale yellow or dark cream to pale brown.<br />
Remarks: So far, <strong>Acropora</strong> tortuosa has <strong>on</strong>ly been<br />
recorded in the western and central Pacific Ocean<br />
(Wallace, 1999). Its presence in the Persian Gulf is therefore<br />
remarkable. For c<strong>on</strong>firmati<strong>on</strong>, a specimen of this species<br />
was shipped to the Museum of Tropical Queensland, where<br />
it was identified as <strong>Acropora</strong> cf. tortuosa by C.C. Wallace.<br />
Later <strong>on</strong>, with more detailed studies <strong>on</strong> the morphology<br />
and other characteristics of the col<strong>on</strong>y, it became clear that<br />
the specimen was indeed <strong>Acropora</strong> tortuosa. C<strong>on</strong>sequently,<br />
this is the first record of A. tortuosa from the Persian Gulf.<br />
Morphologically, the specimens of A. tortuosa from<br />
the Persian Gulf were very similar to those described<br />
by Wallace (1999), except that the branches were a little<br />
l<strong>on</strong>ger and the walls of axial corallites were thinner in the<br />
Persian Gulf specimens.<br />
Figure 10. <strong>Acropora</strong> tortuosa: a) fresh col<strong>on</strong>y out of water; b)<br />
porti<strong>on</strong> of branch; c) porti<strong>on</strong> of col<strong>on</strong>y; d) SEM micrograph<br />
showing terminal part of branch; e) SEM micrograph showing<br />
coenosteum <strong>on</strong> a radial corallite; f) SEM micrograph showing<br />
close up view of coenosteum <strong>on</strong> radial corallite; g) top view of<br />
axial corallite; h) SEM micrograph showing coenosteum between<br />
radial corallites.
4. Discussi<strong>on</strong><br />
Because of the high morphological plasticity of staghorn<br />
corals (Todd, 2008), differences in the estimates of species<br />
diversity in these corals might be expected in any given<br />
area. Such differences, however, should be within a<br />
reas<strong>on</strong>able range. A number of studies carried out <strong>on</strong><br />
the corals of the southern part of the Persian Gulf (Riegl,<br />
1999; Wallace, 1999; Coles, 2003) showed that between<br />
11 and 13 <strong>Acropora</strong> species inhabit the area. That number<br />
for the northern and northeastern parts was <strong>on</strong>ly 5, which<br />
included <strong>Acropora</strong> clathrata (Maghsoudlou, 2008, 2011),<br />
before this <str<strong>on</strong>g>study</str<strong>on</strong>g>. In the present <str<strong>on</strong>g>study</str<strong>on</strong>g>, 9 staghorn corals<br />
were found, excluding <strong>Acropora</strong> clathrata and A. phara<strong>on</strong>is,<br />
which have frequently been reported (Maghsoudlou, 2008,<br />
2011; Mostafavi et al., 2010) from Iranian coasts of the<br />
Persian Gulf. If the records of A. phara<strong>on</strong>is were deemed<br />
valid, the number of <strong>Acropora</strong> species of the northern and<br />
northeastern parts of the Persian Gulf would reach at least<br />
10. It should also be menti<strong>on</strong>ed that from our specimens,<br />
C.C. Wallace identified A. horrida, A. mossambica, A.<br />
muricata, and A. tortuosa as ‘c<strong>on</strong>fer’ (cf.). The provisi<strong>on</strong>al<br />
identificati<strong>on</strong>s were caused by the small sizes of those<br />
specimens. More and closer examinati<strong>on</strong> of specimens<br />
from the same dives and of additi<strong>on</strong>al specimens collected<br />
later <strong>on</strong> allowed us to c<strong>on</strong>firm the identificati<strong>on</strong>.<br />
Nevertheless, taking into account the reports of other<br />
