The most complex seaweeds are composed by holdfast(s), stipe(s) and frond(s).

A
typical example of this morphology is the genus Sargassum (Fig. 41A). The function
of the holdfast is solely attachment (as opposed to roots in higher plants which also
play a role in extracting water and nutrients from the soil). It can be rhizoids (thin
filamentous structures: Caulerpa spp., Fig. 41B). In Avrainvillea erecta (Fig. 41C)
and Halimeda macroloba, these filamentous structures get intricated and hold large
amounts of sand, resulting in a ‘bulbous holdfast’ which is completely sunken in the
soft substratum. Attachment can also be performed by a disc (most Sargassum spp.,
most red algae, Fig. 41D). The stem-like portion (stipe) of the thallus can be cylindrical
or compressed, unbranched or branched, supple or rigid. It bears one or several
blades (the genus Sargassum, Fig. 41G) which are wider than the stipe and are the
main photosynthetic part of the seaweed. At the basis of the stipe, horizontally spread
branches can be present (stolons or rhizomes, Figs 19D, 41E, F), spreading across
the substratum, possibly attaching to the substratum again and giving rise to new
uprights. In some species (Sargassum) the uprights bear air bladders (Fig. 41H) as
‘floaters’, to keep the plant upright and optimalize the surface for photosynthesis.

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 Fig. 41. Seaweed morphology: holdfasts, leaf-like structures, air bladders. A. Sargassum sp.:
a thallus with holdfasts, stipes and blades; B. Caulerpa sertularioides with prostrate rhizomes
attached by numerous rhizoids; C. Avrainvillea amadelpha with a bulbous holdfast composed
of intertwined filaments; D. Halymenia durvillei: discoid holdfast; E. Turbinaria sp. with stolons;
F. Gracilaria corticata, with basal stolons; G. Sargassum crassifolium with leaf-like blades on
the stipes; H. Sargassum crassifolium with air bladders (aerocysts).

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 The growth direction of seaweeds can vary: most are erect (Dermonema virens,
Fig. 42A), at least when they are submerged. Others grow horizontally and mostly
have numerous attachment points to the substratum (Dictyota sp., Fig. 42B): they
are prostrate. Some are horizontally spread in the basal portion, but upwardly curved
towards their apices (Halimeda gracilis, Fig. 42C): they are ascending, or downwardly
curved: they are arcuate (Valoniopsis pachynema, Fig. 42D). Others again are
horizontally spread from a vertical wall (Peyssonnelia spp., some Halimeda spp.
Fig. 42E): they are resupinate. Finally some seaweeds hang down from vertical or
overhanging walls (some Halimeda spp., Fig. 42F): they are pendulous.

Fig. 42. Growth forms. A. Erect: Dermonema virens; B. Prostrate: Dictyota; C. Ascending:
Halimeda gracilis; D. Arched branches of Valoniopsis pachynema; E. Resupinate: Halimeda
sp.; F. Pendulous: Halimeda sp. hanging down from an overhang.

