Calcareous sponges of the Netherlands. (Porifera, Calcarea) Th. van Koolwijk. Abstract. and that a wide intraspecific variability
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1 Bulletin Zoologisch Museum JNVERSTET VAN AMSTERDAM Vol. 8 No Calcareous sponges of the Netherlands (Porifera Calcarea) Th. van Koolwijk Abstract taxonomy sponges of calcareous occurring in the Netherlands is reviewed using field The observations of live individuals microscopical examination of individual skeletons and study of the breeding cycle. This led to the conclusion that a new species had to be erected and other Leucosole- species reidentified. The common calcareous in sponges the Netherlands are found to be: nia variabilis (Haeckel 872) Scypha ciliata (Fabricius 78) and Scypha scaldiensis n.sp. INTRODUCTION and that a wide intraspecific variability exists in shape and size and in the presence or A recent publication on sponges of the Netherlands (Van Soest 977) provides data on calcareous sponges based on the taxonomy of Burton' s monograph (93)- Unfortunately it has been absence of spicule categories. One cannot describe species only by looking at spicule variation; also morphological histological cytological ecological and life history characters shown that Burton has gone too far by lumping have to be used. large numbers of species (e.g. Hartmann 9; Van Soest Jones 9; Tuzet 95; Borojevic 97). These authors do not share Burton's belief that spicules provide the only valid characters (977) mentioned the following calcareous sponges: Leucosolenia botryoides (Ellis & Solander 78); Clathrina coriacea (Montagu l8l8); Scypha ciliata (Fabricius 78);
2 9 Fig.. Spicules of Leucosolenia variabilis; a T-shaped triradiate; b T-shaped quadriradiate; c Triradiates; d Quadriradiates; e Large oxea; f Small réfringent oxea; g Small non refringent oxea. Grantia compressa (Fabricius 78). In view were studied using light microscope (including of the above it was necessary to reexamine his interference contrast and polarized light) and data and to study these sponges in the field. scanning electron microscope. Three different preparation techniques were used. Thin hand made sections mounted in artificial canada bal- MATERIAL AND METHODS sam to study skeletal features. Mere spicule slides were gained after solution of the soft Calcareous sponges were studied in the field tissue of the sponge in sodium hypochlorate. and in the laboratory on variation in morpholo- The résidu was washed in water and mounted in gy and spiculation breeding periods (in the artifificial canada balsam. In order to be able genus Scypha) growth population density and to study the microstructure of spicules dried ecology. pieces of the sponge were covered with a thin These observations were made in different layer of gold to study the material with SEM. localities along the coast of the Netherlands Every two weeks a number of specimens were during a period of 3 months from July 978 to stained in Hemalun-eosin after decalcifying December 98. Five of these localities all with % acetic acid. After staining oocytes along the coast of the Eastern Scheldt (SW part and embryos if présent were visible. of the Netherlands) were chosen as permanent stations to be visited every two weeks: TAXONOMY Flauwersbocht (artificial rocky shore); Wemeldinge (pontoon); Schelphoek (industrial harbour); Kats (marina); Sas van Goes (sluice). Genus Leucosolenia Bowerbank 8 Sampling was done directly from the shore at (Order Homocoelida Family Leucosoleniidae) low tide sometimes using a scraping net or by snorkling; occasionally scuba dive techniques were used for sublittoral sampling. Spicules Leucosolenia variabilis Haeckel 87 (fig. )
