Hydrobiologia (2012) 687:219–226
DOI 10.1007/s10750-011-0774-5
SPONGE RESEARCH DEVELOPMENTS
Growth and regeneration of the elephant ear sponge
Ianthella basta (Porifera)
Sven Rohde • Peter J. Schupp
Received: 15 February 2011 / Accepted: 21 May 2011 / Published online: 8 June 2011
Ó Springer Science+Business Media B.V. 2011
Abstract Sponges are an important component of
the benthic community, especially on coral reefs, but
demographic data such as growth, recruitment or
mortality are notably limited. This study examined the
growth of the elephant ear sponge Ianthella basta, the
largest and in some areas one of the dominating sponge
species on Guam and other pacific reefs. We measured
growth rates of the natural population on Guam over
the course of one year and identified intra-individual
growth patterns. Initial sponge sizes ranged from 200
to 35,000 cm2. Specific growth rates ranged from 0.08
to 6.08 with a mean specific growth rate of 1.43 ± 1.29
(SD) year-1. Furthermore, specific growth decreased
with sponge size. The age estimate for the largest
sponge (1.7 m height 9 9.5 m circumference) was
*8 years. Intra-individual growth was mostly apical.
This study demonstrated high growth rates, which has
Guest editors: M. Maldonado, X. Turon, M. A. Becerro &
M. J. Uriz / Ancient animals, new challenges: developments in
sponge research
S. Rohde (&) P. J. Schupp
Institute for Chemistry and Biology of the Marine
Environment (ICBM), Carl-von-Ossietzky University
Oldenburg, Schleusenstr. 1, 26382 Wilhelmshaven,
Germany
e-mail: sven.rohde@uni-oldenburg.de
P. J. Schupp
University of Guam Marine Laboratory, UOG Station,
Mangilao, Guam 96923, USA
notable implications for environmental assessments,
management and potential biomedical applications.
Keywords Elephant ear sponge Ianthella basta
Porifera Demography Growth
Introduction
Sponges are an important component of benthic coral
reef communities (Diaz & Rützler, 2001). On Caribbean reefs, their importance has begun to be recognized, where sponge assemblages reach similar
diversities and abundances as scleractinian corals
(Targett & Schmahl, 1984; Suchanek et al., 1985).
However, on Pacific reefs, sponges have received
much less attention despite their various ecological
roles. Sponges are benthic filter feeders (Reiswig,
1971; Hadas et al., 2009; Riisgard & Larsen, 2010),
serve as habitat for organisms (Duffy, 1992; Henkel
& Pawlik, 2005; Hultgren & Duffy, 2010) and affect
the benthic community composition by competitive
interactions (Suchanek et al., 1985; Engel & Pawlik,
2000).
Despite their high diversity and abundance,
research just started to investigate life history traits
like growth, life span, or reproduction in more detail
(e.g. Turon et al., 1998; De Caralt et al., 2008;
Koopmans & Wijffels, 2008; McMurray et al., 2010).
While there is no doubt that growth, size, or life span
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affect and explain many ecological interactions and
functions (Peters, 1983), more research needs to be
done to reveal sponge life histories (McMurray et al.,
2008).
There are many factors that may have contributed to
this lack of studies. Sponges are often very slow
growing organisms, which require long-time studies to
estimate growth rates (Reiswig, 1973; Duckworth &
Battershill, 2001). Growth can vary significantly
among seasons, populations, or sites (Garrabou &
Zabala, 2001; De Caralt et al., 2008), and many sponge
species have a highly variable morphology, which
complicates accurate size estimates (but see Koopmans & Wijffels, 2008). Sponges also lack distinct
morphological structures that can be used as age
indicators like otoliths in fishes or growth rings in trees.
Growth rates have been determined for a number
of sponge species, including encrusting, rope-like,
tubular, and massive growth forms (e.g. Turon et al.,
1998; Garrabou & Zabala, 2001; Tanaka, 2002; De
Caralt et al., 2008; Koopmans & Wijffels, 2008;
McMurray et al., 2008). Hoppe (1988) investigated
growth of the flabellate sponge Agelas clathrodes, but
to our knowledge, no data are known from fan-like
growing sponges.
When measuring growth in sponges it should be
recognized that sponges often heal wounds much faster
compared to their normal rate of growth (Ayling, 1983;
Smith & Hildemann, 1986; Hoppe, 1988), and wound
healing/tissue regeneration is therefore not a good
predictor of normal growth rates.
Estimation of the age structure of benthic communities is essential for applied ecological assessments, which require knowledge of the age and
growth of the community to assess recovery rates
after habitat destruction or mitigation projects.
