African freshwater sponges: Makedia tanensis gen. et sp. nov. from Lake Tana, Ethiopia

7 Pages·1999·2.5 MB·English

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AFRICAN FRESHWATER SPONGES; MAKEDIA TANENSJSGEN. ET SP. NOV. FROM
LAKE TANA, ETHIOPIA
RENATA MANCONI, T1ZIANA CUBEDDU & ROBERTO PRONZATO
Manconi, R.,Cuheddu,T.& Pronzato, R. 9990630:Africanfreshwatersponges:Makedia
1
tanensisgen.etsp. nov. from LakeTana, Ethiopia.MemoirsoftheQueenslandMuseum44:
361-367. Brisbane. ISSN 0079-8835.
The new genus Makedia is described and illustrated from shallow waters of Lake Tana,
Ethiopia. Itsmorphologicaldistinguishtraitsarecharacterisedanddiscussedincomparison
with those genera belonging to genera incertae sedis from ancient lakes ofthe world. D
Porifera, Demospongiae, Makedia tanensis, newgenus, new species, taxonomy, scanning
electron microscopy. Ethiopian region, biodiversity, endemism, ancient lakes, geographic
distribution.
Renata Manconi (email: [email protected]) & Tiziana Cuheddu, Dipartimento di
ZoologiaeAntropologiaBiologicadelVUniversita, via Murom 25, 1-07100Sassari. Italy;
Roberto Pronzato, Dipartimento per lo Studio del Territorio e delle sue Risorse
delVUniversita, viaBalbi5, 1-16126 Genova. Italy; 29April 1999,
Africa has a rich freshwater sponge fauna, so Inthispaperwedescribeanew genusMakedia
farcontainingabout60speciesofSpongillidae(8 n.g.,with typespeciesMakediatanemissp. nov.,
NW
genera),Potamolepidae(2genera), Metanidae( from the Pleistocenic Lake Tana in the
genus), and 4 genera with still undefined taxon- marginal areaofthe African Rift Valley. Priorto
omic status (Hilgendorf, 1883ab; Marshall, thepresentstudyonlyArndt(1936)hadreported
1883a,b; Weltner, 1895, 1898, 1913; Evans, on sponges from Abyssinia, recording an
1899;Kirkpatrick, 1906, 1907;Annandale, 1909, unidentified freshwater species.
1914; Jaffe, 1916; Schouteden, 1917; Stephens,
MATERIALS AND METHODS
1919; Burton, 1929ab, 1934, 1938;Seurat, 1930;
Topsent, 1932ab; Amdt, 1933,1936; Schroeder, In a preliminary survey of the freshwater
1934; Tuzet, 1953; Brien & Govaert- sponge fauna ofN EthiopiaandEritrea, in 1988-
Mallebrancke, 1958; Levi, 1965; Brien 1967, 1989 ten water courses and lakes were sampled.
1968abc, 1969abc, 1970ab, 1972, 1973, 1974, Ofthese sites sponges where found only in Lake
1975; Boury-Esnault, 1980; Vacelet et al., 1991; Tana, although sampling was performed under
Gugel, 1993). severe constraints represented by the civil war
and particularly by the endemic schistosomiasis
The most extensive reviews ofthis fauna are in the lake.
the synopsis ofArndt (1936) and the worldwide
revision ofPenney & Racek (1968). The cosmo- Sponges were collected in shallow waters
along the S coast of Lake Tana at Bahir Dar
pwoildietsapnrefaadmiliyn ASfpornigcialnlifdraeeshGwraatye,r 1h8a6b7i,tatiss ((F1i1g°.361')N.,S3p7e°c23i'mEe)n,sNwWereEthpiroepsiearvinedMadryy.19A8l8l
ranging from: wadi in Sahara; large perennial specimens, microscopeslidesandSEM stubs are
rivers such as Nilo, Zambesi and Congo; to presently registered in the senior author's
ancient lakes and man-made basins. Neverthe- collection at the Istituto di Zoologia
less, there are some genera and/or species of delTUniversita, Genova(IZUG),to bedeposited
Potamolepidae Brien, 1967, and Metanidae inthe Museo Civico di StoriaNaturale G. Doria,
Volkmer-Ribeiro, 1986, that are endemic to few Genova(MCSNG), Italy.
hydrographic basins in Western and Central Anentirespecimen (FW250) and fragmentsof
Africa. Finally,Brien(1972, 1973)suggestedthe specimen FW280 were sputtered coated with
erection of the new sub-family gold and observed under scanning electron
Globulospongillinae to define the status of the microscopv (SEM) to define skeletal characters.
endemic genus Malawispongia Brien, 1972, Spicules from FW250, 251, 279, 280 were
known from the Mid-Pleistocenic Lake Malawi/ dissociatedby boiling sponge fragments in nitric
Nyasa. acid, washing in water, and dehydrating in
)
62 MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE I. Morphologicaldiagnostictraitsofsonicfreshwaterspongesfromancientlakesoftheworldthatdonot
produce gemmules. Data basedon original descriptionsand a preliminary study ofholotypes bytheauthors.
