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Phylum Porifera
Chapter · December 2015
DOI: 10.1016/B978-0-12-385026-3.00008-5
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Chapter 8
c0008
Phylum Porifera
Renata Manconi
Dipartimento di Scienze della Natura e del Territorio (DIPNET), Università di Sassari, Sassari, Italy
Roberto Pronzato
Dipartimento di Scienze della Terra, dell’Ambiente e della Vita (DI.S.T.A.V.), Università di Genova, Genova, Italy
Chapter Outline
Introduction133
General Systematics
133
Evolution and Phylogenetics
134
Biogeography and Diversity
135
General Biology
138
Body Bauplan
138
Life History Changes in Morphology
140
Anatomy and Physiology
140
Life History, Life Cycle, and Reproduction
143
s0010
INTRODUCTION
p0010 Sponges are basal Metazoa lacking true organs and a ner-
vous system. Their body architecture, usually displaying
an irregular symmetry, is plastic and characterized by a
continuous morphogenesis. Since ancient times, sponges
have been a source of biomaterials and bioactive compounds for the field of pharmaceuticals, biomedicines, and
cosmetics.
p0015
Porifera, encompassing over 8000 valid species, are
primarily marine invertebrates that have colonized continental water since the Paleozoic era. Species richness of
freshwater sponges is commonly underestimated in both
temperate and tropical latitudes, where new findings often
correspond to the discovery of new species. The present
diversity values doubtlessly appear destined to increase
with further research in unexplored and poorly sampled
areas, and from molecular analyses focused mainly on
cosmopolitan and widespread species that are currently
assumed to consist of cryptic species complexes. Taxonomy and paleontology should support the results of molecular biology on a strong morphological and evolutionary
basis.
General Ecology and Behavior
145
Habitat Selection
145
Feeding Behavior
146
Other Behavioral Adaptive Traits
147
Competition and Cooperation
148
Sponges as a Natural Resource
149
Collecting, Rearing, and Preparation for Identification
149
References151
From an ecological viewpoint, sponges perform key p0020
functional roles in aquatic ecosystems. These sessile filter feeders actively and efficiently pump and clean large
amounts of water in almost all oceans and continental
waters, from the surface to the higher depths. Pumping
activity by their aquiferous system helps circulate the water
column, particularly in lentic conditions, and it also traps
particulate and dissolved organic matter. Sponges provide
a living refuge for a wide array of organisms, and symbiosis with autotrophic microorganisms contributes to primary
production. Moreover, their siliceous skeleton contributes
to the formation of sediments after the sponge’s death.
General Systematics
s0015
The phylum Porifera is subdivided into four recent classes: p0025
Hexactinellida, Demospongiae, Calcarea, and Homoscleromorpha, according the multi-authored World Porifera Database (Van Soest et al., 2012) on sponge taxonomy. All recent
freshwater sponges belong to the suborder Spongillina of
the order Haplosclerida (Demospongiae) currently divided
into six families, 47 genera, and 238 species (Manconi and
Pronzato, 2002, 2007, 2008, 2011; Van Soest et al., 2012).
Thorp and Covich’s Freshwater Invertebrates. http://dx.doi.org/10.1016/B978-0-12-385026-3.00008-5
Copyright © 2015 Elsevier Inc. All rights reserved.
10008-THORP-9780123850263
133
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SECTION | III
134
Systematics is based on morphology, anatomy, and
cytology together with reproductive, developmental, and
molecular biology. A morphological dataset of macroand micro-morphotraits is diagnostic up to the genus and
species level: for example, growth form, consistency,
color, surface traits, distribution of inhalant and exhalant apertures, architecture of ectosomal and choanosomal
skeletal network, topographic distribution of skeletal
megascleres and microscleres (Figure 8.1), gemmular
architecture, and gemmuloscleres traits (Manconi and
Pronzato, 2002).
Most freshwater species produce gemmule with
p0035
­species-specific gemmular traits. These are key diagnostic traits, although gemmules are not always present during the entire life cycle of specimens and are always absent
in some taxa—which substantially complicates taxonomic
investigations. The absence of gemmules in some families
and the possibility of convergence/parallelism at the level
of gemmular morpho-traits have resulted in inconsistencies
with previously well-established systematics (Penney and
Racek, 1968; Manconi and Pronzato, 2002) and have biased
attempts to match phylogenetic relationships at the genus/
species level.
p0030
s0020
Evolution and Phylogenetics
p0040 The oldest fossils remains (spiculites) of Spongillina (gen.
et sp. indet.) are Paleozoic spicules deposits in the PermoCarboniferous of Europe (Cayeux, 1929; Schindler et al.,
2008). More recent taxa refer to the Mesozoic era, e.g.,
Eospongilla morrisonensis and Palaeospongilla chubutensis, and date back to the Colorado Upper Jurassic
and Patagonian Lower Cretaceous (Ott and Volkheimer,
Protozoa to Tardigrada
1972; Volkmer-Ribeiro and Reitner, 1991; Dunagan,
1999; Richter and Wuttke, 1999). However, most fossils
date back to the Eocene, Miocene, Pliocene, and Pleistocene, e.g., the W-Palaearctic Lutetiospongilla heily and
the SW-Afrotropical Ephydatia kaiseri (Rauff, 1926;
Rezvoi et al., 1971; Racek and Harrison, 1975; Harrison
and Warner, 1986; Pisera, 1999, 2006; Richter and Wuttke,
1999; Pisera and Saez, 2003). Fossil evidence suggests
the occurrence of inland waters invasion by sponges from
coastal brackish waters of epicontinental or enclosed seas.
These pioneer sponges might have colonized and spread
over the other still-joined continents (Racek and Harrison,
1975; Manconi and Pronzato, 2007).
Although fossil records are discontinuous (Pisera, p0045
2004; Schindler et al., 2008) and data on present geographic distribution are scattered (see Manconi and
Pronzato, 2002, 2004, 2005, 2007, 2008, 2009; Manconi
et al., 2008; Manconi, 2008), the paleontological
approach, together with paleobiogeography and historical biogeography, could help clarify when, where, and
how colonization processes occurred during the geological and paleo-hydrogeological vicissitudes of continents.
The latter could include splitting and fusion of continental plates, migration of microplates, marine ingression/regression, enclosure of salty/freshwater bodies in
inland depressions or along the coastline, river capture,
and desiccation of hydrographic basins (Manconi, 2008).
Paleontological knowledge is fundamental to clarify, at
least partly, climatic conditions and colonization timing,
to calibrate morphological-molecular trees, and to test
phylogenetic hypotheses (Pisera, 2004).
Hypotheses on inland water colonization range from poly- p0050
phyletism to monophyletism (Brien, 1969, 1970; Bergquist,
f0010
FIGURE 8.1 Skeletal spicules of freshwater sponges (Spongillina) display key diagnostic morphotraits: (a) megascleres; (b) microscleres; and (c) gemmuloscleres (not to scale).
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Chapter | 8
135
Phylum Porifera
1978; Volkmer-Ribeiro, 1990; Volkmer-Ribeiro and de
­Rosa-Barbosa, 1979; Volkmer-Ribeiro and Watanabe, 1983;
Manconi and Pronzato, 1994b; Pronzato and Manconi, 2002).
Brien (1968) and Volkmer-Ribeiro, (1986) hypothesized a
polyphyletic origin from both Haplosclerida and Hadromerida
(Figure 8.2).
p0055
The monophyletic status of Spongillina, based originally on classic taxonomic techniques, has been shown to
be robust from recent syntheses that focused on comparative
analyses of morphotraits by scanning electron microscopy,
the original diagnoses, holotypes, materials from several
historical collections, and new field surveys (Manconi and
Pronzato, 1994a, 2000, 2002, 2007, 2008, 2009; Pronzato
and Manconi, 2002). This is also supported by molecular
approaches (Itskovich et al., 2007, 2008; Efremova et al.,
2002; Schröder et al., 2003; Addis and Peterson, 2005;
Redmond et al., 2007; Maikova et al., 2010, 2012; Harcet
et al., 2010; Erpenbeck et al., 2011). The resolution power
of molecular analyses is, however, unsatisfactory at family/
genus/species levels, probably because the focused genomic
targets are not fully diagnostic.
p0060
Plesiomorphic morphotraits, e.g., skeletal architecture,
monaxial spicules, gemmules, and gametes structure, and
larvae suggest that Spongillina are allied to marine Haplosclerina (order Haplosclerida) from epicontinental and
enclosed seas. Apomorphies of Spongillina are parenchymellas (parenchymella type III sensu Ereskovsky, 1999,
2004) provided by choanocyte chambers, canals, and
spicules as well as gemmules displaying a highly diverse
architecture. The evolutionary history of some Spongillina families (Lubomirskiidae, Malawispongiidae and
Metschnikowiidae) restricted to ancient lakes and characterized by the absence of gemmulogenesis and gemmules is
particularly intriguing (Manconi and Pronzato, 2002, 2007,
2008).