species from the southern part of the Persian Gulf, namely<br />
A. valenciennesi (Milne-Edwards and Haime, 1860 )<br />
(Sheppard and Sheppard, 1991; Wallace, 1999), A. y<strong>on</strong>gei<br />
Ver<strong>on</strong> and Wallace, 1984, A. selago (Studer, 1878), A.<br />
divaricata (Dana, 1846) (Wallace, 1999), A. tenuis (Dana,<br />
1846), and A. florida (Dana, 1846) (Riegl, 1999), the total<br />
number of species of <strong>Acropora</strong> for the Persian Gulf would<br />
be at least 16.<br />
<strong>Acropora</strong> is not <strong>on</strong>ly the richest but also the most<br />
abundant coral genus in the Persian Gulf, so much so that<br />
in some places al<strong>on</strong>g Larak Island the coverage by this<br />
genus is 100% (Rahmani, 2012). Fossil records suggest<br />
that <strong>Acropora</strong> has been the dominant coral <strong>on</strong> this island<br />
at least since the Pleistocene (Samimi-Namin and Riegl,<br />
2012). Regardless of the number of <strong>Acropora</strong> ancestral<br />
species in the Europe-Western Indian Ocean regi<strong>on</strong> during<br />
the Miocene (which were resp<strong>on</strong>sible for the diversity and<br />
dominance of the modern <strong>Acropora</strong> of the Indo-Pacific<br />
during the Plio-Pleistocene), <strong>Acropora</strong> species of the Indo-<br />
Pacific are derived from their ancestral line(s) coming from<br />
the Europe-Western Indian Ocean regi<strong>on</strong> that diversified<br />
and dispersed during the Late Oligocene to the Miocene<br />
(Wallace and Rosen, 2006). Evidence for <strong>Acropora</strong><br />
dominance in the Western Indian Ocean can be found<br />
in the Larak outcrops, too (Samimi-Namin and Riegl,<br />
2012). C<strong>on</strong>sidering the repetitious (Riegl, 1999, 2002)<br />
and periodical (Shinn, 1976) mass mortalities am<strong>on</strong>g the<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
<strong>Acropora</strong> species and their 15–20 years of successi<strong>on</strong> cycles<br />
in the Persian Gulf, and also keeping the young nature of<br />
the Persian Gulf in mind (Wallace and Muir, 2005) (water<br />
began entering the Gulf <strong>on</strong>ly 12,500 years ago (Lambeck,<br />
1996)), evaluati<strong>on</strong>s of ecological factors in additi<strong>on</strong> to<br />
evoluti<strong>on</strong>ary and historical factors are necessary in order<br />
to <str<strong>on</strong>g>study</str<strong>on</strong>g> the reas<strong>on</strong>s behind the relatively high diversity<br />
and abundance of <strong>Acropora</strong> in the Persian Gulf.<br />
However, in this part of the Persian Gulf, A. downingi<br />
and A. arabensis are abundant, and A. valida is observed<br />
to a lesser extent than the former 2 species (Maghsoudlou,<br />
2008; Rahmani, 2012).<br />
Except these 3 species, the rest can be defined as rare<br />
species, accounting for <strong>on</strong>ly about 5% of the <strong>Acropora</strong><br />
coverage (Rahmani, 2012). N<strong>on</strong>e of these corals, however,<br />
are listed as vulnerable (Askari, 2012), and no or very few<br />
special protecti<strong>on</strong> measures have been applied to c<strong>on</strong>serve<br />
these species.<br />
The necessity of c<strong>on</strong>servati<strong>on</strong> acti<strong>on</strong>s <strong>on</strong> behalf of the<br />
coral reefs in this area is obvious c<strong>on</strong>sidering the severe<br />
c<strong>on</strong>diti<strong>on</strong>s the Persian Gulf ecology must bear, especially<br />
its temperature, salinity (Coles, 1988; Coles and Fadlallah,<br />