Ianthella basta (Pallas) is a conspicuous, fan- or
funnel-like shaped sponge reaching heights of up
to 2 m (Bergquist & Kelly-Borges, 1995, personal
observation). I. basta is widely distributed in the IndoPacific (Bergquist & Kelly-Borges, 1995). Its distribution ranges from the Mascarene Islands to Vanuatu,
the Philippines and Guam. However, I. basta is absent
from all intervening Micronesian Islands including the
Federated States of Micronesia and Palau (Kelly et al.,
2003). Therefore, it has been speculated whether
it colonized Guam by jump dispersal or became
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Hydrobiologia (2012) 687:219–226
introduced through anthropogenic transport (Kelly
et al., 2003). The fact that on Guam I. basta only occurs
in Apra Harbor, where the port is situated, may support
the latter. The restriction to Apra Harbor as documented by Paulay et al. (2002) has significant consequences for the population of I. basta on Guam. For
construction of a new aircraft carrier wharf, large reef
areas in Apra Harbor are planned to be dredged (Navy,
2010). This would reduce or potentially eliminate
much of the actual habitat of I. basta on Guam. While I.
basta might be an introduced species, it lacks invasiveness in that it does not seem to compete with other
sessile invertebrates for limited space. It is mainly
found along the edge of coral slopes where the hard
substrate changes to soft bottom sediments with
occasional rocky outcrops to which it is attached. In
this habitat, I. basta adds to the rugosity and provides
shelter for other invertebrates and fishes (Rohde and
Schupp, personal observation). As one of the largest
and most conspicuous sponges inside Apra Harbor, I.
basta has also become an attraction for the local dive
operators. Therefore, I. basta has become a biological
and economical important species. Thus for both,
assessments of ecological damage and potential mitigation demands, an assessment of the population
characteristics (age, growth) of I. basta is essential and
encouraged by local resource agencies (D. Burdick,
personal communication).
Another reason to investigate the natural growth rate
of I. basta is recent studies describing its unusual
chitin skeleton and the potential of such skeletons in
biomedical applications (Brunner et al., 2009; Ehrlich
et al., 2010a, b). However, the use of marine natural
products in general and of the identified chitin scaffold
in particular is restricted by supply limitations (e.g.
there is no synthesis available for chitin scaffolds).
Consequently, a detailed knowledge of growth rates
and growth patterns is necessary to evaluate if aquaculture could be viable to produce enough material for
biomedical applications.
The aim of this study was to determine the growth
and consequently the age of the natural population of
I. basta. Beside the general lack of knowledge on
sponge growth rates, the results could highlight the
consequences of habitat destruction and explore
whether I. basta constitutes a sustainable source for
tissue harvest for extraction of chitin scaffolds.
Hydrobiologia (2012) 687:219–226
Materials and methods
Growth of the natural Ianthella basta population
The study site was at Western Shoals, Apra Harbor,
on the west coast of Guam (13°27.30 N, 144°39.20 E).
This is a very sheltered location with a high density
of I. basta. At depths between 8 and 11 m, we tagged
40 specimen of I. basta using numbered aluminum
washers that were nailed to the reef next to the
sponges. We chose specimen over the entire size
range of the present population. In June 2009 and
June 2010, circumference and slant height were
measured by SCUBA using measuring tapes. The
shape of a cone was used as a model to calculate the
area of the sponge tissue, and growth was calculated
as:
G ¼ ðA2 A1 Þ=A1 ð1=dtÞ;
where G is the specific growth rate (year-1), A1 is the
initial area (cm2), A2 is the final area (cm2), and t is
the time (years).
To assess whether size affects the growth rates of
the sponges, individuals were grouped into 4 size
classes: 1 (from 195 to 1,000 cm2, n = 6), 2 (from
1,001 to 3,000 cm2, n = 18), 3 (from 3,001 to
10,000 cm2, n = 9) and 4 ([10,000 cm2, n = 8).
Specific growth data were log10-transformed to
obtain homogeneity of variances (Levene’s test).
Growth differences among size-class were analyzed
by a one-way ANOVA and Tukey’s posthoc test
(SPSS 17).