Biogeography and Diversity
s0025
Current knowledge on biodiversity and biogeographic p0065
distribution is based mostly on several scattered historical papers; indeed, exhaustive monographies are scarce
and unfortunately only a few biologists focus at present on
freshwater sponges (Potts, 1888; Weltner, 1895; ­Annandale,
1911; Arndt, 1926; Penney and Racek, 1968; Racek,
1969; Volkmer-Ribeiro, 1981; Poirrier, 1982; Frost, 1991;
Ricciardi and Reiswig, 1993; Manconi and Pronzato,
1994a, 2002, 2004, 2005, 2007, 2009, 2011; Pronzato and
Manconi, 1995, 2002; Silva and Volkmer-Ribeiro, 2001;
Efremova, 2001, 2004).
Data on taxonomic richness of freshwater sponges p0070
increased slowly (Figure 8.3), as shown in the framework
of a historical analysis focusing on taxonomy and biogeography. The temporal trend—based on the worldwide
number of new species described per year—highlights
that only two species were described by Linnaeus (1759).
Long-lasting periods of inactivity or very low activity
characterized the following 100 years (1760–1860: <10
new species). Later, a positive trend is evident, with peaks
f0015
FIGURE 8.2 The monophyletic hypothesis (below) for Spongillina among Demospongiae is generally accepted, although a polyphyletic origin by
separate, successful invasions of freshwater has also been hypothesized. The monophyletic hypothesis validates marine haplosclerid ancestors able to
produce gemmules as pioneer invaders during the Mesozoic era, as suggested by Mesozoic fossils. However, Paleozoic fossils (Permo-Carboniferous) of
freshwater sponges apparently lack gemmules. Among the six extant families of Spongillina, the Spongillidae display successful adaptive radiation, as
indicated by several arrows.
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SECTION | III
136
Protozoa to Tardigrada
f0020
FIGURE 8.3 Trend of freshwater sponges increasing species richness starting from the 1759 Linnean Systema Naturae (two species) up to the present
(238 species). The periods in which most species were described are the decades 1880–90, 1910–20, and 1969–70. No new freshwater sponge species
was described between 1770 and 1850.
f0025
FIGURE 8.4 Geographic distribution of freshwater sponges. The greatest number of genera and species occurs in the Neotropical region (numbers in
brackets refer to the genera).
corresponding to the periodical maxima of taxonomic richness increase in the decades: 1888–90, 1910–20, and 1969–70
(Figure 8.3). The following authors described many new
sponge species during their careers: Bowerbank (11 species in 1863), Potts (16 species: 1880–1889), Weltner (15
species: 1893–1913), Annandale (29 species: 1907–1918),
Bonetto and Ezcurra de Drago (15 species: 1966–1973),
Volkmer-Ribeiro (12 species with different co-authors:
1963–1995), and Brien (11 species: 1967–1974).
p0075
In our last global assessment (December 2012), freshwater sponges comprised 238 species, 47 genera, and six
families including a few incertae sedis taxa (four genera and
five species) (Figure 8.4). The highest taxonomic diversity
at the biogeographic scale is recorded from the Neotropical
(73 species and 26 genera), Palaearctic (65 species and 22
genera), Afrotropical (52 species and 17 genera), and Oriental (40 species and 12 genera) regions. Species richness is
lower in other areas. The Australasian Region hosts 35 species (13 genera), and the Nearctic 31 species (14 genera),
whereas Pacific Oceanic Islands apparently harbor only
seven species (6 genera) (Figure 8.4) (See also Manconi and
­Pronzato, 2002, 2007, 2008, 2009; Van Soest et al., 2012).
Spongillina seem to occur worldwide in all biogeo- p0080
graphic regions except the Antarctica Region (Figure 8.4),
and their geographical distribution is related both to geological and climatic vicissitudes of the continents and to
the long-term dynamics of hydrographic basins. Spreading
of sponges in continental waters is constrained by habitat
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Chapter | 8
137
Phylum Porifera
fragmentation, and larvae disperse exclusively downstream
by flooding. In contrast, the dispersal power of gemmules
is apparently higher and favored by a successful adaptive
trend in gemmule morphological radiation (e.g., pneumatic
layer and spiny spicules), providing a potentially efficient
dispersal structure.
p0085
The geographic distribution and taxonomic richness
of Spongillina show a notably variable pattern, with some
taxa being widespread, common, and locally abundant,
whereas others are discontinuously distributed and rare or
monotopic. In this scenario, the crucial potential influence
on biogeographic patterns by gemmular morpho-functional
efficiency as promoter of dispersal power is problematic.
p0090
The families Lubomirskiidae Rezvoj, 1936 (four genera
and 15 species) and Metschnikowiidae Czerniawsky, 1880
(one genus and one species) show an extremely restricted
geographic range; they are found only in Lake Baikal and the
Caspian Sea, respectively. A peculiar case is represented by
Malawispongiidae Manconi and Pronzato, 2002 (five genera and six species) with an extremely disjoined pattern in
ancient lakes from the Great African Rift Valley (Tanganyika
and Malawi) to the Syrian-Palestinian Jordan Rift Valley
(Lake Tiberias), and from the Balkan area (Ohrid Lake) to
the extremely distant Wallacea in the Sulawesi Island (Poso
Lake). The restricted biogeographic pattern of all these
families seems to be related to the shared trait “absence of
dormancy and gemmules” in the life history, with their reproductive mode being exclusively sexual or by fragmentation.
p0095
Potamolepidae Brien, 1967 (six genera and 32 species),
which have both sexual and asexual reproductive modes,
shows a circumtropical range of Gondwanian origin, and
species are endemic to ancient hydrographic basins of the
Neotropical, Afrotropical, Oriental, and Pacific Oceanic
Islands regions. The most speciose genus with 16 species
is Oncosclera (Manconi et al., 2012). Their “gemmules,
usually sessile, with a simple architecture” (no pneuma,
thin gemmular theca) seem to be devoted to persistence
in situ in extreme conditions (low water level to harsh
flooding and heavy silting in rainforest) typical of tropical
areas (Brien, 1970; Volkmer-Ribeiro, 1990; Manconi and
Pronzato, 2002, 2009).
The condition of cosmopolitanism in Spongillidae Gray, p0100
1867 (22 genera and 155 species) matches well the sexual
and asexual reproductive modes, and a wide adaptive radiation of gemmules resulting in the most speciose genera of the
entire suborder Spongillina: for example, Corvospongilla 19
species, Radiospongilla 18 species, and Eunapius 16 species (Penney and Racek, 1968; Manconi and Pronzato, 2002,
2004; Manconi et al., 2008; Ruengsawang et al., 2012).
Metaniidae Volkmer-Ribeiro, 1986 (five genera and 25 p0105
species), which have both sexual and asexual reproductive
modes and complex gemmules, display a Gondwanian pattern, and are spread in the circumtropical rainforests of the
Neotropical, Afrotropical, Oriental, and Australian regions
with an enclave (one genus) in the Nearctic.
However, the assumption that taxa bearing complex p0110
gemmules as perfect dispersal devices reach a wider geographic range does not match biogeographic patterns at
the family, genus, and species levels. The case of the genus
Corvospongilla seems to be emblematic of this condition.
Species belonging to this genus are characterized by “two
gemmular types” with diverging morpho-functional roles. In
Corvospongilla mesopotamica the functional role of “heavy,
sessile gemmule” is to persist in situ, whereas the “free
light gemmule” is devoted to dispersal (Figure 8.5). The
genus Corvospongilla better matches the vicariance than
f0030
FIGURE 8.5 Corvospongilla mesopotamica is characterized by the presence of two gemmular morphs. The sessile heavy gemmule, strictly adhering
to the substratum, lacks a pneumatic layer and is protected by a reinforced spicular cage (□); the free light gemmule shows a well-developed pneumatic
layer (●) and few gemmuloscleres. Schematic reconstruction of the life cycle focuses on the diverging morpho-functional role of gemmules. Circles (●)
indicate dispersal; squares (□) indicate persistence.
