J. Limnol.

, 2013; 72(s2): 313-326 BIODIVERSITY AND BIOGEOGRAPHY

DOI: 10.4081/jlimnol.2013.s2.e15 PORIFERA

Biodiversity in South East Asia: an overview of freshwater sponges (Porifera:

Demospongiae: Spongillina)
Renata MANCONI,1* Nisit RUENGSAWANG,2 Viengkorn VANNACHAK,3 Chutima HANJAVANIT,4,5
Narumon SANGPRADUB,4,5 Roberto PRONZATO6
1
Dipartimento di Scienze della Natura e del Territorio, Università di Sassari, Via Muroni 25, 07010 Sassari, Italy; 2Division of Biology,
Faculty of Science and Technology, Rajamangala University of Technology Krungthep, 2 Nanglinchee road, Bangkok, 10120 Thailand;
3
Department of Biology, Faculty of Science, National University of Laos, Vientiane, 7322 Laos; 4Applied Taxonomic Research Center,
Khon Kaen University, Khon Kaen, 40002 Thailand; 5Department of Biology, Khon Kaen University, Khon Kaen, 40002 Thailand;
6
Dipartimento di Scienze della Terra, dell’Ambiente e della Vita, Università di Genova, Corso Europa 26, 16132 Genoa, Italy
*Corresponding author: r.manconi@uniss.it

ABSTRACT

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Despite the fact that South East (SE) Asia is considered as a biodiversity hotspot, knowledge of sessile invertebrates such as

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freshwater sponges (Porifera: Haplosclerida: Spongillina) in this region is poor and scarcely reported. For this synopsis, diversity
and distribution of SE Asian inland water sponges is reported on the basis of available literature and a recent biodiversity assessment
of the Lower Mekong basin. A diagnostic key of families/genera from SE Asia is provided together with Light Microscopy and Scan-
ning Electron Microscopy protocols to prepare the basic spicular complement for taxonomic identification. So far, SE Asian freshwater

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sponges consist of widespread and/or endemic species belonging to the families Metaniidae, Potamolepidae, and Spongillidae. The
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highest diversity is recorded from Indonesia, Philippines, Thailand, and Myanmar, respectively. Data from the other countries are
necessary for our understanding of their diversity and distribution. Biodiversity in SE Asia is strongly underestimated, as indicated
by recent new records and the discovery of new species of freshwater sponges in Thailand. Further investigations should reveal
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higher values of taxonomic richness, highlighting biogeographic patterns at the family/genus/species levels. A cooperative network
involving Thai, Laotian and Italian researchers, was set up to contribute and fulfil knowledge on taxonomy, ecology and biotechno-
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logical potentialities of these neglected filter feeders, playing a key role in water purification and biomass production in both lentic
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and lotic ecosystems in the tropics.

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Key words: Taxonomic richness, Oriental region, biogeographic patterns, conservation.

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Received: April 2013. Accepted: June 2013.

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INTRODUCTION better the concept of modular organisms because of their

ability to alter body architecture in time and space (Pron-
The earliest naturalists grouped freshwater sponges, and
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zato and Manconi, 1994a, 1994b). The functional units

in general Porifera, with plants because of their sessile na-
ture, greenish-brownish colour, and growth form Spongilla (ramets) of these modular animals are clearly evident dur-
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lacustris (Linnaeus, 1759) was described (as Spongia la- ing freshwater sponge cryptobiosis (aestivation or hiber-
custris) by Linnaeus (1759) as repens, fragilis, ramis nation by resting bodies) and modules (gemmules) can be
teretibus obtusis (creeping, fragile, with cylindrical isolated from one another, unlike those of adult freshwater
branches showing swellings at their ends) in the second sponges (Pronzato and Manconi, 1994a, 1994b, 1995;
volume of Plantae of the Systema Naturae, and the type Manconi and Pronzato, 2013).
material was preserved in the Linnean Herbarium (Man- Sponges are suitable microhabitats and are associated
coni and Pronzato, 2000). Sponges were not recognised as with or host bacteria, unicellular algae, protists, hydro-
animals until 1765, when the internal water current was first zoans, turbellarians, nematodes, oligochaetes, leeches, bi-
described, and only in 1875, Huxley proposed the complete valves, gastropods, decapods, amphipods, copepods,
separation of sponges from other Metazoa (Gaino, 2011). ostracods, hydracarins, and bryozoans, as well as several
At present, sponges are considered as basal Metazoa, orders/families of insects encompassing the typical
lacking true organs and a nervous system. Their body ar- spongillaflies together with fishes and amphibians.
chitecture, usually displaying an irregular symmetry, is Sponges have been reported to be fed upon by some fresh-
very plastic and characterised by continuous morphogen- water turtles, ducks, fish and insects where spicules were
esis. The bauplan of sponges does not match completely found in stomach contents (Pronzato and Manconi, 2001;
the concepts of the colony or the individual. It matches Manconi and Pronzato, 2008).
314 R. Manconi et al.

