I I BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, BIOLOGIE, 70: 235-252, 2000 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN, BIOLOGIE, 70: 235-252, 2000
Ecology of the Leptoconchus spp. (Gastropoda, Coralliophilidae) infesting Fungiidae (Anthozoa, Madreporaria)
by Claude MASSIN
Abstract Le parasitisme est maximum en eau peu profonde (1-5m); il est rare de trouver des Leptoconchus spp. a plus de 20 m de profondeur. Underwater surveys of 27,000 mushroom corals (Fungiidae) at L'emplacement ou Ia larve du mollusque se fixe (partie centrale ou Papua New Guinea (Laing Island, Madang and Motupore), Singa peripherique du corail), l'ouverture de Ia loge (face orale ou aborale pore, the Maldi ves (Ari Atoll and South Male Atoll), the Red Sea du corail) et' fa deformation du squelette du corail varient en (Hurghada), Indonesia (Makassar, Sulawesi) and examination of fonction de I'espece de Fungiidae. Ces variations semblent I ,000 fungi ids from museum collections (mainly, National Museum specifiquement liees aux differentes especes de Leptoconchus of Natural Hi story, Leiden) have shown that 36 coral species were plut6t qu' a I' espece h6te infestee. infested by several species of Coralliophilidae belonging to the ge Mots-clefs: Fungiidae, Leptoconchus, Coralliophi lidae, ecologie, nus Leptoconchus. The rate of infestation of the mushroom coral parasitisme, relations corail-h6te. assemblage varied from 0 to 7%. The two most infested Fungii dae were Fungia ( Fungia) fimgites and F. (Verril/ofimgia) repanda. At the east side of-Laing Island, F. (F.) fimgites reached an infestation rate of 19%, the hi ghest value ever recorded. Introduction The rate of infestation was posi tively correlated with hydrodynam ics and negatively with turbidity. It was, however, not correlated Indo-pacific corals can be infested by many mollusc species with the density of the fun giid assemblage. (ROBERTSON, 1970) among which the boring Corallio Infestation was maximum in shall ow water (1 -5 m); nearly no philidae (Gastropoda). They represent inconspicuous ani Leptoconchus spp. were fo und deeper than 20 m. Place of settle mals that infest mainly corals belonging to the families ment of the mollusc (coral centre or edge), opening of the burrow Fungiidae and Faviidae (MASSIN, 1989). So far, there is little (oral/aboral side of the coral) and deformation of coral skeleton var information on their biology and ecological requirements. ied according to the coral species infested . These variations seemed Taxonomic studies of the Fungiidae (HOEKSEMA, 1989a, b, specifically related to the different Leptoconclws species rather than 1990, 1993a, b; HOEKSEMA & CHANG-FENG DAI, 1991 ) and the infested host species. studies on their ecology and distribution (CLAEREBOUDT, Key-words: Fungiidae, Leptoconchus, Coralliophilidae, ecology, parasitism, coral-host relationships. 1989; KOH & CHOU, 1989; HOEKSEMA, 1990, 199la, b, 1992; HOEKSEMA & MOKA, 1989; CHADWICK-FURMAN & LOYA, 1992) stimulated the study on the biology and ecology of Leptoconchus species infesting these corals.
L' etude in situ de 27.000 Fungiidae en Papouasie Nouvelle-Guinee (Laing Island, Madang et Motupore), a Singapour, aux lies Material and methods Maldives (Ari Atoll et South Male Atoll), en Mer Rouge (Hurghada), en Indonesie (Makassar, Sulawesi) et I' examen de Fungiidae provenant de collections museologiques (principalement The fungiids were observed and collected by SCUBA diving du Natio nal Museum of Natural History, Leiden) montrent que 36 between 0 and 30 m depth at Laing Island, Hansa Bay, especes sont parasitees par plusieurs especes de Coralliophili dae Madang Province (fig. 3), off of Madang, Madang Province appartenant au genre Leptoconchus. Le taux d'in festati on des (fig. 2), and at Motupore Island Research Station, Central populations de Fungiidae etudiees varie de 0 a 7%. Les deux Province (fig. 4), all three in Papua New Guinea (fig. 1); at especes de Fungiidae les plus parasitees sont Fungia ( Fungia) Singapore (fig. 6); at the Ari Atoll and South Male Atoll , the ftmgites et F. (Verrilloftmgia) repanda. Sur le cote Est de !'lie de Maldives (fig. 5), at Hurghada, Red Sea, Egypt (fig. 7) and Laing, 19% des F. (F.) fungites sont parasites, ce qui represente Ie off Makassar, Sulawesi, Indonesia (figs 8-9). The fungiids pl us fort raux jamais observe. collections of the National Museum of Natural History L' intensite du parasitisme est positivement correlee avec !'agitation (Leiden, the Netherlands), the Museum of the National Uni hydrodynamique et negati vemenl avec Ia turbidite. En revanche, versity of Singapore (Singapore) and the Royal Belgian In elle ne presente pas de relati on avec Ia densite de populati on des stitute of Natural Sciences (Brussels, Belgium) were exam Fungiidae. ined for infested Fungiidae. Contribution n° 355 of the Leopold III Biological Stat ion, Laing Island. Most of the infested Fungiidae were measured (diameter or I I
236 CLAUDE MASS IN
1 0
·3.: ,.. ...:~ _·· ..... \ ...... 3 \ '" ··...... , , \ .. ·\_
<>o"f ·ri 500 km A AUSTRALIA
3
LAING ISLAND 1 km
3km
PORT MORESBY
I -;::J"A;;·- ... '' ...... ) -~-- ...... 0 ...... - - - ( .._ .. ··-,., ·-.._,_
__.- ·...... --· , __ MOTUPORE .- .··. ------····-···· ·... ·.·... · ... ·······::::.:~.-:-:. 100m 147"10' E
Figs 1-4. - Papua New Guinea. I. General view; 2. Madang reefs (Madang Province); 3. Laing Island, Hansa Bay (Madang Province); 4. Motupore (Central Province). length/width) and photographed. The distance between the with cumul ative curves of settlement of Leptoconchus li ving openin g of the siphon and the mouth or the centre of the coral in ell iptical Fun gi idae with a very long mouth slit and with was measured. Random cumul ative curves of settlement the bun·ow opening at the oral side. were establi shed (fig. 10). Curve A was calculated from the For most of the infested fungiids, coral thickness was meas centre of a circle or an ellipse and curve B was calc ul ated ured at the centre and on the edge. This height, including sep from the median of a rectangle. Curve A was compared with tal and coastal teeth, was measured at the level of the most cumul ative curve of settlement of Leptoconchus li vin g in cir exsert septae (primary septae). The ratio she ll height/shell cu lar Fu ngi idae with burrow openi ng at both oral and aboral width (H/W) of the Leptoconchus specimens was calc ul ated sides and of Leptoconchus li ving in elliptical Fungiidae with for the females. She ll s are defined as globose when H/W~0.8 burrow openin g at the aboral side. Curve B was compared and lenti cular when H/W::;0.7. II
Ecology of the Leptoconchus spp. infesting Fungiidae 237
5 6 MALAYSIA ,.