Species Lakes Characteristics
Makediatanensis LakeTana s1p)aurnsdeisfpfoernegnitni;at4e)doexcetaossormaneg;i2n)gaflrveoomlastleenidseortrtoopsitcocuth,osatnroasigohmtat!osslkieglhettloyncwuirtvhedp,aufcriosmpiscmuolaorthfibtroevsa;r3i-
gen.elsp.nov. Ethiopia ablyornamented;aceratetips; 151-285/5-22um;
BaUiviaspongia
wirrmanni LakeTiticaea r1e)geucltaroscohmoeananosiormreaglulsakrenleettownorwkitoh(mnunlit-iosrpibciu-lspaircpurliarmamreyshtreascttsaangnedntpiaaulcitsoptihceusluarrfiarcree;g2u)larretsieccuolnatdeariry-
&B1Vo9ou9rl2yk-mEesrn-aRuilbteiro, Peru tsrmaoctost;h3)tospsaprisneeds;poancgeirna;te4)tiopsx;ea1s53r-a4n5g0i/n2g.f6r-o1m3uslmendertostout,fromstraighttoslightlycurved,from
Cortispongilla LakeKinneret p1a)uucnidsipfifceurleanrtisaetcedonedcatroysotmrea;ct2s)mroetriecudlaetnescehtooawnaorsdomtahe!ssukreflaecteo;n3w)itshpamrusletissppiocnuglianr;p4r)imoaxreyastrarcatnsgainndg
bTaorprsoeinsti, 1892 Israel f1r8o0m-3s7l0e/n3d0er-3t3oumstout, from straight to slightly curved, from smooth to granulated; acerate tips;
tkividuspongia 1)undifferentiatedectosome;2)reticulatechoanosomalskeletonwithmultispicularprimarytracts,di-
rotunda LakeOhrid verging in tuftstoward the apical surfacesand paucispicularirregularsecondary tracts; 3)sparse
Macedonia spongin; 4)oxeas ranging from slender to stout, from straight to slightlycurved, from smooth to
Arndt, 1937 spined;aceratetips; 180-367/5-23um
Malawispongia 1)undifferentiatedectosome;2)reticulatemultispicularchoanosomalskeletonwithprimarytractsdi-
echinoides LMaakleawMialawi vraenrggiinnggifnrtoumftsslteonwdaerrdtothsetoapuitcaflrosmursftarcaeisghatntdoisrlriegghutllayrcsuercvoendd,arfyrotmrasctmso;o3t)habtounsdpainnteds;paocnegriant;e4to)obxleuanst
Brien. 1972 tips; 190-240/4-lOum
Ohridospongilla
sGtialnbekrotvi&ciHadzisce, LMaakceedOohnriiad s1p)aurnsdeisfpfeornegnitni;a4t)edoxeecatsosroamneg;in2g)farlovmeosllaetnedeirsottorsotpoiuct,skferloemtosntrawiigthhttpoausclii-gh(tmluyltciu-r?v)esdp,icfurloamrsfmiboroets;h3t)o
1984 spined;aceratetips;V
Pachydictyum 1) undifferentiatedectosome; 2)reticulatemultispicularskeleton withprimary tracts, divergingin
globosum LakePoso tuftstowardtheapicalsurfaces,andmultispicularirregularsecondarytracts;3)sparsesponginwith
Sulawesi abundantforeignmaterial;4)stoutoxeas,straighttoslightlycurved,varyingfromsmoothtospined;
Weltner, 1901 tipsrangefromaceratetoblunt;220-410/30-64um
Lake 1)ectosomeanirregularnetworkofuni~orbi-spicularmeshestangentialtothesurface;2)alveolate
Spinospottgillapotti Tanganika isotropicmultispicularchoanosonalskeletonwithrarescatteredsmalleroxeas(128/2,3urn);3}sparse
Brien, 1974 spongin;4)oxeasrangingfromslendertostout,fromstraighttoslightlycurved,fromsmoothtospined
Zaire andgranulated;tipsrangefromaceratetoblunt; 160-208/1l-12um
alcohol. Suspendedspiculesfrom eachspecimen ectosomal differentiation present. Megascleres
were dropped on slides for light microscopy; are oxeas ranging from completely smooth to
SEM analysis was performed on spicules granulated, tuberculated and/or strongly spined;
sputter-coated in gold sputtering. Seventy spines acutely slanting with a globular base,
spicules, photographed by SEM, were measured named drop-like spines, with an evident axial
from specimens FW251 and FW280; mean and canal. Microscleres and gemmules absent.
standard deviation of measurements were
calculated (in um). Makedia tanensis new species
(Figs 2-3)
SYSTEMATICS
MATERIAL. HOLOTYPE. IZUG-FW251: Bahir Dar,
NW
Class Demospongiae Sollas Lake Tana, Ethiopia, coll. R. Manconi, -.v.1988.
