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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 Ecology and Behavior 145
General Systematics 133 Habitat Selection 145
Evolution and Phylogenetics 134 Feeding Behavior 146
Biogeography and Diversity 135 Other Behavioral Adaptive Traits 147
General Biology 138 Competition and Cooperation 148
Body Bauplan 138 Sponges as a Natural Resource 149
Life History Changes in Morphology 140 Collecting, Rearing, and Preparation for Identification 149
Anatomy and Physiology 140 References151
Life History, Life Cycle, and Reproduction 143
s0010 INTRODUCTION From an ecological viewpoint, sponges perform key p0020

functional roles in aquatic ecosystems. These sessile fil-
p0010 Sponges are basal Metazoa lacking true organs and a ner-
ter feeders actively and efficiently pump and clean large
vous system. Their body architecture, usually displaying
amounts of water in almost all oceans and continental
an irregular symmetry, is plastic and characterized by a
waters, from the surface to the higher depths. Pumping
continuous morphogenesis. Since ancient times, sponges
activity by their aquiferous system helps circulate the water
have been a source of biomaterials and bioactive com-
column, particularly in lentic conditions, and it also traps
pounds for the field of pharmaceuticals, biomedicines, and
particulate and dissolved organic matter. Sponges provide
cosmetics.
a living refuge for a wide array of organisms, and symbio-
p0015 Porifera, encompassing over 8000 valid species, are
sis with autotrophic microorganisms contributes to primary
primarily marine invertebrates that have colonized conti-
production. Moreover, their siliceous skeleton contributes
nental water since the Paleozoic era. Species richness of
to the formation of sediments after the sponge’s death.
freshwater sponges is commonly underestimated in both
temperate and tropical latitudes, where new findings often
correspond to the discovery of new species. The present
General Systematics s0015
diversity values doubtlessly appear destined to increase The phylum Porifera is subdivided into four recent classes: p0025
with further research in unexplored and poorly sampled Hexactinellida, Demospongiae, Calcarea, and Homosclero-
areas, and from molecular analyses focused mainly on morpha, according the multi-authored World Porifera Data-
cosmopolitan and widespread species that are currently base (Van Soest et al., 2012) on sponge taxonomy. All recent
assumed to consist of cryptic species complexes. Taxon- freshwater sponges belong to the suborder Spongillina of
omy and paleontology should support the results of molec- the order Haplosclerida (Demospongiae) currently divided
ular biology on a strong morphological and evolutionary into six families, 47 genera, and 238 species (Manconi and
basis. 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. 133
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134 SECTION | III Protozoa to Tardigrada
p0030 Systematics is based on morphology, anatomy, and 1972; Volkmer-Ribeiro and Reitner, 1991; Dunagan,
cytology together with reproductive, developmental, and 1999; Richter and Wuttke, 1999). However, most fossils
molecular biology. A morphological dataset of macro- date back to the Eocene, Miocene, Pliocene, and Pleis-
and micro-morphotraits is diagnostic up to the genus and tocene, e.g., the W-Palaearctic Lutetiospongilla heily and
species level: for example, growth form, consistency, the SW-Afrotropical Ephydatia kaiseri (Rauff, 1926;
color, surface traits, distribution of inhalant and exhal- Rezvoi et al., 1971; Racek and Harrison, 1975; Harrison
ant apertures, architecture of ectosomal and choanosomal and Warner, 1986; Pisera, 1999, 2006; Richter and Wuttke,
skeletal network, topographic distribution of skeletal 1999; Pisera and Saez, 2003). Fossil evidence suggests
megascleres and microscleres (Figure 8.1), gemmular the occurrence of inland waters invasion by sponges from
architecture, and gemmuloscleres traits (Manconi and coastal brackish waters of epicontinental or enclosed seas.
Pronzato, 2002). These pioneer sponges might have colonized and spread
p0035 Most freshwater species produce gemmule with over the other still-joined continents (Racek and Harrison,
species-specific gemmular traits. These are key diagnos- 1975; Manconi and Pronzato, 2007).
tic traits, although gemmules are not always present dur- Although fossil records are discontinuous (Pisera, p0045
ing the entire life cycle of specimens and are always absent 2004; Schindler et al., 2008) and data on present geo-
in some taxa—which substantially complicates taxonomic graphic distribution are scattered (see Manconi and
investigations. The absence of gemmules in some families Pronzato, 2002, 2004, 2005, 2007, 2008, 2009; Manconi
and the possibility of convergence/parallelism at the level et al., 2008; Manconi, 2008), the paleontological
of gemmular morpho-traits have resulted in inconsistencies approach, together with paleobiogeography and histori-
with previously well-established systematics (Penney and cal biogeography, could help clarify when, where, and
Racek, 1968; Manconi and Pronzato, 2002) and have biased how colonization processes occurred during the geologi-
attempts to match phylogenetic relationships at the genus/ cal and paleo-hydrogeological vicissitudes of continents.
species level. The latter could include splitting and fusion of conti-
nental plates, migration of microplates, marine ingres-
sion/regression, enclosure of salty/freshwater bodies in
s0020 Evolution and Phylogenetics
inland depressions or along the coastline, river capture,
p0040 The oldest fossils remains (spiculites) of Spongillina (gen. and desiccation of hydrographic basins (Manconi, 2008).
et sp. indet.) are Paleozoic spicules deposits in the Permo- Paleontological knowledge is fundamental to clarify, at
Carboniferous of Europe (Cayeux, 1929; Schindler et al., least partly, climatic conditions and colonization timing,
2008). More recent taxa refer to the Mesozoic era, e.g., to calibrate morphological-molecular trees, and to test
Eospongilla morrisonensis and Palaeospongilla chubu- phylogenetic hypotheses (Pisera, 2004).
tensis, and date back to the Colorado Upper Jurassic Hypotheses on inland water colonization range from poly- p0050
and Patagonian Lower Cretaceous (Ott and Volkheimer, 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) gem-
muloscleres (not to scale).