3 9 Synonyms.- Ascandra complicata; Maitland 897: 55. (pers.comm. Van Soest). Elsewhere the species is common along the coasts of Western Europe. Ascartes fabricii; Maitland 897-' 55. Distribution outside Europe (Burton 93): Leucosolenia botryoides; Rousseau 92: ; Arctic Mediterranean South Africa Straits of Van Soest 977: 2 pi- A-B. Magellan Chile. Leucosolenia fabricii; Rousseau 92: 2. Leucosolenia complicata; Rousseau 92: 2. Ecology.- Description.- L. variabilis occurs in a variety of environments and shows no specific preference for substratum although it is commonly found attached Sponge asconoid appearing in two transitional forms either forming a rather compact to algae hydroids etc. bushy mass (bushy fonn) or spreading out over Discussion.- the substratum (spreading form). Sponge connected basally with the substratum by little ana- The taxonomy of the Homocoelidae has been a stomosing basal tubes often bearing diverticu- problem for a long time mainly because of the la. Oscular tubes are long smooth and slightly mistakes made by Bowerbank (8 8 87) curved they show no tendency to anastomose or and Haeckel (872). Minchin (9) revised the form diverticula. Surface minutely hispid vents terminal; colour alive and in spirit: genus Leucosolenia (Ascandra) thoroughly and Sara (953» 95b) completed the taxonomy and white yellow or grey. Skeleton consists of ecology of the genus. triradiates quadriradiates and oxea. L. complicata appears in a bushy and in an Triradiates slender sagittal unpaired arborescent form (the latter form is never angle distinctly larger than 2 unpaired ray 5- ym by 7 nm paired rays 8-2 pm by 7 Mm. T-shaped triradiates are relatively found in L. variabilis). Its habitats are rockpools of the sublittoral and it is very rarely found in situations in which it is liable to be left dry at low tide. The sponge is very scarce unpaired ray 3- \m by 5- Mm paired rays 5-7 torn by 5- pm. fragile. The very long oscular tubes are always provided with diverticula. The oscular rim is Quadriradiates similar to triradiates apical ray - Mm by 7 Mm projecting into long. The unpaired angle of the triradiates is the cloacal cavity. only slightly thicker than the paired angles but always longer than the paired rays. Small Oxea curved at the proximal end lanceheaded 8-32 Mm by 2-9 Mm; short slender réfringent straight 7- Mm by 2- Mm; do. non-réfringent oxea are not present (Minchin 9; Sara b). non-réfringent. L. fabricii is considered a synonym of L. complicata. Distribution.- L. botroides is a very characteristic The potential distribution of sponges in the species (Ellis & Solander 78) it appears Netherlands is restricted to only few areas only in one form and in one situation. The with suitable substratum. L. variabilis is very species is always found growing over algae common in the Oosterschelde (Eastern Scheldt). forming a dense cluster of smooth oscular The species is more difficult to find in the tubes rising from a basal reticulum of fine Westerschelde and appears to be relatively tubes. The sponge resembles a bunch of bananas. scarce on the islands Texel and Terschelling in The oscular rim is short. The spiculation of L. botryoides resembles the spiculation of L. the northern part of the Netherlands. Occasionally the species has been found attached to variabilis however all spicules in L. botryoides cork and seaweed on the beaches along the west are much more robust T-shaped triradiates are coast. The species was found recently in the abundant and large proximally thickened oxea are present (Michin 9; Sarâ a sublittoral of the closed sea arm the Grevelingen (also in the SW part of the Netherlands) 95b).
4 92 Fig. 2. Spicules of Scypha ciliata; a Tubar triradiate; b Oxea; c Subendosomal triradiates; d Endosomal quadriradiates. Genus Clathrina Gray 87 This species has been found washed ashore on (Order Homocoelida Family Clathrinidae) the beach of Camperduin (Van Soest 977); it is apparently an allochthonous species. The ma- Clathrina coriacea (Montagu l8l8) terial conforms to the description of this species in Burton (X93) This species has been found only once in the The family Grantiidae is distinguished from coastal waters of the Netherlands (Van Soest the family Sycettidae by the following characters: Transition between the syconoid and the 977: 2 pl. C). It is apparently an autochthonous species. The material conforms to the description of this species in Burton (93). leuconoid form appearing sacciform compressed; vents apical or dispersed; dermal cortex The family Clathrinidae is distinghuished present consisting of tangential radiates and from the family oxea. Leucosoleniidae by the following characters: Always a clathrate mass of anastomosing tubes vents never markedly tubular; only regular triradiate systems are present; Genus Saypha Gray 82 the larva is parenchymula larva instead of the (Order Helerocoelida Family Sycettidae) Amphiblastula larva found in the Leucosoleniidae. Scypha ciliata (Fabricius 78) (fig. 2) Synonyms.