1991; Coles 1992; Riegl, 1999; Coles, 2003), and red<br />
tides, as well as anthropological activities including wars,<br />
offshore oil drilling, tanker traffic, ballast water discharge,<br />
the c<strong>on</strong>structi<strong>on</strong> of several artificial islands, development<br />
of coastal areas, and destructi<strong>on</strong> of coral reefs by divers.<br />
The need to c<strong>on</strong>serve the corals of the Persian Gulf is even<br />
more urgent c<strong>on</strong>sidering the c<strong>on</strong>tinuous trend of global<br />
warming and the fact that the Persian Gulf is a breeding<br />
ground for warmth-tolerant species of the future.<br />
As menti<strong>on</strong>ed earlier, A. clathrata was not observed<br />
during the present work. This species has been reported as<br />
a comm<strong>on</strong> and dominant species by Maghsoudlou (2008,<br />
2011). Such an abundant, comm<strong>on</strong>, and dominant species<br />
could hardly be missed in an intensive sampling program.<br />
C<strong>on</strong>sequently, it seems that either A. clathrata does not<br />
exist in these parts of the Persian Gulf, or it occurs at a very<br />
low density (see remarks <strong>on</strong> A. downingi). However, the<br />
authors believe that the reports of A. clathrata in this area<br />
were, in fact, the result of a misidentificati<strong>on</strong>/c<strong>on</strong>fusi<strong>on</strong><br />
between the 2 forms of A. downingi. Further studies are<br />
necessary to definitely c<strong>on</strong>firm the presence or absence of<br />
A. clathrata in the Persian Gulf.<br />
Coles (2003) suggested that the coral species of the<br />
Persian Gulf are a small sample of a larger fauna of the<br />
Indo-Pacific area and maintained that the Acroporidae<br />
account for 16% of the total coral species of the Persian<br />
Gulf. This <str<strong>on</strong>g>study</str<strong>on</strong>g> changed that proporti<strong>on</strong> in the Persian<br />
Gulf to 25%. This value is much closer to the proporti<strong>on</strong><br />
of Acroporidae species to all corals species of the Indo-<br />
Pacific (30%). The 5% difference could be the result of the<br />
envir<strong>on</strong>mental stresses in the Persian Gulf.<br />
317
A review <strong>on</strong> the <strong>Acropora</strong> species of the area shows that<br />
the Persian Gulf shares coral species with both the eastern<br />
and western parts of the Indian Ocean. Biogeographical<br />
analyses of the corals of the Persian Gulf, however, point<br />
to the similarities with those elements of the Red Sea fauna<br />
(Sheppard and Sheppard, 1991). Nevertheless, when <strong>on</strong>ly<br />
<strong>Acropora</strong> species are c<strong>on</strong>sidered, the results are different,<br />
clustering the Persian Gulf with Hawaii and the East Pacific<br />
(Wallace, 1999). Repeating the biogeographical analyses<br />
with the new data and new records has not changed the<br />
outcome (Rahmani, 2012).<br />
This <str<strong>on</strong>g>study</str<strong>on</strong>g> indicates that in the area investigated, the<br />
number of <strong>Acropora</strong> species is higher than the previously<br />
reported number. We believe that the real <strong>Acropora</strong> species<br />
diversity may still be higher and that this assumpti<strong>on</strong><br />
could be extended to the other coral genera. A number of<br />
the Iranian islands surveyed in the present <str<strong>on</strong>g>study</str<strong>on</strong>g> where<br />
the <strong>Acropora</strong> diversity is relatively high, such as Larak, are<br />
located at the beginning of a migratory corridor formed<br />
