BULLETIN OF MARINE SCIENCE. 65(1): 227-234.

1999 CORAL REEF PAPER

COMMUNITY STRUCTURE OF THE LIVING FOSSIL CORALLINE

SPONGE POPULATIONS AT THE GROTIO, SAIPAN,

NORTHERN MARIANA ISLANDS

Norman J Quinn and Barbara L. Kojis

ABSTRACT
An extensive hermatypic sponge community at a popular dive site called The Grotto is
described from Saipan, Commonwealth of the Northern Mariana Islands. The commu·
nity is dominated by the coraBine sponge Acanthochaetetes wellsi with a maximum den­
sity of 672 individuals m- 2 • The coralline sponge As/rosclera willeyana, and the lithistid
sponge Leioderll/atiu", sp. are also present but at lower densities. A. 'A.'ellsi and A. willeyana
were most dense at depths >9-16 m. These sponges pref¢r the low illumination found in
specialized cryptic habitats in warm tropical waters, but are not necessarily found on
coral reefs. Two previously unreported hermatypic sponge communities on Saipan and
on the neighboring island ofTinian are briefly mentioned. There was no evidence that
present dive practices are damaging these cryptic communities. However, owing 10 the
slow growth of these sponges, recovery from damage to the community may take a cen­
tury or more.

A large, hcnnatypic sponge community exists on Saipan (15°10'N, 145°45 'E), Com­
monwealth of the Northern Mariana Islands (CNMI) at a popular dive site called The
Grotto. The site was first briefly mentioned by Goreau et al. (1969; 1972) as an " ...unusu­
ally abundant living fauna of hermatypic sponges or geological significance...". Subse­
quently, Hartman and Reiswig (1980) studied the taxonomy of the coralline sponges in
the Grotto. However, no quantitative surveys describing the community have been done.
The coralline sponges Acalllhochaeteles wellsi (Hartman and Goreau, 1975), and
As/rosclera willeyana Lister 1900, and the lithistid sponge Leiodermatium sp. occur in a
very restricted ecological niche in tropical waters. Coralline sponges are considered the
surviving relatives of sponges which contributed to reef building during the Paleozoic
and Mesozoic Eras (Vace1et, 1985). These "relict" sponge communities are considered to
be very similar to those found in fossil reefs in LateAlbian (Cretaceous) times (Reitner,
1993) in their preference for the cryptic environs of caves and similar dimly lit habitats of
tropical fore reefs (Basile et aI., 1984; Reitner and Engeser, 1987).
In this paper the coralline sponge population densities, size, distributions and zonation
in the Saipan Grotto are described. The presence of a large coralline sponge community
offTinian and another near the Saipan Grotto are reported.

MATERIALS AND METHODS

SAIPAN.-Saipan is about 260 km northeast of Guam and is the second largest of the Northern
Mariana Islands. It is 22.4 krn long, between 4 to 8 km wide, and has an area of 182 km 2 . It is a
major tourist destination for Japanese and Koreans with about 0.5 million visitors yr l of which
approximately one-third are scuba divers.
The Grotto, a popular dive and snorkel site, is off the northeast coast near Unai Fanuchulauan
(Fig. IA) and receives the direct effect of the northeast trade winds. The eastern side of the island is
dominated by limestone cliffs with few beaches or extensive reefs (Gareau et aI., 1969). There is no
~2S BULLETIN Of MARINE SCJFJ\CE, VOL 05.:'-10 1. 1999

Unai Fanuchubuan

Figure la. Location orthe Grotto on Saipan. Commonwealth of the Northem Mariana Islands.

Point Lamanibot

Tinian Grouo

Figure 1b. Location of the Grotto on Tinian, Commonwealth of the Northern Mariana Islands.

land for thousands of kilometers to the east and the water in and around The Grotto is generally
exceptionally clear with visibility commonly >30 m. The Grotto is a world class dive site and in
1995 aver<'tged about 60 divers d- I during good sea conditions. It covers a surface area of approxi­
mately 8000 m 2 and has the world's largest reported coralline sponge community. It has a surface
entrance and three underwater entrances at 5, 12 and 20 m. The maximum depth of the Grotto is 24
Ill.
Outside the Grotto, about 70 III to the north, a large cave exisls at a depth of25-32 Ill. The cave
extcnds into the limestone rock <'tbont 50 m and has an extensive, similar cryptic commnnity. Be­
cause of the dcpth and distance from a passage to The Grotto, quantitative measurements were not
done in the cave.
T1NIAN.-Tinian (1 Y'OON, 14°57'E) is approximately 3 km southwest of Saipan, 250 km north­
east of Gnam, and is the largest of the Northern Mariana Islands. It is presently the least populated
(-700 people) of the three major Northern Mariana Islands and has no major tourist or industrial
development. A similar coralline sponge dominated community is present at the Tinian Grotto, 22­
28 In deep, (latitude l5.05°:i'L l45.6°E.), ofl the west coast ofTinian about 2 km south of Point
 QUINN AND KOllS: CORALLINE SPONGE COMMUNITY
229

Table 1. Population statistics of Acallthochaeleles wellsi at the Grotto, Saipan.