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Chapter | 8 Phylum Porifera 135
1978; Volkmer-Ribeiro, 1990; Volkmer-Ribeiro and de architecture. The evolutionary history of some Spongil-
Rosa-Barbosa, 1979; Volkmer-Ribeiro and Watanabe, 1983; lina families (Lubomirskiidae, Malawispongiidae and
Manconi and Pronzato, 1994b; Pronzato and Manconi, 2002). Metschnikowiidae) restricted to ancient lakes and charac-
Brien (1968) and Volkmer-Ribeiro, (1986) hypothesized a terized by the absence of gemmulogenesis and gemmules is
polyphyletic origin from both Haplosclerida and Hadromerida particularly intriguing (Manconi and Pronzato, 2002, 2007,
(Figure 8.2). 2008).
p0055 The monophyletic status of Spongillina, based origi-
nally on classic taxonomic techniques, has been shown to Biogeography and Diversity s0025
be robust from recent syntheses that focused on comparative
analyses of morphotraits by scanning electron microscopy, Current knowledge on biodiversity and biogeographic p0065
the original diagnoses, holotypes, materials from several distribution is based mostly on several scattered histori-
historical collections, and new field surveys (Manconi and cal papers; indeed, exhaustive monographies are scarce
Pronzato, 1994a, 2000, 2002, 2007, 2008, 2009; Pronzato and unfortunately only a few biologists focus at present on
and Manconi, 2002). This is also supported by molecular freshwater sponges (Potts, 1888; Weltner, 1895; Annandale,
approaches (Itskovich et al., 2007, 2008; Efremova et al., 1911; Arndt, 1926; Penney and Racek, 1968; Racek,
2002; Schröder et al., 2003; Addis and Peterson, 2005; 1969; Volkmer-Ribeiro, 1981; Poirrier, 1982; Frost, 1991;
Redmond et al., 2007; Maikova et al., 2010, 2012; Harcet Ricciardi and Reiswig, 1993; Manconi and Pronzato,
et al., 2010; Erpenbeck et al., 2011). The resolution power 1994a, 2002, 2004, 2005, 2007, 2009, 2011; Pronzato and
of molecular analyses is, however, unsatisfactory at family/ Manconi, 1995, 2002; Silva and Volkmer-Ribeiro, 2001;
genus/species levels, probably because the focused genomic Efremova, 2001, 2004).
targets are not fully diagnostic. Data on taxonomic richness of freshwater sponges p0070
p0060 Plesiomorphic morphotraits, e.g., skeletal architecture, increased slowly (Figure 8.3), as shown in the framework
monaxial spicules, gemmules, and gametes structure, and of a historical analysis focusing on taxonomy and bioge-
larvae suggest that Spongillina are allied to marine Hap- ography. The temporal trend—based on the worldwide
losclerina (order Haplosclerida) from epicontinental and number of new species described per year—highlights
enclosed seas. Apomorphies of Spongillina are parenchy- that only two species were described by Linnaeus (1759).
mellas (parenchymella type III sensu Ereskovsky, 1999, Long-lasting periods of inactivity or very low activity
2004) provided by choanocyte chambers, canals, and characterized the following 100 years (1760–1860: <10
spicules as well as gemmules displaying a highly diverse 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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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 rich- (73 species and 26 genera), Palaearctic (65 species and 22
ness increase in the decades: 1888–90, 1910–20, and 1969–70 genera), Afrotropical (52 species and 17 genera), and Ori-
(Figure 8.3). The following authors described many new ental (40 species and 12 genera) regions. Species richness is
sponge species during their careers: Bowerbank (11 spe- lower in other areas. The Australasian Region hosts 35 spe-
cies in 1863), Potts (16 species: 1880–1889), Weltner (15 cies (13 genera), and the Nearctic 31 species (14 genera),
species: 1893–1913), Annandale (29 species: 1907–1918), whereas Pacific Oceanic Islands apparently harbor only
Bonetto and Ezcurra de Drago (15 species: 1966–1973), seven species (6 genera) (Figure 8.4) (See also Manconi and
Volkmer-Ribeiro (12 species with different co-authors: Pronzato, 2002, 2007, 2008, 2009; Van Soest et al., 2012).
1963–1995), and Brien (11 species: 1967–1974). Spongillina seem to occur worldwide in all biogeo- p0080
p0075 In our last global assessment (December 2012), fresh- graphic regions except the Antarctica Region (Figure 8.4),
water sponges comprised 238 species, 47 genera, and six and their geographical distribution is related both to geo-
families including a few incertae sedis taxa (four genera and logical and climatic vicissitudes of the continents and to
five species) (Figure 8.4). The highest taxonomic diversity the long-term dynamics of hydrographic basins. Spreading
at the biogeographic scale is recorded from the Neotropical of sponges in continental waters is constrained by habitat
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fragmentation, and larvae disperse exclusively downstream Islands regions. The most speciose genus with 16 species
by flooding. In contrast, the dispersal power of gemmules is Oncosclera (Manconi et al., 2012). Their “gemmules,
is apparently higher and favored by a successful adaptive usually sessile, with a simple architecture” (no pneuma,
trend in gemmule morphological radiation (e.g., pneumatic thin gemmular theca) seem to be devoted to persistence
layer and spiny spicules), providing a potentially efficient in situ in extreme conditions (low water level to harsh
dispersal structure. flooding and heavy silting in rainforest) typical of tropical
p0085 The geographic distribution and taxonomic richness areas (Brien, 1970; Volkmer-Ribeiro, 1990; Manconi and
of Spongillina show a notably variable pattern, with some Pronzato, 2002, 2009).
taxa being widespread, common, and locally abundant, The condition of cosmopolitanism in Spongillidae Gray, p0100
whereas others are discontinuously distributed and rare or 1867 (22 genera and 155 species) matches well the sexual
monotopic. In this scenario, the crucial potential influence and asexual reproductive modes, and a wide adaptive radia-
on biogeographic patterns by gemmular morpho-functional tion of gemmules resulting in the most speciose genera of the
efficiency as promoter of dispersal power is problematic. entire suborder Spongillina: for example, Corvospongilla 19
p0090 The families Lubomirskiidae Rezvoj, 1936 (four genera species, Radiospongilla 18 species, and Eunapius 16 spe-
and 15 species) and Metschnikowiidae Czerniawsky, 1880 cies (Penney and Racek, 1968; Manconi and Pronzato, 2002,
(one genus and one species) show an extremely restricted 2004; Manconi et al., 2008; Ruengsawang et al., 2012).
geographic range; they are found only in Lake Baikal and the Metaniidae Volkmer-Ribeiro, 1986 (five genera and 25 p0105
Caspian Sea, respectively. A peculiar case is represented by species), which have both sexual and asexual reproductive
Malawispongiidae Manconi and Pronzato, 2002 (five gen- modes and complex gemmules, display a Gondwanian pat-
era and six species) with an extremely disjoined pattern in tern, and are spread in the circumtropical rainforests of the
ancient lakes from the Great African Rift Valley (Tanganyika Neotropical, Afrotropical, Oriental, and Australian regions
and Malawi) to the Syrian-Palestinian Jordan Rift Valley with an enclave (one genus) in the Nearctic.