by the currents entering the Persian Gulf from the Indian<br />
Ocean via the Gulf of Oman. C<strong>on</strong>sequently, all of the coral<br />
species found in the Persian Gulf <strong>on</strong>ce entered through<br />
this corridor. Therefore, we surmise that at least some of<br />
the species observed in the southern Persian Gulf and<br />
not yet found in the northern and northeastern parts of<br />
it, e.g., A. valenciennesi, A. divaricata, A. selago, A. y<strong>on</strong>gei<br />
(Wallace, 1999), A. tenuis, and A. florida (Riegl, 1999), are<br />
either too uncomm<strong>on</strong> to be easily found or have become<br />
locally extinct due to <strong>on</strong>e of the recent devastating events<br />
(such as red tide). More studies are necessary to obtain a<br />
more accurate estimate of the diversity of the corals of the<br />
Persian Gulf in general, and that of <strong>Acropora</strong> in particular.<br />
Tax<strong>on</strong>omic key to the <strong>Acropora</strong> of the Persian Gulf<br />
A: Col<strong>on</strong>y and growth form<br />
A 1 ) table and growth determinate ………...........… B<br />
A 2 ) corymbose to caespitose and growth determinate<br />
or semi-determinate …....................................... C<br />
A 3 ) other growth forms …................…….....….….. H<br />
B: Col<strong>on</strong>y and radial corallites characters<br />
B 1 ) col<strong>on</strong>y flat; sometimes with upward branches;<br />
radial corallites tubular with sharp-pointed<br />
outer walls extending al<strong>on</strong>g the branches .............<br />
.............................................................. A. downingi<br />
B 2 ) sturdy and intertwining branches form stalk;<br />
branchlets present …........................ A. phara<strong>on</strong>is<br />
C: Radial corallites’ shape<br />
C 1 ) nariform or tubo-nariform …………...…..….. D<br />
C 2 ) appressed ………………………..…….....…… F<br />
C 3 ) cochleariform ….........….………............…….. G<br />
D: Col<strong>on</strong>y and axial corallites characters<br />
D 1 ) corymbose; axial corallites large (outer diameter<br />
2.8–4.3 mm) …...............................................….. E<br />
318<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
D 2 ) caespitose-corymbose; axial corallites smaller<br />
than former (outer diameter 1.8–3.0 mm); axial<br />
corallites’ primary septa 1/2R, sec<strong>on</strong>dary septa<br />
up to 1/4R ........................................... A. divaricata<br />
E: Col<strong>on</strong>y, axial, and radial corallites’ characters<br />
E 1 ) axial corallites’ outer diameter 2.8–3.5 mm; axial<br />
primary and sec<strong>on</strong>dary septa 2/3R and 1/3R,<br />
respectively; radial corallites mostly nariform<br />
…............................................................... A. nasuta<br />
E 2 ) branches growing horiz<strong>on</strong>tally, forming plate;<br />
axial corallites larger than former (outer diameter<br />
3.1–4.3 mm); axial primary and sec<strong>on</strong>dary septa<br />
1/3R and 1/5R, respectively; radial corallites<br />
mostly tubo-nariform ……..…... A. mossambica<br />
F: Col<strong>on</strong>y, axial, and radial corallites’ characters<br />
F 1 ) corymbose to caespitose; growth determinate;<br />
axial corallites’ primary septa 2/3R; axial<br />
sec<strong>on</strong>dary septa up to 1/3R; tertiary septa rarely<br />
present; radial corallites’ profile length 2.0–3.3<br />
mm; radial primary and sec<strong>on</strong>dary septa 2/3 and<br />
1/3R, respectively …................................ A. valida<br />
F 2 ) caespitose; growth semideterminate; axial<br />
