Marine Ecology. ISSN 0173-9565
ORIGINAL ARTICLE
Sponges boring into precious corals: an overview with
description of a new species of Alectona (Demospongiae,
Alectonidae) and a worldwide identification key for the
genus
Barbara Calcinai1, Carlo Cerrano2, Nozomu Iwasaki3 & Giorgio Bavestrello1
1 DiSMar, Università Politecnica delle Marche, Ancona, Italy
2 DipTeRis, Università di Genova, Genova, Italy
3 Usa Marine Biological Institute, Kochi University, Kochi, Japan
Keywords
Alectona; boring sponge; Japan Sea; new
species; Paracorallium japonicum.
Correspondence
Giorgio Bavestrello, DiSMar, Università
Politecnica delle Marche, Via Brecce Bianche,
60131 Ancona, Italy.
E-mail: g.bavestrello@univpm.it
Accepted: 24 April 2008
doi:10.1111/j.1439-0485.2008.00246.x
Abstract
Precious corals represent peculiar substrata for several species of boring
sponges that exploit their carbonatic scleraxis, strongly decreasing their commercial value. Here we describe a new species of the genus Alectona from Japan
recorded in a colony of Paracorallium japonicum (Kishinouye, 1903). The spicular complement of the new species consists of diactinal spicules covered by
mushroom-like tubercles, often modified into styloid forms, and fusiform amphiasters with two or sometimes four verticils, each generally made up of six
short tubercled rays. A complete survey of the literature on boring sponges
recorded in precious corals in the Mediterranean Sea and Pacific Ocean indicates that the species of the family Alectonidae are the most strictly associated
to this kind of substratum. Their world distribution, in fact, partially or totally
overlap that of their coral hosts.
Problem
The genera Corallium and Paracorallium (Octocorallia,
Corallidae) are known all over the world due to their
heavy exploitation by the jewel industry. Considering
their generally slow growth rate (Bramanti et al. 2005;
Roark et al. 2006) there are several international projects devoted to their protection and management. The
main areas where the precious coral fishery is strongly
active are the Mediterranean for the red coral Corallium
rubrum (Cicogna et al. 1999) and the Pacific Ocean for
several other species (Parrish & Baco 2007; UICN
2007).
Precious coral colonies are characterised by a scleraxis
composed of Mg-calcite organised in an extremely compact structure of acicular to lamellar calcite crystals that
grow centripetally, forming fan-like structures (Cortesogno et al. 1999). Often the collected colonies cannot be
carved owing to the presence of boring sponges, strongly
decreasing the commercial value of the coral. This is one
of the most economically damaging defects to be found
in commercial stocks (Liverino 1983). When occupied by
boring sponges, the scleraxis of the precious coral shows
a series of chambers interconnected by channels, giving
rise to a ‘‘spongy’’ pattern in the carbonate. Information
about which sponge species are involved in this boring
activity, their relative destructive power, their distribution
and their relationship with the affected coral colonies will
help both the coral industry and coral scientists to draw
together adequate measures for management and exploitation of this resource.
The genus Alectona (Demospongiae) is characterised by
the presence of diactinal, sometimes polyactinal, spicules,
covered with spines and tubercles and fusiform or
branched amphiasters (Rützler 2002). The pitting pattern
of the excavated chambers, characterised by concentric
Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
273
Calcinai, Cerrano, Iwasaki & Bavestrello
Sponges boring into precious corals
grooves overlapped by radiating scars, can be used as an
additional feature in the diagnosis of this genus (Vacelet
1999; Calcinai et al. 2004a).
This group of sponges shows a strong affinity for biogenic substrata. The first two species described for the
genus, Alectona millari Carter, 1879 and Alectona wallichii Carter, 1874, were initially recorded as boring into
calcareous rocks. However, the former was later
observed in Mediterranean red coral (Maldonado 1992)
and the latter in Pacific precious corals (Bavestrello et al.
1998). In 1899, Johnson recorded Alectona verticillata
from Madeira excavating colonies of Corallium, later redescribed by Calcinai et al. (2004b) boring into Corallium elatius Ridley, 1882 from the Japan Sea. Alectona
primitiva Topsent, 1932 was described in the mollusc
Haliotis excavata Lamarck. Pang (1973) described Alectona jamaicensis excavating into a colony of Acropora
and Bavestrello et al. (1998) recorded Alectona triradiata
Lévi & Lévi, 1983 in a colony of C. elatius. The species
Alectona sorrentini Bavestrello et al. (1998) and Alectona
microspiculata Bavestrello et al. (1998) were recorded
respectively in C. elatius and in the hydrocoral Distichopora sp. Vacelet (1999) described the deep Alectona mesatlantica excavating into a calcareous rock that was
probably biogenic.