Depth Density n Mean C. V. (%) of Maximum %

(m) (m-') individuals diameter diameter diameter cover
(em) (em)
39 220 275 2.4 21 5.0 14
>9-16 672 840 3.9 67 8.5 65
>16-23 284 355 5.1 49 10.5 58
Lamanibot and about 150 m from shore (Fig. I B). The entrances are at 10, 16 and 22 m. The Tinian
Grotto is smaller than the Saipan Grotto and covers an area of approximately 2000 m l . Owing to the
logistic difficulty in accessing the Tinian site and the surge within the Grotto, no quantitative mea­
surements were made. Work performed in the presence of strong surges would have resulted in
damage to the community.
CORALLINE SPONGE COMMUNITY SURvEY.-The population density ofcoralline sponges in the main
Saipan Grotto were quantitatively surveyed in July/August/September 1995 using scuba equip­
ment. Density and diversity of the coralline and lithistid sponges on the walls were quantified from
15 0.25 m2 quadrats. Quadrats were located along a vertical transect extending from 3 to 23 m on
the wall in the middle of the Grotto between the two eastern underwater entrances. A metal rcbar
was driven into the wall at 10m to help relocate the transect. The quadrats were assigned to three
depth ranges (five quadrats in each of three depth ranges) (Table J). The numbers of individuals for
each species from each depth range were standardized to number of individuals m- 2 and expressed
as a whole number. The community in shallower water «3 m) was not sampled because of the surge
and the potential for damage to the community during sampling. Each quadrat was photographed
using a Nikonos RS camera with a 20 mm lens. A scale was included in each frame and care W'.ts
taken to insure that the field of view was flat. The relief on the wall was no more than 7 em which
would introduce at most a 3% error in measuring sponges in different planes.
The film was processed and transferred to digital images by Seattle Film Works. Jandel's Sigma
Scan ver. 3.0 program was used on the digital images to determine the length of the colonies (to the
nearest 0.5 em) and percent cover. Colonies with a diameter <1 em were not counted as a reliable

Figure 2. Coralline sponge community at 14 m at the Grotto, Saipan.

230 llLlLLETIN Of I\fARI\iE .~CTE\iCc \'OL 65. -"'0.1,1999

T,lbJe 2. Population statistics of ASfrOsdf!J"o \l·j/leyLma at the Grotto, Saipan.

Depth Density n Mean C.V (%) of Maximum %

( Ill) (m-') individuals diameter diameter diameter cover
(em) (em)
3-9 20 25 2.8 11 4.0 1
>9-16 136 170 3.2 18 5.5 11
>16-23 92 115 3.5 16 4.5 9

identification \vas beyond the reasonable resolution of this technique. Light Je'vels were approxi­
mated at various dcpths using the intcmal1ight meter in a Nikonos RS llnden'iuter C<lmer<l.
In a gcncral survey of the main Saipan Grotto, the largest individunl of e"Kh species \vithin each
depth range was searched tor <lnd dJameter measured. Smaller specimens werc collecled nnd flX-::d
lTl 10% formalin ror later identlfication.

RESULrS

BENTHIC COl\'IMUNITY SURVEY.-Two species of coralline sponges (A. 11'ellsi, A. \',:iIleyana)