.::;_:.:·:·:~. ~.:: ·..... MALDIVES .~~~ ~.·. -~ . ·::.. :( ~ ·\.. NORTH \ , 'j MALE ATOLL 4' N :\.;,,.:,. ~c~: · o· J(~· ·' ·,l M~~~r;im SUMATRA ·~~:.,. .... )=' 105' E
7 ~
CAIRO • 30' N
.... ::·':: .::· :;. RED SEA
73' E 35' E
Fig. 5. - The Maldives. • Fig. 6. - Singapore. • Fig. 7.- Hurgada (Egypt, Red Sea).
Each infested fungiid was broken by cracking it with pincers. above coral surface), contracted (still visible but at level with The molluscs were extracted from their hole, then counted, the coral surface) and witlidrawn (no longer visible). anaesthetized with 4% MgCl? for a few hours and fixed in I 0% buffered formalin. Late!· on, they were preserved in 70% buffered alcohol. Results Population density of mushroom coral assemblages were measured by counting the fungiids larger than 50 mm in di GENERAL DISTRIBUTION OF INFESTED FUNGIIDS AND ameter, present on surface of 25 X 2 or 50 X 2 m at different RATE OF INFESTATION depths: 2-4 or 2-5m and 4-7 or 5- 10 m according to the pro fi le of the studied site. Together, the most common corals A total of 28,000 mushroom coral individuals (Fungiidae) were identified underwater and the presence or absence of were examined (27,000 in the fi eld and I ,000 in museum col Leptoconchus specimens was noted. Prospected sites were lections). Of the 43 known species in this family, 7 had, up to selected on protected, semi-protected and exposed areas. now, not been found infested by Leptoconchus species (table About forty infested Fungia specimens (mainly Fungia I). Four of the non infested fungi ids belonged to the genera ( Fungia) fungites and a few F scutaria, F paumotensis and Podabacia, Lithophyllon and Cantharellus, which included Halomitra pileus) have been observed in situ in a protected only fixed fungiids. Three others i.e. Fungia (Cycloseris) area (zone I , fig. 3) day and ni ght to notice the activity of the hexagonalis, F (Pleuractis) taiwanensis and Halomitra tentacles of the Fungia and of the siphons of the clavator are rare species generally characteristic of soft, Leptoconchus. Siphons were observed extended (well visible sandy or muddy bottoms. I'
238 CLAUDE MASS IN
Table I. Li st of fun giid s observed and/or collected +data of literature. A: opening of Leptoconc/111s burrow at aboral side of coral ; F: record from field collected material; !NO: Indonesia; L: record from literature; M: record from muse um collections; MAL: Maldi ves; nbr: number of infested corals observed; NCAL: New Caledonia; 0: opening of Leptoconclllls burrow at oral side of coral; PNG : Papua New Guinea; RS: Red Sea; SEY: Seyc helles; TAW: Taiwan ; 1: HOE KSEMA & ACHITUV 1993; 2: ZIBROWIUS & ARNAUD 1994; 3: HOEKSEMA 1993a; 4: HOEKS EMA 1993b; 5: VERON 1990.
Coral species origin infest. Oral/ Aboral nbr locality Fungia (Cyclose ris) sinensis u + 0 . 6 !NO Fungi a (Cycloseris) fragilis M+F + 0 3 IND+PNG Fungia (Cycloseris) hexagmwlis M - I 0 IND Funliia (Cycloseris) distorra .. M + 0 I !NO Fungia (Cycloseris) cyclolires M + 0 I !NO Fungia (Cycloseris) vaughani M + 0 4 !NO Fungia (Cycloseris) sommen ·il!ei M+L1 + 0 2 IND+SEY Fwzgia (Cycloseris) costulaw F + 0 5 PNG+I ND Fungia (Cycloseris) remtis F + 0 6 PNG Camharel!us doederleini M - ... I 0 RS Cwllha rel!us noumeae M - I 0 NCAL Canrha rel!zts jebbi LJ + 0 I PNG Funliia (Wel/sofi uzliia) l! IW!lllo.w F + 0 18 PNG+IND Funliia ( Pleuracris) mollucensis M+F + 0 13 !NO Funliia (P leura e/is) Kmvis M+F + A 6 IND+PNG Fungia (Pieuractis) pawtwtensis F + 0 35 PNG+IND Fzingia ( Pleuracris) raiwan ensis M - I 0 TAW Fwzgia ( Pleuracris) seychellensis L ~ + A I SEY Funliia (\le rrillofungia) scabm M + A 3 !NO Fun gia (Ve rrillofiuzgia) spini(er F + A 2 PNG Fungia (\le rrillofun gia) concitulll F + A 16 PNG/IND FunKia (\le rrillofilllgia) repanda F + A 58 PNG+MAL+IND Fungia (Lobacris) scuta ria F + 0 20 PNG Herpolirha !imax F + A 15 PNG/IND Pol)phyl!ia ralpina F + 0 7 PNG+IND Pol_lp/zyllia lwvaehibemiae F + 0 4 PNG Heliofzm gia acriniformis M+F + A 23 IND+PNG Fwzgia ( Dwwfungia) fralinae M+F + A 2 IND Ftmgia (Danq(wzgia) lwrrida M+F + A . 18 PNG+RS+IND Fungia ( Dwwfimg ia) scrupo.w F + 0 37 PNG+MAL+IND Fwzgia (Fungia)fungires F + 0 30 1 PNG+MAL+IND Halomirra pileus F + 0 18 PNG +MAL+IND Halomirra clavaror M - I 0 !NO Cwwcris albirenraculara F + A 6 PNG Crenacris crassa F + A 7 PNG+IND Crenactis echinaw F + A 7 PNG Sandalorirha robusra F + A 3 PNG Sandalorirha denrara M+F + A 2 IND+PNG Zoopilus echinmus M+F + 0 2 IND+PNG Podabacia moruporensis Ls - I 0 PNG Podabacia cruswcea F + 0 3 PNG Lirhophyllon mokai M - I 0 IND Lirlwphyllon wzdulatwu M+F + A I !NO I 1 Ecology of the Leptoconchus spp. infesting Fungiidae 239
8 9
4" 20'. 1"
2" ~ 4" 40' • (/) 1- ~ 1- (/) 3 • • 0:: <( (/) (/) ;2 <( ~ s· oo· •
4" PARE PARE ,.o 0~ """' (:)..:j~
s·. []W~M.M OO s· 2o· • 0 0 (MAKASSAR) KABAENA ( ~ NORTH 6 100km 0 SALAYAR s· 20km
118° 119" 120° 121 . 119" 00' 119" 20' 119" 40'
Figs 8-9. - In donesia. 8. Spermonde Archipelago (Sulawesi); 9. Reefs off Makassar (S ul awesi).
