Porifera research: Biodiversity, innovation and sustainaBility - 2007

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South American continental sponges: state of the art of the research

Ceclia Volkmer-Ribeiro
Museu de Cincias Naturais, Fundao Zoobotnica do Rio Grande do Sul. Rua Dr. Salvador Frana, 1427. 90690-000, Porto Alegre, RS, Brazil. Research fellow of CNPq. cvolkmer@fzb.rs.gov.br Abstract: An intensive survey and study of the South American freshwater sponges started around four decades ago. A large number of results came out which now allows a first overview. An outstanding number of new species of freshwater sponges and even new genera were described. The new materials presented remarkable differences when compared with species from the other continents. This new sponge continental fauna has ever since been subjected to constant checking due to continued and extensive surveys, redescriptions and habitat descriptions. The diversity of the South American sponge fauna is remarkable being possibly the richest in the world. All the review efforts dedicated to this branch of Demosponges are enhancing confidence in the perception of how thrustworthy some characters are and how much their degree of variation is linked to specific habitats, thus enabling the application of this taxonomic knowledge. Keywords: Continental sponges, research, South America, state of the art

Build and rebuild your software

Evolution has been the main word leading the research of all of us as biologists and naturalists in our efforts to understand how living beings have organized themselves and how living processes of all sorts play around us. Lately, with the growing success of replicating life in our labs, evolution includes the idea of how we are setting up life around the world and even around the planetary system. One but not the least of the resulting consequences is that taxonomists are being urged to come up with applied propositions stemming from their taxonomic research. This taxonomic perfectioning takes place and weight as our scientific life goes by and particularly if it stays long enough as that of a large number of us has been staying. Looking back and forth becomes a daily routine with special emphasis on ideas, hypotheses and conclusions, challenged by every new evidence, findings and reading. One starts with grain-size evidences, cements them together, builds up his/her initial operating system and from then on builds and rebuilds oneselfs rationale. Plain common sense, but a painful process And here it is when Science turns into Art and Philosophy. Into art because a scientists always present inquiring impulses are driven by a desire to grasp and reproduce the perfection offered by Nature. Into philosophy because only this endeavour will heal the human mind in its failure to reach that goal.

The gains of the continental approach

A whole new continental fauna of Demosponges, that of the South American continent, was disclosed mainly over the last half of the 20th century. This continental fauna may be comparable in importance to the findings of fossil marine

sponge faunas preserved in rocky outcrops which lately emerged as continental areas. The main difference between these lies on the fact that those marine faunas are a snapshot of the past and not an ongoing movie as the present continental sponge faunas are. And, as such, research on continental sponge faunas benefits from playbacks whenever collecting sites are re-visited to check hypotheses, improve descriptions and study niches (niche sensu Hutchinson 1967). But more than offering easy access to previous collecting sites, continental sponge faunas offer the chance to study biological/ ecological (adaptation),versus geologic (phylogeny) evolution, when continental drift is considered and continental plates are understood as analogous to huge islands. A large isolation degree in respect to time and space causing easier determination and follow up of geographic barriers and vicariance effects (Nelson and Platnick 1981) appears then as important aspects to drive the search for evolving characters in these continental Demosponge faunas aiming to the perfection of species identification, the detection of endemisms and the estimation of the time consumed along all these processes. All such aspects are obviously harder to detect when marine demosponges are considered. Three main realms should be in fact considered in what respects sponges: the marine, the epicontinental and the continental one. The first and third ones need no further explanation. The second one has to do with those seas in the process of continental enclosure, so turning into brackish and then fresh water. This epicontinental marine fraction of the present world waters has seldom been surveyed for sponges, in spite of the fact that other phyla were studied in detail and seen to pass by a drastic reduction in biodiversity, like, for instance the Echinoderms in the Baltic Sea (Hutchinson 1967). The presently known rich marine versus poor freshwater

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poriferan biodiversity (Hooper and van Soest 2002) certainly allows for the expectation that the present epicontinental sponge faunas may also offer very interesting and intriguing selection processes and local extinctions prior to adapting to freshwater.