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SECTION | III
138
the dispersal model because of its notably disjoined range
at the biogeographic scale, where 19 species are extremely
rare and geographically restricted (i.e., endemics). The evolutionary novelty of two diverging gemmular architectures
does not explain well the wide dispersal of the genus Corvospongilla, because each species is rare and endemic to an
ancient coastal basin (Manconi and Pronzato, 2004, 2008).
p0115
Although the definition of endemic taxa sensu stricto
is problematic, about 50% of freshwater sponge species
are endemic to small areas, but endemicity reaches the
highest values (100%) for the families Lubomirskiidae,
Metschnikowiidae, and Malawispongiidae. Also, incertae
sedis species/genera are endemic each to a single hydrographic basin, as probably is the case of Spongillina indet.
from the Towuti lake. Endemic sponges occur in ancient,
old and/or crateric basins such as Titicaca (South America), Yunnan lakes (western China), Tana (Ethiopia), and
Mweru, Luapula, Barombi ma Mbu, and Soden lakes
(Africa). Insular, endemic species are recorded from Japan,
New Zealand, Fiji, New Caledonia, Philippines, Indonesia,
and Cuba. Endemics are reported also from ancient coastal
basins (e.g., Louisiana, Florida, western North America,
Iraq, and Brazil). Main biodiversity hotspots (i.e., highest
species richness and endemicity) are ancient hydrographic
basins in tropical latitudes, such as Amazonian, Orinoco,
Paranà-Uruguay-Paraguay, Zaire, Mekong, and several others (Manconi and Pronzato, 2002, 2009; Van Soest et al.,
2012).
Protozoa to Tardigrada
Currently, it is impossible to define the precise geo- p0120
graphic ranges for several genera/species recorded only
once or exclusively from restricted geographic areas. Moreover, knowledge of sponges in some geographic areas is
scattered and fragmentary if not almost completely lacking
(e.g., Madagascar, central-northern Asia, Indochina, and
Wallacea). Species richness seems to be underestimated but
is destined to increase with further sampling campaigns. On
the other hand, the number of taxonomists on freshwater
sponges worldwide is dramatically low (less than 10), and
without the absolutely necessary recruitment of new sponge
taxonomists, the field is on the brink of extinction.
GENERAL BIOLOGY
s0030
Body Bauplan
s0035
The bauplan of sponges does not perfectly match the con- p0125
cepts of colony or individual. It better matches the concept
of modular organisms because of their ability to alter body
architecture in time and space. However, the functional units
(ramets) of other modular animals such as Bryozoa and some
Cnidaria and Tunicata cannot be isolated from one another,
unlike those of adult freshwater sponges (Pronzato and
Manconi, 1994a,b; Manconi and Pronzato, 2011).
The habitus of freshwater sponges is characterized by p0130
three morphs during the life cycle: namely, the long-life sessile adult (Figure 8.6) devoted to growth and reproduction,
f0035
FIGURE 8.6 Habitus of freshwater sponges (Spongillina). Growth form ranges from small thin crusts to massive or branched-erected with a bush-like
or tree-like shape: (a) encrusting juvenile of Ephydatia fluviatilis; (b) digitate green specimen of Spongilla lacustris; (c) massive rounded specimen of
Baikalospongia bacillifera; (d) specimen of Metania reticulata settled on a branched plant; and (e) gemmular carpet of Ephydatia fluviatilis during dormancy (not to scale).
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Chapter | 8
139
Phylum Porifera
the planktonic short-lived swimming larva, and the dormant
gemmule (restant body/propagule) (Figures 8.6 and 8.7).
Long-term, in situ investigations of gemmulation in Ephydatia fluviatilis as an experimental model demonstrated that
a specimen 5 cm in diameter produces, at the beginning of
the dry season, a gemmular carpet with more than 2000
gemmules (ramets) in less than 1 week. Each gemmule in
this clonal strategy is an individual unit (a ramet of a unique
genet) able to regenerate a modular sponge (mother sponge).
However, after gemmule carpet hatching, the various small
growing sponges become fused into a single functional unit
with a unique aquiferous system (Figure 8.8) (Pronzato and
Manconi, 1994a,b,1995). This phenomenon demonstrates
the modular organization of these sponges and their clonal
strategy. It also shows the potential of gemmules to dispersal within either the same or new habitats via production
of clonal metapopulations/meta-individuals (Figure 8.8)
(Manconi et al., 1995; Pronzato and Manconi, 1995).
The new concept of “meta-individual,” or spatially frag- p0135
mented individuals (Figure 8.8), can be added to the other
definitions that have been formulated to describe individual functional units in the phylum Porifera (Pronzato and
Manconi, 1995). Among clonal animals, sponges are the
only ones that, after the proliferation of the ramets, can
f0040
FIGURE 8.7 Reproductive modes by sexual and asexual elements: (a) spermatic cyst; (b) egg cell surrounded by nurse cells; (c) larva of Ephydatia
fluviatilis; (d) larva of Spongilla lacustris; and (e) gemmule of Spongilla lacustris (Modified from several historical sources).
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SECTION | III
140
Protozoa to Tardigrada
f0045
FIGURE 8.8 The spatial structure of a “freshwater sponge individual/population” is based on the ramet-genet concept in the framework of a clonal
strategy by gemmules. Step 1, (a) a gemmule (G) colonizing a water body hatches and generate (b) an adult active sponge (S) producing (c) by metamorphosis a carpet of dormant gemmules (nG). Step 2, Three hypothetical pathways are possible: (1) cyclic metamorphosis between active sponge (S) and
gemmules carpet (nG) continues without changes in the same microhabitat; (2) dispersal of gemmules from the carpet (nG) within the same water body
generates a new clonal sponge population (cP); or (3) dispersal of gemmules from the carpet (nG) in several distant water bodies generates several clonal
[AU8] populations (cP). The result is a clonal meta-population (cMP). The ramets (dispersed clonal sponges/populations) from a single genet (mother sponge)
match the concept of “meta-individual” (MI).
refuse into a single functional unit (i.e., original mother
sponge), thereby confirming the extreme plasticity of
sponges and focusing on their status of basal taxon with
alternative strategies (Pronzato and Manconi, 1995). Fusion
between sponges in the same gemmular carpet (clone) was
observed also in vitro (Van de Vyver and de Vos, 1978;
Weissenfels, 1989).
p0140
The modular organization of freshwater sponges is supported, although in a different morpho-functional sense,
by the body architecture of non-gemmulating sponges.
This is shown by Lubomirskia baicalensis, which is composed of serial modules arranged along an apical–basal
axis in arborescent-branched growth forms (Wiens et al.,
2006).
s0040
Life History Changes in Morphology
p0145 Earliest naturalists ascribed freshwater sponges and Porif-
[AU5]
era, in general, among plants because of their sessile nature,
greenish-brownish color, and growth form. Spongilla lacustris was described by Linnaeus (1759) as “repens, fragilis,
ramis teretibus obtusis” (“creeping, fragile, with cylindrical
branches showing swellings at their ends,” Figure 8.9) in
the second volume on Plantae of the Systema Naturae, and
its type material was preserved in the Linnean Herbarium
(Manconi and Pronzato, 2000). Sponges were not recognized as animals until 1765, when the internal water current
was first described, and only in 1875 did Huxley propose
the complete separation of sponges from other Metazoa
(Gaino, 2011).
Three morphs characterize the habitus of freshwater p0150
sponges during the life cycle: namely, the sessile growing
adult (sponge-like), the planktonic short-life swimming
larva, and the dormant gemmule (restant body/propagule).
The growth form of adults ranges from thin crusts strictly
adhering to the substratum to massive cushions; they may
also form variably erected finger-like branching and arborescent growth forms (Figures 8.6 and 8.9). Dimensions are
variable from a few millimeters to large irregular patches
and bush-like or arborescent forms of more than 1 m in
diameter/height. Body consistency is from soft to fragile
to very hard (stony). A wide range of colors are displayed,
from whitish to greenish or brilliant emerald green, or grayish to dark brown and black. The colors result from sponge
pigments, or symbiotic bacteria and algae, or englobed thin
detritus.
During seasonal dormancy, most freshwater sponges are p0155
represented only by gemmules (Figure 8.10) or small spherules (0.25 mm to over 1 mm in diameter) that adhere to hard
substratum, float at the water surface, remain within the old
skeleton, or rest on the silty/sandy bottoms.