- Genus Grantia Fleming 828 (Order Heterocoelida Family Grantiidae) Sycandra ciliata; Vosmaer 882: 5. Sycandra ciliata trans.var. coronata; Vosmaer 882: 5. Grantia compressa (Fabricius 78) Sycon ciliatum; Maitland 897: 55; Van
5 Distribution Soest 977; 25 (in part) pi. 2 A. whereas in other parts of the Netherlands the length apparently does not exceed 5 cm. Description.- Discussion.- Sponge syconoid tubular erect cylindrical. Surface papillate and hispid. Distal chambers grouped around central atrial cavity. Vent Dubosq & Tuzet (937 9) considered S. apical usually without fringe sometimes with ciliata and S. coronata as being two separate fringe. Texture soft to firm; colour alive and species based largely on ovogenetical and em- in spirit: white yellow grey or brown. Tubar bryological criteria. However both species are skeleton of basal rays of subendosomal sagittal triradiates and several rows of tubar triradi- morphologically identical. The original description by Ellis & Solander (78) does not pro- ates distal cones ornamented with oxea; endoso- vide distinctive features (they were probably màl skeleton of paired rays of subendosomal unaware of Fabricius' earlier description) and sagittal triradiates and endosomal triradiates Haeckel's criteria (872) are merely based on and quadriradiates. the thickness of the oxea and the length of the Tubar triradiates sagittal or subregular apical ray of the quadriradiates. Above all paired rays -8 um by 5- um basal ray its is not clear on which skeletal citeria -2 um by 5- um. Dubosq and Tuzet identified their material. If Ectosomal oxea 3-3 um by -8 um. we follow Haeckel's description both "species" Subendosomal triradiates sagittal paired occur in the Netherlands as part of a series rays -2 um by 5- um basal ray 22-2 of continuous transitions. um by 5- um. Furthermore they are not geographically dis- Endosomal triradiates sagittal or subregu- tinct and they occupy the same ecological lar paired rays -5 um by 5- um basal niches. Breeding of coronata individuals ray -2 um by 5- um. occurs in the same period of the year as that Endosomal quadriradiates similar to triradiates with apical ray 2-8 ym by 7- vim. of ciliata individuals. Therefore in agreement with Burton S. ciliata and S. covonata are regarded as one being species. The species is very common in all coastal waters of the Netherlands provided suitable Scypha scaldiensis n.sp. (figs. 3-) substratum is present. At present it is not found in the closed sea arm Grevelingen (salini- Synonym.- ty 2-23 / oo S). Elsewhere the species appears Scypha ciliata (in part: fo. villosus); Van Soest 977: 25 pi. 2 B. to be very common in all Western European coastal waters. Distribution outside Europe (Burton 93): Holotype : ZMA POR.5O Kats marina on weeds Arctic Atlantic and Pacific coasts of North attached to pontoons 23-XI-979 America Chili Atalantic Mediterranean and In- coll. Th. van Koolwijk & J. Ver- dian coast of Africa Australia. meulen. Paratypes: Ecology.- ZMA POR.52 2 specimens Wemeldinge on harbour poles 2-VIII- The sponge is able to grow on all suitable 975 coll. R.W.M. van Soest & substrata such as seaweeds wood rock etc. In J. Vermeulen. the littoral S. ciliata is mostly found underneath rocks except on Terschelling where it ZMA POR.SI 8 specimens Schelphoek on pontoon 23-XI-979 coll. was commonly found attached on the upper side Th. van Koolwijk & J. Vermeulen. of rocks. At Terschelling too the species BMNH reg. No speci- reaches a considerable length of up 8 cm mens Schelphoek same data.