Continental surveying of an unknown sponge fauna

The first descriptions of South American continental sponges were produced in the 19th century and were based on a few specimens gathered in the Orinoco, Amazon and Uruguay Rivers by foreign explorers and deposited mainly in English and German Museums. The next main taxonomic efforts date to the middle of the last century, when Bonetto and Ezcurra de Drago started to survey the Argentinean continental sponges (Ezcurra de Drago 1971) and VolkmerRibeiro the Brazilian ones (Volkmer-Ribeiro 1981) resulting at present in a number of twenty nine new species, six new genera and one new extant family described, plus one new fossil species and family defined. At this time significant freshwater sponge collections were initiated by these authors at respectively the Instituto Nacional de Limnologia - INALI, Santa F, Argentina and the Museu de Cincias Naturais, Fundao Zoobotnica do Rio Grande do Sul, Rio Grande do Sul, Brazil. Bonetto and Ezcurra de Drago as well as Volkmer-Ribeiro initially faced a confusing situation in what respected the taxonomy of the worlds freshwater sponges long ago split by Carters taxonomic proposals (Jewell 1952). The problem was however overcome due to the remarkable differences that the new materials presented when compared with the descriptions available for the species already known. In that way an outstanding number of new species was described which is resisting the continued studies and surveys that are being carried out until the present. The appearance of Penney and Raceks (1968) comprehensive revision of the worlds freshwater sponges offered next a sound basis for revisional studies of species and genera and proposition of new genera. The master lines established by Penney and Racek were followed by both of the authors, Volkmer-Ribeiro and Ezcurra de Drago, resulting in the validation and enlightening of prior proposals for diverse continental sponge genera by Gray (1867), according to detailed redescriptions (VolkmerRibeiro and De Rosa-Barbosa 1972, Volkmer-Ribeiro 1984, Volkmer-Ribeiro and Costa 1992, Volkmer-Ribeiro and Tavares 1995, Tavares and Volkmer-Ribeiro 1997) of new materials with South American species, which had been only briefly described before.

South American continental plates. In this way genera with exclusive Nearctic-Neotropical distribution came into light such as Corvomeyenia Weltner, 1913 (Volkmer-Ribeiro et al. 2005) and Anheteromeyenia Schrder, 1927 (VolkmerRibeiro 1986a), or with predominant occurrence in these two continents like Racekiela Bass and Volkmer-Ribeiro, 1998 (Bass and Volkmer-Ribeiro 1998).

One genus and five continental plates drifted apart

The continued concern with the search for trustworthy diagnostic specific characters present in dried preserved materials (so that comparative studies could keep on encompassing old preserved materials) was deepened next with the revision of Metania Gray 1867, which has a tropical distribution. Such an effort sprung from the large number of specimens of this genus collected particularly in the Brazilian and Venezuelan Amazonia. At this time the search for what would become trustworthy diagnostic characters for species identification was centered on the large isolated sponge faunas at the plates of South America, Africa, Australia, India and Indonesia split from Gondwana and drifted apart. The resulting study brought to attention the existence of a gondwanian fauna of freshwater sponges (Volkmer-Ribeiro 1986b, Volkmer-Ribeiro and Costa 1992, 1993, Silva and Volkmer-Ribeiro 2001), which Volkmer-Ribeiro and De Rosa-Barbosa (1979) had already indicated by extending the occurrence of the family Potamolepidae from the Ethiopian to other gondwanian plates. An array of characteristics came to light again which confirmed Grays (1867) insight of the value of gemmoscleres, microscleres and the gemmule structure for the definition of genera and species.