Anatomy and Physiology
s0045
Porifera possess no head and no tail; they are basal metazoans p0160
characterized by the absence of true tissues (with few exceptions), a muscular or nervous system sensu stricto, a digestive cavity, and gonads. The body architecture (Figure 8.11)
is arranged around the aquiferous system, which consists
of a network of canals and chambers (in the complex,
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Chapter | 8
141
Phylum Porifera
(a)
(b)
f0050
[AU9]
FIGURE 8.9 (a) The genus Badiaga is a pre-Linnean taxon established by Buxbaum (1729). Currently, Badiaga is a homeopathic remedy obtained from
freshwater sponges, which is particularly used in Russia. From this artistic representation, we can deduce that the illustrated species habitus is ascribable
to Spongilla lacustris. (Original color plate from Kops et al. (1865).) (b) Linnean holotype of S. lacustris from the Linnean Herbarium.
f0055
FIGURE 8.10 Gemmules of Spongillina. (a) Pectispongilla sp.; (b) Umborotula bogorensis; (c) Spongilla lacustris; (d) Saturnospongilla carvalhoi; (e)
Eunapius carteri; and (f) Trochospongilla horrida (not to scale). The gemmular surface displays a wide array of diagnostic morphotraits at the genus and
species levels: (a) spicules entirely embedded in the ornamented outer spongin layer of the theca; (b) spicules (i.e., birotules) protrude from the surface;
(c) spicules are almost absent; and (d) in a few cases gemmuloscleres belonging to the same gemmule display different morphs. The pneumatic layer of
the gemmular theca ranges from (e) very thick with polygonal chambers and belonging to a single gemmule; to (f) shared by grouped gemmules embedded in a continuous spicular cage; foramen is the variously ornamented aperture (a–c, f) favoring the staminal cells migration from inside the theca during
gemmule hatching.
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SECTION | III
142
Protozoa to Tardigrada
f0060
FIGURE 8.11 The body architecture and the aquiferous system of a sponge. (Figure below: modified from Boury-Esnault and Rützler (1997)).
leucon-type organization of freshwater sponges) with flowing ambient water.
p0165
An external layer of flat cells (pinacoderm), pierced
with small inhalant apertures (ostia) and larger exhalant
apertures (oscula), isolates the sponge internal structure
(mesohyl) from the external environment (Figure 8.11). The
mesohyl includes an extracellular matrix with the consistency of jelly, collagen fibrils and fibers, skeletal structures
with mineral deposits (spicules), and cells. Most body cells
are totipotent, with a high degree of mobility and morphofunctional plasticity. The main sponge cell categories are
cells lining outer and inner surfaces, cells secreting the skeleton (organic and inorganic), contractile cells, totipotent
ameboid cells, and cells with inclusions (Figure 8.11).
The internal flagellated choanoderm bears a monolayer
p0170
of choanocytes lining filtering chambers (Figure 8.12).
Choanocytes are specialized cells characterized by a collar
of cytoplasmic microvilli with a central flagellum actively
driving a unidirectional water current from the inhalant
sponge surface (ostia) through the entire body up to the
exhalant apertures (oscula).
p0175
The aquiferous system (Figure 8.11) is involved in the
production and maintenance of water current flow within
the sponge body. Its morpho-functional role is devoted to
supporting feeding by pinocytosis/phagocytosis, together
with excretion and O2/CO2 exchanges occurring by simple
diffusion. To control osmotic pressure, all cells have to
expel excess water. The system consists of inhalant and
exhalant openings at the sponge surface (ectosome), with
respective canals connecting these openings to the filtering
chambers (choanosome). Incurrent water, with nutrients,
flows from the water column into the inhalant openings
and canal network up to the choanocytic chambers. Filtered
water flows toward the surface along the exhalant canal system and oscules. The direction of the flow of ambient water
is from dermal pores through an inhalant canal to a choanocyte chamber, and then out an exhalant canal and osculum.
A skeleton with inorganic and organic components sup- p0180
ports the sponge body. The inorganic skeleton, as in the entire
class Demospongiae, is made of siliceous (hydrated silicon
dioxide [SiO2]) structures named spicules (Figure 8.1);
their secretion by sclerocytes occurs by silica deposition in
layers around an axial organic filament (evident by transparency upon light microscopy (LM)). The organic skeleton
is composed of an extremely variable amount of collagenous material, which is structured as a network of spongin
fibers or dispersed in the intercellular matrix. Spongin in
the form of fibrils and fibers is an apomorphic trait of the
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Chapter | 8
143
Phylum Porifera
f0065
FIGURE 8.12 The choanocyte is the typical feeding cell of Porifera. These flagellated cells are able to produce water flow within the aquiferous system
and catch very small (a few micrometers) suspended organic particles or bacteria. Ingested food is then transferred to mobile archeocytes.
phylum Porifera. Also, the skeleton of gemmules is made
of spongin armed by siliceous spicules (gemmuloscleres).
Chitin, a polysaccharide, also occurs in the inner layer of
the gemmular theca (Simpson, 1984).
p0185
During cryptobiosis (dormancy), the anatomy of
sponges consists of skeleton remains and gemmules, small
subspherical or emispherical to elliptical structures bearing
a protective theca (Figure 8.10). The gemmular theca architecture is reinforced by species-specific spicules (gemmuloscleres) and functions to protect totipotent/staminal cells
(thesocytes) with their energetically rich yolk platelets. The
theca is mono- to bi- or tri-layered (outer, medium, and inner
layers) and is made of laminar and/or trabeculate to fibrous
spongin to form the pneumatic layer. Two types of gemmules with diverging morpho-functional roles are produced
in the same individual (Manconi and Desqueiroux-Faundez,
1999; Manconi and Pronzato, 2004). Gemmuloscleres are
tangential to the surface or embedded (radially to tangentially) in the gemmular theca. Gemmuloscleres are absent
in some species. At the theca surface, one (foramen) or
more apertures (foramina) permit cells migration toward
the substratum during gemmular hatching (Penney and
Racek, 1968; Volkmer-Ribeiro, 1979, 1986; Weissenfels,
1984; Manconi and Pronzato, 2002,2009). The shape, position, and orientation of foramen are variable.
p0190
Production of gemmules is triggered by environmental
factors such as decreased temperature or low water level
(desiccation) and involves cell aggregation of thesocytes and
the construction of the gemmular theca (Fell, 1974,1995;
Pronzato and Manconi, 1994a,1995; Loomis, 2010).
Thesocytes are characterized by low metabolic activity
p0195
and inhibition of cell division to survive critical environmental conditions. In some species, thesocytes display a
state of diapause, metabolic arrest controlled by endogenous factors. The persistence of high osmotic concentration
(due to polyols) maintains metabolic depression and turns
off cell division. After exposure to unfavorable conditions,
cells shift from diapause into quiescence in which metabolic depression is controlled by environmental factors.
Transition to quiescence requires the conversion of polyols
to glycogen, reducing the osmotic concentration early in the
germination process, and the occurrence of favorable conditions to trigger gemmular germination reestablishes an
active sponge (Fell, 1974,1995; Loomis, 2010). The latter
phase is followed by cell differentiation and production ex
novo of extracellular matrix and the siliceous/proteinaceous
skeleton. Reinvasion of the old skeleton of the mother
sponge, if persistent, could also occur by proliferating cells.
High levels of phenotypic plasticity characterize sponges p0200
(Manconi and Pronzato, 1991; Gaino et al., 1995; Hill and
Hill, 2002), a condition typically restricted to embryonic
development in most Metazoa. The extreme adaptability
and plasticity of sponges are supported by exclusive, typical traits: (a) all cell types are able to change their morphology and physiology and move freely in the extracellular
matrix, being absent cells junctions in most cellular types; [AU6]
(b) gametes derive directly from somatic cells by diffused
gametogenesis; and (c) morphological traits and behavior
are extremely plastic. The concept of plasticity in sponges
matches all organization levels from the cell to the population (Gaino et al., 1995; Pronzato and Manconi, 1995).
Life History, Life Cycle, and Reproduction
s0050
Some aspects of the life history of freshwater sponges have p0205
been largely neglected. Little is known about sexual reproduction in natural conditions and about the effects of ecological factors on the timing of sexual production. The role
of both sexual and asexual reproduction in maintenance of
the population, the amount of reproductive effort, and the
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SECTION | III
144
p0210
p0215
p0220
p0225
tendency of the population to persist or spread are processes
that need long-term in situ studies.