and a heavily silicified lithistid sponge (LeiodermalilllJl sp.) \....crc found in the Saipan
Grotto (Fig. 2). The IllOst conmlon species. A. \9ellsi, repres~ntC'd S2.9\~. <) of coralline sponges
surveyed. Less common wereA. H'illcyana (16.1 %) and LeiodermatiulII sp. (l.O~. .o). Addi­
tional fauna present included gorgonian fnns (Subogorgia sp.), hydrozoans (Stylasler
sp.), bryozo<lns, minute barnacles (Lacazella sp. and Thecidellina sp.), and a few encrust­
ing soH sponges.
The color of A. H'ellsi and A. willeyana ranged from light yellow to orange. Rcitner et
a1. (1996) obsC'rved green specimens at Lizard Island which harbored a boring green alga
(Oslreobiufn sp.). Green specimens were not observed in The Grotto.
The greatest density of each species occurred at >9-16 ill (Table I). A total of 672
individuclls m- 2 ofA. \vellsi occurred at >9-16 m compared with 284 individuals m- 2 at
> 16-23 m and 220 individuals m-1 at 3-9 nl. There were nearly seven times more A.
lFilleyana at >9-16 m than at 3-9 m, but only 1.5 times more than at >16 to 23 ill Cfable
2). Leiodermlltilffn sp. was unconmlOn at all depths (Table 3).
Both A. wellsi and A. wilieyana were spherical, while Leiodermatiwn sp. had the shape
of an elongated uneven trapezoid with rounded corners. The largest diameter of A. well.si
recorded was 10.5 em. Individuals 0/A. weLlsi about 7 em in diameter were often dead
with smaller sponges growing 011 their top most surface. They eithcr altered their growth
form when they achieved this size or the small individuals were ne\v recruits that settled
after the sponge died. The largest individuals of A. HJellsi were found at lhe >16 to 13 m
depth range (Table 1). The maximum and mean diameters of individuals of theA. willt:.~vaJla

Table 3. Population statistics of Leiodenl1l1!iUIII sp. in the Grotto. S<lipan.

Depth Density n !vICJO C.V.(%)of Maximum 0/,.

(m) (m"') individuals diametcr diameter diamc[cr cover
(em) (em)
3-9 4 5 56 35 6.0 1
>9-16 8 11) 4.9 28 5.0 2
>16-23 4 5 6.8 44 9.5 1
 Ql'r'~'\I AND ROJIS CORALU)\'L SPOl\GE COM\1UNlTY 23]

80

70
w
~ 60
u
:2:
u
50

E 40
'0
ill 30
.0
E 20
~
10

0
2 3 5 6 7 8 9 10
Diameter (em)
D !lc;"nlhoc;haeleles wellsl
• Astrosclera wllieyana

Figlln:: 3. Size frequellcy distribulion histogram of Acal1t!Jocli</e[etes we/lsi and Astrosc!era ,villeyana
at the Cirotto, Saipan. using 0.5 em siLc da~:ies.

population were smallcr than those of A. wcfls; (Fig. 3). Thc largest A. ~·vifle. yana observed
was 5.5 cm diameter, with the largest individuals occurring at >9-16 m (Table 2). The
larg~st individual of Leiodermatillm sp. was 9.5 em long.
The illumination of both grottos was so Jow (~1 lux) even in the middle ofa sunny day
that no comBine algae were observed in association with the coralline sponge commu­
nity. There was more variation in illumination between time of day and sky conditions at
the SJme site than betwecn various depths.

DISCUSS10N

This is the first study to quantify the abundance, depth distribntion, and species diam­
eters of a modern hermatypic sponge eommnnity in thc Mariana Islands, western PJcific
Ocean. Most studies have dealt with taxonomic questions (Goreau et al.. 1969, 1972;
Hartman and Gorean, ] 975; Hartman and Reisv...ig, 1980; Vacelet, 1981; Reitner, 1991,
1992), morphological descriptions (Vaeelet and Garrone, 1985: Vacel"" 1990; Reitner,
199]), evolutionary theories (Vacelet 1985), and biocalcification processes (\V6rheide et
aI., 1996), Ecological studies were first published by Ayling (1982) and Reitner (1993) in
the small reef caves off Lizard Island on the GreJt Barrier Rcef. Other thelll these studies,
little ecological information is available on these uncommon cryptic communities.
Mean size of individuals for each of the three species and their density was dcpth de­
pendent. The greatest densities occulTed between >9-16 m. No change in water currents
with depth was observed 'A'hile scuba diving. However, the strong surge at shallower depths
associated with oceanic s\vells mJy dislodge sponges; during intense storms. While Ayling
(1982) noted that large A. willeyanG specimens appeared to develop in strong \vater move­
ment, we hypothesize that after a certain point extremely turbulent watcr movement will
be more detlimental to brger individuals with their greater snrface area.
The Grotto containcd few large individuals. Coralline spongc specimcns 20 em in di­
ameter with individuals up to 25 cm were observed at New Caledonia (Bohm et aL 1996)
and Hartman and Gorcau (1970) reported a specimen (no identification given) from the
..,., .., BIJLLETli\ OF I,,1Af{INF. ~(lEJ\CF. VOL. 65. NO.1. 1999