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 Another vegetative character that can be used in some groups of seaweeds is the way
of cell division. In most cases, the apical cell undergoes a transverse division, the
daughter cells grow longitudinally, elongating the main axes. A successive inclined
division at the apical pole results in a lateral branch. If this cell division process is
repeated, the result is an acropetal organization of the thallus: the side branches are
progressively longer from the apex to the basis (Fig. 43A). In other taxa, intercalary
cell divisions occur: older cells undergo cell divisions. This results in a non acropetal
organization of the thallus: longer side branches alternate with shorter ones (Fig.
43B). In other green algae, the formation of a transverse wall at the basis of the side
branch is delayed (Cladophoropsis sundanensis, Fig. 43C). In some green algae (the
genera Siphonocladus, Struvea, Dictyosphaeria) a special kind of cell division occurs,
called segregative cell division. A multinucleate protoplast divides into several, rounded
daughter protoplasts within the mother cell (Figs 43D, E), which subsequently become
surrounded by a wall (Fig. 43F). The newly formed cells are either released after
rupture of the mother cell (Valonia ventricosa), remain in situ and form parenchymatic
thalli (Dictyosphaeria spp.), or rupture old parental walls and form branches (Struvea
spp., Siphonocladus spp., Figs 43G-H). In the genera Ernodesmis (Fig. 43I) and
Valonia, small, lens-like cells are formed at the apex of the mature cells, growing out
to new cells.
A major problem in describing or identifying seaweeds is their morphological plasticity.
Depending on the ecological conditions, the same species can become larger (in a
sheltered lagoon) or smaller (on the seaward, surf-exposed rock wall), less or more
densely branched, plane or spirally twisted, without or with hook-like branches. An
extreme example is the Caulerpa racemosa-complex, where on the same stolon (thus
the same individual) the erect branches can have a different morphology from the
proximal to the distal part of that stolon. Sometimes the side branchlets of a single
upright can be different from the basis to the tip, being cylindrical at the basis, club-
shaped higher up, becoming turbinate or even peltate at the tip. As the morphology of
these side branchlets has been used in the past to describe taxa (species, varieties
or forms), the presence of a mixture of morphologies creates major identification
problems. Other seaweeds change their morphology by ageing or show sexual
dimorphism (the genus Sargassum, Boodlea composita-Phyllodictyon anastomosans
complex).
On the other hand, molecular systematics frequently points out to ‘cryptic diversity’:
seaweeds with a similar morphology appear to belong to different taxa, based on
the DNA-information. As a result, new species will have to be described, preferably
with (at least) a distinguishing morphological or anatomical character or a different
geographical distribution (the different taxa being present in different oceans).

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 Fig. 43. Cell division. A. Acropetal organization: the branchlets gradually become more
developed proximally; B. Non acropetal organization, with short branchlets alternating with
longer ones (Cladophora sericea); C. Postponed transverse cell wall formation after the
formation of a lateral branch (Cladophoropsis sundanensis); D-G. Segregative cell division
in Siphonocladus sp.; H. Final stage of segregative cell division: numerous side branchlets
growing out of the parental cell; I. Formation of apical lenticular cells from where new cells
grow out (Ernodesmis sp.).

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 8.4. Life histories and reproduction
Life histories in seaweeds are complex; moreover they vary among and even within
groups. Therefore only a general scheme can be given here, although characters of
the reproductive structures can be critical for the identification on species or even on
genus level. In most green and brown algae there is an alternation of two generations:
the haploid gametophytes and the diploid sporophytes. The gametophytes produce
gametes in gametangia, specialized structures which, in general, can only be observed
by microscope. In several brown algae however, where reproductive structures are
often grouped in sori (Dictyota sp., Figs 44A-D) or in receptacles with gametangia in
Sargassum sp. (Fig. 44E) and Turbinaria sp. (Fig. 44F) which can be observed with
the naked eye. The male and female gametangia are mostly produced on different
plants, but in some cases they are both present on the same plant. The gametes
will fuse and produce a diploid zygote which germinates into a diploid sporophyte.
On the sporophyte, meiosis takes place and haploid spores are produced (Dictyota
spp., Figs 44G, H), developing into new gametophytes. In some rare cases (Codium
spp., Caulerpa spp.), the life cycle is reduced to a single diplont generation, the only
haploid stages being the gametes. Moreover, in the genera Halimeda, Caulerpa and
other green algae, the whole cytoplasmic content of the thallus is being transformed
to gametes (= holocarpy, Fig. 44I).
In red algae, the life history is even more complex by the addition of a third generation:
fertilisation of the female gamete (carpogonium) attached on a carpogonial branch, is
performed by a male gamete (spermatium), produced in a spermatangium (Fig. 45A);
spermatangia can be grouped in sori (Fig. 45B). The diploid zygote remains attached
to the haploid female gametophyte and develops in a diploid carposporophyte. This
part of the life history (a generation) usually has only small dimensions, but generally
visible with the naked eye, as globular structures, called gonimoblasts (Fig. 45C) or
as lateral, ball-like structures, called cystocarps (Figs 45D-F). In some cases, the
cystocarps are embedded in the thallus and therefore more difficult to see in the field.
The carposporophytes produce diploid carpospores which germinate after liberation
into tetrasporophytes in which meiosis takes place with the production of haploid
tetraspores (Fig. 45G) which in some cases can be grouped in stichidia (Fig. 45H) or
in sori (Fig. 45I). The tetraspores germinate into haploid gametophytes. In most of the
red algae, the life cycle thus consists of three generations, of which the gametophyte
and the tetrasporophyte are often (almost) identical. In some cases (Asparagopsis
taxiformis), the tetrasporophyte (Falkenbergia hildenbrandii) is markedly different
from the gametophyte (Fig. 45J). In the past, both generations of that seaweed have
been described as different algae, placed in different genera, as phycologists then
were unaware of the fact that they represent two phases of the same seaweed. It
is only after culture experiments in aquaria that this was discovered. In some brown
(Sargassum) and red algal genera (Liagora) one of the phases can be microscopic
or crustose.