(Lake Tiberias), and from the Balkan area (Ohrid Lake) to However, the assumption that taxa bearing complex p0110
the extremely distant Wallacea in the Sulawesi Island (Poso gemmules as perfect dispersal devices reach a wider geo-
Lake). The restricted biogeographic pattern of all these graphic range does not match biogeographic patterns at
families seems to be related to the shared trait “absence of the family, genus, and species levels. The case of the genus
dormancy and gemmules” in the life history, with their repro- Corvospongilla seems to be emblematic of this condition.
ductive mode being exclusively sexual or by fragmentation. Species belonging to this genus are characterized by “two
p0095 Potamolepidae Brien, 1967 (six genera and 32 species), gemmular types” with diverging morpho-functional roles. In
which have both sexual and asexual reproductive modes, Corvospongilla mesopotamica the functional role of “heavy,
shows a circumtropical range of Gondwanian origin, and sessile gemmule” is to persist in situ, whereas the “free
species are endemic to ancient hydrographic basins of the light gemmule” is devoted to dispersal (Figure 8.5). The
Neotropical, Afrotropical, Oriental, and Pacific Oceanic 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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the dispersal model because of its notably disjoined range Currently, it is impossible to define the precise geo- p0120
at the biogeographic scale, where 19 species are extremely graphic ranges for several genera/species recorded only
rare and geographically restricted (i.e., endemics). The evo- once or exclusively from restricted geographic areas. More-
lutionary novelty of two diverging gemmular architectures over, knowledge of sponges in some geographic areas is
does not explain well the wide dispersal of the genus Cor- scattered and fragmentary if not almost completely lacking
vospongilla, because each species is rare and endemic to an (e.g., Madagascar, central-northern Asia, Indochina, and
ancient coastal basin (Manconi and Pronzato, 2004, 2008). Wallacea). Species richness seems to be underestimated but
p0115 Although the definition of endemic taxa sensu stricto is destined to increase with further sampling campaigns. On
is problematic, about 50% of freshwater sponge species the other hand, the number of taxonomists on freshwater
are endemic to small areas, but endemicity reaches the sponges worldwide is dramatically low (less than 10), and
highest values (100%) for the families Lubomirskiidae, without the absolutely necessary recruitment of new sponge
Metschnikowiidae, and Malawispongiidae. Also, incertae taxonomists, the field is on the brink of extinction.
sedis species/genera are endemic each to a single hydro-
graphic basin, as probably is the case of Spongillina indet. GENERAL BIOLOGY s0030
from the Towuti lake. Endemic sponges occur in ancient,
old and/or crateric basins such as Titicaca (South Amer- Body Bauplan s0035
ica), Yunnan lakes (western China), Tana (Ethiopia), and
The bauplan of sponges does not perfectly match the con- p0125
Mweru, Luapula, Barombi ma Mbu, and Soden lakes
cepts of colony or individual. It better matches the concept
(Africa). Insular, endemic species are recorded from Japan,
of modular organisms because of their ability to alter body
New Zealand, Fiji, New Caledonia, Philippines, Indonesia,
architecture in time and space. However, the functional units
and Cuba. Endemics are reported also from ancient coastal
(ramets) of other modular animals such as Bryozoa and some
basins (e.g., Louisiana, Florida, western North America,
Cnidaria and Tunicata cannot be isolated from one another,
Iraq, and Brazil). Main biodiversity hotspots (i.e., highest
unlike those of adult freshwater sponges (Pronzato and
species richness and endemicity) are ancient hydrographic
Manconi, 1994a,b; Manconi and Pronzato, 2011).
basins in tropical latitudes, such as Amazonian, Orinoco,
The habitus of freshwater sponges is characterized by p0130
Paranà-Uruguay-Paraguay, Zaire, Mekong, and several oth-
three morphs during the life cycle: namely, the long-life ses-
ers (Manconi and Pronzato, 2002, 2009; Van Soest et al.,
sile adult (Figure 8.6) devoted to growth and reproduction,
2012).
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 dor-
mancy (not to scale).
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the planktonic short-lived swimming larva, and the dormant Manconi, 1994a,b,1995). This phenomenon demonstrates
gemmule (restant body/propagule) (Figures 8.6 and 8.7). the modular organization of these sponges and their clonal
Long-term, in situ investigations of gemmulation in Ephy- strategy. It also shows the potential of gemmules to disper-
datia fluviatilis as an experimental model demonstrated that sal within either the same or new habitats via production
a specimen 5 cm in diameter produces, at the beginning of of clonal metapopulations/meta-individuals (Figure 8.8)
the dry season, a gemmular carpet with more than 2000 (Manconi et al., 1995; Pronzato and Manconi, 1995).
gemmules (ramets) in less than 1 week. Each gemmule in The new concept of “meta-individual,” or spatially frag- p0135
this clonal strategy is an individual unit (a ramet of a unique mented individuals (Figure 8.8), can be added to the other
genet) able to regenerate a modular sponge (mother sponge). definitions that have been formulated to describe individ-
However, after gemmule carpet hatching, the various small ual functional units in the phylum Porifera (Pronzato and
growing sponges become fused into a single functional unit Manconi, 1995). Among clonal animals, sponges are the
with a unique aquiferous system (Figure 8.8) (Pronzato and 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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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 metamor-
phosis 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 Three morphs characterize the habitus of freshwater p0150
sponge), thereby confirming the extreme plasticity of sponges during the life cycle: namely, the sessile growing
sponges and focusing on their status of basal taxon with adult (sponge-like), the planktonic short-life swimming
alternative strategies (Pronzato and Manconi, 1995). Fusion larva, and the dormant gemmule (restant body/propagule).
between sponges in the same gemmular carpet (clone) was The growth form of adults ranges from thin crusts strictly
observed also in vitro (Van de Vyver and de Vos, 1978; adhering to the substratum to massive cushions; they may
Weissenfels, 1989). also form variably erected finger-like branching and arbo-
p0140 The modular organization of freshwater sponges is sup- rescent growth forms (Figures 8.6 and 8.9). Dimensions are
ported, although in a different morpho-functional sense, variable from a few millimeters to large irregular patches
by the body architecture of non-gemmulating sponges. and bush-like or arborescent forms of more than 1 m in
This is shown by Lubomirskia baicalensis, which is com- diameter/height. Body consistency is from soft to fragile
posed of serial modules arranged along an apical–basal to very hard (stony). A wide range of colors are displayed,
axis in arborescent-branched growth forms (Wiens et al., from whitish to greenish or brilliant emerald green, or gray-
2006). ish to dark brown and black. The colors result from sponge
pigments, or symbiotic bacteria and algae, or englobed thin
detritus.
s0040 Life History Changes in Morphology
During seasonal dormancy, most freshwater sponges are p0155
p0145 Earliest naturalists ascribed freshwater sponges and Porif- represented only by gemmules (Figure 8.10) or small spher-
era, in general, among plants because of their sessile nature, ules (0.25 mm to over 1 mm in diameter) that adhere to hard
greenish-brownish color, and growth form. Spongilla lacu- substratum, float at the water surface, remain within the old
[AU5] stris was described by Linnaeus (1759) as “repens, fragilis, skeleton, or rest on the silty/sandy bottoms.