corallites’ primary and sec<strong>on</strong>dary septa 3/4 and<br />
1/4R, respectively; radial outer walls extended<br />
al<strong>on</strong>g the branches ............................. A. arabensis<br />
G: Branches’ and axial and radial corallites’ characters<br />
G 1 ) branch length up to 90 mm; axial corallites’ outer<br />
diameter 1.8–3.4 mm; radial primary septa 2/3R<br />
..........………......……………….……… A. tenuis<br />
G 2 ) branches shorter than former (up to 40 mm);<br />
axial corallites’ outer diameter smaller than<br />
H 1 above (up to 1.1–2.4 mm); radial corallites<br />
crowded, radial primary septa 1/3R ...... A. selago<br />
H: Radial corallites’ shape<br />
H 1 ) tubular or appressed tubular ………….....……. I<br />
H 2 ) cochleariform ...…….............….......….. A. y<strong>on</strong>gei<br />
H 3 ) labellate …....................…............….….. A. aspera<br />
I: Branches’ appearance and radial corallites’ characters<br />
I 1 ) extremely fragile; branches with ragged<br />
appearance ………..........................….. A. horrida<br />
I 2 ) branches and radial corallites unlike the former<br />
…...….…....................................................……… J<br />
J: Axial and radial corallites’ characters<br />
J 1 ) axial corallite outer diameter 2.0–2.8 mm; axial<br />
sec<strong>on</strong>dary septa absent or 1/4R; round opening;<br />
radial primary septa up to 1/4R, sec<strong>on</strong>dary septa<br />
absent ……………………………… A. tortuosa<br />
J 2 ) axial corallites larger than those of A. tortuosa<br />
(outer diameter 2.5–3.2 mm); axial sec<strong>on</strong>dary<br />
septa up to 2/3R; oval opening; radial corallites’<br />
septa well developed, primary and sec<strong>on</strong>dary<br />
septa up to 3/4 and 2/3R, respectively ..................<br />
............................................................... A. muricata
Acknowledgments<br />
This <str<strong>on</strong>g>study</str<strong>on</strong>g> was made possible thanks to the extensive help of<br />
Dr C.C. Wallace, Museum of Tropical Queensland, Australia.<br />
Without her help the identificati<strong>on</strong>s could not have been<br />
c<strong>on</strong>firmed. We express our gratitude to the administrati<strong>on</strong><br />
of the Hormozgan and Bushehr Envir<strong>on</strong>mental Protecti<strong>on</strong><br />
Agency and the Iranian Offshore Oil Company for their<br />
valuable logistic support during sampling. Mr Hashemi,<br />
University of Tehran, carried out the scanning electr<strong>on</strong><br />
References<br />
Abdel-Moati, M.A. 2011. Coral reef c<strong>on</strong>servati<strong>on</strong> in Qatar. http://<br />
assets.panda.org/downloads/coral_reefs_qatar_mohamed_<br />
alaa_abdel_moati.pdf. Accessed 10 May 2011.<br />
Askari, A. 2012. C<strong>on</strong>served coral species of Iran. http://www.<br />
hormozgandoe.ir/pages.php?number=3.3.2. Accessed 7 July<br />
2012.<br />
Brook, G. 1891. Descripti<strong>on</strong>s of new species of Madrepora in the<br />
collecti<strong>on</strong>s of the British Museum. Ann. Mag. Nat. Hist. 8:<br />
458–471.<br />
Brook, G. 1893. The genus Madrepora. Cat. Mad. Corals Brit. Mus.<br />
(NH) 1: 1–212.<br />
Chen, I.P., Tang, C.Y., Chiou, C.Y., Hsu, J.H., Wei, N.V. and Wallace,<br />
C.C. 2009. Comparative analyses of coding and n<strong>on</strong>coding<br />
DNA regi<strong>on</strong>s indicate that <strong>Acropora</strong> (Anthozoa: Scleractina)<br />
possesses a similar evoluti<strong>on</strong>ary tempo of nuclear vs.<br />
mitoch<strong>on</strong>drial genomes as in plants. Mar. Biotechnol. (NY) 11:<br />