Porifera are primary marine invertebrates which have sponges in this area is poor and scarcely reported. In the
colonised continental water since the Palaeozoic as indi- present study, we compile a report on the diversity and
cated by the Permo-Carboniferous spiculites of freshwater distribution of the SE Asian freshwater sponges, based on
sponges (Haplosclerida: Spongillina) (Schindler et al., available literature and a recent biodiversity assessment
2008). Although records are discontinuous, Mesozoic re- of the Lower Mekong basin in Thailand.
mains date back to the Upper Jurassic and Lower Creta-
ceous, but most fossils refer to the Eocene, Miocene, METHODS
Pliocene, and the Pleistocene (Pisera and Saez, 2003; Pis- Bibliographic investigations were performed in depth
era, 2004; Schindler et al., 2008). Fossil evidence, to- to ascertain previous records in the eastern area of SE Asia
gether with the ability of some recent taxa of Spongillina vs global taxonomic assessments.
to live in estuaries and coastal water bodies with variable New collections of sponges were carried out during
salinity (Annandale, 1915; Evans and Kitting, 2010), sug- visual census by wading, snorkelling and scuba diving in
gests the occurrence of the invasion of inland waters by the Pong river (Mekong Hydrographic basin, Khon Kaen
sponges from coastal brackish waters of epicontinental province, NE Thailand). Sampling was easier during low
and/or enclosed seas. water seasons when dried sponges and/or gemmular car-
Systematics and phylogeny of freshwater sponges pets can be found on substrata although they can be easily
(Porifera: Haplosclerida: Spongillina) are based on mor-

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overlooked in this life cycle phase. Species-level identi-
phology, anatomy, and cytology together with reproduc- fication required collection of gemmules from the basal

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tive, developmental, and molecular biology. A portion of the sponge body or sometimes from the surface
morphological dataset of macro- and micro-morphotraits of hard substrata where they adhere. Gemmules, however,
is diagnostic up to the genus and species level (Manconi may be absent in some phases of the sponge life cycle.

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and Pronzato, 2002). The most representative specimens were pho-
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The monophyletic status of Spongillina, based on clas- tographed in vivo and/or in situ, preserved by air drying,
sic taxonomic techniques, has been shown to be robust ethanol/methanol, or freezing and registered in a voucher
from recent syntheses that focused on comparative analy-
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collection each with a label reporting basic collection data
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ses of morphotraits (Manconi and Pronzato, 2002, 2009). (locality, date, collector name). It is not necessary to col-
This is supported in part by molecular approaches
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lect always entire specimens: representative body frag-

(Meixner et al., 2007; Erpenbeck et al., 2011). The reso-
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ments bearing gemmules is enough. Entire sponges

lution power of molecular analyses in Porifera is not, (possibly with a part of their substratum), however, are
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however, fully diagnostic and needs to be improved and necessary for museum collections. A set of key macro-
compared with morphological analyses. and micro-morphological diagnostic characters consid-
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Plesiomorphic morphotraits, e.g. skeletal architecture, ered for identification at the genus and species levels is
monaxial spicules, gemmules and gametes together with focused on growth form, consistency, colour, surface
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larval (parenchymella) structure suggest that Spongillina traits, topographic distribution of inhalant and exhalant
are allied to coastal marine Haplosclerina (order Hap- apertures, architecture of ectosomal and choanosomal
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losclerida). Apomorphies of the Spongillina are at the skeleton, topographic distribution and traits of skeletal
level of both sexual and asexual propagules such as swim- megascleres, microscleres, and gemmules, gemmular ar-
ming parenchymellas type III provided by choanocyte chitecture (foramen, gemmular cage, gemmular theca,
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chambers, canals, and spicules as well as gemmules dis- spatial arrangement of pneumatic layer and spicules), and
playing a highly diverse architecture. gemmuloscleres morphology (Manconi and Pronzato,
Present knowledge on the biodiversity and biogeog- 2002; but see the present diagnostic key of genera from
raphy of freshwater sponges from Southeast (SE) Asia are SE Asia).
based mostly on few records and scattered papers. Indeed, To characterise morphotraits, representative fragments
exhaustive monographies are scarce and at present only a of sponges were dissected for light microscopy (LM)
few biologists focus on this region (Bowerbank, 1863; and/or scanning electron microscopy (SEM) observation.
Carter, 1881; Potts, 1887; Weber, 1890; Weltner, 1895, A rapid protocol to prepare preliminarily slides of spicular
1913; Annandale, 1909, 1911, 1918; Arndt, 1926; complement is based on the digestion of organic matter
Vorstman, 1927, 1928; Gee, 1930, 1931, 1932a, 1932b; of skeleton and gemmules in a drop of boiling 65% nitric
Schröder, 1935; Penney, 1960; Penney and Racek, 1968; acid on a slide. To obtain perfectly cleaned spicule prepa-
Racek, 1969; Soota, 1991; Volkmer-Ribeiro and Costa, rations for SEM observation and LM slides for a voucher
1993; Banarescu, 1995; Manconi and Pronzato, 2002, collection, sponge fragments were dissolved in boiling
2007, 2008, 2009; Masuda, 2004; Ohtaka et al., 2011). 65% nitric acid or in sodium hypochlorite (ambient tem-
Although SE Asia is considered as a biodiversity perature) in test tubes and then suspended in water. To
hotspot (Myers et al., 2000), knowledge of freshwater achieve total sedimentation of the lighter, smaller
 Southeast Asian freshwater sponges 315