ramis teretibus obtusis” (“creeping, fragile, with cylindrical
branches showing swellings at their ends,” Figure 8.9) in
Anatomy and Physiology s0045
the second volume on Plantae of the Systema Naturae, and
its type material was preserved in the Linnean Herbarium Porifera possess no head and no tail; they are basal metazoans p0160
(Manconi and Pronzato, 2000). Sponges were not recog- characterized by the absence of true tissues (with few excep-
nized as animals until 1765, when the internal water current tions), a muscular or nervous system sensu stricto, a diges-
was first described, and only in 1875 did Huxley propose tive cavity, and gonads. The body architecture (Figure 8.11)
the complete separation of sponges from other Metazoa is arranged around the aquiferous system, which consists
(Gaino, 2011). of a network of canals and chambers (in the complex,
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(a) (b)
f0050
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
[AU9] 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 embed-
ded 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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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 flow- supporting feeding by pinocytosis/phagocytosis, together
ing ambient water. with excretion and O2/CO2 exchanges occurring by simple
p0165 An external layer of flat cells (pinacoderm), pierced diffusion. To control osmotic pressure, all cells have to
with small inhalant apertures (ostia) and larger exhalant expel excess water. The system consists of inhalant and
apertures (oscula), isolates the sponge internal structure exhalant openings at the sponge surface (ectosome), with
(mesohyl) from the external environment (Figure 8.11). The respective canals connecting these openings to the filtering
mesohyl includes an extracellular matrix with the consis- chambers (choanosome). Incurrent water, with nutrients,
tency of jelly, collagen fibrils and fibers, skeletal structures flows from the water column into the inhalant openings
with mineral deposits (spicules), and cells. Most body cells and canal network up to the choanocytic chambers. Filtered
are totipotent, with a high degree of mobility and morpho- water flows toward the surface along the exhalant canal sys-
functional plasticity. The main sponge cell categories are tem and oscules. The direction of the flow of ambient water
cells lining outer and inner surfaces, cells secreting the skel- is from dermal pores through an inhalant canal to a choano-
eton (organic and inorganic), contractile cells, totipotent cyte chamber, and then out an exhalant canal and osculum.
ameboid cells, and cells with inclusions (Figure 8.11). A skeleton with inorganic and organic components sup- p0180
p0170 The internal flagellated choanoderm bears a monolayer ports the sponge body. The inorganic skeleton, as in the entire
of choanocytes lining filtering chambers (Figure 8.12). class Demospongiae, is made of siliceous (hydrated silicon
Choanocytes are specialized cells characterized by a collar dioxide [SiO2]) structures named spicules (Figure 8.1);
of cytoplasmic microvilli with a central flagellum actively their secretion by sclerocytes occurs by silica deposition in
driving a unidirectional water current from the inhalant layers around an axial organic filament (evident by trans-
sponge surface (ostia) through the entire body up to the parency upon light microscopy (LM)). The organic skeleton
exhalant apertures (oscula). is composed of an extremely variable amount of collage-
p0175 The aquiferous system (Figure 8.11) is involved in the nous material, which is structured as a network of spongin
production and maintenance of water current flow within fibers or dispersed in the intercellular matrix. Spongin in
the sponge body. Its morpho-functional role is devoted to the form of fibrils and fibers is an apomorphic trait of the
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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 (due to polyols) maintains metabolic depression and turns
of spongin armed by siliceous spicules (gemmuloscleres). off cell division. After exposure to unfavorable conditions,
Chitin, a polysaccharide, also occurs in the inner layer of cells shift from diapause into quiescence in which meta-
the gemmular theca (Simpson, 1984). bolic depression is controlled by environmental factors.
p0185 During cryptobiosis (dormancy), the anatomy of Transition to quiescence requires the conversion of polyols
sponges consists of skeleton remains and gemmules, small to glycogen, reducing the osmotic concentration early in the
subspherical or emispherical to elliptical structures bearing germination process, and the occurrence of favorable con-
a protective theca (Figure 8.10). The gemmular theca archi- ditions to trigger gemmular germination reestablishes an
tecture is reinforced by species-specific spicules (gemmu- active sponge (Fell, 1974,1995; Loomis, 2010). The latter
loscleres) and functions to protect totipotent/staminal cells phase is followed by cell differentiation and production ex
(thesocytes) with their energetically rich yolk platelets. The novo of extracellular matrix and the siliceous/proteinaceous
theca is mono- to bi- or tri-layered (outer, medium, and inner skeleton. Reinvasion of the old skeleton of the mother
layers) and is made of laminar and/or trabeculate to fibrous sponge, if persistent, could also occur by proliferating cells.
spongin to form the pneumatic layer. Two types of gem- High levels of phenotypic plasticity characterize sponges p0200
mules with diverging morpho-functional roles are produced (Manconi and Pronzato, 1991; Gaino et al., 1995; Hill and
in the same individual (Manconi and Desqueiroux-Faundez, Hill, 2002), a condition typically restricted to embryonic
1999; Manconi and Pronzato, 2004). Gemmuloscleres are development in most Metazoa. The extreme adaptability
tangential to the surface or embedded (radially to tangen- and plasticity of sponges are supported by exclusive, typi-
tially) in the gemmular theca. Gemmuloscleres are absent cal traits: (a) all cell types are able to change their morphol-
in some species. At the theca surface, one (foramen) or ogy and physiology and move freely in the extracellular
more apertures (foramina) permit cells migration toward matrix, being absent cells junctions in most cellular types; [AU6]
the substratum during gemmular hatching (Penney and (b) gametes derive directly from somatic cells by diffused
Racek, 1968; Volkmer-Ribeiro, 1979, 1986; Weissenfels, gametogenesis; and (c) morphological traits and behavior
1984; Manconi and Pronzato, 2002,2009). The shape, posi- are extremely plastic. The concept of plasticity in sponges
tion, and orientation of foramen are variable. matches all organization levels from the cell to the popula-
p0190 Production of gemmules is triggered by environmental tion (Gaino et al., 1995; Pronzato and Manconi, 1995).
factors such as decreased temperature or low water level
(desiccation) and involves cell aggregation of thesocytes and Life History, Life Cycle, and Reproduction s0050
the construction of the gemmular theca (Fell, 1974,1995;
Pronzato and Manconi, 1994a,1995; Loomis, 2010). Some aspects of the life history of freshwater sponges have p0205
p0195 Thesocytes are characterized by low metabolic activity been largely neglected. Little is known about sexual repro-
and inhibition of cell division to survive critical environ- duction in natural conditions and about the effects of eco-
mental conditions. In some species, thesocytes display a logical factors on the timing of sexual production. The role
state of diapause, metabolic arrest controlled by endoge- of both sexual and asexual reproduction in maintenance of
nous factors. The persistence of high osmotic concentration the population, the amount of reproductive effort, and the
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tendency of the population to persist or spread are processes populations of northern Europe, the triggering of gameto-
that need long-term in situ studies. genesis seems to be related to seasonal changes: Oogenesis
p0210 Sexual reproduction occurs by gonochorism and her- and spermatogenesis starts at the beginning of winter and
maphroditism. Sex reversal from one year to the next is, spring, respectively (Van de Vyver and Willenz, 1975).
however, recorded for S. lacustris (Gilbert and Simpson, Endogenous control of gametogenesis in species that pro-
1976). Eggs (clustered oocytes and nurse cells in the meso- duce gametes during the entire year was hypothesized by
hyl) derive from archeocytes, whereas spermatozoa derive Simpson (1980).
from choanocytes (spermatic cyst). Mature sperms (Figure Although sexual reproduction occurs at the beginning of p0230
8.7) are released via an exhalant canal system and the oscu- the vegetative growth phase in northern Europe hibernating
lum in the surrounding water. Fertilization is internal and populations of E. fluviatilis (Weissenfels, 1989), this behav-
occurs when choanocytes recognizing compatible sperms ior is delayed just before dormancy in aestivating popula-
(belonging to the same species) from the inner water flow tions of E. fluviatilis (Pronzato and Manconi, unpublished
capture and drive them toward oocytes in the mesohyl. data) in circum-Mediterranean streams.