6 9 Fig. 3. Scypha scaldiensis n.sp. holotype ZMA POR.5. Description. Distribution.- The species is common in the Oosterschelde but not found elsewhere. Sponge tubular erect cylindrical syconoid. Surface hispid scarcely papillate. Distal chambers grouped around central atrial cavity. Vent apical always with large fringe. Discussion.- Colour alive and in spirit: Grey or brown. Sponge length up to 5 cm. Tubar skeleton of The first record of S. scaldiensis is Wemeldinge 28-VIII-95'> leg. J.H. Stock. Although basal rays of subendosomal sagittal triradiates the sponge is common at the present time it is and several rows of regular tubar triradiates. very rare in preserved collections before 975- Distal cones ornamented with large oxea giving The Oosterschelde where S. scaldiensis the sponge a very "hairy" look. Endosomal skele- appears to be present is a unique environment. ton of paired rays of subendosomal sagittal It contains very clear water with rather high and constant salinity (28-3 / oo S). Each tide triradiates and endosomal triradiates and quadriradiates. % of the total watervolume is renewed. It has Tubar triradiates regular to sagittal rays a rather rich flora and fauna with specific 8-5 um by 9- Mm. sponge endemes such as Mycale micracanthoxea Ectosomal oxea 5-7 nm by 5- om. Subendosanal triradiates sagittal paired Buizer & Van Soest 977 and rarities as Haliclona loosanoffi Hartmann 958 (cf. Van rays 8-2 Mm by 5- nm basal ray 8-35 Soest 97). by 5- nm. Van Soest (977) found among "normal" papil- Endosomal triradiates sagittal or subregular paired rays -28 im by 5- Mm ba- late S. ciliata specimens occasionally samples of greyish very hairy specimens. He sug- sal ray 2-3 pm by 5- Mm. gested that this form might be identical with Endosomal quadriradiates similar to endosomal triradiates with apical ray 8-35 ym by S. villosum (Haeckel 872). From a microscopical slide of one of Haeckel s specimens of 5- pm. Sycandra villosum (from Ireland incorporated Triradiates of distal cones 3- van by in the Muséum national d'histoire naturelle 5- Mm. Paris kindly sent on loan by Prof. C. lévi)
7 Fig.. Spicules of Scypha scaldiensis n.sp.; a Tubar triradiates; b Oxea; c Subendosomal triradiates; d Endosomal quadriradiates; e Endosomal triradiate; f Triradiates of the distal cones.
8 9 (Oocytes embryos or both) Table I: Breeding cycle of Scypha ciliata (S.c.) and Scypha scaldiensis (). May June July August September October November December January February March April S.c. S. s. S.c. S. s. 5. c. S.c. S.c. S. s S.c. S. s. Total No. of specimens No. of specimens with embryos i % Active % Embryos
9 - late 97 it was clear that it is not identical with S. breeding periods of both species were studied. sealdiensis. The conspicuous robustness of the It has already been shown that they occur in oxea and the tubar triradiates in S. villosum (-3 vim) differ from the rather slender the same ecological niche without signs of hybridization so we may conclude that both species are not very closely related. Orton oxea and tubar triradiates in S. scaldiensis (7- urn). In S. villosum we see spicules (9 92) showed that breeding in S. covona- of two clearly different sizes spicules as ta occurred twice a year spring and late mentioned above are two to four times as robust summer and fall. In S. ciliata (table I) breed- as endosomal and subendosomal spicules whereas ing occurred in a continuous period from May to in S. scaldiensis they are of about the same November. The individuals disintegrate late thickness. fall after breeding. During winter the young and small individuals grow. In early spring we Etymology.- see again a regression in population density. The name scaldiensis is derived from the These remaining individuals breed during summer. In S. scaldiensis the breeding period is latin name Scaldia = (Ooster) Schelde (ref. Julius Caesar de Bello Gallico). much longer from June to early February. This period is divided into two by a sexually nonactive period in the month of September. A Breeding cycle in S. ciliata and S. scaldien- large part of the population disintegrates sis (cf. table I fig. 5).- during early fall but the remaining part of the population In order to find other distinctive characters between S. ciliata and S. scaldiensis and the fast growing young individuals continue to live throughout the winter. Fig. 5. Breeding cycle of Scypha ciliata and Scypha scaldiensis n.sp.; a Percentage of sexually active specimens of Scypha ciliata; b Percentage of embryobearing specimens of Scypha ciliata; c Percentage of sexually active specimens of Scypha scaldiensis n.sp.; d Percentage of embryobearing specimens of Scypha scaldiensis n.sp.