141–152.<br />
Coles, S.L. 1988. Limitati<strong>on</strong>s <strong>on</strong> reef coral development in the Arabian<br />
Gulf: temperature or algal competiti<strong>on</strong>. In: Proceedings of 6th<br />
Internati<strong>on</strong>al Coral Reef Symposium (Eds., J.H. Choat, D.<br />
Barnes, M.A. Borowitzka, J.C. Coll, P.J. Davies, P. Flood, B.G.<br />
Hatcher, D. Hopley, P.A. Hutchings, D. Kinsey, G.R. Orme, M.<br />
Pich<strong>on</strong>, P.F. Sale, P. Sammarco, C.C. Wallace, C. Wilkins<strong>on</strong>, E.<br />
Wolanski and O. Bellwood), Vol. 3. Townsville, Australia, pp.<br />
211–216.<br />
Coles, S.L. 1992. Experimental comparis<strong>on</strong> of salinity tolerances of<br />
reef corals from the Arabian Gulf and Hawaii. Evidence for<br />
hyperhaline adaptati<strong>on</strong>. In: Proceedings of 7th Internati<strong>on</strong>al<br />
Coral Reef Symposium (Ed., R.H. Richm<strong>on</strong>d), Vol. 1.<br />
University of Guam Press, UOG Stati<strong>on</strong>, Guam, pp. 227–234.<br />
Coles, S.L. 2003. Coral species diversity and envir<strong>on</strong>mental factors in<br />
the Arabian Gulf and the Gulf of Oman: a comparis<strong>on</strong> to the<br />
Indo-Pacific regi<strong>on</strong>. Atoll Res. Bull. 507: 1–19.<br />
Coles, S.L. and Fadlallah, Y.H. 1991. Reef coral survival and mortality<br />
at low temperatures in the Arabian Gulf: new species-specific<br />
lower temperature limits. Coral Reefs 9: 231–237.<br />
Crossland, C.J. 1952. Madreporaria, Hydrocoralinae, Heliopora<br />
and Tubipora. <strong>Scientific</strong> Reports of the Great Barrier Reef<br />
Expediti<strong>on</strong> 1928-29 British Museum (Natural History) 6: 85–<br />
257.<br />
Dana, J.D. 1846. Structure and Classificati<strong>on</strong> of Zoophytes. Lea and<br />
Blanchard, Philadelphia.<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
microscopy and Mr Mohajeran, Nati<strong>on</strong>al Museum of<br />
Natural History, carried out the microphotography. We wish<br />
to thank them. We also wish to thank Dr Fazel, Dr Shirdam,<br />
Dr M<strong>on</strong>tazami, Captain Mansour, Captain Sorori, Captain<br />
Hoseini, Mr Namdar, Mr Shokri, Mr Moien, Mr Tollab, Mr<br />
Hadi, and Mr Shirzad for their help and support during<br />
sampling. This <str<strong>on</strong>g>study</str<strong>on</strong>g> was partially financed by the Office<br />
of Research Affairs, University of Tehran; their support is<br />
hereby acknowledged.<br />
Eguchi, M. and Shirai, S. 1977. Ecological Encyclopaedia of the<br />
Marine Animals of the Ryukyu Islands Japan. Kyoiku Shuppan,<br />
Tokyo.<br />
Fukami, H., Chen, C., Chiou, C.Y. and Knowlt<strong>on</strong>, N. 2007. Novel<br />
group i intr<strong>on</strong>s encoding a putative homing end<strong>on</strong>uclease in<br />
the mitoch<strong>on</strong>drial cox1 gene of scleractinian corals. J. Mol.<br />
Evol. 64: 591–600.<br />
Fukami, H., Omori, M., Shimoike, K., Hayashibara, H. and Hatta, M.<br />
2003. Ecological and genetic aspects of reproductive isolati<strong>on</strong><br />
by different spawning times in <strong>Acropora</strong> corals. Mar. Biol. 142:<br />
679–684.<br />
Hodgs<strong>on</strong>, G. and Carpenter, K. 1996. Scleractinian corals of Kuwait.<br />
Pac. Sci. 49(3): 227–246.<br />
Kämpf, J. and Sadrinasab, M. 2006. The circulati<strong>on</strong> of the Persian<br />
Gulf: a numerical <str<strong>on</strong>g>study</str<strong>on</strong>g>. Ocean Sci. 2: 27–41.<br />
Klunzinger, C.B. 1879. Die Korallenthiere des Rothen Meeres.<br />
Gutmann, Berlin.<br />
Lambeck, K. 1996. Shoreline rec<strong>on</strong>structi<strong>on</strong>s for the Persian Gulf<br />