100 The most infested species were, by decreasing order, Fungia (Fungia) fungites, F. (Verrillofungia) repanda, F. Random curves 90 ( Pleuractis) pawnotensis, and F. (Danafun gia) scruposa (ta ble I ). However, vari ations occurred and some coral species 80 heavily infested at one locality were free of Leptoconchus at another locality (table 2) 70 I'll The two most prospected sites in Papua New Guinea were ..c:l= 60 Laing Island and M adang (figs 11 , 12). At Madang reefs, =u u= 50 Table 2. Number of in fested specimens/totnl number of infested fungi ids ...c.= Q,l collected x I 00 for the ten most infested fungi ids species according to ~ 40 table I. PNG: Papua New Guinea; IND: Indonesia. ~ 0 30 Coral species PNG IND Fwzgia ( Fun gia) fimgites 39.3 15.0 20 --Random A Fungia (Ve rrillo.fungia) repanda 12 .9 5.0
10 --RandomB Fungia (P!eura ctis) pwull otensis 2.5 25 .0
Fungia (Danqfimgia) scruposa 6.2 12.0 0 10 20 30 40 50 60 70 80 90 100 Helio.fungia actiniformis 4.0 8.0 Fungia (Loba ctis) scutaria 5.0 I DIR X 100 Fungia (Wel!sofilllgia) gran ulosa 2.5 8.0 Fig. 10.- Random cumulative curves of settlement. A: calcul ated from center of a circle or an elli psoid; 8 : calcul ated Fungia (Danafwzgia) horrida 2.0 2.0 from median of a rectangle. D: shorter distance from Halomirra pileus 3.2 1.0 mouth of coral to opening of burrow of the mollusc. R: radius (c urve A) or 1/2 width (c urve B) of coral. Fwzgia (Verrillofun gia) concinna 3.5 3.0 I I
240 CLAUDE MASSI N
700 3,5 Ne 0 600 3 0 ~ "Cl"' D2-5m 500 :~ 2,5 ::!"' D2-5m 5-10m 'S'tJ 400 5-10m ~= 2 "Cl = Q,l :::: 300 .... 1,5 0 ~"' r.. Q,l 200 .5 1 .c :::R e 100 = 0,5 z= 0 0 1 2 3 4 1 2 .)"' 4 Location Location
Fig. II . - Number of fungi ids/1 00 m2 and rate of infes tat ion versus differe nt locali ties th ro ugh Madang's Reef (Pap ua New Guinea) .
(fig . II ) there was a clear decrease in the number of Table 3. Number of fu ngiid species in fes ted (A) and number of specime ns Fungiidae/1 00 m2 and an increase in the rate of infes tati on infested (B) at seve ral coral reefs in the Spermonde Archi pe lago (fig. 9). when sampling from the Mangrove (pro tected area: point I, Localities fig. 2) to the batTier reef (exposed area: point 4, fi g. 2). A Localiti es B Fun gia ( Fungia) fun gires represented 8% of the fungiid as 0 Pan ikiang 0 Panikiang semblage in the mangrove area and 36% on the barrier ree f. I Lae-Lae I Lae-Lae Fi gure II shows also that on the barri er reef, depth (2-5m 2 Gusung 2 Gusung versus 5- 1Om ) had no influence on the number of fun gi id s/ I 00 ·m2. Moreover, the mangrove which CO !Tesponded to 5 Barang Lampo 12 Barang Lampo very turbid waters, lacked any Leptoconchus. 7 Barang Caddi 13 Bone Tambung At Laing Island (fi g. 3), which is sim ilar to a barr ier reef, the 8 Bone Tambung 18 Barang Caddi density of Fun giidae was always hi gher between 4-7 m than between 2-4 m depth in exposed (point 3, fig. 3) and semi 12 Samalona 25 Samalona protected areas (poi nt I , fig. 3). In contrast, the infestati on 12 Kudigareng Keke 32 Kudigare ng Keke rate was simil ar or higher between 2-4 m than between 4-7 m depth. At the exposed side of Lai ng Island , Fun gia ( Fun gia) f ungites reached an infestati on rate of 19%, the hi ghest value At Makassar, reefs cl ose to the coast as Paniki ang, Lae-Lae ever recorded. The num ber of fun gi ids/ I 00 m2 between 0 and or Gusung (fig. 9), with turbid to very turbid waters, were I 0 m depth varied from 0 to 640 wi th an average of 161 nearl y free of Leptoconchus. In offshore directi on, the (Madang +Laing Island). number of infested fun giids increased and the further away
350 8
N 300 7 e 0 6 ~ 250 :s"' 0 2-Sm "?;; · ~ "CC 5 5-lOm :Sn 200 .2 "CC 4 .;:= Q,l ._ 150 ~- 0 "' 3 r.. .5 ~ e 100 =~ 2 = = 50
0 +--'-- 2 3 2 3 Location Location
Fig. 12.- Number of fu ngiids/1 00 m2 and rate of infestatio n vers us different localities arou nd Laing Island (Hansa Bay, Papua New Gui nea). Ecology of the Leptoconchus spp. infesting Fungiidae 241
from the coast, the more infested the fungiid assemblage (ta Fungia (Fungia) fungites ble 3). However, the rate of infestation of the fungiid assem 100 blage for the whole area was still very low: 0 to 0.02%. At Makassar, the three most infested species were, by decreas 90 ing order, Fungia ( Pleuractis) paumotensis, F ( Fungia) 80 fungites and F ( Pleuractis) moluccensis. The most heavily infested fungi ids were found at Papua New 70 Guinea (North Coast) and Indonesia (Sul awesi). In these ar :I"' .c 60 eas, up to now, 35 fungiid species, representing 85 % of the =CJ =CJ species present, were infested by Leptoconchus spp. Around ....= 50 the Maldives and at the Red Sea (Hurghada) only four and ..c. one fungiid species, respectively, were observed infested by ...:l 40 ~ Leptoconchus spp. No Leptoconchus spp. were observed in 0 --Random A 30 Fungiidae around the reefs close to Singapore, where the --61->150mm --61-90mm reefs are situated in muddy water. 20 --o-91-120 mm All the genera and subgenera of F ungiidae were infested by -o--121-> 150mm Leptoconchus spp. which makes the Funoiidae the most in 10 fested coral family. 0 0 16"' 20 30 40 50 60 70 80 90 100 THE POSITION AND THE NUMBER OF LEPTOCONCHUS D/Rx 100 IN THE CORAL