spicules, they were suspended and repeatedly rinsed with RESULTS

distilled water and then dehydrated in graded ethanol se-
Southeast Asia is not a biogeographic region (or sub-
ries, with a gap of 15-20 min between successive washing
region) but a geographic entity of convenience which ex-
(Manconi and Pronzato, 2000). Spicules suspended in al-
cludes India, Southwestern China and Wallacea (Fig. 1).
cohol have been poured from tubes with a slow movement
The freshwater sponge species recorded from SE Asia
to leave the deposit of spicules in 0.5 mL of the medium
add up to a total of 26, belonging to 12 genera with a total
within the tube before dropping some suspended spicules
of 40 records (Fig. 1, Tabs. 1 and 2). The most represented
on slides for LM and SEM analysis (Manconi and Pron-
genus is Eunapius Gray, 1867 with five species (Fig. 1,
zato, 2000). After the total evaporation of alcohol, each
Tab. 2). The most reported species is Eunapius carteri
slide fragment was fixed on a stub with silver glue drops.
The presence of a glass substratum under the spicules (Bowerbank, 1863) with five records (Figs. 1 and 2; Tab.
gives the best results as a perfectly black background in 2). A well represented genus is Radiospongilla Penney
the SEM photographs. Dry body fragments, dissociated and Racek, 1968 with four species, three of which have
spicules, entire gemmules, and their cross-sections were been recorded two times (Fig. 1, Tab. 2). Other, relatively
sputter-coated with gold and observed under SEM (Man- common species, belong to the widespread genus Tro-
coni and Pronzato, 2000). Morphometric data of diagnos- chospongilla Vejdowsky, 1883 with Trochospongilla la-
tic traits were performed by LM and/or SEM. To touchiana Annandale, 1907 (four records), and the

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characterise potential new species, approximately 50 Asiatic-Australian genus Umborotula Penney and Racek,
1968 with the Umborotula bogorensis (Weber, 1890)

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spicules for each diagnostic spicular type were measured
for representative specimens of recorded species. Com- (three records) (Fig. 1, Tab. 2). Indeed, a high number of
parative analyses were performed on new voucher collec- species (18 out of 26) is reported only once from this area.

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tions vs materials from several historical collections and Moreover, in some cases the findings are very old and not
original descriptions.
l usconfirmed by recent field campaigns (Fig. 1; Tab. 2).
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Fig. 1. Distribution of Spongillina species in

freshwater of the Southeast Asia. Numbers
refer to records in the map. 1, Corvospongilla
burmanica; 2, Corvospongilla siamensis; 3,
Dosilia plumosa; 4, Ephydatia fluviatilis; 5,
Ephydatia fortis; 6, Eunapius carteri; 7, Eu-
napius crassissimus; 8, Eunapius fragilis; 9,
Eunapius potamolepis; 10, Eunapius tinei; 11,
Metania pottsi; 12, Metania vesparia; 13,
Metania vesparioides; 14, Oncosclera asiat-
ica; 15, Radiospongilla cerebellata; 16, Ra-
diospongilla crateriformis; 17, Radiospongilla
indica; 18, Radiospongilla philippinensis; 19,
Spongilla alba; 20, Spongilla sp.; 21,
Stratospongilla clementis; 22, Stratospongilla
indica; 23, Stratospongilla sumatrana; 24, Tro-
chospongilla latouchiana; 25, Trochospongilla
phillottiana; 26, Umborotula bogorensis.
316 R. Manconi et al.

Tab. 1. Check-list (39 species, 12 genera, 3 families) of Spongillina (Porifera: Haplosclerida) of the Oriental region.
SPONGILLIDAE Gray, 1867
Corvospongilla Annandale, 1911
C. bhavnagarensis Soota, Pattanayak and Safena, 1984 (Gujarat-India) OL
C. burmanica (Kirkpatrick, 1908) (Burma, India) OL
C. caunteri Annandale, 1911 (India) OL
C. lapidosa (Annandale, 1908) (India) OL
C. siamensis Manconi and Ruengsawang , 2012 (Thailand) OL
C. ultima (Annandale, 1910) (S-India) OL
Dosilia Gray, 1867
D. plumosa (Carter, 1849) (India, Kashmir-Pakistan, Philippines) type species OL
Ephydatia Lamouroux, 1816
E. fluviatilis (Linnaeus, 1759) (cosmopolitan) type species OL-PA-NA-AT-AU
E. meyeni (Carter, 1849) (India, China) OL-PA
E. fortis Weltner, 1895 (Indonesia, Philippines, Japan, Vanuatu) OL-PA-PAC
Eunapius Gray, 1867
E. calcuttanus (Annandale, 1911) (India) OL
E. carteri (Bowerbank, 1863) (S-Asia, India, Thailand, East Europe, Africa, Panama) type species OL-PA-AT-NT

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E. crassissimus (Annandale, 1907) (India, Australia, tropical SE Asia) OL-AU
E. fragilis (Leidy, 1851) (Cosmopolitan) OL-PA-NA-AT-NT-AU

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E. geminus (Annandale, 1911) (India) OL
E. potamolepis (Annandale, 1918) (Thailand) OL
E. tinei (Gee, 1932) (Philippines) OL

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Pectispongilla Annandale, 1909
P. aurea Annandale, 1909 (India) type species OL
P. stellifera Annandale, 1915 (S-India) OL
us
P. subspinosa Annandale, 1911 (India, Japan, Korea) OL-PA
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Radiospongilla Penney and Racek, 1968
R. cerebellata (Bowerbank, 1863) (tropical Africa, India-Pakistan, Indonesia, Philippines, Japan, New Guinea, China, Russia?, SE-Europe, Texas)
c

OL-AU-AT-PA-NA
R. cinerea (Carter, 1849) (Bombay, Himalayas, Pakistan) OL
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R. crateriformis (Potts, 1882) (USA, Canada, Mexico, West Indies, Suriname, Brazil, China, Japan, India, S-Asia, Australia) OL-NA-NT-PA-AU
R. hemephydatia (Annandale, 1909) (India, New Guinea, E-Australia) OL-AU
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R. indica (Annandale, 1907) (India, Indonesia, Philippines, New Guinea?) OL-AU

