Balian E.V., L?v?que C., Segers H., Martens K. (Eds.) Freshwater Animal Diversity Assessment

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Macropodiella pellucida) or vulnerable (Lederman-

niella thalloidea), primarily as a result of habitat loss

(caused by forestry and agricultural expansion) and

water pollution (IUCN, 2004). Many of the threats to

fresh waters (e.g., climate change, eutrophication,

acidiﬁcation, alien species introductions) will lead to

reduced macrophyte diversity and will, as a result,

threaten the faunal diversity of aquatic ecosystems,

favour the establishment and expansion of exotic

species at the expense of native species, and

challenge our abilities to sustain productions of

aquatic macrophytes that are needed to meet human

consumptive demands.

Acknowledgements We thank the Royal Belgian Institute of

Natural Sciences (RBINSc), in particular K. Marten and

E. Balian, for the opportunity to participate in a workshop on

aquatic biodiversity and the invitation tocontribute to this volume.

We also thank S. Jubeau, A. Lewin and D. McGoldrick for

assistance in assembling data on macrophyte distribution, and

Prof. Dr. C. D. K. Cook and Dr. P. Denny for their advice and

reviews of an earlier version of this article. S. M. Thomaz is a

researcher of the Brazilian Council of Research (CNPq) and

acknowledges this agency for long-term funding.

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26 Hydrobiologia (2008) 595:9–26

123

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global diversity of sponges (Porifera: Spongillina)

in freshwater

R. Manconi Æ R. Pronzato

 Springer Science+Business Media B.V. 2007

Abstract Porifera is a primarily marine phylum

comprising more than 15,000 species. The successful

and wide adaptive radiation of freshwater sponges

(Haplosclerida: Spongillina) has resulted in the

colonization of an extremely wide variety of habitats

at all latitudes. Colonization is dated back to the

Mesozoic, and the mono- or poly-phyletism of

Spongillina, and the n umber of potential sponge

invasions into freshwater is still under debate. Living

freshwater sponges belong to 45 genera in six

families for a total of 219 species. The highest

diversity, at the scale of zoogeographic regions, is

recorded from the Neotropical (65 species), Palae-

arctic (59 species), and Afrotropical regions (49

species). Endemic freshwater sponge species are 103

(47%) out of 219. All species belongin g to the

families Lubom irskiidae, Metschnikowiidae, and

Malawispongiidae are endemic. Endemic species

among the other families are 72% for Potamolepidae,

38% for Spongillidae, and 32% for Metaniidae. Data

on some wide geographic areas are scattered and

fragmentary if not almost completely lacking. Spe-

cies richness is probably underestimated and

doubtless destined to increase with further research.

Keywords Taxonomic richness 

Geographic distribution  Endemicity 

Habitat

Introduction

The successful and wide adaptive radiation of

freshwater sponges has resulted in the colonization

of an extremely wide variety of habitats at all

latitudes, from cold deserts of the Arctic Circle and

Patagonia, to the tropical and equatorial rain forests.

Freshwater sponge s are dispersed in both lentic and

lotic habitats, in continental and insular waters with

perennial or temporary regimes, from coast lines to

high plain and from high mountains to subterranean

environments.

An extremely wide variety of habitats have been

colonized by sponges such as springs, streams, rapids,

falls, swamps, rivers, estuaries, lakes, thermal vents

and springs, caldera lakes, tectonic lakes, alpine lakes,

ancient lakes, salt lakes, karstic caves, anchialine

caves, ephemeral water bodies in both temperate and

strictly arid climates (pools, billabongs, oued in the

Guest editors: E. V. Balian, C. Le

que, H. Segers &

K. Martens

Freshwater Animal Diversity Assessment

R. Manconi (&)

Dipartimento di Zoologia e Genetica Evoluzionistica,

Universita

di Sassari, Via Muroni 25, Sassari 07100, Italy

e-mail: rmanconi@uniss.it

R. Pronzato

Dip.Te.Ris. (Dipartimento per lo Studio del Territorio e

delle sue Risorse), Universita

di Genova, Corso Europa

26, Genova 16132, Italy

e-mail: pronzato@dipteris.unige.it

123

Hydrobiologia (2008) 595:27–33

DOI 10.1007/s10750-007-9000-x

Sahara, and pans in the Namibia deserts), man-made

basins from pools in zoological–botanical gardens to

fonts in archaeological sites, to water tanks, pipelines,

reservoirs, and channels.

