Acta Bot. Need. 23 October-December (5)-(6), Vaucheria J. A compacta: estuarine algal 613-626. 1974, p. euryhaline species Simons Vakgroep Plantensytematiek, Biologisch Laboratorium,Vrije Universiteit, Amsterdam SUMMARY Data are A presented mass occurrence. races” “salinity ridial of but that At sociation a As a a of ecology of V. Vaucheria compacta, in salinity compacta estuarine an of consequence this, a new monoecious filaments Vaucheria species the it is concluded ranges observed were Netherlands. often showing that there is present in is described: variety in community, non-modificative morphological variation occasions some and experiments From culture characters. dalcis. var. distribution on description is given Vaucheria within are no anthe- some compacta this dioecious species. 1. INTRODUCTION the Investigating that V. Dutch coastal From the records few are to it is named types be must, in any 1956, & Conover to it Denmark 1939); Behre 1953, appeared in the 1968). that seems and N. America. Europe (Dangeard as (1900) V. compacta Rosenvinge by to as be Although dioecious species, “It is can of it species & Vroman (Simons distribution, the European and England American records 1961). Webber a V. piloboloides distinct (Collins) 1968) and Quebec be hoped arranged case, treat in So history. V. sphaerospora var. compacta. After his so in Taylor Christensen (1937) prefers to Taylor. ex to in form and especially and correct, Collins as var. species strictly not (1879) paper Christensen also drew attention same widespread information about the nomenclatural Collins by compacta. name within this states: (Blum (1952) gives V. coasts Germany (Rieth Collins considered it description In the geographical Netherlands, the and 1958). dioica. and later quote the its has been described first species in common in estuarine habitats France from Massachusetts & Wilce on most the Atlantic from 1952); Christensen the records a.o. (Christensen (Blum of the one region, especially is restricted species are brackish compacta is morphological variation size of the antheridia. He that further research will decide whether the different mutually the complex distinct as a groups or not, but at present one whole”. The present paper deals with variation in form and arrangement of antheridia, and based on these dulcis. Another V. compacta var. has not oogonia, data variety a new has been koksoakensis. variety is described named distinguished by This variety, V. compacta Blum & Wilce characterized been recorded in the Netherlands so far. (1958) var. named by extremely long 614 Fig. J. SIMONS 1. relate lines Survey to the are of the S.W. Dutch situation drawn according surface, midtide, and : isohaline -I —I —h: l°/oo. before estuarine the to dams data average river 0,3°/o 0 - On the map of the Netherlands: D: Dollard, Z.F.: polder Z. from The position Haringvliet (1970) (1967). Peelen The of isohalines and and Volkerak isohalines were tidal (1969). limit Isohaat measured discharge. 16.5°/00 Cl", + + +: region. in the : = = T: 15°/o„, 10°/oo =: designed Terschelling, Flevoland, G: : , 5.5°/o 0 , 3 0 /00 , construction of dams. S: Grathem. Schiermonnikoog, L: Lauwerszeepolder, VAUCHERIA 2. COMPACTA: A AND DISTRIBUTION 615 EURYHALINE ESTUARINE ALGAL SPECIES PERIODICITY OF V. COMPACTA IN THE NETHERLANDS The distributional pattern of of habitats variety The vertical range MHW in freshwater marine V. plex in the consisting tufted compacta in the Netherlands includes about or about 30 1 the typical rivers zone, muddy (figs. greater part of 1971). Fig. tufted aspect. a areas V. compacta is the often forming sand 3 and (figs. 4). 3). Dutch Luxurious Meuse reaches 2. Mass occurrence of V. Along V. species occurrence as compacta on a algal far as sandy of the flat at V. and subsimplex in salt marshes and most These “nopjeswier” below euhalinic and important algal mats tidal Vaucheria especially mats algal mat (Nienhuis oligohalinic often The function of these tufts may be name m. dark green Vaucheria extensive 2 and great 1). In such habitats V. subsim- mesohalinic Along a MHW in brackish MHW. littoral part of creek banks high Biesbosch the Rhine and below thuretii). V. mode of aeration of the otherwise compact area the map in fig. from either about 2 m. above cm. and V. velutina (= littoral appearance see V. compacta grows intermixed with the the of names compacta extends areas and in freshwater tidal species a V. Nienhuis & Simons 1970, V. geographic V. velutina make up the carpeting waters flats waters sphaerospora) and often of tidal localities towards polyhalinic (= local (for to on show a provide felts. In the freshwater tidal has been derived from this compacta upstreams of the tidal influence is measurable. Hellegatsplein(Haringvliet). 