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The Sea-Grasses of the World

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The Sea-Grasses of the World VERHANDELINGEN DER KONINKLIJKE NEDERLANDSE AKADEMIE VAN WETENSCHAPPEN, AFD. NATUURKUNDE TWEEDE REEKS, DEEL 59, No. 1 9 THE SEA-GRASSES OF THE WORLD C. DEN HARTOG NORTHHOLLAND PUBLISHING COMPANY AMSTERDAM,, LONDON 1970 GEOGRAPHICAL DISTRIBUTION ORIGIN, EVOLUTION AND OF THE SEA-GRASSES treatment meant to be a taxonomic this paper is primarily such Although aspects will be discussed, Angiosperms, some other of the marine origin, fossil record and for life in the sea and the as the requirements ecological aspects I of these plants. For the geographical distribution the ecology of paper (DEN HARTOG, 1967) on want to refer to my earlier and succession of which growth-forms, zonation sea-grass communities in ecosystem structure and function of their the sea-grasses as well as the were discussed. a. Requirements for life in the sea. four properties which are indis­ According to AjBER (1920) there are it must be adapted pensable for a marine water plant. In the first place, it must be able to grow when complete­ to life in a saline medium. Secondly, system at its disposal an anchoring Thirdly, it must have tidal cur­ ly submerged. withstand wave action and sufficiently developed to which is for hydrophilous pollination, it must have the capacity rents. Finally, to failure, except in halcyon method must be doomed since "any aer-ial plants must be able to sea". In other words, the weather in a non-tidal cycle in a saline me­ as well as their generative achieve their vegetative securely anchored in subncrged, and must be dium, while completely It is obvious that sea­ attached to the substratum. the substratum or be they possess more first two properties. Furthernmore, grasses possess the gregarious growth; this and show a tendency for or less strong rhizomes the fourth Finally, all sea-grasses satisfy fulfills the third requirement. cd for hydrophilous pollination. requirement, as they are well-cquiP Adaptations takes place in most genera. Entirely subunerged pollination and Halodule, are the long styles of Halophila type of pollination d i n and Thalasso­ to this Syringo w Amphibolis the long stigmata of Cymodocca, and the disc­ of Thelassia and lalophila, dendroni, the elongate hypanthia the confervoid stigma of Posidolia.Moreover, shaped, lobate or laciniate of the Potamoge­ of all marine representatives shape of the pollen-grains their floating ca­ as an adaptation increasing may be regarded have spherical tonaccae ilalophila and Th(lassia pacity. The llydrocharita(,ol!s moniliforni chains ark arranged in coherent, pollen-grains, but as these a similar result is reached. on the wat­ whereby the pollen drifts Semi-aquatic surface pollinatiun,l with the water are at least partially in contact, er surface and the styles although entirely for Phyllospadix and Zostera, surface, has been recorded just.as often in these genera. submerged pollination occurs TIE SEA-GRASSES OF THE WORLD 13 Among the marine Angiosperms Enhalus acoroides is the only species which shows aerial surface pollination. Hereby the pollen as well as the styles remain dry. The species is further peculiar in having large globular pollen-grains and by the fact that its male flowers break off in the spathe and rise to the surface, where they have a short but independent exist­ ence. The latter featare is also known from some other Hydrocharitaceae, e.g. Vallis)i2ria species and Elodea nuttallii (Planch.) St John, and is also shown by Lepilacna australis Drummond ex Harv. in Hook.. a rep­ resentative of the Potamogetotiaceae. The flowering biology of Enhalus is completely adapted to the tidal cycle. Most sea-grasses are dioccious and those which are monoecious show proterogyny (Zostera, Heierozostera, Halophila decipiens). Thus cross­ fertilization is a rule. Although all sea-grasses satisfy the 4 criteria of Arber, there are a few other taxa which also fulfill these conditions, viz. Zannichellia,Lepilaena, Althenia, Ruppia and Potatnogetion subgen. Cologton (with P. pectinatus L.), all of which are representatives of the Potamogetoaaceae.The range of salinity which they can tolerate is even greater than that of the sea­ grasses, as they are to be found in fresh water, in mixohaline and hyper- Ialine brackish waters and in continental salt waters. As far as the chem­ ical composition of the salt water is concerned they are not quite so restricted as the sea-grasses. Iuppia, for example, occurs not only in waters where NaCl is the dominant salt but also in waters where Na2 SO 4, .1lgSO 4 or CaSO. dominate. Zawnichellia and Polamoyclon pectinatus have been recorded friom waters where Na 2C0 3 is the dominant salt. Furthermore, ;ll these taxa can tolerate very sudden and very large fluctuations in the salt content. From an ecological point of view they are in fact ubiq­ nitous. These extremely euryhaline taxa, nevertheless, seldom penetrate into the purely marine environment, although their salt tolerance cer­ lainly" does not prevent them from doing so. They sometimes occur to­ _, lhe:' with the most. eur'haiine sea-grasses in brackish waters, such as estuaries and lagoons, but are generally restricted to l)oikilohaline waters. The fact that these taxa. in spite of their enormous tolerance with r'slp'ct to salinity, are apparently unable to inhabit the marine environ­ iniit permanently must be ascribed to a low competition capacity. It is probably a basic rule in ecology that a wide tolerance with regard to ('uv'ironnental fluctuations is coupled with a reduced capacity to compete with i more stenobiontie taxa inl more or less stable habitats. i.