Sexual reproduction occurs by gonochorism and hermaphroditism. Sex reversal from one year to the next is,
however, recorded for S. lacustris (Gilbert and Simpson,
1976). Eggs (clustered oocytes and nurse cells in the mesohyl) derive from archeocytes, whereas spermatozoa derive
from choanocytes (spermatic cyst). Mature sperms (Figure
8.7) are released via an exhalant canal system and the osculum in the surrounding water. Fertilization is internal and
occurs when choanocytes recognizing compatible sperms
(belonging to the same species) from the inner water flow
capture and drive them toward oocytes in the mesohyl.
Trapping of a sperm cell in a vesicle by a choanocyte is
followed by the disappearance of both collar and flagellum
and migration of the carrier cell. Noncompatible spermatozoa are presumably digested as food particles.
Viviparity is the rule, in Spongillina and larvae are
brooded within the mother sponge body up to the completion
of the larval development. At that point, the mature and delicate larvae swim out through the oscules within the exhalant flow. Larval growth is supported by nurse cells. During
development, cells of different types are present among the
blastomeres, and small cells migrate to the periphery of the
embryo and become ciliated. After cell differentiation, ciliate
pinacocytes protect the larval surface, allowing swimming;
whereas in the solid, central region of the larva, large cells
such as archeocytes, sclerocytes, and choanocytes are present
together with a few spicules (Figure 8.7).
The mature larvae (parenchymellae), ready for release,
leave their follicle and cross the mesohyl to an exhalant
canal. Released larvae actively swim and are transported by
water flow for a short time. As settlement time approaches,
the larvae enter a short creeping phase. Once initiated, settlement and early metamorphosis are rapid. The larva come
to rest vertically on the anterior pole, and the cells of this
region spread evenly over the substrate shaping the young
sponge as a cone on which the apex is occupied by the posterior pole. The apex collapses and the sponge becomes
hemispherical; the canal system, with its single apical osculum, is functional soon afterward. The sponge at this time
is about 600 μm in diameter. It retains the single osculum
and hemispherical shape until it reaches 2.0 mm in diameter
after a few days (Bergquist, 1978).
The regulatory aspects of sexual and asexual reproduction are poorly known because sponge species are
differently affected by environmental factors such as temperature, salinity, age, size, peak of vertical fluxes of particles (Witte, 1996), and other metabolic features. Egg
and sperm development (gametogenesis) and their release
involve a high percentage of specimens within a population (synchronization) around the year, with a short period
of pause (Annandale, 1911; Corriero et al., 1994; Bisbee,
1992; Ruengsawang et al., in press). In nondormant sponge
Protozoa to Tardigrada
populations of northern Europe, the triggering of gametogenesis seems to be related to seasonal changes: Oogenesis
and spermatogenesis starts at the beginning of winter and
spring, respectively (Van de Vyver and Willenz, 1975).
Endogenous control of gametogenesis in species that produce gametes during the entire year was hypothesized by
Simpson (1980).
Although sexual reproduction occurs at the beginning of
the vegetative growth phase in northern Europe hibernating
populations of E. fluviatilis (Weissenfels, 1989), this behavior is delayed just before dormancy in aestivating populations of E. fluviatilis (Pronzato and Manconi, unpublished
data) in circum-Mediterranean streams.
The reproductive strategy of freshwater sponges fits
both r and K strategies (Figure 8.13). Although sexual
reproduction (K) represents a source of genetic variability of the population, the extremely vulnerable larvae have
few survival opportunities. On the contrary, gemmules (r)
are strong and resistant propagules that can colonize new
substrates and habitats with a large number of new (clonal)
individuals (Figure 8.13).
Environmental constraints for the active vegetative
phases are dry-up and freezing. In warm, arid climates
such as the Mediterranean area, populations of E. fluviatilis display aestivating behavior to survive summer low
water levels and/or total dry-up as a cryptobiotic biomass
by dormancy. Aestivating dry gemmules may persist on the
substratum as lichenoid patches. Gemmules are cyclically
produced, and upon the reestablishment of suitable environmental conditions they undergo hatching. In such conditions the phases of the life cycle are more or less regularly
timed, and gemmulation seems to be controlled by a synergism of endogenous and exogenous factors (Pronzato et al.,
1993). Dephasing of the life cycle in distant populations
together with the fragmented conditions of hydrographic
basins could have driven speciation by reproductive isolation (e.g., by life cycle inversion) and genetic divergence.
Several species of freshwater sponges cannot tolerate
desiccation, and their geographic range is restricted to areas
where the limiting climatic factors are floods or freezes
and dormancy is displayed as short- to long-term hibernation (Frost et al., 1982; Pronzato et al., 1993; Pronzato and
Manconi, 1994a,1995; Fell, 1995). Among the species characterized by extremely plastic gemmules able to tolerate
long periods of both extremely cold and arid climates, distant populations of E. fluviatilis display hibernating or aestivating life cycles (Figure 8.14) (Penney and Racek, 1968;
Poirrier, 1969; Van de Vyver and Willenz, 1975; Pronzato
et al., 1993; Manconi et al., 1995; Manconi, 2008; Manconi
and Pronzato, 2011). Moreover, gemmulation is not perfectly synchronous within a population, and dephasing is
notable (up to some months), possibly to avoid population
extinction resulting from stochastic events (Manconi and
Pronzato, 1991; Pronzato and Manconi, 1995).
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p0230
p0235
p0240
p0245
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Chapter | 8
145
Phylum Porifera
f0070
FIGURE 8.13 Reproductive strategies of most Spongillina display both K and r phases. The persistence in time of a population matches the K phase
with an almost regular metamorphosis (middle) between the active (vegetative sponge) and the dormant (gemmules) habitus of each sponge in a pluriannual life cycle (A–A–A–A); sexual reproduction (S–S) also is the rule in this strategy (left). On the contrary, gemmule dispersal to colonize new habitats
(A′–A′′–A′′′–A′′′′) matches the r phase when unsuccessful events (x) occur (right).
s0055
GENERAL ECOLOGY AND BEHAVIOR
s0060
Habitat Selection
p0250 Processes of freshwater colonization offered sponges new
niches (i.e., functional roles) in benthic communities subjected to unforeseeable fluctuations of environmental conditions such as floods, freezes, and droughts. Despite their
sessile condition, freshwater sponges face these environmental constraints by means of extreme plasticity in body
bauplan, life cycle, and reproductive behavior. Chronic
morphogenesis supports the regeneration of the sponge
both after cryptobiosis by dormant bodies and after various types of injury (e.g., harsh flooding, predation) by body
fragmentation.
Colonization of continental waters occurred at all latip0255
tudes from equatorial rainforests to both cold and hot deserts and at all altitudes from the coastline to high plains and
mountains. Sponges have even colonized some subterranean environments. An extremely wide variety of lotic and
lentic habitats were colonized, ranging from springs, water
courses (rapids and falls), and brackish waters in estuaries
and enclosed seas to thermal vents, caldera lakes, alpine
lakes, and water bodies in subterranean caves (karstic caves
and anchialine caves). Ephemeral water bodies (pools,
marshes, swamps, billabongs, widians in the Sahara, and
pans in the Namibia deserts) are also suitable habitats,
together with extremely isolated water bodies in both tectonic and oceanic islands and manmade basins such as
pools in gardens, zoological-botanical gardens, and pools
to fonts in archaeological sites (see Manconi and Pronzato,
1994a, 2002, 2007, 2008, 2011; Pronzato and Manconi,
2002). Reservoirs, channels, water tanks, dams, and pipelines are also suitable, although demographic blooms or
massive growth in key positions within pumping devices or
industrial plants are problematic (Potts, 1888; Annandale,
1911; Manconi et al., 1995; Fromont, in litt.).