The place of settlement on a fungiid and the number of Fig. 14. - Cumulative curves of settlement of Leploconclws individuals infest ing Fun gia ( Fungia) fungiles. Leptoconchus in a fungiid vari ed greatly depending on the coral species. These differences were obvious mainly be tween species that had the opening of the burrow either at the oral side or at the aboral side of the fungiid. pawuotensis, F (Lobactis) scutaria, F (Danafungia) scruposa, and F (F)fungites. Small Fungia ( Fungia) fungites ( <60 mm in diameter) were Opening ofLeptoconchus burrows at the oral side of never in fested. In corals ranging from 60 to 120 mm in diam monostomatous fungi ids eter, there were nearly always 2 Leptoconchus individuals per coral (8 1% of the observations) whereas more than 2 or 3 Sixteen monostomatous infested fungiids had the opening of Leptoconchus individuals in a single coral were exceptional the Leptoconchus bun·ow at the oral side i.e. Fungia (< I 0% of the observations) (fig. 13). In large corals (> 120 (Cycloseris) sinensis, (C.) fragili s, (C.) distorta, F (C) F. F. mm in diameter), 2 Leptoconchus snails, per coral were less cyclolites, F (C) vaughani, F (C) sommervillei, F (C) frequent than in small corals ( <60% of the observations), (C) costulata, F tenuis, Cantharellus j ebbi, F whereas the observati ons deali ng with more than 2 or 3 (Wellsofungia) granulosa, F (Pleuractis) mollucensis, F (P) Leptoconchus specimens in a single coral were more abun dant (>40%) (fig. 13). Six Leptoconchus individuals (three 90 well separated pairs) per F (F) fungites was the maximum observed. 80 The cumulati ve curve of settletnent of Leptoconchus infest :g 70 ing F (F) f ungites (fi g. 14) clearly shows that the molluscs ..c "' were concentrated near the mouth opening: 90% of the e 60 ~ Leptoconchus specimens were near the coral mouth on a sur fr 50 -+-FFF 2Lep face representi ng 50% o f the coral surface. T he smaller the ...l ... -e-FFF >2Lep " coral, the more concentrated were the Leptoconchus indi IS. 40 -+-FFF> 3Lep viduals near the coral mouth (fig. 14 ). These observations " "~ 30 were valid for F (F) fun gites populations from the Maldives and Papua New Guinea. ~ 20 Measurements of F (F) f ungites corals show that their thick 10 ness varied wi th increasing size. Up to 120 mm in diameter, the peripheral thickness increased sl owly whereas the central thickness remained nearl y constant (fig. I 5). Beyond 120 60-69 70-79 80-89 90-99 100- 110- 120- 130- 140- 150- 109 11 9 129 139 149 159 mm in diameter, the thickness increment was restricted to the Coral diameter (mm) central part o f the corals whereas the edge remained constant (fi g. I 5). Fig. 13. - Number of Lep10conchus indi vidu als present versus T he Leptoconchus specimens in festi ng Fungia ( Danafungia) co ral diameter: Fungia ( Fungia) f ungiles. scruposa and F ( Lobactis) scutaria presented very simi Jar ''
242 CLAUDE MASSIN
28 The Leptoconchus specimens infesting Fwzgia (Wellso --F. fungites centre fungia) granulosa, F (Pleuractis) paumotensis and all the F 26 (Cycloseris) spp. presented very similar cumulative curves --F. fungites edge of settl ement (fig. 16). The Leptocpnchus specimens were 24 somewhat less concentrated near the coral mouth than those ofF (D.) scruposa and F (L.) scutaria. a 22 The single infested specimen of Cafltharellus jebbi had the '-'a opening of the mollusc burrow cl ose to the mouth and resem "':l 20 12... bled the settlement pattern of the Fungia (Cycloseris) spp. ~ 18 In all the monostomatous fungiids with the Leptoconchus f opening at the oral side, there was generall y no or only a 8 16 weak defom1ation of the coral skeleton (figs 17a, b, e, 18a). Only occasionall y strong deformati ons, with sometimes for 14 mation of secondary mouths (figs 17c, d, f ; I8b, c), were observed. Eighty to 90% of the Leptoconchus individuals in 12 festi ng the monostomatous fungiids with the opening at the oral side were single or occurs in pairs. Clusters of 3 10+---.---,---.---.---.---.---.---.---· Leptocon.chus specimens represented more or less 10% of 60-69 70-79 80-89 90-99 100- 110- 120- 130- 140- the ol;>.~ e r va ti o n s and 4 Leptoconchus snail s or more together 109 119 129 139 149 were exceptional. Moreover, the shell of Leptoconchus was Coral diameter (mm) always globose. Tentacles of Fungi a (F) fun.g ites, F ( Lobactis) scutaria and Fi g. 15 . - Coral thickness versus coral di ameter: Fun gia ( Fungia) F ( Pleuractis) pawnotensis came out around 6 p.m. at sun fungites. set. At 9 p.m. they were all full y extended. At 6 a.m. nearl y all the tentacles of the Fungia spp. were withdrawn. During day time, 20% of the Leptoconchus siphon were completely settlement curves (fig. 16): 95% of the molluscs were con withdrawn (fig. 19). This valu e decreased at sun set to reach centrated near the coral mouth on a surface representing I 0% its lowest value (5%) at 3 a.m. Between 6 p.m. and 6 a. m. of the coral surface. Inside F (D.) scruposa, up to 80% of the siphons were parti cul arl y visible, 75% bein g in full extension molluscs had the siphon in the mouth opening. M oreover, the (fig. 19). Around midnight the percentage of extended si number of Leptoconchus snail s infesting a coral was re phons already decreased (fig. 19). During day and ni ght stri cted to one or two (88 and 79% of the observations fo r F d ives, proboscides of the Leptoconchus spp. were never ob (D.) scruposa and F (L.) scutaria, respectively). served protruding through the opening of the bun·ow.