R. philippinensis (Annandale, 1909) (Philippines to N-Australia) OL-AU
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Spongilla Lamarck, 1816

S. alba Carter, 1849 (India, Thailand, SE-Asia, Turkey, Africa, Madagascar, Australia, Papua-New Guinea, USA, S.Salvador, S America) OL-AT-
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AU-NT-PA-NA
Spongilla sp.
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Stratospongilla Annandale, 1909

S. bombayensis (Carter, 1882) (Mumbay, Natal) type species OL-AT
S. clementis (Annandale, 1909) (Philippines, China, Japan, tropical W-Africa) OL-PA-AT
N

S. gravelyi (Annandale, 1912) (Mumbay, India) OL

S. indica (Annandale, 1908) (Thailand, India, Africa) OL-AT
S. sumatrana (Weber, 1890) (Indonesia, India, Africa) OL-AT
Trochospongilla Vejdowsky, 1883
T. latouchiana Annandale, 1907 (India, Burma, China, SE-Australia, SW-Africa) OL-PA-AU-AT
T. phillottiana Annandale, 1907 (India, S-China, Philippines, Africa) OL-AT
Umborotula Penney and Racek, 1968
U. bogorensis (Weber, 1890) (N-China, Korea, Thailand, SE-Asia, Andaman islands, E-Australia) OL-PA-AU
METANIIDAE Volkmer-Ribeiro, 1986
Metania Gray, 1867
M. pottsi (Weltner, 1895) (Congo basin, Angola, Borneo, Indonesia) OL-AT
M. vesparia (von Martens, 1868) (Borneo, Indonesia, Australia) OL-AU
M. vesparioides (Annandale, 1908) (Tenasserim, Burma, Australia) OL-AU
POTAMOLEPIDAE Brien, 1967
Oncosclera Volkmer-Ribeiro, 1970
O. asiatica Manconi and Ruengsawang, 2012 (Thailand) OL
OL, Oriental region [modified and updated from Manconi and Pronzato (2007)]; PA, Palaearctic region; NA, Nearctic region; AT, Afrotropical region;
AU, Australian region; PAC, Pacific islands; NT, Neotropical region; S, South; SE, Southeast; E, East; N, North; W, West; SW, Southwest.
 Southeast Asian freshwater sponges 317

Only four species are considered endemic to SE Asia, mm in diameter; Fig. 2d) adhering to hard substrata, float-
namely three from Thailand [Corvospongilla siamensis ing at the water surface, or within the old skeleton of the
Manconi and Ruengsawang, 2012, Eunapius potamolepis sponges. Morphological characters of gemmules are
(Annandale, 1918), and Oncosclera asiatica Manconi and species-specific, diagnostic at genus/species level and
Ruengsawang, 2012] and one [Eunapius tinei (Gee, therefore fundamental for taxonomic identification.
1932)] from the Philippines (Figs. 3 and 4). As for sexual reproduction, viviparity is the rule, and
Among the 26 recorded species only 4 were recorded larvae are brooded as suboval, soft, whitish bodies pro-
from both continental and insular freshwater of SE Asia, tected within the mother sponge body up to the comple-
indeed 13 species are present exclusively in continental tion of the larval development (Manconi and Pronzato,
freshwater, whereas 9 were recorded exclusively from is- 2013). During late sexual reproductive phases of the life
lands (Fig. 1, Tab. 2). cycle, larvae were observed in recently studied species
At the level of biogeographic regions, the freshwater from SE Asia only after sampling, swimming just during
sponge species known at present for the Oriental region releasing from the oscules in the excurrent water flux, or
are 39 (Tab. 1) with a notably high number of endemics brooded larvae as suboval, soft, whitish bodies protected
(42.5%) (Manconi and Pronzato, 2013). The latter values within sponge bodies up to the complete development.
slightly differ from what reported by Manconi and Pron- The natural resource represented by freshwater
zato (2007, 2008) as consequence of taxonomic revisions, sponges have been used since ancient times (Neolitic) by

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new records and/or discovery of new species. As for bio- humans, and at present some African and Amazonian pop-

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geographic patterns at species level, they range from en- ulations produce ceramics tempered by siliceous sponge
demic sensu stricto (one water body) or endemic sensu spicules (Manconi and Pronzato, 2013). Other practical
lato (large hydrographic basins, i.e. Mekong), to wide- uses are in the field of cosmetics and medicines. For ex-

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spread (two-three geographic regions), up to apparently usample, dried spongillids were used in the 19th century by
cosmopolitan. Russian young ladies to scrub their faces to have rosy
In contrast, the value of endemicity is higher in the In- cheeks. Some cosmetics today exploit the scrubbing ac-
dian Subregion (ca. 45%) harbouring 10 endemic species tion of siliceous spicule powder, and recently this appli-
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out of 23 recorded species (C. bhavnagarensis, C. caun- cation was patented in the United States in the field of
teri, C. lapidosa, C. ultima, E. calcuttanus, E. geminus, dermatology (Manconi and Pronzato, 2008, 2013). In the
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P. aurea, P. stellifera, R. cinerea, S. gravelyi) (Annandale, 17th century, Samuel Hahnemann enclosed freshwater
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1911; Soota, 1991; Manconi and Pronzato, 2007, 2013). sponges (as Spongilla fluviatilis and Spongia palustris) in
his Materia Medica as a homeopathic remedy for psoria-
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The Spongillina species of SE Asia share gross mor-