616 Fig. J. 3. Close up of V. compacta mat at Krimpen a/d Lek, with Callitriche stagnalis panyingphanerogamic species. Fig. 4. Net-like appearence of tufted V. compacta mat in Lauwerszee polder. as SIMONS accom- VAUCHERIA Fig. 5. COMPACTA: Parts Salicornia of province of displaced V. only a one of the few records exist. The to due 200 km. mats Haringvliet, of the most With the (1969). Hollands Diep and most been rolled up behind a compacta is nowadays restricted The tidal influence at these Rotterdam. For at example, Krimpen a/d to places one vanishing theseriver systems. only a comes compacta be observed can towards Apparently the beyond species (Dangeard 1969 along level where 1939). limit of this belt stones tidal was withstand a quays determined In the northern part of the is one of freshwater tidal with locality region by much V. occurrence of marshes called the river Western Scheldt. heavy pollution. as Bremen more were the rich (mesohalinic) or V. 1961), Bordeaux and V. compacta in Paris in less continuous belt above covered with splash Netherlands, This is in accordance (Behre The present author has observed of the longer occurrence via the Nieuwe Waterweg canal from the brackish the river Seine forming the streams Lek East of Rotterdam. with other European records from towns Paris no Mass few remaining freshwater Antwerp along can Haringvliet of the tides in the V. compacta In the southern part of the Dutch S.W. estuarine Saaftinge distant record is from the the mouth of the Biesbosch, prominent algal species along compacta is left V. has inland). the construction of dams in and in the Volkerak called near mats, 617 ALGAL SPECIES the tidal influence in the greater part of the Rhine-Meuse estuary has Recently areas. compacta Limburg (about disappeared V. EURYHALINE ESTUARINE plant. Farther upstream, (1970) A some silt. water The upper zone. compacta occurs in the Wadden 618 J. Sea in salt region creek optimum growth along and of the the Dollard substratum. The soil This of compacta A Saaftinge dominant assumed species. ”Vaucherietum” . actually tion is presumed compacta be to shores of brackish appear Z. polder be to an and the Sea. In these draining an inland can mass is up. Similar vast over really and a V. During ditches. cases of up dominant one as association an species a estuaries. but was visit mats non-tidal in to studied be May polders the namely 1969 from the Wadden present within areas can also reclaimed Newly could relatively the along it polder was be lifted from the can after the year a V. compacta showed polder this weather. In fig. 5 it rolling as but species, compacta 1968 from the freshwater lake May sandy which mat rainy of V. by from Schiermonnikoog (Simons 1974), in the Lauwerszee Especially contains European on brackish reclaimed in of creeks. during stormy be rolled plain Vaucheria aspect development Vaucheria beach pools, (1961) Kötter by environments, Lauwerszee reclaimed in the water. gently sloping edges water a 1), polder polders of the abundant the (fig. mass. developed mention any not growth consisting ideal environment. Two Flevoland Ijsselmeer, in tidal occurs are: in mesohalinic Vaucheria compacta sociation. This seda- in the N.W. common environments. Examples richly most Vaucheria aspect community sociation: a only not Vaucheria described this This be called should V. the the of lutum large very develop to nature mentioned here has also been described as that this Kdtter the relatively sandy places. habitat in the river Elbe. Kötter does be safely where Scheldt), a shown in from the tidal part of the river Weser and comparable named (Western mixed again appearance that mass contains also shows its oligohalinic parts explained by environment for algae is is compacta show the not of the (part compacta between salt marshes zone V. sea This may be composition aspect of V. compacta (1961) can the Dollard marshes an be found in mats can mass Behre it of the influence of substratum marshes transitional a to of salinity. unstable too in Closer Dollard V. V. velutina. In the