Recorded bathymetric distribution ranges from

habitats at hundreds meters of depth in some lakes

(Baikal, Tanganyika, Poso) to the surface of water

bodies exposed to direct sun irradiation during low-

water levels (e.g. tropical swamps, oueds, and coastal

lines of rivers and lakes).

Freshwater sponge s are able to tolerate and survive

extreme chemico-physical conditions ranging from

permafrost, thermal waters, long-lasting dry-up,

ﬂuctuating water levels, stagnant to high speed

waters, anoxy, oligotrophy to eutrophy, high levels

of chemicals, and natural or man-made pollution by

hydrocarbons and heavy metals (Harr ison, 1974;

Pronzato & Manconi, 2002).

Colonised substrata include rocks, boulders, peb-

bles, shells of bivalves and gastropods, wood debris,

roots or branches of riparian trees and bushes, aquatic

plants, and various man-made substrata such as glass,

cement, plastic, and metallic objects (Pronzato &

Manconi, 2002).

Freshwater sponges d isplay highly variable body

shape and dimensions, consistency and colour. Spec-

imens range from thin whitish crusts a few mm thick

strictly adhering to the substratum, to dark brown

massive cushions, to branching or erected growth

forms. In most species the body texture is soft and

fragile while other species are hard and massive. In

unfavourable conditions, such as hard climate, fresh-

water sponges are represented on the substratum

exclusively by carpets of small spherules or resting

bodies known as gemmules.

The main diagnostic traits that enable us to identify

freshwater sponges are skeletal architecture, range of

spicule geometry, size and shape of spicules, and

gemmular traits. Skeletal network is a reticulum of

siliceous spicules associated to a notably variable

amount of spongin. Spicules, megasclere and micros-

cleres, are typically monaxial. Gemmules of

freshwater sponges are subspherical, 100–1200 lm

in diameter , and bear a structured coat of spongin to

protect a mass of totipotent cells contained inside.

Morphological characters of this resting stage (such as

gemmular cage, gemmular theca, gemmular foramen,

arrangement of spicules, architecture of spongin

structures, shape, and ornamentations of spicules) are

notably diversiﬁed and diagnostic at the genus and

species level (Manconi & Pronzato, 2002). Identiﬁca-

tion of g emmule-producing sponges, however, is not

possible when gemmules are absent from specimens

according to their life cycle phase. Moreover, a number

of freshwat er sponges, mostly belonging to taxa

endemic to ancient lakes, do not produce gemmules.

Diversity

The knowledge on diversity and distribution of

sponges from inland waters is reported in a number

of historical synopses (Potts, 1887; Weltner, 1895;

Annandale, 1911; Arndt, 1926) and in more recent

syntheses (Penney & Racek, 1968; Racek, 1969;

Volkmer-Ribeiro, 1981; Poirrier, 1982; Frost, 1991;

Ricciardi & Reiswig, 1993; Silva & Volkmer-Ribe-

iro, 2001; Manconi & Pronzato, 1994, 2002, 2004,

2005; Pronzato & Manconi, 2002; Efremova, 2004).

Living freshwater sponges are ascribed, at present,

to 219 species belonging to 45 genera in six families,

namely Lubomirskiidae Rezvoi, 1936 (4 genera, 10

species), Malawispongiidae Manconi & Pronzato,

2002 (5 genera, 6 species), Metaniidae Volkmer-

Ribeiro, 1986 (5 genera, 25 species), Metschnikow-

iidae Czerniawsky, 1880 (one genus, one species),

Potamolepidae Brien, 1967 (6 gener a, 29 species),

and Spongillidae Gray, 1867 (21 genera, 145 species)

(Tables 1, 2). The family Palaeospongillidae Volk-

mer-Ribeiro & Reitner, 1991 contains exclusively

one monotypic genus of fossil sponges. Three

monotypic genera of living sponges, namely Balliv-

iaspongia, Makedia, and Ohridospongilla are

incertae sedis. The most speciose genera contains

15–17 speci es as in the case of Corvospongilla,

Radiospongilla, and Eunapius.

Species richness of freshwater sponges is high

when compared to that of the other freshwater sessile

invertebrates belonging to Cnidaria and Bryozoa. The

total value of 219 species, considered valid at present

by the authors, is emended since the Systema Porifera

(Manconi & Pronzato, 2002). Species richness is,

however, probably underestimated in both temperate

and tropical latitudes, where new ﬁndings often

correspond to the discovery of a new species or

genus (Manconi & Pronza to, 2004, 2005). The

present diversity values appear doubtless destined to

increase with further research on unexplored or

28 Hydrobiologia (2008) 595:27–33

123

poorly sampled areas, and on the basis of morpho-

logical-molecular analyses focused mainl y on

cosmopolitan and widespread species presently

assumed to consist of complexes of cryptic species.