PARATYPES. IZUG-FW250, IZUG-FW279,
Order Haplosclerida Topsent
Family incertae sedis IZUG-FW280: same locality,coll. R. Pranzato, -.v.1988.
ETYMOLOGY. Named forthetype locality,LakeTana.
Makedia newgenus
DESCRIPTION. FW250. Whitish encrusting
TYPE SPECIES. Makedia tanensis sp. nov., sponge(0.5x0.2x0.1cm)onapebbleatadepthof
monotypic. 15cm. FW251. Whitish encrusting sponge
ETYMOLOGY. Named for the Abyssinian Queen of a(0.d5exp0t.h8xo0f.55cmc)m.onFthWela2t7e9ra.lTswidoeowfhaitbiosuhldetrhiant
Saba, Makeda.
contiguous crusts (0.5x0.7cm and 0.3x0.3cm),
DIAGNOSIS. Spongillid-like sponge with within the same concavity of a cobble, on the
skeleton shaped as an alveolate isotropic pauci- dried shoreline. FW280. Brown crust (lx2x
spicular network with sparse spongin. No 0.2cm) covered by unidentified epi- and
LAKE TANA SPONGES
3150km" and a maximum depth of 14m (mean
8m), is tributary ofthe Nilo basin with its single
outlettheBlueNile(RiverAbbay); locatedinthe
N highlands of Ethiopia at an altitude ofabout
1800m the lake is characterised by a strong
seasonality in rainfall and water level; the rainy
and wet seasons occur in summer and winter,
respectively, with recurrent long lasting drought
periods;waterlevelrangeisabout0.4-2.30mand
reaches its maximum about 2 months after the
peak of the rainy season (Bini, 1940a;
Nagelkerke & Sibbing, 1996).Waters are
characterised as oligotrophic; watertemperature
range is 15.6-20°C; silica is 9-16 mg/1 (Bini,
1940a; Rzoska, 1976). The lake is^isolated
downstream by the Tis Issat Falls, and hosts a
FIG. I. Geographical position ofthe type locality of scarcely diversified fauna with the exception of
Makedia tanensis along the coast ofTana Lake in fishes and nematods (Bini, 1940b; Brunelli,
NW
Ethiopia. 1940; Brunelli & Cannicci, 1940; Nagelkerke et
al., I995;Abebe, 1996).
endobionts on the lateral surface ofa cobble al
20cm depth. All specimens share the following DISCUSSION
traits. Encrustingbody shape small in size, up to
0.5cm thick, 2cm diameter (Fig. 2A). Cyclic disturbances produced by seasonal
Consistency was soft and fragile. Surface was waterlevelfluctuationsinthelittoralzonelargely
hispid(Fig.2B,C), osculeswerenotconspicuous influences sponge populations, notably inducing
(Fig. 2A). Ectosome was undifferentiated from stressed conditions and small body size of
choanosome macroscopically. Isotropic specimens. In spite of the suboptimal habitat,
alveolate choanosomal skeleton with where unfavourable conditions are linked to the
paucispicular fibres and scanty spongin (Fig. alternationofthewetanddryseasons,gemmules
2B.C). Megascleres range from slenderto stout, wereabsentinallspecimensofM. tanensisonthe
straight or slightly curved oxeas, smooth to driedshoreline,orinveryshallowwatersofLake
variably spinedwithapicesfrom smoothtospiny Tana. The absence ofgemmules in May, at the
acerate (Fig. 3A). Spicules have a wide endofthedryseason,stronglysupportsthehypo-
morphological variety of irregularly scattered thesis that this species is not able to produce
sculpturing, ranging from granules (Fig. 3C-H), resistant bodies. Several data, however, suggest
granulated tubercules (Fig. 3C-E), drop-like that the production ofgemmules isnotnecessary
granulated spines (Fig. 3C),to large acute spines obligatory in the life history of all freshwater
(Fig. 3E). Two or more ofthese sculptures are sponges, as shown in Lubomirskiidae from Lake
associated on the same or in different oxeas. Baikal (Rezvoi, 1936), which have lost their
Atypicalapiceswerealsofrequentlyobservedon ability to reproduce asexually by means of
spicules (Fig. 3B). Lengths/widths of gemmules(Efremova, 1994),and inothergenera
megascleres are as follows: FW251, from ancient lakes ofthe world.