phological characters, typical of all freshwater sponges. sis with the common pre-linnean name of Badiaga (Man-
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Three morphological characters distinguish the habitus of coni and Pronzato, 2013). Spongilla lacustris was also
freshwater sponges during their life cycle, namely the ses- used in China for more than 500 years as a traditional
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sile growing adult (sponge-like), the planktonic short-life medicine for reinforcing the kidney and supporting yang
swimming larvae, and the dormant gemmules (restant (aphrodisiac), although the chemical research on this
species is poor (Hu et al., 2009; Manconi and Pronzato,
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body/propagule) (Manconi and Pronzato, 2013). The

growth form of adults (active sponges) ranges from thin 2013). Unfortunately, the exact identification
crusts in irregular patches strictly adhering to the substra- (genus/species) of the sponges used as cosmetics and
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tum to massive cushions to bush-like (Figs. 3 and 5); more medicines in both the past and present is questionable be-
rarely they may also display variably erected finger-like cause of inadequate taxonomic expertise.
to branching and arborescent growth forms. Dimensions As for indicators of palaeoenvironmental changes,
are variable from a few millimetres to more than one freshwater sponges spicule remains (spiculites bed) play
metre in diameter. Body consistency is from soft, to frag- a key role in sediment analysis in lake beds and soils. The
ile to very hard (stony) according with the amount of time interval needed for the formation of a spiculite of a
spongin in the skeleton. A wide range of colours are dis- few centimetres in thickness is estimated to have ranged
played, from white to greenish or brilliant emerald green, from years to decades (Schindler et al., 2008). Variations
or greyish to dark brown and black. The colours result in species composition and amount of siliceous microfos-
from sponge pigments, or symbiotic bacteria and algae, sils (sponges spicules and diatoms-diatomites) are indi-
or englobed detritus. All these characters are largely not cators of water quality changes and past salinity
species-specific and are strictly related to environmental conditions (Manconi and Pronzato, 2008, 2013).
conditions, seasonal rhythms and/or life cycle phases. Freshwater sponge spicules are at present considered a
The habitus of SE Asia freshwater sponges during sea- potential agent of severe ocular disease among children
sonal dormancy is very different from that of the active swimming or diving underwater in water bodies rich of
sponges, i.e. as gemmules (small spherules up to over 1 sponge assemblages. Risk factors include dislodging of
318 R. Manconi et al.

Tab. 2. Taxonomic richness (genera, species) and endemic taxa of Spongillina from continental and insular areas of Southeast Asia, on
the basis of forty recent and old records.
Genera Species Records Species validation SE-Asia Continental Insular Remarks
endemics
Corvospongilla C. burmanica 1 Penney and Racek (1968); Manconi and 1 Scanty data; needs
2 species Pronzato (2004, 2007); Manconi et al. (2012) revision
C. siamensis 1 Ruensawang et al. (2012) Thailand 1 Recently described
Dosilia D. plumosa 1 Penney and Racek (1968); Manconi and 1 Recent detailed
1 species Pronzato (2007) description in
Jakhalekar and
Ghate (2013)
Ephydatia E. fluviatilis 1 Manconi and Pronzato (2007); Manconi (2008) 1 Common worldwide;
2 species needs revision;
potential species
complex
E. fortis 2 Penney and Racek (1968); Manconi and 2 High variable
Pronzato (2007) morphology; needs
revision

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Eunapius

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5 species E. carteri 5 Penney and Racek (1968); Manconi and 4 1 Widespread; potential
Pronzato (2002, 2007, 2009) species complex;
recent detailed

e
description by
Manconi and
us Pronzato (2009);
Jakhalekar and Ghate
(2013)
l
E. crassissimus 1 Penney and Racek (1968); Manconi and 1 Easily identifiable
ia

Pronzato (2007)
c

E. fragilis 1 Penney and Racek (1968); Manconi and 1 Widespread; potential

Pronzato (2007, 2009) species complex
er

E. potamolepis 1 Penney and Racek (1968); Manconi and Thailand 1 Uncommon; needs
Pronzato (2007) revision
m

E. tinei 1 Penney and Racek (1968); Manconi and Philippines 1 Needs revision
m

Pronzato (2007)
Metania M. pottsi 1 Manconi and Pronzato (2007, 2009) 1 Recent detailed
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3 species description in
Manconi and
Pronzato (2009)
on

M. vesparia 1 Penney and Racek (1968); Manconi and 1 High variable

Pronzato (2007) morphology
M. vesparioides 1 Penney and Racek (1968); Manconi and 1 Very close to
N

Pronzato (2007) M. vesparia

Oncosclera O. asiatica 1 Manconi et al. (2012) Thailand 1 Recently described
1 species
Radiospongilla R. cerebellata 2 Penney and Racek (1968); Manconi and 2 Recent detailed
4 species Pronzato (2007) description in
Jakhalekar and Ghate
(2013); potential
species complex
R. crateriformis 2 Penney and Racek (1968); Manconi and 1 1 Widespread; high
Pronzato (2005, 2007) variable
gemmulosclere
morphology;
potential species
complex
R. indica 2 Penney and Racek (1968); Manconi and 2 Typical megascleres
Pronzato (2007)
R. philippinensis 1 Manconi and Pronzato (2007) 1 Needs revision
To be continued on next page.
 Southeast Asian freshwater sponges 319

Tab. 2. Continued from previous page.