80 100 --extended Siphonal activity 70 90 --contracted 60 __._withdrawn 80
=Q 50 70 .9--= "' ....."' 40 Q -==(,1 60 ~ =Q 0 30 (,1 Q 50 ....c. 20 Ill ...;! 40 ~ 0 --Random 30 --F. (C.) spp...... _F. granulosa 9H 12H 15H 18H 21H 24H 3H 6H 9H · 20 -e- F. scruposa Time 10 -e-F. paumotensis --6- F. scutaria Fig. 19. - Siphonal ac ti vity. 0 0 10 20 30 40 50 60 70 80 90 100 Opening of Leptoconchus burrows at the oral side of D/R x 100 polystomatous fungi ids Fi g. 16.- Cumulative curves of settl ement of Leptoconchus indi viduals in fe sting Fun gia (Wellsofungia) granulosa, Five polystomatous infested fu ngii d species had the F (Pieuractis) pawnotensis, F (Cycloseris) spp. , F Leptocon.chus opening at the oral side, i.e. Halomitra pileus, (Danafungia) scruposa and F (Lobactis) sot/aria. Po/yphyllia talpina, P novaehiberniae, Zoopi/us echinatus II
Ecology of the Leptoconchus spp. infesting Fungiidae 243
Fig. 17.- a: oral side of Fungia (Fungi a) fungites without deformation of coral skeleton. Arrow indicates a pair of siphons of Leptoconchus specimens; b: oral side ofF (F) fun gites without deformation of coral skeleton. Arrows indicate siphons of Leptoconchus specimens; c & d: oral side ofF (F) fwzgites with deformation of coral skeleton . Arrows indicate openin g of Leptoconchus burrows; e: oral side of Fun gia ( Danafungia) scruposa without deformation of coral skeleton. Arrow indicates opening of a Leptoconchus burrow ; f: close up of oral side (central part) of a F (D.) scruposa with strong deformation of coral skeleton around the mouth. I I
244 CLAUDE MASSIN
Fig 18. - a: oral side of Fzm gia ( Lobactis) scuraria without deformation of coral skeleton. Arrow indicates siphon of a Leptoconchus specimen; b: oral side of Fungia (Pleurae/is) pawno1ensis with strong deformation of coral skeleton. Arrow indicates opening of a Lep10conchus burrow; c: oral side of Fungia (Wellsofungia) granulosa with strong deformation of coral skeleton. Arrows indicate opening of Leptoconchus burrows; d: oral side of Halomitra pileus without deformation of coral skeleton. Arrows indicate siphons of Leptoconclws specimens; e & f: gall formed on the oral side of Holomitra pileus. Arrows indicate siphons of Lep!Oconchus specimens; g: inside gall showing a cluster of globose Leptoconchus indi viduals; h: Heliofungia actinifon nis split and showing inside a cluster of lenti cular Lep10conchus specimens. No deformation of coral skeleton is visible; j: aboral side of Ctenactis crassa without deformati on of coral skeleton. Arrows indicate openings of a Leploconchus burrow. 1\
Ecology of the Leptoconchus spp. infesting Fungiidae 245
and Podabacia crustacea. Except H. pileus, few of them 80 were found infested (table I) and cumulative curves of infes 70 tation could not be established. Leptoconchus infesting H. pileus presented a cumulative "' .E... 60 curve of settlement close to the Leptoconchus infesting F (F) =Q ~ fungites (fig. 20): 95 % of the settlement occurred on 50% of c. 50 the coral surface starting from the centre. ...l.. '<: 40 The polystomatous fungiids infested with the Leptoconchus "'c. opening at the oral side generally presented a low (Zoopilus e 3o echinatus, Podabacia crustacea) or a strong (Pol)phyllia ..... spp.) deformation of the coral skeleton. Tlie shell s of the 'f. 20 --+-FVspp 2Lep Leptoconchus specimens infesting those corals were lenticu -+- FVspp >2Lep 10 -+- FVspp >3Lep lar. Sometimes, as for Halomitra pileus, there was formation of a gall (fig. 18e-f) containing a cluster of globose 0 +---.-A-,---,---.---.---.---.---.---.---- Leptoconchus (fig. 18g). 60-69 70-79 80-89 90-99 100- II 0- 120- 130- 140- 150- 109 119 129 139 149 159 Coral diameter 100 N~;nber 90 Fig. 21 .- of Leproconchus specimens present versus coral diameter: Fungia (Verri!lofungia) spp. 80 The number of Leptoconchus specimens m Fungia 70 (Verrillofungia) repanda and F (V) concinna populations "' was poorly correlated with the coral di ameter (fig . 21 ). Two -=....= 60 = Leptoconchus specimens per coral represented onl y 42 % of .... = 50 the observations, whereas more than 2 or 3 Leptoconchus ...c.= Gl specimens in a single coral represented 40 % of the observa ...:l 40 ti ons. The situation was the same for Heliofungia ..~ actiniformis in which up to ten Leptoconchus snail s, fonning 30 --Random A a cluster, were observed in a single coral (fig. 18h). --RandomB The Leptoconchus individuals infesting F (V) repanda, F 20 -6- Ctenactis sp. (V) concinna, H. actinifonnis, and F. ( P.) gravis presented very similar curves of settlement (fig. 22), which were close 10 --Halornitra pileus -e-Herpolitha limax 0 100
0 10 20 30 40 50 60 70 80 90 100 90 DIR X 100 80 Fig. 20.- Cumulative curves of settlement of Leptoconchus indi vidual s infesting Ctenactis spp, Halomitra pileus 70 and Herpolitha limax. "'= -=.... 60 Cl Q "' 50 .sc. Opening of Leptoconchus burrows at the aboral side of ~ ,...;] 40 monostomatous fungi ids ~ 0
Twelve monostomatous infested fungiids had the 30 -e-H. actiniforrnis Leptoconchus opening at the aboral side, i.e. Fungia 20 ( Pleuractis) gravis, F. ( P.) seychellensis, F. (Verrillofungia) -.1r- F. concinna --+-F. repanda repanda, F. (V) concinna, F. (V) spinifei; F (V) scabra, 10 Heliofungia actiniformis, F. (Danafungia) fralinae, F. (D.) --e- F. gravis horrida, Ctenactis albitentaculata, C. echinata, and C. 0 crassa 1• 0 10 20 30 40 50 60 70 80 90 100 D/R x 100 1. C. crassa is a po lystomatous fun giid . However, all the mouths res ulted of intrastomodeal budding inside the ax ial groove. I consider this subdi vi Fig. 22.- Cumulative curves of se ttl ement of Leptoconchus sion of the mouth not impo1tant fo r the settl ement of a Leptoconcllll s. indi vidual s in fest ing Heliofungia actinifonnis, Fungia Co nsequentl y, in the present work , C. crassa will be dealt wi th the (Ve rrillofimgia) concinna , F (V) repanda and F monostomatous fungi ids. (Pleurae/is ) gravis. ''
246 CLAUDE MASSJN
40 cumulative curve of settlement (fig. 20) very similar to the one of those infesting H. pileus (fig. 20) and clearly distinct --F. repanda centre 35 --F. repanda edge from a random curve. No cumulative curves of settlement could be established for the Leptoconchus infesting the two Sandalolitha species and Lithophyllon undulatum because of 30 the scarcity of the observation (table 1). ,-... 8 In all the polystomatous fungiids mentioned above, one or .! 25 two Leptoconchus specimens were present per specimen in .."' 82% of the observations. If more Leptoconclws specimens ~.. 20 occurred in a single coral, they occurred in pairs or isolated. :s Only once a cluster of 3 Leptoconchus individuals was ob ;; ... 15 served. u= The presence of Leptoconchus individuals did not induce a deformation of the skeleton (thickening) for Sandalolitha 10 robusta and 5. dentata (figs 24g, 25a) but did so for H. limax (fig 25b). For the latter, 73 % of the specimens infested pre 5 sented a strong deformation of the septa and a thickening of the coral (fig. 25b). The deformation was always at the oral side. At die aboral side the deformation was weak (fig. 25c) or absent (fig.25e). The shell s of the Leptoconchus infesting 60- 70- 80- 90- 100- 110- 120- 130- 140- 150- 160- 69 79 89 99 109 119 129 . 139 149 159 169 H. limax were globose and those infesting Sandalolitha spp. were lenticular. Coral diameter (mm)
Fi g. 23.- Coral thickness ve rsus co ral diameter: Fun gia SHELL HEIGHT VERSUS CORAL THICKNESS (Verrillofun gia) repanda. From table 4 it appeared that there was a relationship be tween H 2 ffc (H 2: height of shell of 2 Leptoconchus; TC: to a "B" random curve. The Leptoconchus individuals infest thickness of the coral near the center) and the presence/ab in g Ctenactis spp. had a curve of settl ement (fig. 20) very sence of a deformation of the coral skeleton. All the studied close to an "A" random curve. corals (except Fungia ( Pleuractis) gravis) could be divided For the Leptoconchus infesting the species F. (V) spinifet; F. among 3 categories: (V) scabra, F. (D.) fralinae, F (D.) horrida, and F. (P) I ) H Cjl rrc X I 00<60: coral defomlation appeared in 0-30% of seychellensis data were too scarce to establish cumulative the observations, curves of settlement. 2) 60
Ecology of the Leptoconchus spp. infesting Fung iidae 247
Fig. 24.- a: aboral side of Ctenactis echinata without deformation of coral skeleton. Arrows indicate openings of Leptoconchus burrows; b: aboral side of Fungia (Verrillofimgia) concinna with slight deformation of coral skeleton. Arrows indicate opening of Leptoconchus burrows; c: aboral side ofF (V.) concinna wi thout deformation of coral skeleton. Arrow indicates opening of a Leptoconchus burrow; d: aboral side of Fun gia (Verrillofimgia) repanda with slight deformation of coral skeleton. Arrows indicate opening of Leptoconchus burrows; e: aboral side of Ctenacti s echinata wi th slight deformation of coral skeleton. Arrows indicate opening of Leptoconchus burrows; f: aboral side of Heliofimgia actiniformis without deformation of coral skeleton. Arrows indicate opening of Leptoconclws burrows; g: oral side of Sandalolitha robusta without deformation of coral skeleton. II
248 CLAUDE MASSIN
Fig. 25.- a: aboral side of Sandalolitha robusta (specimen of figure 24g) without deformation of coral skeleton. Arrows indicate opening of Leptoconchus burrows; b & c: oral and aboral side of Herpolitha limCLr with stro ng deformation of coral skeleton. Arrows indicate opening of Leptoconchus burrows; d & e: oral and aboral side of H. li111ax without deformation of coral skeleton. Arrows indicate opening of Leptoconchus burrows.