Bathymetric distribution ranges from hundreds of meters p0260
in some lakes (e.g., Baikal) to surfaces exposed to hot air
and direct sunlight during low water levels (Crane, 1991; de
Ronde et al., 2002; Manconi and Pronzato, 1994b,2002,2009;
Manconi et al., 1995; Manconi, 2008; Tabachnick, pers.
comm.). Freshwater sponges can survive extreme environmental constraints ranging from ice to cold and hot water,
desiccation, silty floods, anoxy, eutrophy, and high concentrations of chemicals, hydrocarbons, and heavy metals (Old,
1932; Jewell, 1935,1939; Sarà and Vacelet, 1973; Harrison,
1974, 1977; Rader, 1984; Rader and Winget, 1985; Van Soest
(Capital) and Velikonja, 1986; Willenz et al., 1986; Francis
and Harrison, 1988; Ricciardi and Reiswig, 1993; RichelleMaurer et al., 1994a,b; Vacelet, 1994; Reiswig and Miller,
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SECTION | III
146
Protozoa to Tardigrada
f0075
FIGURE 8.14 Life cycle. Phases of dormancy and activity in the annual cycle of freshwater sponges able to produce gemmules. On the top: (a) active
sponge; (b) gemmule production; (c) gemmule carpet dormancy; and (d) gemmule hatching. Light gray indicates vegetative activity by sponge; dark
gray indicates dormancy by gemmules. On the bottom: life cycle annual rhythm and time shift according to the climate in overwintering (A), short-term
aestivation (B), and long-term aestivation (C) populations of the same species. A: in a very cold climate the short active phase (1) occurs in summertime,
whereas overwintering dormancy (3) is long; in this case, gemmulation (2) is longer (about 2 months) than hatching (about 1 month). B: in a warm
favorable climate, the active phase (1) is very long, whereas summertime dormancy (3) is short; gemmulation (2) and hatching (4) are quick processes
occurring in less than 1 month. C: in a hot, arid climate, the length of aestivation (3) and the winter active phase (1) is similar, whereas gemmulation (2)
and hatching (4) last for about 1 month.
1998; de Ronde et al., 2002; Rota and Manconi, 2004; M
­ üller
et al., 2007). However, despite their apparently tolerant
behavior, sponges are at risk in water bodies with increasing
environmental alterations and pollution.
p0265
Substrates colonized by sexual and asexual propagules
include rocks, boulders, pebbles, shells, wood debris, roots
or branches of riparian trees and bushes, aquatic plants,
living bivalves and gastropods, and various manmade substrata such as glass, cement, boats, floating devices, plastic
(nylon lines, fishing nets, and bottles), and metallic objects
[AU7] (Manconi and Pronzato, 2002, 2008, 2009).
s0065
Feeding Behavior
p0270 Sponges display a low-cost feeding strategy as active filter
feeders and their pumping activity has a key role in water
purification (Sarà, 1973; Reiswig, 1974). Feeding activity
is performed by phagocytosis on a high fraction of particles
ranging in size from zoo-phyto-pico-plankton to bacteria
and by absorption of dissolved organic matter (Frost, 1978,
1980a, b; Willenz and Rasmont, 1979; Francis and Poirrier, 1986; Willenz et al., 1986; Van de Vyver et al., 1990;
Pile et al., 1996, 1997; Savarese et al., 1997; Ramoino
et al., 2011; Ledda et al., 2014). The feeding current is controlled by contracting both inhalant and exhalant apertures
and canals in the aquiferous system. Bacteria represent a
notably high nutritional resource for freshwater sponges
able to retain about 80% of the microbial component, and
pinacocytes can also perform phagocytosis on bacteria at
the body surface and along canals. In E. fluviatilis, capture
by choanocytes of tagged bacteria (30 min) is followed by
their transfer (30 min) into the mesohyl. Wastes begin to
be extruded 24 h later, and after 48 h no further fluorescent material is evident in the sponge body (Schmidt, 1970;
Weissenfels, 1975, 1976; Reiswig, 1975; Harrison, 1977;
Mank and Kilian, 1979; Frost, 1978, 1980a,b; Willenz and
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147
Phylum Porifera
Rasmont, 1979; Willenz, 1980; Willenz and Van de Vyver,
1982; Francis and Poirrier, 1986; Willenz et al., 1986;
Richelle et al., 1988; Van de Vyver et al., 1990; Imsiecke,
1993).
p0275
The pumping rate of sponges is impressive. A massive
sponge can pump about 1000 times its own volume of water
over 24 h (Reiswig, 1971a,b). Among freshwater sponges,
S. lacustris can filter more than 6 ml/h/mg dry mass (Frost,
1980a); at this rate, a finger-sized sponge could filter more
than 125 l in a day (Frost, 1991). Dense sponge communities act as efficient natural biofilters.
p0280
Sponges have a basic role in the recycling of organic
matter in natural processes of freshwater purification and
contribute to the energetic equilibrium of lentic and lotic
ecosystems (Annandale, 1911; Manconi and Pronzato,
2007, 2008, 2009). They are also centers of biological
association in which a cooperative activity is displayed
by several associated benthic taxa ranging from heterotrophic organisms to grazers, filter-feeders, detritivors, and
predators. Conservation of freshwater sponge fauna and an
increase in their presence by means of the culture of sponge
mesocosms in waters enriched of organic matter would represent a sustainable approach to maintaining biodiversity
and improving the rational management of freshwater natural resources (Manconi and Pronzato, 2007, 2008, 2009).
s0070
Other Behavioral Adaptive Traits
p0285 Freshwater sponges are capable of coordinated behavioral
responses and rhythmic contractions, both endogenously and
in response to external stimuli. This may involve part of the
sponge body or the whole individual, and it occurs despite the
absence of true muscles and neurons. This behavior has repeatedly been observed in species of the genus Ephydatia (De Vos
and Van de Vyver, 1981; Weissenfels, 1984, 1990; Elliott and
Leys, 2007). Chemical messengers are able to induce or modulate the rhythm of contraction and feeding, thus proving that
a relatively complex system regulates this phenomenon by
acting upon specific receptor systems (Ellwanger and Nickel,
2006; Ellwanger et al., 2007; Ramoino et al., 2007, 2011).
p0290
The lifespan of species producing gemmules is pluriannual, with an annual cycle according to the length and the
chronology of seasonal rhythms with four phases: vegetative
growth, gemmulation/sexual reproduction, cryptobiosis,
and hatching of gemmules and regeneration (Weissenfels,
1989; Pronzato and Manconi, 1994a, b; 1995).
p0295
The wide adaptive radiation of Spongillina seems to
be strictly related to the behavior of cryptobiosis by dormancy of asexual gemmules with a double functional role
as resistant bodies to persist in situ and propagules for dispersal. The presence of resistant asexual bodies (gemmules)
sharing most traits with recent Spongillidae in one of the
best preserved fossils, P. chubutensis, indicates that gemmules have been extremely conservative structures since the
Cretaceous period (Racek and Harrison, 1975; Manconi and
Pronzato, 2002, 2007).
Cryptobiosis occurred in the evolutionary history of different marine lineages of Porifera, i.e., Astrosclera, Petrobiona,
and a few species of Suberitidae, Clionaidae, and Haliclonidae. This preadaptive strategy clearly favored dispersion in
continental waters. Key adaptive characters favoring the colonization of continental freshwaters are strictly related to the
phenomenon of cryptobiosis: (1) the persistence in the adult
body of totipotent cells (continuous morphogenesis), (2)
the inclusion of staminal cells in a gemmular theca, (3) the
appearance of a protective multilayered proteinaceous theca,
and (4) the production of stout, ornamented gemmuloscleres.
These characters allow the sponge to overcome stressful environmental-climatic conditions, e.g., riverbed drying, strongly
adhering to the substratum (persistence in situ). Other ecological strategies are to float during floods and potentially adhere
to the body of carriers for dispersal (Figure 8.15).
A cryptobiotic phase characterized by simple gemmules
with a monolayered theca of spongin protecting totipotent
cells is displayed in the life cycle of only some species
of euryhaline Haplosclerina living in the marine/brackish
water of epicontinental seas (Fell, 1974, 1978; Fell et al.,
1979; de Weert, 2002). This supports the hypothesis that
marine organisms could adapt to freshwater environments
only via an intermittent brackish stage (“estuary effect”)
(Park and Gierlowski-Kordesch, 2007).
Among the recent Spongillina, the trait “gemmules with
simple to complex architecture” is shared by families with
the highest taxonomic richness: namely Spongillidae, Metaniidae, and Potamolepidae; but it is absent in the families
Lubomirskiidae, Malawispongiidae, and Metschnikowiidae (Manconi and Pronzato, 2002, 2007, 2008, 2009).
Absence or extreme scarcity of gemmules also occurs in
populations of Spongillidae that do not undergo complete
yearly regression with gemmule formation (Manconi and
Pronzato, 2002, 2007, 2008). Mechanical and physiological
performance of gemmules supports survival and persistence
of sponges under extreme conditions.