The aim of this paper is to describe a new species of
the genus Alectona recorded in a colony of Paracorallium
japonicum (Kishinouye, 1903) from the Japan Sea. The
new species is discussed in the framework of a complete
bibliographic survey of all the boring species recorded in
precious corals in the Mediterranean and the Pacific
Ocean.
Methods
The material consists of portions of a branch of a colony
of P. japonicum collected by Mr Yoshihiko Niiya off Ashizuri Cape (Kochi, Japan) at 120 m depth.
Spicule preparations were made by dissolving sponge
fragments in boiling nitric acid, and after rinsing in water
and ethanol, mounting the spicules on microscope slides.
The same preparation was followed for the scanning electron microscope observation. In this case, spicules were
put on stubs and sputtered with gold.
Ranges and mean spicule size (in brackets) were calculated from 15 measurements for each spicule type.
The type material of the new species is deposited at the
Museo Civico di Storia Naturale di Genova (MSNG).
Results
The examined sample of P. japonicum was collected alive,
as suggested by the covering of intact coenosarc on the
274
mineral scleraxis. The coral is excavated by discrete,
subspherical chambers (0.9–1.8 mm in diameter, 1.3 mm
on average) present mainly in the central part of the
scleraxis. The chambers communicate by numerous small
foramina (0.2–0.4 mm, 0.3 mm on average). These
chambers were produced by the boring activity of
Holoxea excavans Calcinai et al. 2001, a common borer
of Japanese precious corals. This attribution is due both
to the large number of the spicules of this species (oxeas
of two sizes) filling the boring chambers and to the pitting pattern with a fingerprint-like surface (Calcinai et al.
2004a).
Some of the peripheral chambers produced by
H. excavans were filled with densely packed groups of diactinal spicules that we attribute to a new species of the
genus Alectona described below.
Family Alectonidae Rosell, 1996
Genus Alectona Carter, 1879
Alectona sarai n. sp.
Material examined
Holotype MSNG 54326. Portions of P. japonicum, dry
preserved and one spicule slide, 5 ⁄ 05 ⁄ 2007, Ashizuri
Cape, Kochi, Japan; other material: two slides in the
authors’ collection.
Skeleton
Proper erosion chambers of Alectona sarai n. sp. were not
identified by SEM analysis. In fact, all the examined excavated portions showed scars with a fingerprint-like surface
characteristic of Holoxea (Calcinai et al. 2004a), whereas a
radial pattern overlapping deep concentric lines was not
observed (Vacelet 1999; Calcinai et al. 2004a).
The skeleton of the new species of Alectona is made
up of:
1 Diactinal spicules (Fig. 1A–C), 175–250 lm long and
12.5–20 lm wide (207.5 ± 27.3 · 17 ± 3 lm); they are
generally flexuous, sometimes with a central bend
(Fig. 1B). The axial canal is often evident. The tips are
sharp and often end with an apical single spine (Fig. 1A–
C). These spicules are covered with long mushroomshaped tubercles 5–7.5 lm long, arranged in alternate
rows (Fig. 1D). On a few spicules the tubercles are transformed into spines. Several spicules, about half of the
total, have a rounded extremity and resemble stout papillose, sometimes bent, styles (Fig. 1E–H).
2 Fusiform, straight amphiasters with a long and generally blunt axis (Fig. 1I–M); 20–40 lm (27 ± 4.9 lm) long
and 2.5–3.7 lm thick. In the middle of the axis, two
Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
Calcinai, Cerrano, Iwasaki & Bavestrello
Sponges boring into precious corals
A
B
E
I
C
F
J
D
G
K
H
L
M
Fig. 1. Spicules of Alectona sarai n. sp. (A–C)
Diactinal tubercled spicules. (D) Magnification
of the surface of a diactinal spicule showing
the mushroom-shaped tubercles arranged in
alternate rows. (E–G) Spicules modified into
styles. (H) Magnification of the rounded
extremity of a style. (I) Fusiform amphiaster
with two verticils of six short tubercled rays.