Five commonly used growth models were fit to
size-increment data to determine the best model
describing growth of I. basta (McMurray et al.,
2008): the general von Bertalanffy growth formula
(gVBGF) (von Bertalanffy, 1938; Beverton & Holt,
1957; Pauly, 1981), specialized von Bertalanffy
growth formula (sVBGF) (Richards, 1959; Pauly,
1981), Gompertz (Gompertz, 1825; Winsor, 1932),
Richards (Richards, 1959; Ebert, 1980), and Tanaka
(1982) growth functions. The square root of area
estimates was used as an average linear size to model
growth of I. basta. The difference equations of the
models were fitted to final and initial linear sizes on a
Walford plot by nonlinear regression (SOLVER, MS
Excel 2007). To produce size-at-age curves, we used
parameter estimates using the integrated forms of the
growth functions, which were subsequently squared
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to obtain area-at-age plots. The sum of squared error
(SSE), coefficient of determination and Akaike
information criterion (AIC) (Akaike, 1973) were
used to evaluate model fit. Because the AIC evaluates
the trade-off between model biases that results from
too few parameters versus variance that results from
too many, it can be used to evaluate any difference
between the 2- and 3-parameter models that may
otherwise be neglected through use of the SSE as the
sole fitting criteria (Burnham & Anderson, 2002).
The model with the lowest SEE and AIC is sought to
select the best fitting model. Because sample size
(n) was small with respect to the number of model
parameters, K, (n/K \ 40), the second-order correction (AICc) was used (Burnham & Anderson, 2002).
As a measure of each model relative to the best
model, the model selection criteria AIC difference,
Di, was calculated.
Regeneration and intra-individual growth patterns
To observe the intra-individual growth and regeneration patterns, we tagged ten additional specimen of
I. basta with slant heights of 60–90 cm at the same
site at 12 m depth in May 2010. Of these sponges, all
the tissue above 50 cm slant height was cut off. At
every 10 cm height, cable ties were pierced through
the sponge tissue to determine whether growth
occurred throughout the entire sponge body, or
whether apical regeneration of tissue dominated.
Additionally, two round holes (3 cm diameter) were
cut at 10 and 30 cm height to identify and measure
regeneration in response to injuries.
Sponge growth was remeasured two and five
months later by measuring the distances between the
cable ties and the diameter of the holes to estimate
growth at the different portions of the thalli.
Results
Initial size estimates of the sponges ranged from 200 to
35,000 cm2 with a mean value of 5,702 cm2. Specific
growth rates ranged from 0.08 to 6.08 with a mean
specific growth rate of 1.43 ± 1.29 (SD) year-1.
Comparisons among size-classes revealed that smaller
sponges had higher growth rates (Fig. 1). Sponges in
size-class one showed specific growth rates of 3.21 ±
2.16 year-1 (mean ± SD), sponges in size-class two
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Hydrobiologia (2012) 687:219–226
Table 2 Fitting criteria of growth functions
Model
K
r2
SSE
AICc
Di
Tanaka
3
0.922
8483.7
216.6
0.00
Gompertz
2
0.914
9303.9
217.8
1.25
Richards
gVBGF
3
3
0.914
0.914
9303.9
9310.0
220.2
220.2
3.60
3.63
sVBGF
2
0.892
11730.3
226.9
10.29
Fig. 1 Specific growth (±SD) of I. basta for the 4 size-classes.
Letters indicate significant differences between size-classes
(Tukey’s posthoc comparison, P \ 0.05)
1.45 ± 0.88 year-1 (mean ± SD), sponges in sizeclass three 0.86 ± 0.47 year-1 (mean ± SD) and
sponges in size-class 4 grew 0.70 ± 0.51 year-1
(mean ± SD). Differences among size-classes were
significant (P \ 0.001, ANOVA). Figure 1 depicts the
significant differences from the posthoc comparisons.
The parameter estimates of the growth models
are presented in Table 1. All models showed a high
model fit. The models in order of best fit are:
Tanaka [ Gompertz [ Richards [ gVBGF [ sVBGF.
The SSE values were in agreement with the AICc and
revealed a similar fit of Tanaka, Gompertz, Richards
and gVBGF with a slightly lower fit of the sVBGF
model (Table. 2). According to the models, the
largest measured sponges in this study were *8
years old (Fig. 2).
Specimen of I. basta that were cut off at 50 cm
height regenerated apical tissue at rates of 1.03 cm/
month (Fig. 3). The basal and middle parts of the
sponge body did either not grow at all, or only
slightly (0.25 cm/month). The holes cut in the center
of the sponge (3 cm diameter) healed within 8 weeks.
Fig. 2 Size at age curves from parameter estimates in Table 1.
Line at 82,000 cm2 refers to the largest measured sponge in this
study
Discussion
Ianthella basta is a common member on reefs in Apra
Harbor, Guam, and reefs in the tropical Western
Pacific. While there has been renewed interest in this
species for biomedical applications (Bergquist &
Kelly-Borges, 1995; Brunner et al., 2009), little is
known about its ecology and demography. Because
the US NAVY has large scale dredging plans for
Apra Harbor, more detailed knowledge of I. basta
ecology and demography is needed to adequately
assess mediation for destroyed coral reef areas.