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 Fig. 44. Reproduction structures in brown and green algae. A. Sori of female gametangia
(oogonia) on the haploid gametophyte of a Dictyota; B. Detail of a sorus of oogonia of a
Dictyota; C. Sori of male gametangia (spermatangia) on the haploid gametophyte of a
Dictyota; D. Detail of a sorus of spermatangia of a Dictyota; E. Receptacles of a Sargassum,
containing the gametangia; F. Receptacles of a Turbinaria, containing the gametangia; G.
Tetrasporangia on the diploid sporophyte of a Dictyota; H. Detail of a cruciately divided
tetrasporangium of a Dictyota; I. A Caulerpa-plant in which the formation of gametes is taking
place (the yellowish part of the thallus).

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 Fig. 45. Reproduction structures in red algae (mainly shown by African examples).
A. Spermatangia in Sciurothamnion stegengae; B. A sorus of spermatangia in Platysiphonia
delicata; C. Gonimoblasts (groups of diploid carpospores) in Sciurothamnion stegengae;
D. A cystocarp on Rhodomelopsis africana; E. Cystocarps as wart-like protrusions on
Gracilaria corticata; F. Cystocarps (mainly) on the margin of the female blade of Sarcodia
montagneana; G. Tetraspores in Sciurothamnion stegengae, produced after meiosis in
tetrasorangia on the diploid sporophyte; H. Tetrasporangia in a stichidium of Platysiphonia
delicata; I. Sori of tetrasporangia in Augophyllum marginifructum; J. Asparagopsis taxiformis:
the large gametophyte with cystocarps and the filamentous tetrasporophyte (Falkenbergia
hildenbrandii) in the centre (arrow).

Reproductive structures, or even the presence of a particular life history phase, are
generally seasonal. It is therefore imperative to carry out collecting in different seasons
as reproductive characters are mostly needed for correct identification (as flowers are
in higher plants).

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 Many seaweeds also reproduce asexually (without formation of gametes), by the
production of asexual spores. Some even multiply vegetatively, by fragmentation
(some branches break off, stay alive, attach to the substratum and go on growing
to new plants) or by production of propagules (Sphacelaria spp.: branchlets with a
special morphology, detaching from the mother plant and each of them producing
a new juvenile; Fig. 104). Others again, growing in soft substratum, can produce
underground, horizontally growing bundles of rhizoids from which new erect plants
develop (genera like Udotea, Halimeda, …).
8.5. Biodiversity of seaweeds
In most biodiversity studies the algae are omitted, probably because they are
‘invisible’ as a result of their submersed habitat. The total number of species of algae
is difficult to assess: the important environmentally induced morphological plasticity
and variability results in major identification problems: some entities are classified on
different taxonomic levels, depending on the author (species, variety, form). The total
number of Algae [including (freshwater) microalgae] would be approximately 350 000
spp. (Brodie & Lewis, 2007).
Some areas in the world are more species rich than others. In the Pacific Ocean,
species-rich areas are the Philippines and Japan; in the Atlantic Ocean: Europe
(N-Spain, France, United Kingdom), the Caribbean Sea. The Red Sea and the Indian
Ocean are still understudied, but South Africa and southern Australia seem to have
a high seaweed diversity. In South Africa this could be the result of the presence of
different climate zones.
Maximum seaweed endemism is present in Antarctica, southern Australia and New
Zealand.
Based on data from the literature, Silva et al. (1996) mention 455 taxa belonging to
410 species and 161 genera for Sri Lanka. However, as already mentioned in the
chapter on the history of seaweed research in Sri Lanka, this island is absolutely
understudied. Historical collections from sublittoral biotopes are sporadic, and recent
ones are still under study. A study by Mallikarachchi (2004) shows that a large
percentage of the known seaweed flora of the island is present along a limited SW-
shoreline. This conclusion is partly biased as this part of Sri Lanka has been most
frequently visited by phycologists, whereas collections from the N and East coast are
scarce and therefore even more fragmentary.
Smaller species (such as turf-forming algae and epiphytes) are more numerous,
adding to the species diversity, but they are not readily observable/identifiable and
therefore most of these are not included in this Field Guide.