Phylogeny and historical processes

Porifera is a primarily marine phylum comprising

more than 15,000 marine species ascribed to three

classes, Hexactinellida, Calcarea and Demospongiae

(Hooper & van Soest, 2002). Only the suborder

Spongillina of the highly evolved order Haplosclerida

of the Demospongiae is represented in freshwater

(Manconi & Pronzato, 2002).

The oldest fossil records of freshwater sponges are

so far known only from the genera Palaeospongilla

and Eospongilla dating back to the Cretaceous and

Jurassic from Patagonia and Colorado (Manconi &

Pronzato, in press). Palaeospong illa chubutensis

shares most gemmular traits with living Spongillidae

(Volkmer-Ribeiro & Reitner, 1991) strongly suggest-

ing that gemmular architecture is highly conservative,

since the Mesozoic.

The successful colo nization of inland waters by

these primitive invertebrates seems to be strictly

related to cryptobiosis and to the evolutionary

novelty represented by gemmules. These peculiar

survival devices have a double functional role as

resistant resting bodies to persist in situ, and propa-

gules for dispersal in the same or in distant

hydrographic basins. Gemmules allow sponges to

overcome critica l or extreme environmental condi-

tions and to re-establish an active sponge by the rapid

proliferation of the totipotent cells contained within

them. The pluriannual life cycle of gemmule-pro-

ducing sponges is characterised by four steps:

vegetative growth phase, gemmulation/sexual repro-

duction, cryptobiosis, hatching of gemmules, and

regeneration (Pronzato & Manconi, 1994).

Table 1 Species richness of Spongillina and distribution at the scale of zoogeographic regions

PA NA NT AT OL AU PAc ANT World

Lubomirskiidae 10 0 0 0 0 0 0 0 10

Malawispongiidae 3 0 0 2 0 1 0 0 6

Metaniidae 0 2 17 3 3 3 0 0 25

Metschnikowiidae 1 0 0 0 0 0 0 0 1

Potamolepidae 0 0 11 15 0 0 2 0 29

Spongillidae 44 30 35 28 34 29 3 0 145

Incertae sedis 1011000 0 3

Total 59 32 65 49 37 33 5 0 219

PA: Palaearctic, NA: Nearctic, NT: Neotropical, AT: Afrotropical, OL: Oriental, AU: Australasian, PAc: Paciﬁc oceanic islands,

ANT: Antartic

Table 2 Genera diversity of Spongillina and distribution at the scale of zoogeographic regions

PA NA NT AT OL AU PAc ANT World

Lubomirskiidae 4 0 0 0 0 0 0 0 4

Malawispongiidae 2 0 0 2 0 1 0 0 5

Metaniidae 0 1 5 1 1 1 0 0 5

Metschnikowiidae 1 0 0 0 0 0 0 0 1

Potamolepidae 0 0 3 4 0 0 1 0 6

Spongillidae 13 12 14 9 10 11 3 0 21

Incertae sedis 1011000 0 3

Total 21 13 23 17 11 13 4 0 45

PA: Palaearctic, NA: Nearctic, NT: Neotropical, AT: Afrotropical, OL: Oriental, AU: Australasian, PAc: Paciﬁc oceanic islands,

ANT: Antartic

Hydrobiologia (2008) 595:27–33 29

123

Spongillina share many reproductive characters

with marine Haplosclerida. Sex ual reproduction

occurs by gonochorism, with the development of

brooded parenchymula larvae with a short planktonic

life. The adaptation to inland fragmented and ephem-

eral habitats constrained sponges to particular

changes resulting in differences between marine and

freshwater haplosclerids at the level of sperms, eggs,

embryos and larvae supporting an evidence of an

early divergence (Ereskovskii, 2004).

Sponge phylogeny at the level of higher taxa

(orders) has been outlined based on morphological

traits by van Soest (1991). The mono- or poly-

phyletism of Spongillina , and the number of potential

sponge invasions into freshwater is still under debate

(Brien, 1969; Manconi & Pronzato, 2002; Pronzato &

Manconi, 2002). The absence of gemmules in some

families of freshwater sponges and the high possibil-

ity of convergence/parallelism at the level of

gemmular morpho-traits resulted in inconsistencies

with the nearby well-settled systematics (Penney &

Racek, 1968; Manconi & Pronzato, 2002), and biased

the attempts to match phylogenetic relationships at

the genus/species level.