mesohalinic and account on also in the of the Wadden isles V. compacta does marshes, expected fraction, causing a and subsimplex up with V. and plains, systems (meso-oligohaline). dunes could be beach marshes, In the salt marshes estuary). Eems SIMONS be bottom that such seen moist, observed that has also been mentioned by the by a mat Kotter (1961). One species of the is explanations the aplanospores. The this left behind envelope aplanospore production an or liberation, (fig. 6d-g). the Sometimes before being liberated. of this occurrence of asexual aplanospores pioneering propagation takes place at by algal means the filament of tips. the walls of the terminal part of the filament laterally be formed and after mass empty envelope. The filament (fig. 6a) can of aplanospore (fig. 6c). so on has been observed in V. Immediately filaments as an the capacity production After liberation of are for well-developed can Shortly (fig. 6a). sescuplicaria, grow after A similar V. are way subsimplex aplanospore germinates aplanospores out either growing into observed of a new aplanospore and V. velutina. one to through out or more new germinate even VAUCHERIA Fig. 6. COMPACTA: A EURYHALINE different modes of Sexual reproduction place the the whole over mesohalinic to As the occurrence can be and summer THE 3.1. In order gical the of sea to with to environment, compacta, it in never an ways: a: a in optimum be can great stated optimum an that of the periods and quantities, propagation September AND in this it does paper occur in the year optimum the species often more is restricted in to and October. MORPHOLOGICAL clearly over out to the which species, alga was were extent VARIATION demonstrate the the whole char- euryhaline salinity traject of the cultivated freshwater basis 12°C, c: many a 25°/ salinity estu- or basis. ranges. of Salinity Cl“. Culture condi- were Salinity L/D: 12/12, and light Erdschreiber solutions was with Erdschr./seawater, dilution of physiolosalinity 00 photoperiod Media used or a seawater mixing Erdschr./freshwater and morphological times in situated between 0 and temperature of Erdschr./freshwater there exists which could be correlated with the intensities between 1000 and 2000 Lux. with either much smaller scale. It takes freshwater, with freshwater. values in the cultures a aplanospore produc- character presented make involved on a in non-brackish situations. Sexual variation within this tions V. well, although euryhaline from sea to and autumn, but in other summer of V. compacta as aries, be it also, range from EURYHALINE CHARACTER The acter observed was salinity autumn The field data a-c: areas. as than germination of aplanospores. germinationof aplanospores. periodicity is in found brackish 3. and Aplanospore production tion; d-g: 619 ESTUARINE ALGAL SPECIES Erdschr./seawater varied in different b: adding NaCl with distilled to water. 620 Within each the number of range about sample (after clonal rial of V. Biesbosch and From showed 11 growth Cl 5°/ could with respect Sexual range. is Aplanospores 00 - Cl be salinity to ten. Each collected freshly from or have tested mate- the Western Scheldt on localities concluded the as 16°/ 00 Cl that (mostly freshwater occurs both the V. V. ; 20°/ often was Cl oo - Thus . mesohalinic part and the no compacta. optimum an of the between 0.5 and about only brackish 10°/oo freshwater material. conditions tested, salinity are compacta - and oligohalinic the there the brackish Growth stops between 16 and under all produced are . reproduction similar for again , a laboratory) we freshwater salinity: to between 0 and found in was growth and - Waarde locality it experiments races occurs found between 1 and optimum the In this way from several similar behaviour towards a taken from of acclimatization in aplanospores. and Cl”) 0 o culture physiological Vegetative salinity °/ were SIMONS Krimpen a/d Lek). these different quantities compacta from the brackish of about (salinity weeks two cultivated from material, steps varied from four salinity lasted about one month. Small test J. without any apparent regularity. it Resuming penetrates 3.2. can into be stated that marine and V. compacta is a true brackish species, which into freshwater environments. especially variation Morphological 3.2.1. Variation in antheridium characters In material from different localities tinuities could be found in size of Oggel value salinity ments were 140 364 - (mean examples sometimes three terminally plantation constant (fig. in For a character the following figures 128) conclusion [x; filaments: 25-70 lie within these the of this at least was are more material the two are be drawn that this is a in in measure- variation. obtained: oogonia; All jx. sizes - 224 given in of however, some & Vroman 1968) antheridia in brackish shown infig. 