Genera Species Records Species validation SE-Asia Continental Insular Remarks
endemics
Spongilla S. alba 1 Penney and Racek (1968); Manconi and 1 Widespread;
1 species Pronzato (2007) potential species
complex; needs
revision
Spongilla sp. 1 1
Stratospongilla S. clementis 1 Penney and Racek (1968); Manconi and 1 Needs revision
Pronzato (2002, 2007)
3 species S. indica 1 Penney and Racek (1968); Manconi and 1 Typical
Pronzato (2007) megascleres
S. sumatrana 1 Penney and Racek (1968); Manconi and
Pronzato (2007) 1 Several ecomorphs
Trochospongilla T. latouchiana 4 Penney and Racek (1968); Manconi and 2 2 Recent detailed
2 species Pronzato (2007) description in
Jakhalekar and Ghate
(2013)

ly
T. phillottiana 2 Penney and Racek (1968); Manconi and 2 Needs revision

on
Pronzato (2007)
Umborotula U. bogorensis 3 Penney and Racek (1968); Manconi and 2 1 Needs revision
1 species Pronzato (2002, 2007)

e
SE, Southeast.
l us
ia

sponges determining the suspension of spicules in the water ......................................................................Metaniidae

column as reported from South America (Araguaia river) (4) Lubomirskiidae, Metschnikowiidae, Malaw-
c

(Volkmer-Ribeiro et al., 2006, 2008; Mendes et al., 2009). ispongiidae, Spongillina incertae sedis (not reported at
er

present from SE Asia).

m

Key to families and genera of Spongillina from South-

east Asia Family Spongillidae Gray, 1867
m

(1) Gemmules (resting stages) present ........................2 (1) Birotules and pseudobirotules present ..................5
co

- Gemmules absent ..................................................4 - Birotules and pseudobirotules absent ....................2

(2) Gemmular theca with pneuma absent to scantly de- (2) Presence of microscleres and/or gemmuloscleres
veloped. Gemmuloscleres, when present, as tangentially (not pseudobirotules and/or birotules) ..........................4
on

arranged, short, irregular strongyles; gemmules when (4) Microscleres absent; megascleres are spiny oxeas;
present in the basal portion. Skeletal network typically gemmuloscleres are spiny strongyles radially oriented ..
N

multispicular alveolate-reticulate, scanty spongin; megas- ................................................................Radiospongilla

cleres strongyles to oxeas; microscleres rarely recorded - Microscleres absent; megascleres are spiny and
................................................................Potamolepidae smooth oxeas to strongyles; gemmuloscleres are spiny
- Gemmular theca bi- to tri-layered (pneumatic layer oxeas of two morphologies: strongyles stout, short and
generally present) ..........................................................3 spined, tangentially embedded in the outer layer of gem-
(3) Gemmuloscleres as oxeas, strongyles, birotules, mular theca; spiny oxeas abruptly pointed, radially
pseudobirotules, botryoidal, club-like (radially to tangen- arranged in between the inner and outer layer ................
tially arranged); skeletal network paucispicular, irregularly ................................................................Stratospongilla
reticulate; megascleres oxeas to strongyles, smooth/vari- - Microscleres absent; megascleres smooth oxeas;
ably ornamented; microscleres usually present.................. gemmuloscleres oxeas to strongyles, usually smooth, tan-
........................................................................Spongillidae gentially arranged in the gemmular theca ........Eunapius
- Gemmuloscleres tubelliform, parmuliform, pseudo- - Microscleres spiny oxeas; megascleres smooth
birotules (almost exclusively radially arranged); skeletal oxeas; gemmuloscleres, when present, spiny oxeas to
network typically multispicular alveolate-reticulate with strongyles often very curved ............................Spongilla
scanty spongin; smooth to variably ornamented megas- (5) Birotules with rotules showing smooth edges,
cleres (oxeas to strongyles); microscleres usually present pseudobirotules absent ..................................................6
320 R. Manconi et al.

ly
on
e
l us
c ia
er
m
m
co
on
N

Fig. 2. Eunapius carteri from the Lower Mekong hydrographic basin (Khon Kaen province, Northeast-Thailand), apparently cosmo-
politan, is at present the most common species in the Southeast Asia. a) Skeletal network with scattered subspherical gemmules; b, c)
oxeas (megascleres) of the skeletal network; d) gemmules with surface armed by tangential gemmuloscleres; e) close up on the gemmule
surface with ornamentations of polygonal compact spongin and some gemmuloscleres (microxeas) on the outer gemmular layer); f)
cross section of a gemmule showing the multilayered theca with the foramen, the thick pneumatic layer of chambered spongin, and the
inner layer of compact spongin protecting a dense mass of staminal cells in the internal chamber.
 Southeast Asian freshwater sponges 321