(J–L) Amphiasters with nodules arranged in
two accessory distal verticils and scattered
along the axis. (M) Amphiaster with a short
axis ending with groups of spiny knobs. Scale
bars: (A–C) 50 lm; (D,H–M) 10 lm; (E–G)
70 lm.
verticils, each generally made up of six (sometimes fewer)
short tubercled rays, are present (Fig. 1I). Numerous nodules are often arranged in two other accessory distal verticils or are scattered along the axis (Fig. 1J–L). Several of
the latter spicules have one extremity of the axis which is
shorter and thinner, ending with groups of spiny knobs
(Fig. 1M).
Etymology
The species was named in memory of our mentor Prof.
Michele Sarà.
Taxonomic Remarks
The species of the genus Alectona are characterised by
amphiastrose microscleres which are very variable in
shape: they include amphiasters with a small centre and
slender long rays, amphiasters with a stout diactine axis
(Rützler 2002) and nodular amphiasters with a short axis
and microspinations mainly grouped at the extremities.
We suggest the term ‘nodular amphiasters’ for the microscleres described as small amphiasters by Rützler
(2002). They often have a thick axis and may resemble
the nodular amphiasters typical of Thoosa (compare for
Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
275
Calcinai, Cerrano, Iwasaki & Bavestrello
Sponges boring into precious corals
example Fig. 16, b page, 184 and Fig. 1, I page, 282 in
Rützler 2002).
We suggest the term ‘fusiform’ for the amphiasters
with a stout diactine axis and two verticils of spiny knobs,
and the term ‘oxyamphiasters’ for the third kind of microsclere with hispid long oxyform axis (Fig. 1, H page
282 in Rützler 2002).
Alectona sarai n. sp. is characterised by short diactines,
often transformed into styloid forms, with long flattened
tubercles and by fusiform amphiasters with two or often
four verticils of spined knobs which vary in shape.
Six Alectona species have been recorded in the IndoPacific area (Table 1): Alectona microspiculata, Alectona
primitiva, Alectona sorrentini, Alectona triradiata, Alectona
verticillata and Alectona wallichii. The species most closely related to A. sarai n. sp. is A. wallichii, as the shape
of the diactinal spicules is characterised by mushroomlike tubercles (Vacelet 1999), although it differs in the
size of these spicules, which are longer in A. wallichii
(Bavestrello et al. 1998; Vacelet 1999), and in the shape
and arrangement of the tubercles, which are flat and
closely spaced in A. wallichii. The latter species is also
characterised by fusiform amphiasters with two regular
verticils different in shape from those of A. sarai n. sp.
Moreover, the smooth diactines regularly present in
A. wallichii have never been observed in A. sarai n. sp.
The other species known in the area are also different in
the shape and size of the spicules (Table 1). Alectona
microspiculata has short, sausage-shaped diactines (30–
60 · 7–10 lm) and two kinds of ramose and short amphiasters (nodular and oxyamphiasters). Alectona primitiva has longer, spined diactines (680 · 30 lm) and short
amphiasters with tylote extremities (nodular amphiasters) (Vacelet & Vasseur 1971). Alectona sorrentini has
diactines covered with short spines and spiny, ramose
amphiasters (oxyamphiasters), different in shape from
those of A. sarai n. sp. Alectona triradiata has spiny
triactines and two kinds of non-fusiform amphiasters,
very different in shape from those of A. sarai n. sp.
Alectona verticillata is characterised by three kinds of
amphiasters (nodular, fusiform and oxyamphiasters) and
by the lack of diactinal spicules (Calcinai et al. 2004b).
This new species is also similar to A. millari from the
Mediterranean Sea and North Atlantic Ocean for the size
of the diactinal spicules and fusiform amphiasters.
Alectona sarai n. sp. differs for the presence of tubercles
instead of long spines on the diactinal spicules and the
presence of irregular amphiasters; moreover, we have not
observed the modified diactines that are always present in
A. millari.
This new record brings the number of Alectona species
to 10 (Table 1) and we propose the following key for
their identification:
276
(1) Triactinal spicules
Not as above
(2) Diactinal spicules absent (no
macroscleres)
Diactinal spicules present
(3) Diactinal spicules covered with
mushroom-shaped tubercles
Not as above
(4) Diactinal spicules long (>300 lm),
up to 800 lm; fusiform
amphiasters with regular verticils;
smooth diactines may be present
Diactinal spicules short (<300 lm),
irregular fusiform amphiasters, no
smooth diactines
(5) Very short diactinal spicules
(<100 lm), sausage-shaped
Not as above
(6) Only fusiform amphiasters as
microscleres
Not as above
(7) Fusiform amphiasters characterised
by long rays bearing small spines
at the top
Not as above
(8) Only nodular amphiasters
Only oxyamphiasters
(9) Long diactines (>400 lm), no
polyactinal spicules, large pits
(up to 100 lm)
Shorter diactines (<400 lm),
polyactinal spicules, small pits
(about 50 lm)
Alectona triradiata
(2)
Alectona verticillata
(3)
(4)
(5)
Alectona wallichii
Alectona sarai n. sp.