Trapping of a sperm cell in a vesicle by a choanocyte is The reproductive strategy of freshwater sponges fits p0235
followed by the disappearance of both collar and flagellum both r and K strategies (Figure 8.13). Although sexual
and migration of the carrier cell. Noncompatible spermato- reproduction (K) represents a source of genetic variabil-
zoa are presumably digested as food particles. ity of the population, the extremely vulnerable larvae have
p0215 Viviparity is the rule, in Spongillina and larvae are few survival opportunities. On the contrary, gemmules (r)
brooded within the mother sponge body up to the completion are strong and resistant propagules that can colonize new
of the larval development. At that point, the mature and deli- substrates and habitats with a large number of new (clonal)
cate larvae swim out through the oscules within the exhal- individuals (Figure 8.13).
ant flow. Larval growth is supported by nurse cells. During Environmental constraints for the active vegetative p0240
development, cells of different types are present among the phases are dry-up and freezing. In warm, arid climates
blastomeres, and small cells migrate to the periphery of the such as the Mediterranean area, populations of E. fluvia-
embryo and become ciliated. After cell differentiation, ciliate tilis display aestivating behavior to survive summer low
pinacocytes protect the larval surface, allowing swimming; water levels and/or total dry-up as a cryptobiotic biomass
whereas in the solid, central region of the larva, large cells by dormancy. Aestivating dry gemmules may persist on the
such as archeocytes, sclerocytes, and choanocytes are present substratum as lichenoid patches. Gemmules are cyclically
together with a few spicules (Figure 8.7). produced, and upon the reestablishment of suitable envi-
p0220 The mature larvae (parenchymellae), ready for release, ronmental conditions they undergo hatching. In such condi-
leave their follicle and cross the mesohyl to an exhalant tions the phases of the life cycle are more or less regularly
canal. Released larvae actively swim and are transported by timed, and gemmulation seems to be controlled by a syner-
water flow for a short time. As settlement time approaches, gism of endogenous and exogenous factors (Pronzato et al.,
the larvae enter a short creeping phase. Once initiated, set- 1993). Dephasing of the life cycle in distant populations
tlement and early metamorphosis are rapid. The larva come together with the fragmented conditions of hydrographic
to rest vertically on the anterior pole, and the cells of this basins could have driven speciation by reproductive isola-
region spread evenly over the substrate shaping the young tion (e.g., by life cycle inversion) and genetic divergence.
sponge as a cone on which the apex is occupied by the pos- Several species of freshwater sponges cannot tolerate p0245
terior pole. The apex collapses and the sponge becomes desiccation, and their geographic range is restricted to areas
hemispherical; the canal system, with its single apical oscu- where the limiting climatic factors are floods or freezes
lum, is functional soon afterward. The sponge at this time and dormancy is displayed as short- to long-term hiberna-
is about 600 μm in diameter. It retains the single osculum tion (Frost et al., 1982; Pronzato et al., 1993; Pronzato and
and hemispherical shape until it reaches 2.0 mm in diameter Manconi, 1994a,1995; Fell, 1995). Among the species char-
after a few days (Bergquist, 1978). acterized by extremely plastic gemmules able to tolerate
p0225 The regulatory aspects of sexual and asexual repro- long periods of both extremely cold and arid climates, dis-
duction are poorly known because sponge species are tant populations of E. fluviatilis display hibernating or aes-
differently affected by environmental factors such as tem- tivating life cycles (Figure 8.14) (Penney and Racek, 1968;
perature, salinity, age, size, peak of vertical fluxes of par- Poirrier, 1969; Van de Vyver and Willenz, 1975; Pronzato
ticles (Witte, 1996), and other metabolic features. Egg et al., 1993; Manconi et al., 1995; Manconi, 2008; Manconi
and sperm development (gametogenesis) and their release and Pronzato, 2011). Moreover, gemmulation is not per-
involve a high percentage of specimens within a popula- fectly synchronous within a population, and dephasing is
tion (synchronization) around the year, with a short period notable (up to some months), possibly to avoid population
of pause (Annandale, 1911; Corriero et al., 1994; Bisbee, extinction resulting from stochastic events (Manconi and
1992; Ruengsawang et al., in press). In nondormant sponge Pronzato, 1991; Pronzato and Manconi, 1995).
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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 plurian-
nual 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 pans in the Namibia deserts) are also suitable habitats,
together with extremely isolated water bodies in both tec-
s0060 Habitat Selection tonic and oceanic islands and manmade basins such as
p0250 Processes of freshwater colonization offered sponges new pools in gardens, zoological-botanical gardens, and pools
niches (i.e., functional roles) in benthic communities sub- to fonts in archaeological sites (see Manconi and Pronzato,
jected to unforeseeable fluctuations of environmental con- 1994a, 2002, 2007, 2008, 2011; Pronzato and Manconi,
ditions such as floods, freezes, and droughts. Despite their 2002). Reservoirs, channels, water tanks, dams, and pipe-
sessile condition, freshwater sponges face these environ- lines are also suitable, although demographic blooms or
mental constraints by means of extreme plasticity in body massive growth in key positions within pumping devices or
bauplan, life cycle, and reproductive behavior. Chronic industrial plants are problematic (Potts, 1888; Annandale,
morphogenesis supports the regeneration of the sponge 1911; Manconi et al., 1995; Fromont, in litt.).
both after cryptobiosis by dormant bodies and after vari- Bathymetric distribution ranges from hundreds of meters p0260
ous types of injury (e.g., harsh flooding, predation) by body in some lakes (e.g., Baikal) to surfaces exposed to hot air
fragmentation. and direct sunlight during low water levels (Crane, 1991; de
p0255 Colonization of continental waters occurred at all lati- Ronde et al., 2002; Manconi and Pronzato, 1994b,2002,2009;
tudes from equatorial rainforests to both cold and hot des- Manconi et al., 1995; Manconi, 2008; Tabachnick, pers.