151-285/6-16um (mean 210/11; standard The recognition of a new species, Makedia
deviation 26/2.5); FW280, 184-289/5-22^m tanensis. in anewmonotypicgenus, is supported
(233/13; 27/3). Microscleres and gemmules are by the possession ofpeculiarornamentations on
absent. oxeas. This isolated monotypic taxon fits the
trend shown by several authors, such as Topsent
HABITAT.Dryandlivingspongeswerefoundon (1892), Weltner (1901), Arndt (1937), Brien
the dried shoreline and in shallow standing (1974), Gilbert & Hadzisce (1984), Boury-
watersup to 20cm depth, associatedwith gastro- Esnault & Volkmer-Ribeiro (1992), indicating
pods,bivalvesandtricladsonthelowerorlateral thatmost freshwatersponges in isolated, ancient
surfaces oflittoralvolcanic pebbles, cobbles and lakes belongto monotypic genera - with the sole
boulders. The Pleistocenic volcanic Lake Tana, exception ofOchridaspongia Arndt (O. rotunda
&
the largest in Ethiopia with a surface of about Arndt, 1937, and O. interlithonis Gilbert
)
364 MEMOIRS OF THE QUEENSLAND MUSEUM
100 pm
FIG. 2. Makedia tanensis sp. nov. (paratype IZUG-FW250). A, Entire specimen. B, Choanosomal alveolate
skeleton. C, Surface hispidation.
Hadzisce, 1984), presently assigned to oneincertaesedisgroupbecauseoftheirdisjunct
Lubomirskiidae. These genera have some distribution and the high possibility ofconverg-
morphological affinities, but their life histories ence as occur in othertaxa ofancient lakes.
arepoorlyknown,theyarehighlydisjunctintheir Some othergenera and species incertae sedis,
distributions. Thequestion,therefore, remainsas such as Metschnikowia and Nudospongilla (in
to their highertaxonomic affinities. part), could also be included in this group in the
The new genus Makedia is characterised by a aftutiuorneobfutthtehiirs rteyqpueirmeastemroirael daestatioledtheeixramtirnu-e
spongiIlid-like skeleton, with apeculiarrange of morphological characters (Manconi & Pronzato,
ornamentations on spicules, and the absence of
microscleres and gemmules. A comparative in preparation).
analysis, in the framework ofa general revision ACKNOWLEDGEMENTS
of freshwater sponge taxa, ofthe type material
and original descriptions ofgenera from ancient We thank all colleagues of the Asmara
lakes, show that these taxa share the following University (Eritrea) for their kind cooperation
traits (Table 1) with the genus Makedia: 1 ) they during our activities ofteaching and research in
are monotypic; 2) known only from the type 1988-89. We are indebted to Clare Valentine of
locality;3)inhabittectonicorvolcaniclakeswith the British Natural History Museum (London),
high levels of endemism; 4) the skeleton is a Dr. F. Puylaert ofthe Musee Royal de L'Afrique
network of ornamented oxeas with multi- or Centrale (Tervuren), Dr. Peter Bartsch of the
pauci-spicular choanosomal tracts or fibres; 5) Museum fur Naturkunde der Humboldt
ectosomalskeleton,ifpresent,uni-orbi-spicular; Universitat (Berlin), and Prof. Claude Levi,
6) they do not produce microscleres; 7) they do Museum National d'Histoire Naturelle (Paris)
not produce gemmules. However diagnostic for providing assistance and access to their
skeletal traits highlight a morphological collections, particularly to type material. This
divergence within this group of sponges from project was funded by MURST, MAE and
ancient lakes. Makedia, Spinospongilla and Interreg CEE.
Ohridospongilla share the spongillid-like
alveolate skeletal trait; on the other hand LITERATURE CITED
Cortispongilla, Ochridaspongia, Pachydictyum, ANNANDALE, N. 1909. Freshwater sponges. In
Malawispofigia and Balliviaspongia (Table 1 BeitragezurKenntnisderFaunavonSud-Afrika.
share the reticulate choanosomal skeleton. All Ergebnisse einer Reise von Prof. Max Weber im
these genera are amalgamated provisionally into Jahre 1894. Zoologische Jahrbuecher Abteilung
LAKE TANA SPONGES 365
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FIG. 3. Makedia tanensis sp. nov. (holotype IZUG-FW251). A, Dimensional and morphological variety of
megascleres ranging from smooth to granulated, tuberculatedand spinedoxeas. B, Atypical apex ofanoxea.
C-D, Surfaceofoxeaswithgranules,tuberculesanddrop-likespines.E,Isolatedlargespineandtuberculeona
finelygranulatedoxea. F, Surfaceofafinelygranulatedoxea. G-H, sectionofagranulatedoxeawithanaxial
canal extendingtoward agranule(arrow).
366 MEMOIRS OF THE QUEENSLAND MUSEUM
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