Timing the birth of a continental sponge fauna

Also, from the studies on Metania it was possible for the first time to envision the time elapsed in attaining speciation and generic diversification for a group of continental sponges i.e. the Cretaceous drifting apart of the Gondwanian plates. Given the established paradigm that geologic time scales are those required to produce measurable change, particularly of gemmoscleres and microscleres shape and size, as well as on number of spicule categories present, a continued revisional effort was extended to all genera of South American continental sponges, allowing a more confident redefinition of monospecific genera (Acalle, Volkmer-Ribeiro and De Rosa-Barbosa 1972, Uruguaya and Sterrastrolepis, VolkmerRibeiro and De Rosa-Barbosa 1979, and the recognition of new genera (Oncosclera Volkmer-Ribeiro, 1970, Saturnospongilla Volkmer-Ribeiro, 1976, Corvoheteromeyenia, Ezcurra de Drago, 1979, Racekiela Bass and Volkmer-Ribeiro, 1998) as well as the description of new species, among which those where monospecific genera descriptions had been based on: Saturnospongilla carvalhoi, Sterrastrolepis brasiliensis. At present the largest number of records in the South American freshwater sponges are from Brazil and Argentina but reports have also been produced for Suriname (Ezcurra de Drago 1975), Venezuela (Bonetto and Ezcurra de Drago 1973, Volkmer-Ribeiro and Pauls 2000), Chile (Ezcurra de Drago 1974, Kilian and Wintermann-Kilian 1976) and Uruguay

Two neighboring continental plates

A milestone mark at this point was the study of the Edward Potts collection of type materials of the species he described for the United States and Canada (Potts 1887) and deposited at the Academy of Natural Sciences of Philadelphia (VolkmerRibeiro and Traveset 1987). That collection had not been taken in consideration by Penney and Racek (op. cit.). The idea was that comparative descriptions of species restricted to the Nearctic/Neotropical regions would favor any future vicariance studies due to the vicinity of the North and the

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(Berroa Beln 1968). The South American continental plate has so been crossed from north to south and east to west leading to the idea that a large number of habitats has yet to be surveyed for continental sponges in this remarkably diverse continent. Now, quite a different picture has emerged of this South American fauna showing that it is one of the richest, if not the richest in the world.

Applied sponge taxonomy

Continuous checking of habitat characteristics versus species occurrence is a tool never discarded by specialists in their search for the confirmation of a species status and the description of ecomorphic variations of characters (VolkmerRibeiro 1973, Poirrier 1974). The performance of the abovementioned procedures, besides submitting species and genera to constant revisional efforts generates confidence in species definitions and provides a series of applications for this taxonomic knowledge. The first concerns the monitoring of freshwater habitats with respect to the integrity of their biodiversity or their restoration with environmental recovering practices. The knowledge now available encompasses the detection of several sponge assemblages in some typical South American habitats, such as Coastal ponds, lakes and lagoons (VolkmerRibeiro and Machado 2007), Cerrado (Savannah) ponds where an assemblage of five species thrive or where they formed spongillite deposits in the past (Volkmer-Ribeiro et al. 1998). Also, particular sponge assemblages have been detected in large South American rivers, as for instance the rocky bottoms of the middle Uruguay river (Ezcurra de Drago and Bonetto 1969), the rocky tributaries of the middle Paran river, as well as in the macrophyte stands of its middle floodplain (Ezcurra de Drago 1993, 2003), in Amazonian river rocky bottoms and in their marginal seasonally flooded forests (Batista et al. 2003). The application of this taxonomic tool is also proving to be rewarding onto environmental and climatic paleointerpretations, following the identification of sponge species based on the spicules detected in columns of recovered lake sediments of quaternary age (Siffeddine et al. 1994, Volkmer-Ribeiro and Turcq 1996, Turcq et al. 1998, Cndido et al. 2000, Volkmer-Ribeiro et al. 2007, Parolin et al. 2007). While the surveys are being extended across the continent some restricted local endemisms remain unchallenged. As a result the first official State and National recognitions and red listings of sponge species under threat were attained, upon the following of IUCN standard procedures. Oncosclera jewelli, Anheteromeyenia ornata and Drulia browni integrate the red list of endangered fauna of Rio Grande do Sul State, Brazil (Volkmer-Ribeiro 2003). Oncosclera jewelli, A. ornata, Uruguaya corallioides, Sterrastrolepis brasiliensis, Corvoheteromeyenia australis, Corvoheteromeyenia heterosclera, Corvospongilla volkmeri, Heteromeyenia insignis, Houssayella iguazuensis, Racekiela sheilae and Metania kiliani are listed with the Brazilian endangered freshwater invertebrates and fishes (Brasil 2004). This fact offers support to national and regional policies aiming at the