erts and at all altitudes from the coastline to high plains and comm.). Freshwater sponges can survive extreme environ-
mountains. Sponges have even colonized some subterra- mental constraints ranging from ice to cold and hot water,
nean environments. An extremely wide variety of lotic and desiccation, silty floods, anoxy, eutrophy, and high concen-
lentic habitats were colonized, ranging from springs, water trations of chemicals, hydrocarbons, and heavy metals (Old,
courses (rapids and falls), and brackish waters in estuaries 1932; Jewell, 1935,1939; Sarà and Vacelet, 1973; Harrison,
and enclosed seas to thermal vents, caldera lakes, alpine 1974, 1977; Rader, 1984; Rader and Winget, 1985; Van Soest
lakes, and water bodies in subterranean caves (karstic caves (Capital) and Velikonja, 1986; Willenz et al., 1986; Francis
and anchialine caves). Ephemeral water bodies (pools, and Harrison, 1988; Ricciardi and Reiswig, 1993; Richelle-
marshes, swamps, billabongs, widians in the Sahara, and Maurer et al., 1994a,b; Vacelet, 1994; Reiswig and Miller,
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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 ranging in size from zoo-phyto-pico-plankton to bacteria
et al., 2007). However, despite their apparently tolerant and by absorption of dissolved organic matter (Frost, 1978,
behavior, sponges are at risk in water bodies with increasing 1980a, b; Willenz and Rasmont, 1979; Francis and Poir-
environmental alterations and pollution. rier, 1986; Willenz et al., 1986; Van de Vyver et al., 1990;
p0265 Substrates colonized by sexual and asexual propagules Pile et al., 1996, 1997; Savarese et al., 1997; Ramoino
include rocks, boulders, pebbles, shells, wood debris, roots et al., 2011; Ledda et al., 2014). The feeding current is con-
or branches of riparian trees and bushes, aquatic plants, trolled by contracting both inhalant and exhalant apertures
living bivalves and gastropods, and various manmade sub- and canals in the aquiferous system. Bacteria represent a
strata such as glass, cement, boats, floating devices, plastic notably high nutritional resource for freshwater sponges
(nylon lines, fishing nets, and bottles), and metallic objects able to retain about 80% of the microbial component, and
[AU7] (Manconi and Pronzato, 2002, 2008, 2009). 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
s0065 Feeding Behavior
their transfer (30 min) into the mesohyl. Wastes begin to
p0270 Sponges display a low-cost feeding strategy as active filter be extruded 24 h later, and after 48 h no further fluores-
feeders and their pumping activity has a key role in water cent material is evident in the sponge body (Schmidt, 1970;
purification (Sarà, 1973; Reiswig, 1974). Feeding activity Weissenfels, 1975, 1976; Reiswig, 1975; Harrison, 1977;
is performed by phagocytosis on a high fraction of particles Mank and Kilian, 1979; Frost, 1978, 1980a,b; Willenz and
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Rasmont, 1979; Willenz, 1980; Willenz and Van de Vyver, Cretaceous period (Racek and Harrison, 1975; Manconi and
1982; Francis and Poirrier, 1986; Willenz et al., 1986; Pronzato, 2002, 2007).
Richelle et al., 1988; Van de Vyver et al., 1990; Imsiecke, Cryptobiosis occurred in the evolutionary history of differ- p0300
1993). ent marine lineages of Porifera, i.e., Astrosclera, Petrobiona,
p0275 The pumping rate of sponges is impressive. A massive and a few species of Suberitidae, Clionaidae, and Halicloni-
sponge can pump about 1000 times its own volume of water dae. This preadaptive strategy clearly favored dispersion in
over 24 h (Reiswig, 1971a,b). Among freshwater sponges, continental waters. Key adaptive characters favoring the colo-
S. lacustris can filter more than 6 ml/h/mg dry mass (Frost, nization of continental freshwaters are strictly related to the
1980a); at this rate, a finger-sized sponge could filter more phenomenon of cryptobiosis: (1) the persistence in the adult
than 125 l in a day (Frost, 1991). Dense sponge communi- body of totipotent cells (continuous morphogenesis), (2)
ties act as efficient natural biofilters. the inclusion of staminal cells in a gemmular theca, (3) the
p0280 Sponges have a basic role in the recycling of organic appearance of a protective multilayered proteinaceous theca,
matter in natural processes of freshwater purification and and (4) the production of stout, ornamented gemmuloscleres.
contribute to the energetic equilibrium of lentic and lotic These characters allow the sponge to overcome stressful envi-
ecosystems (Annandale, 1911; Manconi and Pronzato, ronmental-climatic conditions, e.g., riverbed drying, strongly
2007, 2008, 2009). They are also centers of biological adhering to the substratum (persistence in situ). Other ecolog-
association in which a cooperative activity is displayed ical strategies are to float during floods and potentially adhere
by several associated benthic taxa ranging from heterotro- to the body of carriers for dispersal (Figure 8.15).
phic organisms to grazers, filter-feeders, detritivors, and A cryptobiotic phase characterized by simple gemmules p0305
predators. Conservation of freshwater sponge fauna and an with a monolayered theca of spongin protecting totipotent
increase in their presence by means of the culture of sponge cells is displayed in the life cycle of only some species
mesocosms in waters enriched of organic matter would rep- of euryhaline Haplosclerina living in the marine/brackish
resent a sustainable approach to maintaining biodiversity water of epicontinental seas (Fell, 1974, 1978; Fell et al.,
and improving the rational management of freshwater natu- 1979; de Weert, 2002). This supports the hypothesis that
ral resources (Manconi and Pronzato, 2007, 2008, 2009). marine organisms could adapt to freshwater environments
only via an intermittent brackish stage (“estuary effect”)
(Park and Gierlowski-Kordesch, 2007).
s0070 Other Behavioral Adaptive Traits Among the recent Spongillina, the trait “gemmules with p0310
p0285 Freshwater sponges are capable of coordinated behavioral simple to complex architecture” is shared by families with
responses and rhythmic contractions, both endogenously and the highest taxonomic richness: namely Spongillidae, Meta-
in response to external stimuli. This may involve part of the niidae, and Potamolepidae; but it is absent in the families
sponge body or the whole individual, and it occurs despite the Lubomirskiidae, Malawispongiidae, and Metschnikowi-
absence of true muscles and neurons. This behavior has repeat- idae (Manconi and Pronzato, 2002, 2007, 2008, 2009).
edly been observed in species of the genus Ephydatia (De Vos Absence or extreme scarcity of gemmules also occurs in
and Van de Vyver, 1981; Weissenfels, 1984, 1990; Elliott and populations of Spongillidae that do not undergo complete
Leys, 2007). Chemical messengers are able to induce or modu- yearly regression with gemmule formation (Manconi and
late the rhythm of contraction and feeding, thus proving that Pronzato, 2002, 2007, 2008). Mechanical and physiological
a relatively complex system regulates this phenomenon by performance of gemmules supports survival and persistence
acting upon specific receptor systems (Ellwanger and Nickel, of sponges under extreme conditions.