8.6. Nomenclature, taxonomy and classification of seaweeds

The nomenclature of algae (giving scientific names to organisms and groups to which
they belong), similarly to higher plants, follows the International Code of Botanical
Nomenclature (ICBN, 2006). Macroscopic seaweeds belong to 4 Divisions (or Phyla)
if the blue-greens are included: Cyanophyta (Cyanobacteria) - Blue-green algae
(prokaryotic), Chlorophyta - Green algae, Phaeophyceae - Brown algae, Rhodophyta
- Red algae. The divisions are subdivided in classes, which names end on -phyceae
(exclusively for Algae). The classes contain orders, ending on -ales, subdivided in

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 families, ending on -aceae. The nomenclature in botany (including flowering plants,
ferns, mosses, algae and fungi) is binomial, meaning that the name is composed of
two words: the genus name (e.g. Rhodymenia), with a capital initial followed by the
species epithet (e.g. Rhodymenia tripinnata) written in lower case. The genus and
species names are usually written in italics. They are followed by the name of the
author who described the species (e.g. Rhodymenia tripinnata Hering). Sometimes,
further research shows that the original author placed the species in a wrong genus.
In this case, the name of the first author is placed between brackets and the name of
the author who makes the new combination (putting the species in the correct genus)
is added (e.g. Portieria tripinnata (Hering) De Clerck). In some species, subentities
(infraspecific taxa) are distinguished which are called varieties (var.) or forms (f.), the
names of which are again usually written in italics.
When proposing a new species, a type specimen is designated after which the
species is described. For seaweeds this is generally a herbarium specimen, which
is then deposited in a registered herbarium. Quite frequently isotype specimens are
deposited in other important herbaria. These isotype specimens were collected at the
same type locality (place where the type specimen is coming from), on the same day
as the type specimen and were regarded as ‘duplicates’ by the original author (= form
part of a single collection). Type specimens are extremely important for subsequent
studies of the species (checking for new characters, for DNA-analysis, …). Preferably,
several specimens should be mounted on a ‘type sheet’, with the indication of the real
type specimen, the holotype, as to show the morphological variation of the species
(sometimes gametophytes and sporophytes are (slightly) different, or different
ecological situations induce a morphological change).
The description of a new species of seaweed has to include the reference to the type
specimen as well as a diagnosis in Latin (what are the characters of this species,
distinguishing it from other species of the same genus). Illustrations also have to be
added.
Subsequent analysis sometimes indicate that two ‘species’, each with their own name,
described from different areas (even from different oceans) are identical. Only a single
name can be applied for that species, and the name from the oldest description has
to be chosen; the other name then becoming a synonym. Opposite to this, molecular
analysis sometimes proves that a species, present in different oceans (with a similar
morphology at the different locations), belongs to different species according to the
locality (cryptic species). The specimens from the type locality then keep the original
name, whereas the other ones have to be described as new species. A thorough study
of morphological and anatomical characters then ‘hopefully’ leads to discriminating
characters for each species. All this means that names of seaweeds change in time
and that the same taxon can have different names in different books, depending on
the time of publication. If one is compiling a species list from a region, he should be
aware of these synonymies for not including the same species several times under
different names.
As a result of ongoing molecular research, the higher rank classification of seaweeds
also changes on a regular basis. We here follow the Index Nominum Algarum (http://
ucjeps.berkeley.edu/INA.html) and Guiry & Guiry (2009; www.algaebase.org) which
are both excellent sources for keeping up with recent taxonomic revisions as they are