The infes tation rate was not correlated with the density of the ily infested (39.3 % of the fun giid s infested) whil e Fun gia fungiid assemblage. This was observed at different localiti es , ( Pleuractis) paumotensis was onl y ra nked at the 11th place such as the North Coast of Papua New Guinea and the wi th 2.5% of the fun giids infested. At Sul awesi, F (P) Spermonde Archipelago (S ul awesi, Indonesia). Both areas paumotensis, with 25 % of infestation rate, was the fun gii d have a very high pop ul ation de nsity [161 fun gi id s/1 00 m2 most infested. The same was shown by Fungia
(present stud y) and I 06 fun giids/100 m1 (HOEKSEMA , 1990)] (Verrillofimgia) repanda: 12.9% of the fun giids infested in but showed quite di!ferent in festatio n t:ates (0-7% versus 0- Papua New Guinea ve rsus 5% in the Spermonde Archi 0.02%). pe lago. Thi s was not correlated to local ab un dance of the spe At the species level, the same disc repancies were observed cies because F (F) fun gites, F (V) repanda and F (P) between Papua New Gu in ea and Sul awesi. At Papua New paumotensis were the three most ab undant spec ies at both Gu in ea, Fungia ( Fungia) fun gites was the species most heav- Papua New Guinea (CLAEREBOUDT, 1989) and the Table 4 . 1-1<( : slu: ll he ight or Lcptoconc hus: Tc: thickness or coral skeleton at cent rc:Tc: thickness or coral skeleton at edge: Del": height or deformation or cora l skeleton: IJIO: nu1nbcr or corals with dcf"ormation versus total number o r corals observed.
Coral H <;l(mm) Tc (1nm) Te (mm) H Fuugiu (Cycloseris) spp 6.1 ± 1.6 (n= I 0) 12.7±2.9 (n= 9) 6.5± J.:l (n= 9) 94± 16 (n= 12) 50±1 0 (n= l2) I 1/12 F (Wellso{illlg iu) grwwlo.w 5.9±1.5 (n=ll ) 15.4±5 .7 (11= 13) 9.6±].7 (11= 13) 79±15 (n= IO) 49± S(n = IO) I 0/10 F (Pieumclis) gmvis 9.3±2.3 (n= 7) 2 1. 1±2.9(n= 5) 14.2±3.7 (11= 5) 61±9 (n= 7) 43± 7 (n= 7) 34± 7 (n= 5) 517 F (Pieumclis) pm1111olellsis 14.4±2.7 (n= 6) 22.9±4.5 (n= 9) I 5.4±2.6 (11= 9) 96± 15 (n= 6) 63±11 (n= 6) 50±12 (n= 3) 316 F (\ferriffl~{i111giu)Sflilli( er 5.7±1.5 (11= 7) I I 68±12 (n= 7) 24± 5 (n= 7) I 017 F (\ferriffo{i111giu) CIJIIC:iiiiiU 11.4±:l.l (n= 12) 17.7±6.5 (n= I 0) 11.2±2.4 (n= I 0) 94±21 (n= l l) r (\lerri/!IJ{illlgiu repuudu 9.3±2.4 (n=58) 24.5±5.1 (n='i2) 14.1±3. 1 (n=52) 68± 18 (n='i6) 38± I 0 (n=56) 39± 6 (n= 8) 8156 F (Lo/J/Ic:tis) scuwriu 11.6±2.7 (n= 19) 20.9±4.4 (n= l7) 11.5±2.5 (n= l7) 104± 17 (n= l7) 58± 9 (n= l7) I 0/17 H ert}(J/itlw limax 13.8±3.6 (n=l 4) 22.3±3.9 (n= II ) 12.9±3.1 (n= i3) 113±24 (n= l3) 64±16 (n= IO) 'i'i± 'i (n= 5) 'i/1 3 Polypliylliu Sflf'. 8.2±2.2 (n= 12) I 0.8±2.8 (n= I 0) 8.2±2.4 (n= I 0) 90± 12 (n=l2) 70± 10 (n=l2) 47± 6 (11= 11) 11/1 2 tTl 0 0 0 Polypliylliu tu!tJillu 9.3±2.2 (n= 7) 11.8±3.0 (n= 6) 9.0±2.5 (11= 6) I I I I [fQ '< 0 Polypliylliu uoJ·ueliibemiue 6.6±0.7 (n= 'i) 9.3± 1.5 (n= 4) 6.9± 1.5 (11= 4) I I I I .....,., :;:. (V He/io{i111gia uctiuifimu is 14.4±3.7 (n= l9) 27.4±3 .6 (n= l2) 19.0±2. 1 (11= 12) 70±17 (n= IS) 50±12 (n=IS) I 2/18 1:"-- .g C) Fuugia (Danq{i111giu) lior rida 7.2± l .'i (n= 7) 19 .5±4.9 (n= 7) I 1.6±3.6 (11= 7) 67±17 (n= 7) 38± 4 (n= 7) I 217 ~'"' F (D auu/illlg iu) SCI"llf'O.\"ll 14.4±3.4 (n=30) 23.9±6.7 (n=26) 12.0±2.9 (n=27) 119±32 (11=30) 58± 12 (n=30) 54± 9 (n= 9) 9/30 '"'~ t:; Vl F (Fuugia)./i lllgites 12.8±2 .4 (n=26'i) 22.3±3.7 (11=71) 11.7±1.8 (n=69) I I 5±22 (n= 79) 59± II (n=82) 58± 7 (n= l4) 14182 "0 "0 10.4±0.6 (n= 1 8) 74±21 (n 7 (n= l 5) 1 :l Hulomitra pi/eu.1· 13.5±4 .6 (n=29) 18.6±2.9 (n= ll ) 6) 37±28 (n= =25) 46± 25 25 ~ IO 2121 :l Ctenactis spfJ 13. 1±3.0 (n= 12) 26.5±3.6 (n= IO) 16.5±1.5 (n= ) 86±18 (n=2 1) 53± I 0 (n=20) I (fQ=· 'Tl c: C. ec:liina w 18.9±2.4 (n= 7) 31 .7±2.7 (n= 6) 18.8±1.7 (n= 6) 101±14 (n= 7) 6 1±12 (n= 6) I 216 :l r!3. 0.: C. c:rassu 12.9±3.5 (n= 8) 26.5±4.3 (11= 6) 16.9±1.5 (11= 6) I I I . I I» (V C. albit entar:ulata 13.6± 1.7 (n= 4) 26.5±2 .3 (11= 4) 15.9±1.4 (11= 4) I I I I 1'-.l I I 2 11± 16 (n= 4) 95±27 (n=5) 5R± 5 (n= 3) 515 .j>.. Poda/J/Ic:iu cmsta c:eu 7.2± 1.1 (n= 5) \0 '' 250 CLAUDE MASS IN Spermonde Archipelago (HOE KSEMA & MOK A, 1989; fungiid 's food into the coelenteron and/or on zooxanthellae HOEKSEMA , 1990). Local differences in fungiids infested as do the mytilid Fungiacava eilatensis (GOREAU et a/., were already observed in other molluscs, such as the mytilid 1970). Fungiacava eilatensis (see GOREAU eta/., 1969). Within a For the Leptoconchu s snails with the bunow openin g at the fungiid species, some populations seemed to be more "sensi oral side of pol ys tomatous fungiids, which induce a strong tive" to parasi tes than others. deformation of the coral skeleton (gajl), the locati on of the Going fro m a mangrove area to a barri er reef (w ith increasing burrow should be random. Here the thickness of the coral is di stance from the shore) there is an increase in hydrodynam no longer a limiting factor and the mouths of the coral are no ics (wave exposure) and a decrease in turbidity (TOM AS CIK et longer centrally located. Nevertheless, locati on of the al. , 1997). The rate of infestati on was positively correlated Leptoconchus specimens in Halomitra pileus was not ran with hydrodynami cs and negati vely with turbidity (fig. II). dom ; since they remained relati vely