since the last glacial maximum. Earth Planet. Sci. Lett. 142:<br />
43–57.<br />
Linnaeus, C. 1758. Systema Naturae. Editio decima reformata<br />
Laurentii Salvii, Holmiae.<br />
Maghsoudlou, A. 2008. Hard Corals of the Iranian Coastal Waters of<br />
the Persian Gulf. Iranian Nati<strong>on</strong>al Institute for Oceanography<br />
Press, Tehran.<br />
Maghsoudlou, A. 2011. Hard coral species in some Iranian coastal<br />
areas of the Persian Gulf. Iranian Nati<strong>on</strong>al Institute for<br />
Oceanography. http://inio.ac.ir/coralreefs/hard%20coral%20<br />
species%20of%20Iran2.htm. Accessed 10 April 2011.<br />
Meyer, C. and Paulay, G. 2000. Indo-west Pacific diversity:<br />
phylogenetic evidence from cowries for a mozaic of causes.<br />
Proceedings of 9th Internati<strong>on</strong>al Coral Reef Symposium, 23–<br />
27 October 2000, Bali, Ind<strong>on</strong>esia, p. 45.<br />
Milne-Edwards, H. and Haime, J. 1860. Histoire Naturelle des<br />
Coralliaires,Vol. 3. Roret, Paris.<br />
Mostafavi, H., Kiabi, B., Liaghati, H., Abdoli, A., Mehrabian, A.R.,<br />
Khoshbakht, K., Najmizadeh, S., Ahmadzadeh, F., Samimi<br />
Namin, K., Bakhtiari, M., Jozaei, J., Delshab, H., Rezaei, M.,<br />
Moradi, A. and Mahdaviani, A. 2010. Management Plan of<br />
Kharkoo Wildlife Refuge. Maaref Publishers, Tehran.<br />
319
Nezlin, N.P., Polikarpov, I.G. and Al-Yamani, F. 2007. Satellitemeasured<br />
chlorophyll distributi<strong>on</strong> in the Arabian Gulf: spatial,<br />
seas<strong>on</strong>al and inter-annual variability. Int. J. Oceans Oceanogr.<br />
2: 139–156.<br />
Nemenzo, F. 1967. Systematic studies <strong>on</strong> Philippine shallow-water<br />
scleractinians. VI. Suborder Astrocoeniina (M<strong>on</strong>tipora and<br />
<strong>Acropora</strong>). Nat. Appl. Sci. Bull. Univ. Philipp. 20: 1–141.<br />
Oken, L. 1815. Steinkorallen. Lehrbuch Naturgesch. 3: 59–74.<br />
Ortmann, A. 1889. Beobachtungcn an Steinkorallen v<strong>on</strong> der<br />
Sudkuste Ceyl<strong>on</strong>s. Zool. Jahrb. Abt. Syst. Geogr. Biol. Tiere 4:<br />
493–590.<br />
Quelch, J. 1886. Report <strong>on</strong> the reef corals collected by H.M.S.<br />
Challenger during the years 1873–76. Reports of the <strong>Scientific</strong><br />
Results of the Voyage of HMS Challenger Zoology 3: 1–203.<br />
Rahmani, M.R. 2012. Biosystematic, biogeographic and biodiversity<br />
studies of genus <strong>Acropora</strong> (Cnidaria: Anthozoa) in the Larak,<br />
Farur and Khark Islands. PhD. University of Tehran, Tehran.<br />
Riegl, B. 1995. Descripti<strong>on</strong> of four new species in the hard coral<br />
genus <strong>Acropora</strong> Oken, 1815 (Scleractinia: Astrocoeniina:<br />
Acroporidae) from south-east Africa. Zool. J. Linn. Soc. L<strong>on</strong>d.<br />
113: 229–247.<br />
Riegl, B. 1999. Corals in a n<strong>on</strong>-reef setting in the southern Arabian<br />
Gulf (Dubai, UAE): fauna and community structure in<br />
resp<strong>on</strong>se to recurring mass mortality. Coral Reefs 18: 63–73.<br />
Riegl, B. 2002. Phoenix from the ashes? Repetitive mass mortality<br />
and the biogeology of southern Arabian Gulf (United Arab<br />
Emirates) coral systems. In: The Vienna School of Marine<br />
Biology: A Tribute to Jörg Ott (Eds., M. Bright, P. Dworschak<br />
and M. Stachowitsch). Facultas Universitätsverlag, Vienna, pp<br />
137–159.<br />
Samimi-Namin, K. and Riegl, B. 2012. Raised reef <strong>on</strong> Larak suggests<br />
<strong>Acropora</strong> dominance <strong>on</strong> the Persian Gulf coral reefs since the<br />
Pleistocene. Coral Reefs 31: 763.<br />