GrOlto "Up to 18 cm". However, the largest specimen (A. 1-Fellsi) that we observed in 15
dives at the Grotto was only 10.5 cm maximum diameter. Considering that Goreau col­
lected specimens, it is possible that he coUected the largest individnals of these 510\'1'
gro\\'lug organIsms.
Although sclerllctiui<ln cor"ls occurred just outside the entranccs to the Grotto, and
coral planula scttled on clay tiles in the Grotto (Quinn and Kojis. 1997), no sc1eractinian
corals \\'ere seen :Jrnoug the coralline sponge community. Corals need sunlight for sur­
vival (Kojis and Quinn, 1984) and the low light conditions likely explain their absence.
The hlck of scleractinian corals reduces the sponge community's competition for space
with this group of organisms aml likely docs so for other light dependent taxonomic
groups.
Coralline and lithistid sponges were <lbundant in the three caves mentioned in this p<l­
per. They were found from spring 1mv tide to a depth of32 m, depths readily accessible to
both snorkelers anJ novicc divers. The geological importance of these communities and
their accesibility by divers and snorkelers <llikc, make these sites ideal candidates for
marine reserve status. A large number ofproteted areas for coastal, reefal. and estuarine
ecosystems have bcen established aronnd the world (Agardy, 1994; Rowley, 1994); none
have incorporated these specialized communities.
Both Grottos are popular dive sites and are potentially threatened by an increasing
human presence. Fortunately, at preseut, most of the divers are led by dive gnides who
havc little knO\v1edge of the cryptofallna on the walls and, therefore, take llO notice of the
sponge community. fortuitously guiding divers away fro/11 the walls. However. dislodged
coralline sponges and broken pieces of Stylaster sp. \vcre present on the sea floor. particu­
larly in the Tinian Grotto. Additionally, air trapped on the cave roof from divers exhaust
may last long enough to kill some individuals. The construction of a road to the Grotto
has resultcd in increased erosion of fine limestone particles from the road shoulders dur­
ing periods of heavy rain. This causes the water to become milky and visibility drops to
<5 m for several days. It is uncertain what effect this periodic disturbance will have on the
cOlmnullity.
Coralline sponges (A. wellsi) are slow grm.ving, 450 J..U11 yr- 1, and are currently being
studied to assess their ability to record long-term climatic changes (Bobm et aL ]996).
Growth rates of <5 cm 100 yr- 1 suggest that the community is unlikcly to recover from
damage for decades. It is recommended that regular monitoring be nndertaken to detect
subtle changes to the community. Furthermore, it is suggested that dive leaders be given
an awareness training program and be enlisted in qualitatively monitoring the sites for
initial indications of disturbance. Finally, government agencies need to consider the de­
velopment of a management plan. including marine reserve status. for this unique,
tropical, shallow water 11abitat.

ACKWJWLEDGMEl'ITS

Identification of sponge species was made by 1. Rcitner, InstitLit und ~·1useum fUr Geologie und
Pali:iontogie, George-Aug.u~t Univcrsit~t. G6ttingen. We gratefully ;KknO\vlcdgc the support pro­
vided by M. Sablan and J. Furey. the par1ia] financial support from the U.S. Environmental Protee­
tionAgency through the Division ofCoJ~tJl Resource M<.11l:lgcment. Commonwealth of the North­
ern Mariana Islands, a gr;mt from Tropic;)] Disco\'eries, ;)nd the assistance from the Council for
International Exchange of Scholars. This study was written while N.J.Q. VdS a J W Fulbright
 QUINN AND KOJIS Cl1RAlLl\E SPO"<GE COi\1i\IUNITY 233

FdlO\v at the Biology Departmcnt, School ofPorc and Applied Scicnccs, University of the South
Pacific. P. Bergquist. M. Furland, B. Gla~by, P. Newell, 1. Reitna, J. Vacclct, and two anonymous
refer..::es are grJteflllly acknowledged for their constructive comments on the mannscript. Vole thank
S. Bm'iess for his diving assistance

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DAT[ SUn/l.·IlTTm: December 11, 1997. DATE ACCEPTED: :ivI,lfCh 2x, ] 99x.

ADDRESS: (N.J.Q.)Department of Coastal Resource lvfanagcmcnl, Division ofNatural Resources,

Saipan, Commonwealth ofthe Northern Mariana Jylands 96950. (RL.K.) Division ofScience and
Math, University a/the Virgin I1'lands, St. Thomas, Us. Virgin Islands 00803. PRESE'\T ADDRESS:
Biology Department, University a/Papua New Guinea, Port Moresby, Papua New Guinea.