close to the coral center Mangrove areas in front of Madang (Papua New Guinea), (fig. 20). No explanation is avail able fo r this kind of distribu shelf zo nes close to the coast at the Spermonde Archipelago ti on. (S ul awesi), reefs close to Singapore, and deep (>20m) The Leptoconchus specimens with the bunow opening at the muddy environments which were all charac teri zed by turbid aboral side of monostomatous fungiids were more wide waters had never, or excepti onall y, been observed with in spread on the coral surface and coral thickness seemed not to fested fungi id s. Obviously, turbid waters were not favourable be a limiting factor. No relati onships were observed between to Leptoconchus settling and/or development. shell fonns and coral thickness. Moreover, neither globose Cumulative curves of settl ement showed that there was a nor lenticular shell s induced a prominent deformati on of the great difference in the distribution on the coral surface be coral skeleton. tween the Leptoconchus individu als with an oral bun·ow For the Leptoconchus specimens with the bun·ow opening at openin g and those with an aboral one (at least for the aboral side of polystomatous fungi id s there seemed to be monostomatous fungiids). a relationship between the shell form (g lobose/lenti cul ar) Leptoconchus with an oral burrow openin g were not se ttling and the presence/absence of a deformation of the coral skel randomly on the coral. They we re always concentrated close eton. Furthemore, the Leptoconchus infestin g Herpolitha to the mouth slit, occupyin g max imum 50% of the coral sur limax, even if a coral deformati on (thickening) occurred, face (s tarting from the center). For most of them, (Fungia were concentrated near the center, where the coral thickness (Danafungia) scruposa, F (Lobactis) scutaria, F was nearl y twice as at the edge. (Wellsofungia) granulosa, F ( Pleuractis) paumotensis, and With or without deformati on of the coral skeleton and what F (Cycloseris) spp. ), the opening of the Leptoconchus bur ever the coral, Leptoconclws specimens occupied half of the rows was located very close to the mouth slit if not in the avai lab le total coral thickness. This represents nearl y the en mouth slit itself (figs 17e, 18a) as it is the case with the tire coral thi ckness th at can by occupied. More or less 40% of mytilid Fungiacava eilatensis (GOREAU et al. , 1969). Only F the total co ral thickness is represented by the spines of the (F) fun gites presented openings of Leptoconchus burrows costae and by the septa of lower orders that are exsert more widespread on it oral surface. The smaller the corals, (HOEKSEMA 1990: fig . 39). The 60% that can be occupied by the closer to the mouth slit were the positions of the Leptoconc!ws spec imens corresponded to the height of the Leptoconchus burrows (fig. 14). Moreover, there was also a compound sy napticulae located in between the septa. clear relationship between the num ber of Leptoconchus Presence/absence of a deform atio n of the coral skeleton was specimens present in a coral and the coral size, 120 mm not linked to the location of the Leptoconchus snail s (see cu ac ross being a hin ge diameter above which more Lepto mulative curves of settl ement) in the coral. Corals like conchus specimens were present (fig. 13 ). This phenomenon Fungia (W) granulosa, F ( L.) scuta ria, F (D.) scruposa with was probably linked to the growth of the coral thi ckness: be molluscs hi ghly concent-rated near the mouth openin g (fig. yond 120 mm across the thickness near the ce nter of the coral 16) or coral s like F (V) repanda, Heliofungia actiniform.is, in creased significantl y (fig. 15), all owin g more Lepto Ctenactis spp. with molluscs randoml y di stributed (figs 20, conchus individuals to establi sh. The thickness at the edge of 22) were evenly present in categori es 1 (H 9 /Tc x 100 <60) the coral remained nearl y constant (fig. 15 ) and most prob and 2 (60< H 9 /Tc x 100<70) deduced from table 4. ab ly never reached the value th at wo uld all ow the full growth Shell shape of Leptoconchus indi viduals (globose or len ti cu of the Leptoconchus shell s. lar) was not necessaril y lin ked to the coral thickness nor to For the Leptoconchus spp. with the burrow openin g at the the fact that the burrow opening was at the oral or aboral side oral side of the coral , the prominent extension of the siphons of the coral. Globose shell s were found in thin corals at ni ght suggested an increase in the respirato ry function and (Halomitra pileus) whereas lenticular shells were found in hence of the mollusc metabo li sm. Since their movements in thick corals (Fungia (V) repanda, F (D .) horrida, the burrow must be very limited, the increasing ni ght ac ti vi Sandalolitha robusta). Lenti cular shell s had the burrow ti es could be related to feeding activities. With a proboscis openin g as we ll at the oral side (Zoopilus echinatus, that never protrudes from the coral surface, the Leptoconchus Polyphyllia spp.) as at the aboral side (F (V) spp, F(D.) spec imens are unable to browse the polyps as do many free hon·ida, Sanda/olitha robusta) of the coral. li vin g Coralli ophilidae (see WARDS, 1965; ROBERTSON , The most heavil y in fested