Geographic distribution

At present it is impossible to deﬁne the precise

geographic ranges for several genera/species without

ﬁrstly undertaking a system atic revision of materials

in both historical and unstudied collections, and a

critical analysis of taxonomic data from the literature.

Several species and some genera have been recorded

only once or excl usively from very restrict ed geo-

graphic areas and the knowledge on some geographic

areas is scattered and fragmentary if not almost

completely lacking (e.g. Madagascar, Central Asia,

Wallacea, West Indies).

Freshwater sponges occur worldw ide except so far

in the Antarctica region, and their g eographical

distribution is related to both geological and climatic

vicissitudes of the continents, and to the long-term

dynamics of hydrographic basins. Different biogeo-

graphic patterns are evident, with some species being

very common and widespread, as in the case of the

Holoarctic Spongilla lacustris (Manconi & Pronzato,

2000), while other species are apparently discontin-

uously distributed, rare or monotopic, as for

Spongilla prespensis and Spongilla stankovici ende-

mic to the Balkanian area (Pronzato & Manconi,

2002).

The highest diversity, at the scale of zoogeo-

graphic regions, is recorded from the Neotropical

with 65 species (23 genera, 3 families), Palaearctic

with 59 species (21 genera, 4 families), and Af ro-

tropical regions with 49 species (17 genera, 4

families). Diversity is lower in the other regions,

namely the Oriental with 37 species (11 genera, 2

families), the Australasian with 33 species (13

genera, 3 families), and the Nearctic with 32 species

(13 genera, 2 families). The lowest diversity is known

from the Oceanic Paciﬁc Islands with ﬁve species (4

genera, 2 families) (Tables 1, 2; Fig. 1).

The geographic range of families varies from a

cosmopolitan diffusion (Spongillidae), to a condition

of an extremely restricted area (Lubomirskiidae in the

Baikal, and Metschnikowiidae in the Caspian Sea). A

peculiar case is represented by Malawispongiidae

recorded only from ancient lakes (e.g. Tanganyika,

Malawi, Tiberias) along the African Great Rift Valley

in SE-Africa to the Sirian-Palestine Jordan Rift

Valley, to the Ohrid lake in the Balkania n area and

Poso lake in the Sulawesi microplate (Manconi &

Pronzato, 2002).

Potamolepidae are present in the rainforests of the

Neotropical, Afrotropical and Paciﬁc Oceanic

Islands, whereas Metaniidae shows a true Gondwa-

nian pattern being spread in the circum-tropical rain-

forests of the Neotropical, Afrotropical, Oriental and

Australian regions with an enclave (one genus) in the

Nearctic. Incertae sedis genera are endemics to old

lakes scattered in South America, Afri ca and Asia

(Titicaca, Tana and Ohrid lakes) (Manconi & Pronz-

ato, 2002).

Endemic freshwater sponge species sensu stricto

(endemic to small areas) are 103 (47% ) out of 219.

All species belonging to the families Lubomirskiidae

(10) from Lake Baikal, Metschnikowiidae (1) from

the Caspian Sea, and Malawispongiidae (6) from

Tanganyika, Malawi, Tiberias, Ohrid and Poso Lakes

are exclusively endemic. Also, the three incertae

sedis species are endemic each to a single lake. The

highest value of endemic species among the most

speciose families is 72% for Potamolepidae, 38% for

Spongillidae, and 32% for Metaniidae.

The highest values of endemicity s.s. are known

for the Oceanic Paciﬁc Islands (60%), Palaearctic

30 Hydrobiologia (2008) 595:27–33

123

(58%) and Afrotropical (51%) regions. Lower values

are shown by the Neotropical (35%), Oriental (24%),

Australasian (15%), and Nearctic (16%) regions.

Endemic sponges also occur in ancient, old and/or

crateric basins such as Titicaca (South America),

Yunnan lakes (W-China), Tana (Ethiopia), Mweru

and Luapula (E-Africa), Barombi ma Mbu and Soden

(W-Africa).

