Porifera Research: Biodiversity, Innovation and Sustainability - 2007
117
South American continental sponges: state of the
art of the research
Cecília Volkmer-Ribeiro
Museu de Ciências Naturais, Fundação Zoobotânica do Rio Grande do Sul. Rua Dr. Salvador França, 1427. 90690-000,
Porto Alegre, RS, Brazil. Research fellow of CNPq. [email protected]
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 oneself’s rationale. Plain common
sense, but a painful process… And here it is when Science
turns into Art and Philosophy. Into art because a scientist´s
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
118
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 Fé, Argentina and the Museu de Ciências Naturais,
Fundação Zoobotânica 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 world’s freshwater sponges long ago split
by Carter´s 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 Racek’s (1968) comprehensive revision of the world’s
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.
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
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 Schröder, 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 Gray´s (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 gemmosclere’s and microsclere’s 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
119
(Berroa Belén 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
Paraná 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,
Cândido 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 Earth´s 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 “várzea” 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
120
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 (Armação dos Búzios, 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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