7. In freshwater ; only together occasionally on relatively than three antheridia (fig. 7d). forms fifty overlap (Simons observed implanted Clusters of freshwater salinity ranges. clustering never a until sufficient of antheridia; 72 (mean 40) earlier paper phenomenon clustering 7e). an sympodial antheridia culturing must habitat, were length This way in both the brackish and freshwater material while medium with showed considerable (mean different localities in ten value of the each discon- no range, oospores and antheridia. cultivated in was salinity observed. In to abundant lateral branches only salinity the form and arrangement of the antheridia were drawn was material such 98- 168 x greater part features material. A few and measurements considering attention mean appeared. diameter of ;x; literature for the interesting the sample The characters (mean 226) 135) When to structures taken. up all his Summing Each range. corresponding reproductive whole measured these organs in material from (1971) the whole the over filaments, oogonia, was are two short formed of antheridium imobserved different salinities. to remain Therefore the non-modificative character. VAUCHERIA COMPACTA: EURYHALINE ESTUARINE feature important Another the especially A be to number of pores. terminal pore and mostly the number of lateral pores can pointed amount five to figs. see V. compacta the number of lateral pores is two, while in freshwater material there is at all. On classes the and pores and fig. the classes than are 1.2, 7. 1.0 and Clustering oogonium, in brackish antheridia water Fig. diagram 1.1 taken a-c: clustering material; d: in together, with two terminal antheridia 8. Form variation two samples, than none 1.2:2 lateral from localities sample from each classes. or antheridial following 1.1:1 lateral pore, mentioned pore more lateral pore, one the material of sometimes or The were result is are clearly larger (mostly localities the classes together. In freshwater twice 1.0 and or more) 1.1 together material, for example and form cluster from of e: a of symfresh- oogonium. of antheridia. than two lateral pores. one in brackish material. For 7 and 8. In brackish In cultivated a-e; antheridia spermatozoids accompanied by plasma expulsion; more with However, of antheridia freshwater material; sample;b; general form papillae. shows that in material from freshwater localities brackish/marine of antheridia conical 100 antheridia above smaller than 1.2 and 1.3 of podium 9. This and that in clearly Fig. in the grouped - provided usually only lateral pores. range, 50 salinity antheridium, always (fig. 8g) mostly 1.0:0 lateral pores, (or more) whole investigated shown in discerned: 1.3:3 the along on basis of the number of lateral pores be can is the form of the lateral pores two variation in antheridia morphological at An antheridium is one or 621 ALGAL SPECIES of freshwater f-h: antheridia material; d: liberation of brackish material of with 622 J. dot ed brackish The estuaries. the of range salinity whole the along samples from clas es antheridal of under cultivated were localites these of samples when distrbuion frequency the to apply distrbuion blocks frequency with Diagram 9. Biesbosch and Grathem the in Fig. lines conditons. SIMONS VAUCHERIA A COMPACTA I from Grathem (non-tidal in class belonging but conditions, into There does not 3.2.2. To such the or be lowered an extent as to to seem be a of the salinity the classes case the material under brackish the change quantitative the although the 1.0 + 1.1 less are question than twice salinity values by were Lek Krimpen a/d the three methods tested; The Waarde material the clonal material sexual following takes In C1“). the clonal - 00 Cl", circumstances tested, and 1.1 are much upon different two In the terial values and clearly than 1.2 + tested, aplanospore On there are no lasted from experiments Krimpen material. in tables 1 and 2. As up : sexual was salinity to the in the reproduction Cl”, contrary to wild of salinity (0.5- 