2006; Ellwanger et al., 2007; Ramoino et al., 2007, 2011). Gemmules produced by most species of Spongillina show p0315
p0290 The lifespan of species producing gemmules is plurian- wide architectural diversity of the gemmular theca made by
nual, with an annual cycle according to the length and the spongin strengthened by species-specific spicules (gem-
chronology of seasonal rhythms with four phases: vegetative muloscleres). These protect totipotent cells (thesocites) rich
growth, gemmulation/sexual reproduction, cryptobiosis, in energetic resources and characterized by low metabolic
and hatching of gemmules and regeneration (Weissenfels, rhythms. Growth after hatching to produce an active sponge
1989; Pronzato and Manconi, 1994a, b; 1995). occurs by means of a proliferative process, followed by cell
p0295 The wide adaptive radiation of Spongillina seems to differentiation and production ex novo of extracellular matrix
be strictly related to the behavior of cryptobiosis by dor- together with a siliceous/proteinaceous skeleton. Most gem-
mancy of asexual gemmules with a double functional role mules of widespread Spongillidae and Metaniidae bear a vari-
as resistant bodies to persist in situ and propagules for dis- ably structured mono-bi- or tri-layered theca of laminar and/
persal. The presence of resistant asexual bodies (gemmules) or trabeculate (pneumatic layer) spongin with one or more
sharing most traits with recent Spongillidae in one of the foramina favoring cell migration on the substratum during
best preserved fossils, P. chubutensis, indicates that gem- gemmular hatching (Volkmer-Ribeiro, 1979, 1986; Penney
mules have been extremely conservative structures since the and Racek, 1968; Manconi and Pronzato, 2002).
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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.
p0320 Simple gemmules lacking pneumatic layer and fora- specimens and is probably under the control of endogenous
men are produced by a few species belonging to the fam- factors (Potts, 1888; Simpson and Fell, 1974; Pronzato
ily Potamolepidae endemic to ancient hydrographic basins et al., 1993; Pronzato and Manconi, 1995). This suggests
of the Afrotropical and Neotropical regions (Brien, 1970; that in spongillids, as in many other organisms, dormancy is
Volkmer-Ribeiro, 1990; Manconi and Pronzato, 2002, an adaptive strategy under the control of a biological clock.
2007, 2008, 2009). In contrast, a perennial life cycle lack- In E. fluviatilis, the production of gemmules could be less
ing gemmules and degenerative/regenerative phases is typi- frequent, as occurs in some crateric lakes of central Italy, or
cal of families endemic to some ancient lakes and of some it could be extremely abundant, as occurs in several water
spongillids (Potts, 1888; Manconi and Pronzato, 2002). bodies of Sardinia and also in the estuarial areas (Pronzato
p0325 The ability to produce gemmules might have been lost and Manconi, 1995; Manconi et al., 1995).
several times in different lineages and in habitats charac- The evolutionary success of freshwater sponges was sup- p0335
terized by a persistent volume of water and by long-term ported by adaptive life history strategies encompassing clonal-
environmental stability, as in Lake Baikal (Efremova and modular organization, cryptobiosis, sexual reproduction by
Goureeva, 1989; Efremova, 2004). An evolutionary shift larvae, asexual processes of fragmentation and gemmulation,
from cryptobiotic strategies toward a perennial vegetative passive dispersal of gemmules, and K and/or r strategies.
phase channeled toward sexual reproductive modes by pro-
gressive reduction in the length of the dormant phase in
Competition and Cooperation s0075
the life cycle and a decreased gemmulation rate may have
occurred at different times and in different geographic areas. Freshwater sponges host a notably diverse assembly of p0340
p0330 Dormancy of sponges, as standing biomass, does not organisms involved in interspecific relationships ranging
seem strictly necessary in stable habitats, because the con- from endocellular symbiosis, to inquilinism, commensal-
tinuous production of a few gemmules occurs only in some ism, or predation. Highly specialized predation is well
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known in the case of some species of associated Trichop- 1997; Dembitsky et al., 2003; Glyzina et al., 2007; Müller
tera, Neuroptera, and Diptera. Caddisfly larvae in the family et al., 2007).
Leptoceridae (Heiman and Knight, 1969) and neuropteran
larvae of the family Sisyridae (Pennak, 1978) eat sponges. Sponges as a Natural Resource s0080
Sponges have been reported to be fed upon by some fresh-
water turtles, ducks, and fish, including darters in which Freshwater sponges have been used since ancient times p0360
spicules were found in stomach contents (Volkmer-Ribeiro (Neolithic) by humans, and some African and Amazonian
and Grosser, 1981; Dominey and Snyder, 1988; McCauley populations produce ceramics tempered by siliceous sponge
and Longcore, 1988; Dostine and Morton, 1989; Seigel and spicules (Linnè, 1925; Serrano, 1933; Brissaud and Hou-
Brauman, 1994; Kennett and Tory, 1996; Armstrong and dayer, 1986; Adamson et al., 1987; McIntosh and MacDon-
Booth, 2005). A special case of exclusive spicule ingestion ald, 1989).
from sediments rich in spicules with blunt tips was recorded Other practical uses are in the field of cosmetics and medi- p0365
for Neotropical oligochaetes (Omodeo and Coates, 2001). cine (Manconi and Pronzato, 2008). For example, dried spon-
p0345 Sponges are suitable habitats and are associated with or gillids were used in the nineteenth century by young Russian
host bacteria, unicellular algae, protists, hydrozoans, turbel- ladies to scrub their faces to have rosy cheeks (Kuznetzow,
larians, nematodes, rotifers, oligochaetes, leeches, bivalves, 1898). Some cosmetics today exploit the scrubbing action of
gastropods, decapods, isopods, amphipods, copepods, cla- siliceous spicule powder, and recently this application was
docerans, ostracods, hydracarins, and bryozoans, as well as patented in the field of dermatology (Villani, 2009, 2010).
several orders/families of insects (Ephemeroptera, Odonata, In the seventeenth century, Samuel Hahnemann enclosed
Plecoptera, Hemiptera, Megaloptera, Neuroptera, Trichop- freshwater sponges in his Materia Medica as a homeopathic
tera, Lepidoptera, Coleoptera, and Diptera) encompass- remedy for psoriasis with the common pre-Linnean name of
ing typical spongillaflies (Pavesi, 1881; Annandale, 1911; Badiaga (Eulenburg and Afanasiev, 1891; Schröder, 1942;
Schröder, 1935; Arndt and Viets, 1938; Berg, 1948; Brown, Allen, 1986). Although the species used were supposed to be
1952; Brønsted and Brønsted, 1953; Brønsted and Løvtrup, Spongilla fluviatilis and Spongia palustris (Figure 8.6), some
1953; Parfin and Gurney, 1956; Steffan, 1967; Heiman and confusion exists in their identification. Spongilla lacustris was
Knight, 1969; Robak, 1968; Poirrier and Arceneaux, 1972; also used in China for more than 500 years as a traditional
Volkmer-Ribeiro and De Rosa-Barbosa, 1974; Resh, 1976; medicine to reinforce the kidney and support yang (aphro-
Resh et al., 1976; Pennak, 1978; Frost and Williamson, 1980; disiac), although pharmacological research on this species is
Moretti and Corallini-Sorcetti, 1980; Kahl and Konopacka, poor (Manconi et al., 1988; Hu et al., 2009). Unfortunately, the
1981; Konopacka and Sicinski, 1985; Crowell, 1990; Melao exact genus and species of the sponges used previously and
and Rocha, 1996; Gugel, 2001; Zitzler and Cai, 2006; Gorni currently as cosmetics and medicine are questionable because
and Alves, 2008; Fusari et al., 2012). The association of some of inadequate taxonomic expertise.