Sheppard, C.R.C. and Sheppard, A.L.S. 1991.Corals and coral<br />
communities of Arabia. Fauna of Saudi Arabia. 12: 1–170.<br />
Shinn, E.A. 1976. Coral reef recovery in Florida and the Persian Gulf.<br />
Envir<strong>on</strong>. Geol. 1: 241–254.<br />
Spalding, M.D., Ravilious, C. and Green, E.P. 2001. World Atlas of<br />
Coral Reefs. University of California Press, H<strong>on</strong>g K<strong>on</strong>g.<br />
Studer, T. 1878. Zweite Abtheillung der Anthozoa Polyactinia,<br />
welche wahrend der Reise S.M.S. Corvette Gazelle um die<br />
Erde gesammelt wurden. M<strong>on</strong>atsberichte der Königlich<br />
Preussischen Akademie der Wissenschaften zu Berlin, Berlin.<br />
320<br />
RAHMANI and RAHIMIAN / Turk J Zool<br />
Todd, P.M. 2008. Morphological plasticity in scleractinian corals.<br />
Biol. Rev. 83: 315–337.<br />
van Oppen, M.J.H., McD<strong>on</strong>ald, B.J., Willis, B. and Miller, D.J.<br />
2001. The evoluti<strong>on</strong>ary history of the coral genus <strong>Acropora</strong><br />
(Scleractinia, Cnidaria) based <strong>on</strong> a mitoch<strong>on</strong>drial and a<br />
nuclear marker: reticulati<strong>on</strong>, incomplete lineage sorting, or<br />
morphological c<strong>on</strong>vergence? Mol. Biol. Evol. 18: 1315–1329.<br />
Vaughan, T.W. 1906. Report <strong>on</strong> the scientific results of the expediti<strong>on</strong><br />
to the eastern tropical Pacific VI Madreporaria. Bull. Mus.<br />
Comp. Zool. 50: 59–72.<br />
Ver<strong>on</strong>, J.E.N. 2000. Corals of the World. Australian Institute of<br />
Marine Science Publisher, Townsville.<br />
Ver<strong>on</strong>, J.E.N. and Wallace, C.C. 1984. Scleractinia of Eastern<br />
Australia. Part V. Family Acroporidae. Aust. Inst. Mar. Sci.<br />
M<strong>on</strong>ogr. Ser. 6: 1–485.<br />
Verrill, A.E. 1902. Notes <strong>on</strong> corals of the genus <strong>Acropora</strong> (Madrepora<br />
Lam.) with new descripti<strong>on</strong>s and figures of types, and of several<br />
new species. Trans. C<strong>on</strong>n. Acad. Arts. Sci. 11: 207–266.<br />
Wallace, C.C. 1978. The coral genus <strong>Acropora</strong> (Scleractinia:<br />
Astrocoeniina: Acroporidae) in the central and southern Great<br />
Barrier Reef Province. Mem. Queensl. Mus. 18: 273–319.<br />
Wallace, C.C. 1999. Staghorn corals of the world: a revisi<strong>on</strong> of the coral<br />
genus <strong>Acropora</strong> (Scleractinia; Astrocoeniina; Acroporidae)<br />
worldwide, with emphasis <strong>on</strong> morphology, phylogeny and<br />
biogeography. CSIRO Publishing, Victoria, Australia.<br />
Wallace, C.C. and Muir, P.R. 2005. Biodiversity of the Indian Ocean<br />
from the perspective of staghorn corals (<strong>Acropora</strong> spp). Indian<br />
J. Mar. Sci. 34(1): 42–49.<br />
Wallace, C.C. and Rosen, B.R. 2006. Diverse staghorn corals<br />
(<strong>Acropora</strong>) in high-latitude Eocene assemblages: implicati<strong>on</strong>s<br />
for the evoluti<strong>on</strong> of modern diversity patterns of reef corals. P.<br />
Roy. Soc. B 273: 975–982.<br />
Wolstenholme, J. 2004. Temporal reproductive isolati<strong>on</strong> and gametic<br />
compatibility are evoluti<strong>on</strong>ary mechanisms in the <strong>Acropora</strong><br />
humilis species group (Cnidaria; Scleractinia). Mar. Biol. 144:<br />
567–582.<br />
Wolstenholme, J.K., Wallace, C.C. and Chen, C.A. 2003. Species<br />
boundaries within the <strong>Acropora</strong> humilis species group<br />
(Cnidaria; Scleractinia): a morphological and molecular<br />
interpretati<strong>on</strong> of evoluti<strong>on</strong>. Coral Reefs 22: 155–166.