Table 1 Parameter estimates for growth functions fitted to square root of final and initial areas; n = 40
gVBGF
Richards
Tanaka
sVBGF
Gompertz
S? = 1950.8
S? = 1590.5
a = 0.0001
S? = 108684.6
S? = 1590.4
k = 0.0002
k = 0.117
k = 0.109
k = 0.117
f = 0.00003
d = 69166.6
d = -96618.5
d = 1981.4
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Fig. 3 Intra-individual growth and regeneration patterns of
I. basta. The drawing shows which sections of the sponge body
were tagged with cable ties and subsequently measured. Bars
represent mean growth of the respective sponge section
This study demonstrated that I. basta is a remarkably fast growing sponge species, which reaches sizes
of up to 2 m height in less than 10 years. To our
knowledge, there is only one other study, which
estimated growth and age of a sponge species that
reaches similar sizes. The Caribbean sponge Xestospongia muta also grows to heights of 170 cm, but
growth measurements indicated that specimens of
this size are around 250 years old (McMurray et al.,
2008). Even though X. muta is a massive species that
needs to build up much more biomass to reach these
large dimensions, it is an interesting fact that X. muta
needs around 25 times longer to reach similar
dimensions as I. basta.
The specific growth rate of I. basta was with
1.43 year-1 around threefold higher than that of X.
muta, but similar to growth of four Mediterranean
sponge species with mean rates from 1.08 to 2.4 year-1
(Garrabou & Zabala, 2001). However, individuals of
size-class 1 grew as much as 3.21 year-1 leading to a
relative fast transition in higher size-classes. Specific
growth rates are a relative measure; therefore, the
absolute biomass production is higher in large sponges
compared to small sponges with similar specific
growth rates. Consequently, growth rates of 0.7–1.45
in size classes 2 to 4 are comparable to Mediterranean
sponge species (Garrabou & Zabala, 2001), but due to
the large size of I. basta specimen, the biomass
production is much higher. This could be one reason
supporting the establishment of many large sponge
individuals.
223
Specific growth rates decreased with increasing
size. This pattern has also been found for many sponge
species (e.g. Reiswig, 1973; Garrabou & Zabala, 2001;
De Caralt et al., 2008; McMurray et al., 2008; but see
Duckworth & Battershill, 2001). The negative correlation of size and specific growth of I. basta is
especially not surprising, since absolute growth was
similar among all sizes. Growth was apical with 1
(±0.69 SD) cm month-1 along the upper edge, resulting in lower specific growth with increasing size.
All measured specimen showed positive growth, but
the intraspecific variation was very high and this
seems to be characteristic for many sponge species
(Reiswig, 1973; Dayton et al., 1974; Wulff, 1985;
Duckworth & Battershill, 2001; Garrabou & Zabala,
2001; McMurray et al., 2008).
Holes that were cut in the center of the sponge
healed within 2 months. This rate is very similar to
the apical regeneration rates and much higher than the
growth measured at this part of uninjured sponge
specimens. This further demonstrates that growth is
almost entirely restricted to edges of the sponge,
whether they are the apical edge, edges of holes from
predation or, in this case artificial injury. Some sponge
species showed wound healing processes that generated tissue much faster than their normal growth
(Ayling, 1983; Smith & Hildemann, 1986; Hoppe,
1988). Assuming that wound healing exerts the highest
physiologically possible growth rate and the fact that
natural apical growth in I. basta showed similar rates,
we suggest that the rate of 1 cm month-1 is the upper
limit of growth under the conditions of this study.
The question arises, what limits the size and the
age of I. basta? The largest specimens of the
population were estimated to be around 8 years old,
but still growing with average rates of 1 cm month-1,
indicating infinite growth for this species. Since no
larger and therefore older specimen could be found
on Guam, other biotic or abiotic factors must restrict
the sponge to a maximal size and age.
Predation can significantly affect a sponge community (e.g. Wulff, 1997; Pawlik, 1998) and consequently
also affect growth measurements by consuming
sponge tissue. During our study, we observed no
evidence of predation, such as bite marks or removal of
significant amounts of biomass. The crude extract of I.
basta deterred feeding by various predators (Becerro
et al., 2003). Therefore, the observed growth seems not
or only minimally restricted by predation.