As far as insular freshwater are concerned,

endemics are recorded from Japan, New Zealand,

Cuba, Fiji, New Caledonia, Philippines, and Indone-

sian islands. Endemics are reported also from coastal

basins such as those of Louisiana, Florida, Western

North America and Brazil. Ancient hydrographic

basins such as Amazonian, Orinoco, Parana

-Uru-

guay-Paraguay, Zaire, and several others host a rich

endemic sponge fauna. Main biodiversity hotspots

(i.e. highest species richness) are ancient hydro-

graphic basins in tropical latitudes (Manconi &

Pronzato, 2002).

Sponges as a natural resource

Sponges represent a natural resource for their func-

tional role involved in natural processes of water

puriﬁcation in freshwater ecosystems. These active

ﬁlter feeders and their pumping activity play a key-

role in the re-cycling of organic matter and contri bute

to the energetic equilibrium of freshwater ecosys-

tems. The feeding activity is performed by

phagocytosis on a high fraction of organic particles

and bacteria, and by absorption of dissolved organic

matter. Pumping rate is high, S. lacustris can ﬁlter

more than 6 ml/h/mg dry mass; at this rate, a ﬁnger-

sized sponge could ﬁlter more than 125 l in a day

(Frost, 1980; Pronzato & Manconi, 2002).

Sponges are centres of biological associations,

representing a suitable but selective refuge micro-

habitat, and are host to a notably diverse assemblage

of organisms ranging from other metazoans and

protists to bacteria and algae. Representatives of most

freshwater invertebrate taxa are recorded in sponges,

namely hydrozoans, turbellarians, nematods, oligo-

chaetes, leeches, bivalvs, gastropods, amphipods,

copepods, ostracods, hydracarina, and bryozoans to

several famili es of insects (Pronzato & Manconi,

2002). The inter-speciﬁc relationships range from

endocellular symbiosis, to commensalism or highly

specialized predation as in the case of Neuroptera

(spongillaﬂies or Sisyridae), Tricoptera, and Diptera.

Some ﬁshes and amphibians were also reported to

nest their fertilized eggs in sponges (Pronzato &

Manconi, 2002).

Freshwater sponges have been used also by Homo

sapiens, since ancient times. Some Amazo nian tribes

perform sponge farming usin g gemmules as seeds to

obtain conspi cuous quantities of sponges to str ength

Fig. 1 Distribution of

Porifera species and genera

per biogeographic region.

PA: Palaearctic; NA:

Nearctic; NT: Neotropical;

AT: Afrotropical; OL:

Oriental; AU: Australasian;

PAC: Paciﬁc Oceanic

Islands, ANT: Antartic

Hydrobiologia (2008) 595:27–33 31

123

pottery. An unusual practice is known in cosmetics as

in the case of young ladies in Russia that, in the 19th

century, used dried spongillids to scrub their skin to

have rosy cheeks, but some modern cosmetics are

based on the same material. At present bioactive

products extracted from sponges are considered one

of the most promising source of natural compounds

for the pharmacological and biomedical ﬁelds. It is

well known that S. lacustris and Ephydatia ﬂuviatilis

are homeopathic remedy, since 1700 with the com-

mon prelinnean Russian name of Badiaga (Pronzato

& Manconi, 2002). No freshwater sponges are

currently listed on the ofﬁcial threatened species

lists, although in some cases they are indirectly

protected being sympatric or syntopic with more

‘‘important’’ taxa such as ﬁshes and amphibians.

Successful experimental transplants of freshwater

sponges in natural habitats have also been attempted.

Conservation of the freshwater sponge fauna and

their increase in prevalence and abundance in waters

receiving high inputs of organic matter would

represent an approach to maintain biodiversity, and

to improve the sustainable management of freshwater

natural resources (Pronzato & Manconi, 2002). A

potential role of sponges to control invasive alloc-

tonous species (e.g. dreissenid bivalves) has been also

highlighted (Ricciardi et al., 1995).

Acknowledgements We thank the anonymous referees for

helpful comments on the manuscript. R. Manconi is grateful to

the organizers of the workshop ‘‘A global Assessment of Animal

Diversity in Freshwater’’ for their kind invitation and ﬁnancial

support. Research supported in part by the Italian Ministero

dell’Istruzione, dell’Universita

e della Ricerca Scientiﬁca e

Tecnologica (MIUR-PRIN 2004057217 ‘Zoogeography of

Mediterranean-Southern African disjunct distributions by a

multimethod approach’), the European program INTERREG

Sardinia-Corsica-Tuscany on Biodiversity, Universita

di Sassari

and Universita

di Genova.

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