preservation of particular freshwater habitats and the species they contain. Lately, surveys for the South American continental sponges have also focused on river, lake, lagoonal and pond waters contained in preserved areas such as State and National Parks and Ecological Stations with the aim of establishing parameters for biomonitoring and bioindication, at the same time improving the knowledge of the species they contain and the use of such preserved areas as biodiversity banks (Volkmer-Ribeiro et al. 1988, 1999, 2005, Tavares et al. 2005, Volkmer-Ribeiro and Almeida 2005). A further application of this taxonomic tool is within the context of archeological studies. Spicules (cauxi), present in archeological Amazonian pottery are revealing unsuspected native technologies and histories of the sustainable management of natural resources (in this case biosilica produced by the sponges), besides allowing the tracing of cultural trends and past native population migrations within the continent (Volkmer-Ribeiro and Gomes 2006, VolkmerRibeiro and Viana 2006).

What next?
Homo sapiens may be producing a more extensive modification of the Earths surface than any other animal species did before. One such profound environmental change is the damming of large rivers in order to produce hydroelectric power, particularly throughout the Tropical and Sub-tropical realms. South America is a continent where the damming of large rivers has boomed over the last thirty years. Huge lakes have been formed in areas where this permanent freshwater habitat was previously absent, such as in the Amazonian Region, famous for its seasonal vrzea lakes. In regard to the rich Amazonian sponge fauna, surveys have extended to this new habitat in order to detect the invasion by sponges and the exclusion/adaptation forces in action. Results have shown that colonization is being carried by some species previously detected in the riverine rocky bottoms when the prior Impact Assessment surveys were done. All harder substrates located in the lake waters (excluding the anoxic ones), including the trunks of the forest flooded by the lake, are being used by those sponges that had occupied more extensively the original river bottoms (Volkmer-Ribeiro and Hatanaka 1991). The monitoring of the occupation of these dammed waters by sponges is being continued bearing in mind to offer taxonomic substrate for further research purposes encompassing from basic sponge biology and ecology to the production of biosilica or biocompounds by sponges. The mapping of substrates occupied by sponges in these dammed waters, allied to their continued recruitment as a consequence of the permanent ingression of upstream gemmules, renders these living stocks ideal for continued observation/monitoring and experimentation. These natural systems are better than laboratory aquaria, where freshwater sponge species other than those belonging to Ephydatia are barely kept alive for a few days. Another area of research being pursued based on the taxonomic knowledge currently available is the area of medicine, as there are several historical and some current records of dermal diseases caused by contact with sponge

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spicules, particularly in the Amazonian Region. The discovery of freshwater sponge spicules acting as agents of ocular pathology in the Araguaia River area (Brazilian Amazonia) has only recently been reported (Volkmer-Ribeiro et al. 2006, Volkmer-Ribeiro and Batista 2007). Other pathologies related to freshwater sponge spicules inferred from archeological work have also been compiled and discussed in the aforementioned publications. The spreading of the geographic surveys of the South American continental sponges is obviously an ongoing process. Hopefully at the same speed as global economic enterprises are reaching them and their habitats. Renewed efforts should, from now on, focus on the aquatic habitats contained in preserved areas which, as a rule, benefit of previous selections aiming the protection of continental biomes. The invertebrate faunas of such biomes are yet poorly known and their study will certainly come up with the detection of new species.

Acknowledgments
The author is indebted to the organizers of the 7th International Sponge Symposium (Armao dos Bzios, RJ) for the invitation to present this opening speech as well as to two anonymous referees for the suggestions presented. She heartily thanks Dr. Eduardo Hajdu for a minutious reading of the MS and valuable improvements indicated. She acknowledges the continued support CNPq. has provided to the research projects proposed along the last three decades.

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