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 continuously updated (see remark at introduction of Rhodophyta). Silva et al. (1996,
http://ucjeps.berkeley.edu/rlmoe/tioc/ioctoc.html) is also an excellent basis to find
synonymies, taxonomic remarks, and a systematic classification of the seaweeds of
the Indian Ocean. Be aware, however, that since 1996 a surprisingly large number of
names have changed already.
As opposed to terrestrial plants, seaweeds rarely have common (vernacular) names.
Moreover, they sometimes induce confusion, such as ‘Ceylon Moss’ which is not a
moss at all, but a red alga, Hydropuntia edulis (not included in the present book).
8.7. Identification of seaweeds
If possible, one should start by following a training course where specialists can
introduce you to the most common genera and species of the area. If this is not
possible, Field Guides on the area (such as this book) or from the same ocean or
from an adjacent tropical region should be used. They are becoming more numerous
nowadays. Useful recent guides for Sri Lanka are: Huisman (2000) on Marine Plants
of Australia, Littler & Littler (2003) on South Pacific reef plants, De Clerck et al.
(2005a) on the seaweeds of Kwazulu-Natal, Oliveira et al. (2005) on Marine Plants
of Tanzania, Huisman et al. (2007) on Hawaiian Reef Plants, Ohba et al. (2007) on
Marine Plants of Palau, Skelton & South (2007) on Samoan Benthic Marine Algae.
For the identification of red turf algae, Price & Scott (1992) is very useful. Anyway,
one should remain cautious with identifying organisms solely based on field guides:
as opposed to a real ‘Flora’ they only contain the dominant species! The possibility
that a different, closely related species was collected cannot be excluded. Therefore,
the next step is the use of (preferably recent) monographs of a group (e.g. De Clerck,
2003 on the genus Dictyota in the Indian Ocean, or Leliaert & Coppejans, 2006 on
the genus Cladophoropsis) or detailed regional publications (e.g. Van den Heede &
Coppejans, 1995 on the genus Codium from Kenya, Tanzania and the Seychelles;
Kraft, 2007 on the marine green algae of the Lord Howe Island area), as well as
comparison with specimens from existing herbaria with trustworthy identifications.
Anyway, for the identification of macroalgae on species-level, morphological and
anatomical characters are needed (e.g. in the genus Codium, measurements of utricles
have to be made; in Ulva spp., the number of pyrenoids per cell have to be counted,
…). In brown and red algae, quite often the analysis of reproductive structures is
important for identification on genus and/or species level (just like flowers in higher
plants!). The analysis of these characters can only be carried out in a laboratory,
with the use of a microscope with a calibration plate. Sterile specimens therefore
frequently remain unidentified because critical characters for species (or even genus)
distinction are absent.
8.8. Seaweed resources from Sri Lanka
Natural populations of the red alga Ceylon Moss (Hydropuntia edulis (S.G. Gmelin)
Gurgel et Fredericq) were harvested in the past for extraction of agar from its cell walls.
This seaweed is quite abundant in Puttalam lagoon, but it is not collected anymore
and not included in this guide.

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 9. Survey methods for seaweeds

For this chapter we also refer to Leliaert & Coppejans (2004); http://www.persga.org/.
9.1. Qualitative assessment of the macroalgal flora of an area
Qualitative assessment of the marine flora of a coastal area implies general collecting
in a specific area, resulting in a more or less complete list of species. Depending on
the study, the coastal area can vary from a small area (e.g. a coastal strip of 10 m, a
rock outcrop, etc.) to a large area (e.g. one to several km of coastline, a small offshore
island, etc.). When comparing species numbers or biodiversity indices of different
coastal areas, these areas should be of comparable size. The resulting species list
is important for calculating biodiversity indices of an area. A major disadvantage of
qualitative collection data is that species abundance is not taken into account. This
can partially be corrected by making the sampling method semi-quantitative. This
implies that each species is ranked based on its abundance, evaluated by visual
observations. An example of such a ranking is the Tansley scale (Table 1 in Appendix).
The growth form (sociability) of the seaweeds can also be taken into account; here the
Braun-Blanquet’s sociability scale can be used for each species (Table 2 in Appendix).
As a matter of fact, Braun-Blanquet’s cover-abundance scale is most used (Table 3 in
Appendix). These data can be added on the herbarium labels.
9.1.1. Getting ready for fieldwork
It is evident that adapted clothing (protection against the sun/rain) is needed. In this
respect good footwear is extremely important. The use of booties (tight, ankle-high,
rubber boots with a thick sole and a zipper) is advisable, as well on rocky as on sandy
or even muddy substratum, because they completely protect the feet against sharp
obstacles (barnacles, oysters, coral fragments, …). If snorkeling is planned, a (thin)
rubber wetsuit is useful for protection against sharp walls or irritating animals (jelly
fishes, siphonophores, …), or at least knee pads. The availability of a towel also
comes out handy.
The value of a report/publication on the biodiversity of an area largely depends on
the presence of reference (voucher) specimens which allow ulterior control of the
identifications. On its turn, the value of these specimens depends on the field data
which are added to them. Therefore, a notebook (intertidal work) or a white plexiglass
plate (in the subtidal and in intertidal pools) and a pencil are indispensable (Fig. 46A).
Collecting gear includes a bucket, plastic vials, plastic bags, prenumbered labels
on hard paper. Many algae and some seagrasses can be removed by hand, but a
scraper or a stout knife may be handy or even necessary. Some thick encrusting
algae can be removed with a knife, but many (especially the crustose coralline algae)
must be collected along with the substratum. This can only be done by use of a heavy
instrument such as a hammer and a chisel.