16°/ 00 range material where is vegetative growth (table 2) does not optimal under vary much under all mentioned in from fifteen records reproduction only difference 1.0 + table 2 of the values in (five salinity aplanospore rates being again are material are the material, not only salinity the but also with the which is a rate based : of sexual one to with value ma- those reproduc- under all the classes case Lek conditions. The constant shows the freshwater pattern was Krimpen a/d almost similar seawater remain com- salinity 1.0+ 1.1 are the fifteen values (at 8°/ 00 Cl") from very strong indication for the constancy of V. compacta. dulcis above mentioned data the different salinity broader values of the 1.3. This material, of the morphological The be made which is lower under Description of variety account Cl". (wild) 8°/ 00 Cl~). this character in the freshwater 3.2.3. Variation in mentioned in 3.1. Five 1.3. The values compiled the material, obtained from the wild the 1.2 + Growth and material, collected shows the brackish pattern: the classes of the antheridialclasses does larger 0 a material while the 5 and freshly tables I and 2 the results of the freshwater tion of the wild much (at presented. of the Waarde position are media), records same are can a matter studied in cultures which is in accordance with the wild material. The clearly smaller than wild Waarde material three over as summed are of the antheridium classes composition were 4f months for the 12°/o place aplanospore 12°/ 16°/ 00 to remarks from 5- occurs reproduction salinities from 5 experiments c (fig. 1.3. of the localities Waarde aplanospores b and 12 and 0.5, 5, 8, results of these 1.2 + as ranges (tidal freshwater). a, months for the Waarde material 2| large as has been done both with material and with clonal material from and material situation whether implantation and form of antheridiais with different salinities. This obtained these antheridial oligohalinic brackish of environmental modification or not, these characters (brackish) freshwater pattern typical relation between environment, Variation of antheridial characters in answer the number of antheridia Biesbosch, by culturing transition between the freshwater and the some In this 9). freshwater) can (fig. 9). one characters and shows 1.0 not to brackish a 623 EURYHALINE ESTUARINE ALGAL SPECIES conclusion must be drawn that “races” within V. compacta, but that there entities which differ from each other in two are two antheridium char- 624 J. Table 1. Summary of results and of the freshwater The results are of symbols: : presence ous, three: and spores V. compiled +: poor from compacta from the growth, from good, + + reproductive numerous, four: series of the brackish test Krimpen a/d data obtained + +; of male and female very salinity abundant, in +: three very organs. c?: the V. Lek. Salinities compacta from Waarde expressed are different media good, One + + + symbol: same, but only SIMONS +; as °/ 00 Cl rich. some organs, antheridia. two: o: some numer- aplano- so on. 0.5°/oo o 5°/oo 00 ++++ ++ + + oo 12°/oo 16°/oo + + + + + + Waarde wild material : + + + vegetative growth sexual reproduction apian ospores clonal S mm ooo o m oo o o material : vegetative growth sexual +++ + reproduction + + + aplanospores +++ <M - o OO ++ <5 oo - OO o Lek Krimpen a/d wild material ; vegetative growth sexual + + aplanospores clonal + + +++ + + m m m ? o o o o oo material: + vegetative growth sexual Table 2. The figures ++++ o distribution between o of antheridium classes +++ ++ <?(?(? - o Frequency medium. ++ ■* reproduction aplanospores were + + + reproduction <? ooo at different brackets relate to the number ooo salinity of records o values of culture from which the values compiled. Antheridium 1.0 classes 1.1 1.2 1.3 Waarde wild material: clonal material : wild material clonal namely The to the 71% 5% (15) 24% 70% 5% ( 2) 42% 46% 12% (15) 42% 44% 14% ( sympodial arrangement the brackish entity. account pacta 20% 1% type material of which antheridia belongs tinct : material : acters 4% Lek Krimpen a/d so occurrence dulcis var. entity in freshwater nov. (fig. shown by that the freshwater form As taxonomic rank of this of its var. form, and the number of lateral are 7b, d, this e; the rank of variety fig. 8a-e). is be named variety named: Latin is O pores. Christensen can . (a, b, c). Explanation (1952), as a proposed. Voucheria diagnosis: A disOn mm- typo VAUCHERIA 10. Fig. COMPACTA: A Morphological lateral common non V. compacta can a monoecious lateralibus plerumque a: poris in disposita dalcis var. be found in sympodiis prefers oligohalinic with that in the variety recorded, was while in the normal brackish form var. the Paris record (1.0:50%, b: oogonia two on one are 1 vice 2 aut plus aut plusquam 2, aut interdum disposita in quam 3. either tidal be it in a less among others: non-tidal. or pronounced Z. Flevoland. It is form. Molenbeek Grathem, interesting to note Z. Flevoland which has been reclaimed from freshwater the It is assumed that example 0 habitats, well, as slowly running water), polder polder dulcis character, non freshwater waters Non-tidal records for the Netherlands (brook filament; lateralibus brevibus lateralibus antheridiis sympodiis It aberrations, 625 ALGAL SPECIES branch. differt in antheridiis antheridia EURYHALINE ESTUARINE dulcis the was as outside the occurs with polder Lauwerszee, recorded a a Netherlands well, as showed the antheridialcharacters of clearly brackish pioneering alga. as var. for dulcis 1.1:44%, 1.2:6%). 3.2.4. Aberrations At some occasions sample. This calities : Saaftinge, aberration monoecious filaments has phenomenon was so De Heen and first in seen 1968 Terschelling (see and, curiously, cious filaments the antheridia and an antheridium and (fig. oogonia oogonium originate map.,fig. the a dioecious 1). stand apart, but same lo- At De Heen this also in 1973. On such mostly on within from three brackish monoe- occasionally sympodial lateral branch 10a). Another one observed were far been observed abnormality lateral branch planted on reduced to be pointed at (fig. 10b) suggesting is the that the occurrence of oogonia two single oogonia are on in fact im- sympodia. ACKNOWLEDGEMENT The author is manuscript, greatly indebted to Mr. G. W. H. to Dr. M. van den Vroman for his advice and the critical Berg for the drawings, to Mr. A. P. van reading Beem for of the general 626 J. assistance, Mr. to calling attention A. for correction Dop J. to the Vaucheria growth in of the text, and English the Lauwerszee Drs. to W. SIMONS Joenje for polder. REFERENCES (1961): K. Behre, & J. L. Blum, marshes. Blum, J. T. & Vaucheria Christensen, R. T. T. F. S. Dangeard, 105 notes Studies Le New or noteworthy Vaucheriae distribution (1958): Description, on from North America. on the ihrer Zufliisse. from New England salt Algae, II. Rhodora ecology of three species of 1. A list of finds from Denmark and Bot. Tidsskr. 49:171-188. species. 2: 11-14. Vaucheria, spécialement genre and Rhodora 60:283-288. genus Vaucheria submarine on some (1939): Beriicksichtigung 395-401. unknown (1900): Notes P. (1953): (3): Wilce (1952): Unterweser unter Bremerh. 7: 71-263. Conover previously England with Collins, Algenbesiedlung der Meeresf. Biol. Bull. L. J. Die Inst. Veroffentl. dans la du région sud-ouest de la France. Le Bolaniste 29:183-254. F. Kötter, Suppl. Nienhuis, P. H. lingen, a Nienhuis, with A. & in R. J. Simons (1967); A. 3 (1956): Rosenvinge, L. on im Tidegebiet der on Unterelbe. Arch.f. Hydrobiol. Vaucheria oecologie en in the delta Neth. J. Sea and species periodicity. their of flats and salt marshes Netherlands. van Res. some other Acta. Bot. Neerl. Vaucheria 20 compacta subject, Vrije Universiteit, area of the rivers Rhine, 5(1): algae in the Greve- 20-49. on a Dutch salt ( 0:107-118. in Z. W. Nederland. Amsterdam. Meuse and Scheldt. Neth. 575-597. Zur Kenntnis K. communities doctoral research Isohalines (4): algal (1971); notes (1971): Morphologic J. Sea Res. Rieth, The benthic the South-Western ecological Typewritten report Peelen, Pflanzengesellschaften (1970): sea-arm P. H. marsh, Oggel, Die (1961): Bd. XXVI: 106-184. (1879): halophiler Vaucherien Vaucheria sphaerospora I & II. Flora Nordst. v. 143: dioica 127-160, n. var. 281-294. Bot. Not.: 190-191. Simons, in J. (1974): Vaucheria birostris the Netherlands. Simons, J. & M. Vroman Acta Bot. Neerl. 17 Taylor, Sc., W. R. (1937): n. sp. Acta Bot. Neerl. 23 (6): (1968): and (4); some further remarks on the genus Vaucheria 399-413. Some remarks on the genus Vaucheria in the Notes on North Atlantic Marine Algae I. Papers of the Mich. Acad, Arts and Letters 22:225-233. Webber, E. E. Netherlands. 461-466. (1968): New England salt marsh Vaucheriae. Rhodora 70: 274-277. of
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