insects with sponges is obligatory, as in the case of Ceraclea Siliceous microfossils of freshwater sponges are also p0370
(Trichoptera), Demeijerea, Oukuriella, and Xenochironomus useful indicators of paleoenvironmental changes through
(Chironomidae), Climacea, and Sisyra (Neuroptera). Among analysis of spicule remains in sediments (spiculites bed)
the few predators of spongillinas, the larvae of spongillaflies in lake beds and soils. The time interval needed to form a
(Sisyridae) live exclusively in association with several spe- spiculite a few centimeters thick is estimated to have ranged
cies of spongillinas and spend their life (three instars) inside from years to decades (Schindler et al., 2008). Variations in
the sponge body, on which they feed using mouthparts highly the species composition of sponges and diatoms together
specialized for cell piercing and sucking (Figure 8.16). with the amount of siliceous spicules and frustules (diato-
p0350 Sponges give shelter to some fishes and amphibians, mites) are indicators of water quality changes and past salin-
serving as a nesting site for fertilized eggs in several cases, ity conditions (Harrison et al., 1979; Harrison and Warner,
e.g., Eunapius carteri and Gobius alcockii in India or S. 1986; Harrison, 1988; Schwandes and Collins, 1994;
lacustris and Euproctus platycephalus in Sardinia (Annan- Volkmer-Ribeiro and Turcq, 1996; Candido et al., 2000;
dale, 1911; Kilian and Campos, 1969; Pronzato and Man- Frost, 2001; Machado et al., 2012; Pearce et al., 2012).
coni, 2002), and other sponges are reported to function as
a refuge for terrestrial invertebrates, including millipedes COLLECTING, REARING, AND s0085
during inundation of Amazonian floodplains (Adis, 1992).
PREPARATION FOR IDENTIFICATION
p0355 Microbial diversity, interactions, and evolutionary
traits within complex symbiotic communities (endocel- Mature sponges can be collected by visually surveying p0375
lular microalgae and bacteria) associated with freshwater hard substrata, wading, snorkeling, or scuba diving, or by
sponges have also been analyzed based on biochemical using submersible devices in very deep water. Sponges
and molecular relationships (Williamson, 1979; Frost prefer shaded microhabitats, where they may be found
and Williamson, 1980; Latyshev et al., 1992; Frost et al., around or under stones, boulders, or wood snags. Sponges
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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 corduroy. Loose gemmules should then be washed in cold
operations. Sampling is easier during low water periods distilled water (4 °C) to remove debris, sterilized with a 1%
when dried sponges and/or gemmular carpets can be found hydrogen peroxide (H2O2) solution for 5 min, and rinsed
on substrata. Species-level identification requires collect- with cold distilled water to remove excess H2O2. Then,
ing gemmules from the basal portion of the body or some- gemmules can be transferred with sterile pipettes to Petri
times from hard substratum to which they are adhered. dishes containing Strekal’s growth medium (Strekal and
However, gemmules will not be present in some portions McDiffett, 1974) or M-medium (Funayama et al., 2005).
of the life cycle. For whole-mount preparations, single gemmules are p0385
p0380 Elliott and Leys (2007) described techniques for cultur- placed on an ethanol-washed, flamed glass or a plastic
ing sponges, as follows. Gemmules and pieces of the fresh- 22-mm2 coverslip in Petri dishes. For sandwich prepara-
water sponge should be stored in unfiltered lake water at tions, one 18-mm2 coverslip can be mounted with dental wax
4 °C in the dark until use (Ricciardi and Reiswig, 1993). (Hygenic Corporation, Arkon, OH, USA) at the corners on
When sponge pieces were stored in bags and aerated a coverslip-bottom culture dish (Willco Wells B. V., Amster-
monthly, gemmules were viable for at least 1 year. Gem- dam, The Netherlands) that has been sterilized in 30% H2O2
mules can be removed from the spicule skeleton by gen- and rinsed with 100% ethanol before use. Two gemmules
tly rubbing sponge fragments between two pieces of wet are then placed at the edge of the raised coverslip, and dishes
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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.
are left undisturbed at room temperature (21 °C) in the dark. achieve total sedimentation of light small spicules, the spic-
The growth medium should be replaced every 48 h. ules should suspended and repeatedly rinsed with distilled
p0390 In situ transplants were attempted in Curacao by Debrot water and then dehydrated in graded ethanol series, with
and Van Soest (Capital) (2001) and in the Mekong River a gap of 15–20 min between successive washings. Slowly
by Ruengsawang et al. (2012). Recently, long-term cultiva- pour the alcohol from tubes to leave the deposit of spic-
tion of Lubomirskia baikalensis primmorphs was attempted ules in 0.5 ml of the medium within the tube before drop-
in vitro (Chernogor et al., 2011). ping some suspended spicules onto slides for LM analysis
p0395 Sponge samples can be preserved by air drying, ethanol/ or on cover-slide fragments (Figure 8.17) for SEM analysis
methanol, or freezing. The most representative specimens (see Manconi and Pronzato, 2000). After the total evapo-
should be photographed in vivo and/or in situ when possible. ration of alcohol, fix each cover-slide fragment on a stub
It is not necessary to collect an entire specimen; a representa- with silver glue drops. The presence of a glass substratum
tive fragment bearing gemmules is enough. Samples should under the spicules gives the best results as a perfectly black
be registered in a voucher collection, each with a label report- background in SEM photographs. Dry body fragments, dis-
ing basic collection data (locality, date, and collector name). sociated spicules, entire gemmules, and their cross-sections
However, entire sponges (possibly with a part of their substra- should be sputter-coated with gold and observed under
tum) are necessary for museum collections. For methods to SEM. Morphometric data of diagnostic traits should be per-
study reproductive cycles and cytology, see Simpson (1963). formed by LM and/or by SEM. To characterize potential
p0400 General traits for identification are growth form, consis- new species, approximately 50 spicules should be measured
tency, color, surface traits, distribution of inhalant and exhalant for each diagnostic spicular type. p0410
apertures, architecture of ectosomal and choanosomal skele-
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10008-THORP-9780123850263

THORP: 08
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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