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The effect of abiotic factors on the growth and
mortality of I. basta has not been studied. Apra Harbor
is a geographically protected bay with low current
and wave dynamics. These conditions have been
described as similar to other locations where I. basta
occurs (Bergquist & Kelly-Borges, 1995; Kelly et al.,
2003). It therefore seems likely that high water
movement restricts the distribution of I. basta and
may also restrict its size. Large sponges with an area
of over 80,000 cm2 offer a large resistance to waves or
currents. One possibility is that this size represents a
threshold where water movements rip off the sponges
and in this way limit their size and age distribution.
Extreme wave action, as it occurs during typhoons,
could topple and kill the sponges (when they subsequently decompose in the fine sediment). With the last
typhoon occurring on Guam on December 8, 2002 it
seems unlikely that typhoons are the sole event
restricting the maximum size, as medium size sponges
would have had over 8 years to grow and therefore should have had reached a total age of 10 to
15 years, with corresponding sizes (100,000 cm2 to
500,000 cm2, depending on the model). However, the
maximum size we observed was 82,000 cm2, making
such a scenario unlikely. Another factor restricting the
size could be the fiber dominated skeleton (chitin and
spongin), which might be too flexible to support a
larger ([82,000 cm2) fan-shaped skeleton against
currents and wave action.
Other sponge species have shown to regrow at
their bases after detachment by storms or anthropogenic effects (Schmahl, 1999; McMurray et al.,
2010). This effect has not been shown for I. basta,
but could contribute to the preservation of the
abundance of I. basta.
The population of I. basta in Apra Harbor is very
isolated. The sponge does not occur on other reefs
around Guam and can also not be found on the
surrounding Micronesian Islands (Kelly et al., 2003).
Consequently, the loss of the population in Apra
Harbor would extinct this species from the entire
region. The planned dredging of large reef areas in
Apra Harbor to build an aircraft carrier berthing will
reduce the habitat of I. basta to a great extent. Many
surveys have been done to assess the marine community, to evaluate environmental consequences and to
determine the appropriate quantity of the compensatory mitigation measures that will be recommended
for the project (Navy, 2010). However, to evaluate
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Hydrobiologia (2012) 687:219–226
how the loss of significant parts of the I. basta
population could be compensated, one requires
demographic data such as growth, recruitment and
mortality. None of these data have been available so
far. The results of this study allow the assessment of
the population structure, i.e. age-distribution curves,
or size-at-age analyses, which describe the present
population. In order to estimate population recovery
rates after disturbances, data on recruitment and
mortality are essential but non-existent so far. However, the fact that I. basta reaches high abundances
despite its relative ephemerality indicates that recruitment could be high and regular. Since I. basta tissue is
relatively tough and does not tear or fragment easily,
asexual reproduction by fragmentation seems less
likely. If the recruitment is accomplished through selfseeding by the current population, large scale dredging could diminish the larval producing population to
the point that recruitment is disrupted and the
population would eventually die off.
Within the last years, several studies investigated
I. basta with regard to its chitin-based skeleton
(Ehrlich et al., 2007a, b, 2010a, b; Brunner et al.,
2009). These chitin-based scaffolds are of high
interest for many biomedical applications like tissue
engineering and biomedicine (e.g., Maeda et al.,
2008; Jayakumar et al., 2010). However, one major
obstacle that also natural product chemists face is the
supply problem (Faulkner et al., 2000). The source
organisms of bioactive compounds often need to be
collected in large quantities to supply the necessary
amount for the industry. This can often not be
justified ecologically (Munro et al., 1994). The
farming of marine organisms is sometimes an
alternative to collecting specimens from the wild.
But this can only be an economically relevant
alternative if the organisms lend themselves to a
cost-effective cultivation (Schupp et al., 2009). Using
I. basta as source for chitinous scaffolds requires the
harvest of large amounts of sponge tissue. Both,
ecologically sustainable wild harvest and farming
therefore rely on high growth rates that provide a
sufficient supply. The growth rates of I. basta are
high compared to other sponge species and preliminary experiments revealed that cultivation of
I. basta can be suitable (Rohde and Schupp, in
preparation). However, this would only be suitable if
a healthy natural population is present to support any
aquaculture settings.
Hydrobiologia (2012) 687:219–226
Further studies on the demography of I. basta (e.g.
reproduction and mortality) together with this study
could provide the necessary framework to enable
effective management and potential mitigation.
Acknowledgments We like to thank Gitta Rohde, Ciemon F.
V. Caballes and the UOG Marine Lab Techs for assistance in
the field. This research was in part supported by NIH MBRS
SCORE grant S06-GM-44796 to PJS. Comments of two
anonymous reviewers improved the manuscript. SR was
supported by a fellowship within the Postdoc-Program of the
German Academic Exchange Service (DAAD).
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