Fungiidae were li vi ng together in 1970; OTT & LEWIS, 1973 , BRAWLEY & ADEY , 1982, the same habitat and were supposed to have the same eco GUZMAN , 1988; PAGE, 1988; HAYES, 1990; BRUCKNER et logical requirements. Consequentl y, openm gs of al. , 1997). The Leptoconchus individu als have to feed on the Leptoconchus burrows at oral/aboral side are not lin ked to a I I Ecology of the Leptoconchus spp. infesting Fungiidae 25 I special habitat but rather would depend on Leptoconchus CLAEREBOUDT 1 M. 1989. Spatial distribu tion of fungii d coral popu species. Shell shape, and shell distribution in the coral seem lation on exposed and sheltered reef slopes in Papua New Guinea. also to be linked to the Leptoconchus species and not to the Proceedings of the 6'" International Coral Reef Symposium, coral species. All these criteria can be used in the taxonomy Townsville 1988, 2: 653-660. of the Leptoconchus. GOREAU, T.F. , N. l. GOREAU, T. SOOT-RY EN & C.M. YONGE. 1969. They are most welcome since Leptoconchus shells present On a new commensal mytilid (Mollusca: Bivalvia) O]Jening into the very few characters to distinguish them one from each other. coelenteron of Fungia scataria (Coelenterata). Journal ofZoolog;~ According to the present observations, several species be London 15 8: 17 1- 195. longing to the genus Leptocon chus infest the family GUZMAN, H.M. 1988. Feeding acti vity of the coralli vorous gastro Fungiidae and not a single species as first beli eved (BOUIL pod Quoyula monodonta (BLAINY ILLE). Review Bioi. Tropical LON et a!., 1983). The Leptoconchus species found in 36(2A): 209-212. Fungiidae by BOUILLON et a!. (I 983) was identified as HAYES J.A. 1990. Distribution, movement and im pact of the Leptoconchus peronii (LAMARCK, 18 I 8) (= L. striatus coral li vorous gastropod Coral/iophila abbreviata (LAMARCK) on a ROPPELL, I 835). However, studies on coral/mollusc Panamani an patch reef. Journal of E.1perimenta/ Marine Biology specificity (MASS IN, I 983, I 989, I 990) and general shell and Ecology, 142: 25-42. shapes are not in agreement wi th this early identi ficati on. HOEKSEMA, B.W. 1989a. Taxonomy, phylogeny and biogeography The taxonomy of Lep toconchus species infesting Fungiidae of mushroom corals (Scleractini a: Fungiidae). Zoologische will be dealt in a separate paper. Ve rhandelingen Lqirlen, 254: 1-295. HOEKSEMA, B.W. 1989b. Mobility of free li ving fungiid corals (Scleractini a) a dispersion mechanism and survival strategy indy Acknowledgements nami c reef habitats. Proceedings of!he 6'" In lerncaional Coral Reef Symposium , Townsvil le 1988, 2: 715-720. It is a pleasure to thank Dr B.W. HOEKSEMA for reviewing HOEKSEl'viA, B.W. 1990. Syslematics and ecology of mushroom cor the manuscript and Dr Ph. WILLENZ fo r improving the Eng als (SiemCiinia: Fun giidae). Doctoral thesis, Uni versity of Leiden, li sh of the first draft; Prof. J. BOUI LLON, Dr M. }EBBS and Netherl ands, 471 pp. Prof. C. PERNETTA fo r facilities at Laing Island Biological HOEKSEMA, B.W. 199 1a . Evolution of body size in mushroom cor Station, Chri stensen Research Institute and Motupore Island als (Scleractini a: Fungiidae) and its ecomorphological conse Research Department, respecti vely. Mr SHUJ AU, ISM AEL, quences. Netherlands Journal of Zoology, 41(2 -3): 11 2- 129. JOH N & ADAM for facilities at the Maldive Islands; Dr D. HOEKSEM A, B.W. 1991 b. Control of bleach ing in mushroom LANE for facilities at Singapore; Dr B.W. HOEKSEMA and his popul ati ons (Scleracti ni a: Fungiidae) in the Java Sea: stress toler family for their hospitality and assistance in Indonesia; the ance and interference by life history strategy. Marine Ecology many people who assisted me during my numerous dives and Progress Series, 74: 225-237. more especially Mrs A. LIEYROUW, S. LUXFORD, M. HOEKSEMA, B.W. 1992. The Position of Northern New Guinea in CLAEREBOUDT, J.M. 0UIN, 1. MALLEFET, H. PILATE and the Center of Marine Benthic Di versity: a Reef Coral Perspective. B.W. HOEKSEMA ; Dr B.W. HOEKSEMA for the identificati on Proceedings of !he 7'" Illfernclfional Coral Reef Symposium, Guam, of some corals. This work was fin ancially supported by the 2:710-717. Belgian Government, the King Leopold III Foundation for HOEKSEMA, B.W. 1993a. Mushroom corals (Scleractini a: Explorati on and Conservation of Nature and several projects Fungiidae) of Madang Lagoon northern Papua New Guinea: an an of the Belgian Fund for Joint Basic Research. notated check-list with description of Ca111harellus jebbi spec. nov. Zoologische Mededeelingen, Leiden 67(1): 1-19. HOEKSEMA , B.W. 1993b. Historical biogeography of Fungia References (Pleurae!i s) spp. (Scleractini a: Fungiidae), including a new species from the Seychell es. Zoo/ogische Mededeelingen, Leiden 67(43): BOUILLON, J., Cl. MASS IN & J. VAN GOETHEM. 1983. Fungiacava 639-654. eilatensis SOOT-RYEN, 1969 (Bivalvia, Mytilidae) et Leptoconchus striatus ROPPELL, 1835 (Gastropoda, Coralliophil idae), mollusques HOEKSEMA, B.W. & Y. ACHITUY. 1993. First Indonesian record of perforant des Fungia (Anthozoa, Fungiidae) recoltes en Papouasie Fungiacava eila!ensis GOREAU et a/., 1968 (Bivalvia: Myti li dae), Nouve ll e-Guinee. 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