- Birotules with spiny rotules and indented edges gen- ......................................................................Oncosclera

erally associated with pseudobirotules ..........................7 (2) Sterrastrolepis, Echinospongilla, Potamolepis,
(6) Microscleres absent; megascleres are oxeas very Potamophloios, Uruguaya (not reported at present from
spiny; gemmuloscleres are smooth birotules radially SE Asia).
arranged ................................................Trochospongilla
(7) Birotules and/or pseudobirotules are present, exclu- DISCUSSION
sively as microscleres ....................................................8 Global diversity of Spongillina
- Birotules and/or pseudobirotules are present, exclu-
sively as gemmuloscleres ..............................................9 At a global level, the biodiversity assessments, al-
(8) Gemmuloscleres spiny/smooth strongyles tangen- though in a dynamic status and continuously under revi-
tially arranged in the gemmular theca; megascleres sion and updating, focus at present on 7 families, 47
smooth to microgranulate strongyles to oxeas; microscle- genera, and 237 species of freshwater sponges (Manconi
res pseudobirotules usually with smooth shaft ................ and Pronzato, 2013; Van Soest et al., 2013). The Neotrop-
................................................................Corvospongilla ical region shows the highest species richness of Spongil-
(9) Gemmuloscleres birotules of a single morphology lina (72 species) followed by Palaearctic (65 species),
......................................................................................10 Afrotropical (52 species), Oriental (39 species), Aus-
(10) Birotules with a short (usually less than the rotule di- tralian (34 species), Nearctic (31 species), and Pacific (7

ly
ameter) shaft ................................................................11 species) regions; no freshwater sponges are reported from

on
- Birotules with a long spiny shaft (2-3 times the rotule the Antarctica. Taxonomic richness values in the Oriental
diameter) ......................................................................12 region are not among the highest (Manconi and Pronzato,
(11) Gemmuloscleres are birotules with a short shaft and 2013), although this may be related to the small size of

e
rotules of identical diameter; microscleres are absent; the region.
megascleres are microspined and/or smooth oxeas ............
us
Biogeography of Spongillina in Southeast Asia
........................................................................Ephydatia
(12) Gemmuloscleres are birotules with long spiny shaft;
l
A comparison among different areas within the Ori-
ia

microscleres are absent; megascleres are microspined and ental region, shows that the Indian subregion is less rich
rarely smooth oxeas ....................................Umborotula
c

(23 species) than SE Asia (26 species); in the latter, the

- Gemmuloscleres are birotules with a long spiny continental species (13 species) slightly overcome the in-
er

shaft; microscleres range from acanthoxeas to typical eu- sular ones (9 species). Conversely, the Indian subregion
m

asters with spiny tips; megascleres are smooth oxeas .... hosts a larger number of endemic taxa (ten species) when
..............................................................................Dosilia compared to SE Asia (4 species). The present diversity
m

(13) Corvoheteromeyenia, Heterorotula, Pectispongilla, values doubtlessly appear destined to increase with further
Uruguayella, Saturnospongilla, Heteromeyenia, Race- research on unexplored and poorly sampled areas, and
co

kiela, Pachyrotula, Anheteromeyenia, Nudospongilla, from molecular analyses focused mainly on cosmopoli-
Duosclera, Sanidastra (not reported at present from SE tan/widespread species that are presently assumed to con-
on

Asia). sist of cryptic species complexes (Tab. 2).

The major affinities of the Oriental region are with the
Family Metaniidae Volkmer-Ribeiro, 1986 Australian (7 shared species) and Afrotropical regions (5
N

(1) Gemmuloscleres of a single tubelliform morph, ra- shared species). Focusing on SE Asia, 6 species are ex-
dially arranged with larger rotule embedded in the inner clusively Oriental and another 6 are cosmopolitan, al-
layer of the gemmular theca; microscleres strongyles though the latter probably represent species complexes.
The species shared with the other biogeographic regions
smooth to spiny; megascleres oxeas to strongyles, smooth
are 7 with the Afrotropical region, 6 with the Australian
to spiny ..............................................................Metania
region, and 4 with the Palaearctic region.
(2) Drulia, Acalle, Corvomeyenia, Houssayella (not re-
The Oriental-Australian boundary is one of most stud-
ported at present from SE Asia).
ied and discussed regions from a biogeographic point of
view; the presence of Wallacea represents an area of over-
Family Potamolepidae Brien, 1967
lapping and vagueness in still unsolved distribution pat-
(1) Megascleres stout oxeas to strongyles, rarely bearing terns. As for freshwater sponges, by simply considering the
tubercles/granules, with acerate tips; microscleres absent; Wallace (1859) or the Weber lines (1937) in alternative, it
gemmules when present, with gemmuloscleres tangen- is possible to include or exclude for example Sulawesi
tially arranged; gemmuloscleres highly variable from true (Celebes) in the Oriental region; this involves the inclusion
strongyles to irregular ovoid strongyles, from smooth to or exclusion of other two species of freshwater sponges viz.
ornamented with short spines/tubercles dense at tips .... Nudospongilla vasta (Weltner, 1901) from Sulawesi and
322 R. Manconi et al.

ly
on
e
l us
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er
m
m
co
on
N

Fig. 3. Freshwater sponges from the Lower Mekong hydrographic basin (Khon Kaen province, Northeast-Thailand) in their natural
surroundings (a-e). a) Corvospongilla siamensis from shallow water; b) Oncosclera asiatica, type specimens in vivo from type locality
near Ubolratana dam, Pong river; c) Radiospongilla sp. with encrusting growth form; d) C. siamensis with symbiotic algae, as indicated
by brilliant green, growing on a buoy in well illuminated habitat from type locality near Ubolratana dam, Pong river; e) Corvospongilla
sp. growth form associated to bivalves, in running water on rock; f) C. siamensis dry holotype from type locality near Ubolratana dam,
Pong river.
 Southeast Asian freshwater sponges 323