Alectona microspiculata
(6)
(7)
(8)
Alectona jamaicaensis
(9)
Alectona primitiva
Alectona sorrentini
Alectona mesatlantica
Alectona millari
Ecological Remarks
In the Mediterranean, during a series of surveys conducted on different red coral populations, 14 boring species were identified (Melone 1965; Barletta & Vighi 1968;
Corriero et al. 1988; Calcinai et al. 2002, 2004a, 2007).
The same number was recorded during our studies
(Bavestrello et al. 1995, 1996, 1998; Calcinai et al. 2001,
2002, 2004b) on the Pacific species of precious corals.
Twenty six species were involved in boring activity in the
precious corals, with only two species, Dotona pulchella
Carter, 1880 and A. verticillata, being common to both
geographic areas, and Delectona ciconiae Bavestrello,
Calcinai & Sarà, 1996 being recorded only in the Mediterranean coral (Fig. 2, Table 2).
There are eight genera involved: Holoxea, Alectona,
Delectona, Thoosa, Spiroxya, Cliona, Dotona, Aka. The most
represented genera are Alectona and Spiroxya (family Alectonidae), both with six species, whereas Cliona (family
Clionaidae), which is very common in calcareous, organic
substrata such as coral reefs, is poorly represented. There
are two species of Alectona in the Mediterranean and five
Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
Species
Diactinal
spicules
(lm)
Alectona
jamaicaensis
Alectona
microspiculata
Alectona
mesatlantica
a) 186–237 · 9.6–12
b) 186–287 · 6.1–8.5
30–60 · 7–10
Alectona millari
215–370 · 15–20
Alectona primitiva
170 · 8
Alectona sorrentini
Alectona triradiata
148–239 · 14–18
Alectona wallichii
a) 380–500 · 40–50
b) 280–490 · 10–28
Poliactinal Nodulose
Fusiform
spicules
amphiasters amphiasters
(lm)
(lm)
(lm)
31–74
· 1.4–4.3
7–10 · 1
410–530 · 21–28
127 · 6.3
12–14 · 5
23–37
49–58
15–35
7.5–13
32.5–70
· 3.7–7.5
20–40
· 2.5–3.7
Substratum
Chambers Chamber
size (mm) shape
Depth (m)
Ref.
Jamaica
Porites furcata
5.8–7.9
14
(7)
Philippines
Disticophora sp.
not stated
(1)
Equatorial
organic,
up to 50
Mid-Atlantic
calcareous rock
Ridge
Mediterranean, calcareous rock,
North Atlantic, coral,
Red Sea
Corallium rubrum
10–70 · 1–5
7–10 · 2.5
Distribution
15–30
(rays)
Australia
Haliotis excavata
(Cangaroo
Island)
· 1.5–3 Japan
Corallium elatius
·2
New Caledonia, Corallium elatius;
Japan
calcareous
conglomerate
limestone rocks,
Indian Ocean,
South
Corallium elatius
Australia
· 2.5
Mediterranean, Corallium elatius
Japan
Japan
Paracorallium
japonicum
0.6–0.8
spherical,
ellipsoidal
hemispherical
single,
2030
subspherical,
irregular
500–660 (3);
202 (8);
70–120 m (2);
shallow water
(9, 10)
small, closed
3 (11)
spaced
(1)
(1), (2), (3),
(8), (9), (10)
(1), (11)
large, irregular
1–6
irregularly
· 1.5–3.5 spherical,
deep
5–6
large,
subspherical,
irregular
20 · 25
large, rounded
not stated
290–350
(1)
(1), (4)
10; 146–182
(1), (5)
195–300
(6)
not stated not stated
not stated
(12)
(1) Bavestrello et al. 1998; (2) Maldonado 1992; (3) Topsent, 1920; (4) Lévi & Lévi, 1983; (5) Vacelet 1999; (6) Calcinai et al. 2004b; (7) Pang 1973; (8) Carter, 1879; (9) Sarà, 1959; (10) Rützler,
1966; (11) Vacelet & Vasseur 1971; (12) present paper.