10 The 98 This group is forming oocytes in late spring. signification.- Arch. zoôl. exp. gen. 8: The severe winter of 978/979 reduced the popu (Notes et Revue). ELLIS J. & SOLANDER D. 78. Natural History lation but the winters of 979/98 and 98/98 apparently had no influence on the population. The winter breeding period of the species might indicate that S. scaldiensis originates from more southern regions and the recent occurrence of the species might be common due to introduction via imported oysters. curious and uncommon zoophytes collected of many of the globe.- from various parts London: -9 tab. 58. HAECKEL E Die Kalkschwâmme eine Monographie 3 Vol. (G. Reimer Berlin). HARTMANN W.D. 9- Review of "A revision of the classification of the calcareous sponges" by M. Burton.- Science : JONES W.C. 9. Review of "A revision of the classification of the calcareous sponges" by M. Burton.- Nature 2:. MAITLAND R.T Prodrome de la faune des ACKNOWLEDGEMENTS Pays Bas et de la Belgique flamande.- I-X -2 (Brill Leiden). MINCHIN E.A. 9. The characters and synonymy of the British species of sponges of the I would like to thank Dr. R.W.M. van Soest for guiding and criticising this project and Mr. genus Leucosolenia.- Proc. zool. Soc. London 2: ORTON J.H. 9- Preliminary account of a J.J. Vermeulen for the practical help and the contribution to an evaluation of the sea.- many trips we made together. J. mar. biol. Ass. K): Sea temperature breeding and distribution in marine animals.- J. mar. biol. REFERENCES Ass. 2: ROUSSEAU E. 92. Note monographique sur les spongiaires de la Belgique.- J. Ann. Soc. male. Belgique 37: -2. BOROJEVIC R. 97- Importance de l'étude de SARA M Variabilità delle Leucosolenie la repartition écologique pour la taxonomie del Golfo di Napoli e nuove vedute sulla des éponges calcaires.- Helgolânder wiss. Meeresunters. _L5: 9* BOWERBANK J.S. 8. A monograph of the British Spongiadae vol. I: i-xx pis. (Ray Society London). 8. do. vol. II: -388 (Ray Society London). 87- do. vol. Ill: i-xviii pis. (Ray Society London). BUIZER D.A.G. & van SOEST R.W.M Mycale micracanthoxea nov. spec. (Porifera Poecilosclerida) from the Netherlands.- Neth. Journ. Sea res. J_ (3/) : sistematica del gruppo.- Ann. st. Mus. zool. Univ. Napoli a. Sulla presenza e significato di un nuovo tipo di oxee in Leucosolenia botryoides (Ell. et Soil.) Ann. (Calcispongie).- st. Mus. zool. Univ. Napoli 8 (5) : -. 95b. Aspetti genetici ed ecologici dell ibridazione naturale fra differenti specie di Leucosolenia (Calcispongie) a Roscoff.- Boll zool. 23 (2): -3. SOEST R.W.M. van 97ÏÏT First European record of Haliclona loosanoffi Hartmann 958 (Porifera Haplosclerida)Beaufortia 2 (3): BURTON M. 93. A revision of the classification of the calcareous sponges (I-V): -93 (British Museum Nat. Hist. London). DUBOSQ. & TUZET L'ovogenèse la 977- Marine and freshwater sponges of the Netherlands.- Zool. Meded. 5 (): TUZET. 9. Remarques fécondation et les premiers stades du dé- veloppement des éponges calcaires.- Arch. Zool. exp. gen. 79: Sur les cellules en croix des Sycon (Sycon cilia tum Fabr. Sycon coronatum Ellis et Soil. Sycon elegans Bower) et leur sur la classification des Sycon telle qu'elle a été conçue par Burton (93).- Arch. Zool. exp. gen. (l): 8-82 (Notes et Revue). VOSMAER G.C.J Sponzen.- Tijdschr. Ned. dierk. Ver. (l): li-lii. Th. van Koolwijk Instituut voor Taxonomische Zoologie (Zoblogisch Museum) Postbus 225 received :.VIII HC Amsterdam Netherlands distributed :
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