(tiqin (4 tIc ialrine A)giosperiiis In her dist ussionj of the re(Iiuii'ements which have to be fulfilled by I,!llts in order to be able to thrive in the marine environment, ARBER (19t20) followed tle then prevailing opinion that the marine Angiosperms lad penetrated the sea fi'om the fresh water. This opinion is still held by sonie authors (SCULTHOIPE, 1967). It is based on the following facts: 14 TIHE BEA.GRASSES OFrnE WOULD 1. Both the Potamogeonaceae and the Hydrocharitaceaeare typical water­ 2. Not a single widely distributed in fresh water. plant families, and areis closely related to a terrestrial plant. .marine Angiosperm evolved that 'h sea-grasses have indicate, according to Arber, She These facts adapted to life in the water. ancestors which were already from water to sea, water may the transition from fresh even suggested that freoh-vater species -+ salt­ in the following sequence: have been bridged species -- marine species. species -- brackish-water to have tolerant fresh-water are supposed by her and Hydrocharitaceac The Polonwgdonaceae other families, e.g. the stages of this series, while passed through all the the Raminculaceae Callitrichaccae, the Xaiadaceae, Ceratophyllaceae, the producing a few salt- not go further than Halorrhagidaceac(lid and the her essay with the follow­ spepies. Arber concluded toleranting optimistic fresh-water expect atio: "Conceivably in future ages, if the evolution present lines, a greater number may of fresh-water pla;nt- proceeds on its u and some of these of hydrophilo s pollination, reach the specialised stage and exclusive circle democratising the narrow may colonise the sea, thus Angiosperms are diametric­ of the marine Angiosperms." t arine s a d t the origin of the mi My own ideas about templing to regard the opinion. Although it is ally opposed to the above of the as a side-track in the development small group of the sea-gresses arguments which can plants. there are several large group of fresh-water the Potanlogco the, opposite. The fact that put forward to prove just fresh water may be are mainly distributed in and the Itydrochariacea' i picture arises aceae a corl ctely different species level; h(wever, be true at belonging to these the distriltion of the subfamnilies and when one considers is rather heterogeneous the ]Iotamogeldo1aceae families. The family of the Three of these, the Zosteroidae, into fi suhfamllies. are com­ can be subdivided with together 9 genera, and thep Cymodoccoid(a(u RUppioidWca Posidonioidae with 3 genera and thit' marine. The Zalnichllioidea( unable to thrive pletely to a high degree, but are with 1 genus are salt-tolerant with 2 genera The I-Jotamogcho)ioid:ae undler marine (.ond;t ions. t permanently of P1offlio oiL subgen. fresh water, with tile exeeption are restricted to to that of the Zamnichd­ an e(eologiacll range similar ColeogCIon, which has and Ruppioidewu. of which lioideae consists of 3 subfanilies of the IHydrochalactaa are coin­ The family both with one genus, and the llalophiloid(ac, the Thalassioidea l7allisnCr'oideaconly the 13 gen(,ra of the subfamily pletely marine. Of undisputedly fresh­ tile otiler twelve arc genus, Edhahl,. is niarhw; one sailt-toleant rcprescntative. genera with an ocazional Fossil re­ water from the fossil records. argument (an be advanced the works of Another have been described in referred to as sea-grasses, (1878), mains, (1886), SAPORTA &' MARION (1828, 1849), ]3VUIlEAU names BRONGNART etc., mainly under the 1851), WA'rELET (1866), UoER (1847, greater part in Aseherson's They are listed for the Zostcriles and Caulivihs. TIM SEA-GRASSES OF THE WORLD 15 Monograph of the Polamogetonaceae (1907, p. 33, 39, 151). SCHENK (in ZITTEL, 1890, p. 379) rightly remarked with respect to the fossils described as Naiadaceaeand Potaniogelonaceaethat these families Lre "in demselben Verhiiltniss reich an Arten, als diese fraglich sind, sowohl hinsichtlieh ihres Werthes als Arten, wie ihres Werthes als Glieder der Gruppe." While admitting that a part of the material had been correctly identified, lv remarked that many of the fossil remains are too fragmentary or too Imidly preserved to allow even their recognition as monocotyledons. He sumgested the possibility that sonm remains described as pine-needles or as Poacitescould just as well be remains of Naiadaceaeor Potanmogetonaceae.
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