Gemmules produced by most species of Spongillina show
wide architectural diversity of the gemmular theca made by
spongin strengthened by species-specific spicules (gemmuloscleres). These protect totipotent cells (thesocites) rich
in energetic resources and characterized by low metabolic
rhythms. Growth after hatching to produce an active sponge
occurs by means of a proliferative process, followed by cell
differentiation and production ex novo of extracellular matrix
together with a siliceous/proteinaceous skeleton. Most gemmules of widespread Spongillidae and Metaniidae bear a variably structured mono-bi- or tri-layered theca of laminar and/
or trabeculate (pneumatic layer) spongin with one or more
foramina favoring cell migration on the substratum during
gemmular hatching (Volkmer-Ribeiro, 1979, 1986; Penney
and Racek, 1968; Manconi and Pronzato, 2002).
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p0300
p0305
p0310
p0315
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SECTION | III
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Protozoa to Tardigrada
f0080
FIGURE 8.15 In the evolutionary history of sponges, the key adaptive characters favoring the colonization of continental freshwater are strictly related
to the phenomenon of cryptobiosis: (a) persistence in the adult body of totipotent cells (continuous morphogenesis); (b) inclusion of staminal cells in a
gemmular theca; (c) appearance of a protective multilayered proteinaceous (spongin) theca; and (d) production of stout, ornamented gemmuloscleres.
These characters allow the sponge to overcome stressing environmental-climatic conditions. For example, they can persist in situ in a drying riverbed by
strongly adhering to the substratum. During floods, they can float downstream or adhere to the body of potential carriers for dispersal.
Simple gemmules lacking pneumatic layer and foramen are produced by a few species belonging to the family Potamolepidae endemic to ancient hydrographic basins
of the Afrotropical and Neotropical regions (Brien, 1970;
Volkmer-Ribeiro, 1990; Manconi and Pronzato, 2002,
2007, 2008, 2009). In contrast, a perennial life cycle lacking gemmules and degenerative/regenerative phases is typical of families endemic to some ancient lakes and of some
spongillids (Potts, 1888; Manconi and Pronzato, 2002).
p0325
The ability to produce gemmules might have been lost
several times in different lineages and in habitats characterized by a persistent volume of water and by long-term
environmental stability, as in Lake Baikal (Efremova and
Goureeva, 1989; Efremova, 2004). An evolutionary shift
from cryptobiotic strategies toward a perennial vegetative
phase channeled toward sexual reproductive modes by progressive reduction in the length of the dormant phase in
the life cycle and a decreased gemmulation rate may have
occurred at different times and in different geographic areas.
p0330
Dormancy of sponges, as standing biomass, does not
seem strictly necessary in stable habitats, because the continuous production of a few gemmules occurs only in some
p0320
specimens and is probably under the control of endogenous
factors (Potts, 1888; Simpson and Fell, 1974; Pronzato
et al., 1993; Pronzato and Manconi, 1995). This suggests
that in spongillids, as in many other organisms, dormancy is
an adaptive strategy under the control of a biological clock.
In E. fluviatilis, the production of gemmules could be less
frequent, as occurs in some crateric lakes of central Italy, or
it could be extremely abundant, as occurs in several water
bodies of Sardinia and also in the estuarial areas (Pronzato
and Manconi, 1995; Manconi et al., 1995).
The evolutionary success of freshwater sponges was sup- p0335
ported by adaptive life history strategies encompassing clonalmodular organization, cryptobiosis, sexual reproduction by
larvae, asexual processes of fragmentation and gemmulation,
passive dispersal of gemmules, and K and/or r strategies.
Competition and Cooperation
s0075
Freshwater sponges host a notably diverse assembly of p0340
organisms involved in interspecific relationships ranging
from endocellular symbiosis, to inquilinism, commensalism, or predation. Highly specialized predation is well
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Chapter | 8
149
Phylum Porifera
known in the case of some species of associated Trichoptera, Neuroptera, and Diptera. Caddisfly larvae in the family
Leptoceridae (Heiman and Knight, 1969) and neuropteran
larvae of the family Sisyridae (Pennak, 1978) eat sponges.
Sponges have been reported to be fed upon by some freshwater turtles, ducks, and fish, including darters in which
spicules were found in stomach contents (Volkmer-Ribeiro
and Grosser, 1981; Dominey and Snyder, 1988; McCauley
and Longcore, 1988; Dostine and Morton, 1989; Seigel and
Brauman, 1994; Kennett and Tory, 1996; Armstrong and
Booth, 2005). A special case of exclusive spicule ingestion
from sediments rich in spicules with blunt tips was recorded
for Neotropical oligochaetes (Omodeo and Coates, 2001).
p0345
Sponges are suitable habitats and are associated with or
host bacteria, unicellular algae, protists, hydrozoans, turbellarians, nematodes, rotifers, oligochaetes, leeches, bivalves,
gastropods, decapods, isopods, amphipods, copepods, cladocerans, ostracods, hydracarins, and bryozoans, as well as
several orders/families of insects (Ephemeroptera, Odonata,
Plecoptera, Hemiptera, Megaloptera, Neuroptera, Trichoptera, Lepidoptera, Coleoptera, and Diptera) encompassing typical spongillaflies (Pavesi, 1881; Annandale, 1911;
Schröder, 1935; Arndt and Viets, 1938; Berg, 1948; Brown,
1952; Brønsted and Brønsted, 1953; Brønsted and Løvtrup,
1953; Parfin and Gurney, 1956; Steffan, 1967; Heiman and
Knight, 1969; Robak, 1968; Poirrier and Arceneaux, 1972;
Volkmer-Ribeiro and De Rosa-Barbosa, 1974; Resh, 1976;
Resh et al., 1976; Pennak, 1978; Frost and Williamson, 1980;
Moretti and Corallini-Sorcetti, 1980; Kahl and Konopacka,
1981; Konopacka and Sicinski, 1985; Crowell, 1990; Melao
and Rocha, 1996; Gugel, 2001; Zitzler and Cai, 2006; Gorni
and Alves, 2008; Fusari et al., 2012). The association of some
insects with sponges is obligatory, as in the case of Ceraclea
(Trichoptera), Demeijerea, Oukuriella, and Xenochironomus
(Chironomidae), Climacea, and Sisyra (Neuroptera). Among
the few predators of spongillinas, the larvae of spongillaflies
(Sisyridae) live exclusively in association with several species of spongillinas and spend their life (three instars) inside
the sponge body, on which they feed using mouthparts highly
specialized for cell piercing and sucking (Figure 8.16).
p0350
Sponges give shelter to some fishes and amphibians,
serving as a nesting site for fertilized eggs in several cases,
e.g., Eunapius carteri and Gobius alcockii in India or S.
lacustris and Euproctus platycephalus in Sardinia (Annandale, 1911; Kilian and Campos, 1969; Pronzato and Manconi, 2002), and other sponges are reported to function as
a refuge for terrestrial invertebrates, including millipedes
during inundation of Amazonian floodplains (Adis, 1992).
Microbial diversity, interactions, and evolutionary
p0355
traits within complex symbiotic communities (endocellular microalgae and bacteria) associated with freshwater
sponges have also been analyzed based on biochemical
and molecular relationships (Williamson, 1979; Frost
and Williamson, 1980; Latyshev et al., 1992; Frost et al.,
1997; Dembitsky et al., 2003; Glyzina et al., 2007; Müller
et al., 2007).
Sponges as a Natural Resource
s0080
Freshwater sponges have been used since ancient times p0360
(Neolithic) by humans, and some African and Amazonian
populations produce ceramics tempered by siliceous sponge
spicules (Linnè, 1925; Serrano, 1933; Brissaud and Houdayer, 1986; Adamson et al., 1987; McIntosh and MacDonald, 1989).
Other practical uses are in the field of cosmetics and medi- p0365
cine (Manconi and Pronzato, 2008). For example, dried spongillids were used in the nineteenth century by young Russian
ladies to scrub their faces to have rosy cheeks (Kuznetzow,
1898). Some cosmetics today exploit the scrubbing action of
siliceous spicule powder, and recently this application was
patented in the field of dermatology (­Villani, 2009, 2010).