Indonesia, and Pachidictyum globosum Weltner, 1901 from make taxonomic and zoogeographical studies on the
Sulawesi, increasing the endemicity level of the area in the Spongillina extremely problematic. Nevertheless, pauci
case of preference of the Weber line. sed indomita scientists continue to persist in their studies.
The life cycle encompassing phases of cryptobiosis
Biodiversity hotspots further complicates the study of the geographic distribu-
tion of Spongillina, as well as the fragmentary nature and
Although SE Asia is considered a biodiversity hotspot
paucity of space-time data.
for many vertebrate and invertebrate taxa (Myers et al.,
As regards cosmopolitanism of freshwater sponges
2000), knowledge on freshwater sponges in this area
recorded from SE Asia, and in general the Oriental region,
seems poor and scarcely reported. The famous axiom, re-
the presence of 6 (out of 26; Tab. 1) cosmopolitan/wide-
peatedly reported by biogeographers reading The distri- spread species suggests the need of investigations in depth
bution and abundance of species characteristic of a to ascertain if they belong to species complex.
geographical region is directly proportional to the distri- The dataset indicates that species richness of freshwa-
bution and abundance of researchers who work and have ter sponges is commonly underestimated in tropical lati-
worked in the same area seems to be confirmed (Manconi tudes where new findings often correspond to the
and Pronzato, 2007). Indeed, a high number of species (18 discovery of new secies as recently focused in Thailand
out of 26) is reported only once from this area. Moreover, (Manconi et al., 2012; Ruengsawang et al., 2012). Infor-

ly
in same cases findings are very old and not confirmed by mation from other countries of SE Asia is needed for a
recent field campaigns. The evidence that during the last

on
better understanding of diversity and geographic distribu-
investigations (Manconi et al., 2012; Ruengsawang et al., tion. More investigations by field surveys in unexplored
2012) on a small area of the Lower Mekong hydrographic and/or poorly sampled areas should reveal higher values

e
basin new species have been discovered, shows how the of taxonomic richness highlighting biogeographic patterns
freshwater sponge diversity is dramatically underesti- usat family/genus/species levels. Cooperative research net-
mated and probably related to undersampling. The recent work involving Thai, Laotian and Italian teams was set
discovery of O. asiatica from Northeastern Thailand up to contribute and extend knowledge on taxonomy,
l
(Pong river enlarges the disjunct geographic range of the
ia

ecology and biotechnological potentialities of these neg-

genus Oncosclera Volkmer-Ribeiro, 1970 to the Oriental lected filter feeders playing a key role in water purifica-
c

region and represents the single record from the entire tion and biomass production in both lentic and lotic
er

continental Asia (Manconi et al., 2012). On the other ecosystems of the tropics.
hand, the discovery of C. siamensis from Northeastern
m

Thailand extends the easternmost range of the disjunct CONCLUSIONS

range in the Oriental region of the genus Corvospongilla
m

Annandale, 1911. Protection of these sessile invertebrates will support the

conservation of the key resource represented by freshwater
co

Endemicity, cosmopolitanism and species rarity

Eunapius potamolepis and Eunapius tinei are the two
on

historical SE-Asiatic endemics being described from

Lampam (Thailand) by Annandale in 1918 and from
N

Philippines by Gee in 1932, respectively. These two

species were no longer reported and are still known only
for their type localities. Penney and Racek (1968), who
studied the available material (slides) of the two species,
expressed many doubts about their taxonomic status.
Also, the illustrations of the two species, lacking detail
(Fig. 4) from these last authors, did not show convincing
distinctions.
This situation is very common for freshwater sponge
species, e.g. although 58 species have been hitherto Fig. 4. Spicular complement of Eunapius potamolepis (left and
recorded from African inland water (Manconi and Pron- a) and Eunapius tinei (right b). These species are endemic to the
zato, 2009), a large number of them (25) is known only Southeast Asia (Thailand and Philippines, respectively) and were
for one sample from the type locality and is not reported discovered in the first half of the 20th century. Left panel modi-
over periods longer than 50 or 100 years. Little material fied from the original low quality drawing by Annandale (1918);
available, old and incomplete descriptions and illustra- right panel modified from the original drawing by Penney and
tions, absence of any other findings and confirmations, Racek (1968).
324 R. Manconi et al.

ly
on
e
l us
Fig. 5. Massive specimens of Corvospongilla sp. on nets from Thailand (Lower Mekong hydrographic basin).
c ia
er

bodies for the future at a global level. From an ecological Getwongsa, Prayut Udonphimai for their field assistance,
m

viewpoint, sponges perform key functional roles in ecosys- and Dr. Amir Shah Ruddin Md Sah for valuable reference
m

tems. Pumping activity by their aquiferous system helps to from Malaysia. Nisit Ruengsawang and Renata Manconi
circulate the water column, particularly in lentic conditions, were supported by the Office of the Higher Education
co

also favouring particulate and dissolved organic trapping. Commission, Thailand. Funds to Renata Manconi were
Sponges also provide a living refuge for a wide array of or- partly provided by the Italian Ministero dell’Università e
ganisms, and symbiosis with autotrophic microorganisms della Ricerca Scientifica e Tecnologica (MIUR-PRIN) and
on

contributes to primary production. Moreover, their siliceous the Fondazione Banco di Sardegna. We acknowledge the
skeleton contributes to the formation of sediments after the Organizing Committee of FISA Conference, and the Maha
N

sponge’s death to form spiculites beds. Sarakham University and Khon Kaen University for par-
Basic research coupled with a sustainable manage- ticipation in the Proceedings.
ment planning of this bioresource will result in a valori-
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