277
Sponges boring into precious corals
Alectona sarai n. sp. 175–250 · 12.5–20
20–30 · 1
20–60 · 3–7
79–84.6
· 28
Alectona verticillata
Oxy
amphiasters
(lm)
Calcinai, Cerrano, Iwasaki & Bavestrello
Marine Ecology 29 (2008) 273–279 ª 2008 The Authors. Journal compilation ª 2008 Blackwell Publishing Ltd
Table 1. Principal features of the species of the genus Alectona (Bavestrello et al. 1998 mod.).
Calcinai, Cerrano, Iwasaki & Bavestrello
Sponges boring into precious corals
Table 2. Boring sponge species recorded in species of the coral
genus Corallium from the Mediterranean Sea and the Pacific Ocean.
Species
Fig. 2. Number of species per genus recorded in the precious corals
from the Mediterranean Sea (white bars) and Pacific Ocean (grey bars).
in the Pacific, and three species of Spiroxya both in the
Mediterranean and in the Pacific Ocean (Table 2). Alectonidae (Alectona, Dotona, Delectona and Spiroxya) account
for 14 species, suggesting a strong affinity of this family for
the compact calcite of the precious coral scleraxis.
The affinity of the species of this family for precious
corals could be related to their typical bathophily. The
deepest known boring sponges are Spiroxya levispira Topsent 1898 (2165 m, Topsent 1928) and Alectona mesatlantica, recorded at a depth of 2030 m (Vacelet 1999). Pang
(1973) underlined that records for A. millari were generally for greater depths and that all the shallow-water
records were from underwater cavities or shaded sites.
This is also true for the shallow records of A. wallichii
(Vacelet 1999). It is possible to hypothesise that species of
this genus, together with the genus Spiroxya, are strongly
stenoecious, limiting their presence to relatively stable
environments such as marine caves and deep bottoms.
Nevertheless, whereas Spiroxya is a genus with a strong
affinity for precious corals (with six species out of nine
recorded in coral colonies), Alectona is common in other
organic carbonates. This is also indicated by the geographic distribution of these genera; Alectona has a circumtropical distribution and the genus Spiroxya has
exactly the same Tethyan relict distribution as the coral
genus Corallium (Calcinai et al. 2004b).
Acknowledgements
This paper is dedicated to Klaus Rützler to remember his
fundamental contribution to the taxonomy and ecology of
boring sponges both in the Mediterranean and in the tropics. He has highlighted, more than any other, the great
278
Genus Holoxea
Holoxea furtiva Topsent, 1892
Holoxea excavans Calcinai et al., 2001
Genus Alectona
Alectona millari Carter, 1879
Alectona wallichii Carter, 1874
Alectona triradiata Lévi & Lévi, 1983
Alectona verticillata (Johnson, 1899)
Alectona sorrentini Bavestrello et al., 1998
Alectona sarai sp. n.
Genus Delectona
Delectona ciconiae Bavestrello et al., 1996
Genus Cliona
Cliona janitrix Topsent, 1932
Cliona lobata Hancock, 1849
Cliona desimoni Bavestrello et al., 1995
Genus Spiroxya
Spiroxya acus (Bavestrello et al., 1995)
Spiroxya corallophila (Calcinai et al., 2002)
Spiroxya heteroclita (Topsent, 1988)
Spiroxya levispira (Topsent, 1898)
Spiroxya macroxeata (Calcinai et al., 2001)
Spiroxya sarai (Melone, 1965)
Genus Thoosa
Thoosa armata Topsent, 1888
Thoosa bulbosa Hancock, 1849
Thoosa midwayi Azzini et al., 2007
Thoosa mollis Volz, 1939
Genus Dotona
Dotona pulchella Carter, 1880
Genus Aka
Aka labyrinthica (Hancock, 1849)
Aka coralliirubri Calcinai et al., 2007
Aka insidiosa (Johnson, 1899)
Mediterranean
Sea
Pacific
Ocean
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
ecological importance of boring sponges, focusing the
attention of new generations of researchers on this fascinating group. The authors wish to thank Mr Yoshihico
Niiya, Kosaitsuno, Kochi, Japan for sampling the material.
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