In the seventeenth century, Samuel Hahnemann enclosed
freshwater sponges in his Materia Medica as a homeopathic
remedy for psoriasis with the common pre-Linnean name of
Badiaga (Eulenburg and Afanasiev, 1891; Schröder, 1942;
Allen, 1986). Although the species used were supposed to be
Spongilla fluviatilis and Spongia palustris (­Figure 8.6), some
confusion exists in their identification. Spongilla lacustris was
also used in China for more than 500 years as a traditional
medicine to reinforce the kidney and support yang (aphrodisiac), although pharmacological research on this species is
poor (Manconi et al., 1988; Hu et al., 2009). Unfortunately, the
exact genus and species of the sponges used previously and
currently as cosmetics and medicine are questionable because
of inadequate taxonomic expertise.
Siliceous microfossils of freshwater sponges are also p0370
useful indicators of paleoenvironmental changes through
analysis of spicule remains in sediments (spiculites bed)
in lake beds and soils. The time interval needed to form a
spiculite a few centimeters thick is estimated to have ranged
from years to decades (Schindler et al., 2008). Variations in
the species composition of sponges and diatoms together
with the amount of siliceous spicules and frustules (diatomites) are indicators of water quality changes and past salinity conditions (Harrison et al., 1979; Harrison and ­Warner,
1986; Harrison, 1988; Schwandes and Collins, 1994;
­Volkmer-Ribeiro and Turcq, 1996; Candido et al., 2000;
Frost, 2001; Machado et al., 2012; Pearce et al., 2012).
COLLECTING, REARING, AND
PREPARATION FOR IDENTIFICATION
s0085
Mature sponges can be collected by visually surveying p0375
hard substrata, wading, snorkeling, or scuba diving, or by
using submersible devices in very deep water. Sponges
prefer shaded microhabitats, where they may be found
around or under stones, boulders, or wood snags. Sponges
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SECTION | III
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Protozoa to Tardigrada
f0085
FIGURE 8.16 The spongillaflies are Diptera, belonging to the family Sisyridae, which spend their larval stages protected inside the body of several
species of freshwater sponges. (Original color plate from Joutel (1900)).
can occasionally be collected during dredging or dragging
operations. Sampling is easier during low water periods
when dried sponges and/or gemmular carpets can be found
on substrata. Species-level identification requires collecting gemmules from the basal portion of the body or sometimes from hard substratum to which they are adhered.
However, gemmules will not be present in some portions
of the life cycle.
p0380
Elliott and Leys (2007) described techniques for culturing sponges, as follows. Gemmules and pieces of the freshwater sponge should be stored in unfiltered lake water at
4 °C in the dark until use (Ricciardi and Reiswig, 1993).
When sponge pieces were stored in bags and aerated
monthly, gemmules were viable for at least 1 year. Gemmules can be removed from the spicule skeleton by gently rubbing sponge fragments between two pieces of wet
corduroy. Loose gemmules should then be washed in cold
distilled water (4 °C) to remove debris, sterilized with a 1%
hydrogen peroxide (H2O2) solution for 5 min, and rinsed
with cold distilled water to remove excess H2O2. Then,
gemmules can be transferred with sterile pipettes to Petri
dishes containing Strekal’s growth medium (Strekal and
McDiffett, 1974) or M-medium (Funayama et al., 2005).
For whole-mount preparations, single gemmules are p0385
placed on an ethanol-washed, flamed glass or a plastic
22-mm2 coverslip in Petri dishes. For sandwich preparations, one 18-mm2 coverslip can be mounted with dental wax
(Hygenic Corporation, Arkon, OH, USA) at the corners on
a coverslip-bottom culture dish (Willco Wells B. V., Amsterdam, The Netherlands) that has been sterilized in 30% H2O2
and rinsed with 100% ethanol before use. Two gemmules
are then placed at the edge of the raised coverslip, and dishes
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Chapter | 8
151
Phylum Porifera
f0090
FIGURE 8.17 The identification of sponges is based on the morphology and diversity of the siliceous skeleton (spicular complement). The protocol for
spicule preparation, to be observed by scanning electron microscopy (SEM), is as follows: (a) first, a coverglass should be cut in smaller parts to be laid
on a microscope slide to drop on the spicules suspended in alcohol; (b) to obtain perfectly cleaned spicules, the skeleton fragment should be previously
cleaned by boiling nitric acid and rinsing in water and then alcohol several times; (c) each coverglass fragment should be fixed on a stub (metallic support)
for SEM by means (d) of electrically conductive silver glue; and (d) then sputter-coated by gold.
p0390
p0395
p0400
p0405
are left undisturbed at room temperature (21 °C) in the dark.
The growth medium should be replaced every 48 h.
In situ transplants were attempted in Curacao by Debrot
and Van Soest (Capital) (2001) and in the Mekong River
by Ruengsawang et al. (2012). Recently, long-term cultivation of Lubomirskia baikalensis primmorphs was attempted
in vitro (Chernogor et al., 2011).
Sponge samples can be preserved by air drying, ethanol/
methanol, or freezing. The most representative specimens
should be photographed in vivo and/or in situ when possible.
It is not necessary to collect an entire specimen; a representative fragment bearing gemmules is enough. Samples should
be registered in a voucher collection, each with a label reporting basic collection data (locality, date, and collector name).
However, entire sponges (possibly with a part of their substratum) are necessary for museum collections. For methods to
study reproductive cycles and cytology, see Simpson (1963).
General traits for identification are growth form, consistency, color, surface traits, distribution of inhalant and exhalant
apertures, architecture of ectosomal and choanosomal skeleton, topographic distribution, traits of skeletal megascleres and
microscleres, topographic distribution of gemmules, gemmular architecture (foramen, gemmular cage, gemmular theca,
pneumatic layer, and spatial arrangement of spicules), and
gemmuloscleres morphology (Manconi and Pronzato, 2002).
Skeletal and gemmular spicules bear key diagnostic
traits. To characterize morphotraits, representative fragments of sponges need to be dissected for LM and/or scanning electron microscopy (SEM) observation. To obtain
spicule preparations, dissolve sponge fragments in boiling
65% nitric acid or in sodium hypochlorite (ambient temperature) in test tubes and then suspend them in water. To
achieve total sedimentation of light small spicules, the spicules should suspended and repeatedly rinsed with distilled
water and then dehydrated in graded ethanol series, with
a gap of 15–20 min between successive washings. Slowly
pour the alcohol from tubes to leave the deposit of spicules in 0.5 ml of the medium within the tube before dropping some suspended spicules onto slides for LM analysis
or on cover-slide fragments (Figure 8.17) for SEM analysis
(see Manconi and Pronzato, 2000). After the total evaporation of alcohol, fix each cover-slide fragment on a stub
with silver glue drops. The presence of a glass substratum
under the spicules gives the best results as a perfectly black
background in SEM photographs. Dry body fragments, dissociated spicules, entire gemmules, and their cross-sections
should be sputter-coated with gold and observed under
SEM. Morphometric data of diagnostic traits should be performed by LM and/or by SEM. To characterize potential
new species, approximately 50 spicules should be measured
for each diagnostic spicular type.
p0410
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typesetter TNQ Books and Journals Pvt Ltd. It is not allowed to publish this proof online or in print. This proof copy is the copyright property of the publisher and is confidential until formal
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publication.
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Non-Print Items
Abstract
Freshwater sponges, at present considered monophyletic, belong to the suborder Spongillina (Demospongiae, Haplosclerida) and date
back to Paleozoic and Mesozoic. Spongillina consists of seven families containing 47 genera, and 238 species with a geographic range
from widespread to sensu stricto endemic. Growth form varies from encrusting to massive and branched; color from green to pale or
dark brown, and consistency from soft to hard firm or fragile. Skeletal network is pauci- to multi-spicular, alveolate to reticulate with a
variable amount of spongin. Skeleton spicules are smooth or variably ornated megascleres ranging from oxeas to styles and strongyles.
Microscleres, usually present, are oxeas, strongyles, aster-like, pseudobirotules. Gemmules, when present, are spherical to ovoid with
a protective theca. Gemmular theca, typically bi- or tri-layered, usually bears a pneumatic layer and it is armed by gemmuloscleres.
Gemmuloscleres are boletiform (tubelliform), parmuliform, pseudobirotules, oxeas, strongyles, birotules, pseudobirotules, club-like,
botryoidal. Gemmules functional role is as resistant or dispersal bodies. Larvae are always parenchymella. The present is work is a
relatively critical synthesis of the literature, however, a critical phylogenetic revision of established taxa is still in progress.
Keywords: Spongillina; freshwater sponges; global diversity; biogeography; life history; morphology; natural resources.
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