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 Fig. 46. Field work. A. Using a plexiglass plate and a pencil for taking notes in the water;
B. Collecting by wading at low tide; C. Even with a seemingly smooth sea, a sudden big wave
can emerge; D. Fully equipped for SCUBA-diving and underwater photography; E. Putting
specimens in zip-lock bags during SCUBA-diving; F, G. Sorting out specimens on the field.

If available, a camera, a map, and a Global Positioning System (GPS) can be extremely
useful. Be careful in this wet environment: put them in a watertight camerabox or
(ziplock) bags!
Intertidal habitats can be sampled by wading (Fig. 46B) during (extreme) low tide or
by snorkeling at high tide. Therefore, check the time of low tide as to get organized
for the sampling. If snorkeling is planned (deep intertidal pools or subtidal) mask,
snorkel, fins, mesh bag, plastic collecting bags and labels shouldn’t be forgotten. For

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 extensive collecting or observation in the subtidal, SCUBA diving is advisable. Next to
the snorkeling gear, a rubber suit, a belt, weights, the full air-cylinder, regulator, diving
watch, depth gauge, inflatable backpack (= BCD, buoyancy control device), and a
buoy should be brought or hired at a diving center (Figs 46D, E). These generally
check the diving license, so don’t forget to bring it.
Freshly collected specimens should be processed as soon as possible to minimize
decay. If the way back to the laboratory is long, the specimens might decay under
way. It then is preferable to prepare the collected specimens in the field (Figs 46F,
G) or to store them in a cool box. If specimens are sorted at the collecting site, bring
sorting and preparation trays, a floater, herbarium paper, a plant press with straps,
card board, newspaper and fleeces, jars (Eppendorfs) and silicagel, formalin, zip-lock
bags and hermetically closed jars (e.g. ice cream boxes).
9.1.2. Arriving in the field
Note the date, the locality (name of the closest town or village + eventually local name
of the collecting site). If you have a GPS-system: add the GPS and longitude-latitude
coordinates.
Make a general description of the site: is it a peninsula (Fig. 6B), a straight coastline,
eventually with a beachrock platform (Fig. 4), a wide bay (Fig. 3B), an enclosed bay
(Fig. 3C), isolated rocky outcrops (Figs 7C, D), an island (Figs 7B, C), a lagoon (Fig.
3D)? Describe the substratum: solid rock (Fig. 6A), boulders (Fig. 6C), sand, mud.
Rate the general coast inclination: overhanging, vertical, sloping, subhorizontal. Give
a general description of the biotope(s): seaweed vegetation (Fig. 6C), seagrasses
(Figs 12A, B), mangrove (Figs 13A-C), coral reef (Fig. 7E).
Eventually add pictures.
9.1.3. Field collecting
Extensive and well-prepared collections are the basis of diversity based studies of
(marine) organisms. The importance of good collections for taxonomic studies is
evident, but it is equally important that representative collections - often referred to as
voucher specimens - be kept of each species recorded in ecological surveys. Without
such specimens, there is little or no possibility of later checking on the basis of names
used in publications. Such specimens should be numbered, labeled and be deposited
in a recognized herbarium (Womersley, 1984).
Take the time of low tide into account, certainly if you want to collect by wading. If it is
already low tide upon arrival, go to the lowermost part first and come up with the tide.
Take care not to get encircled by water. If the tide is still going down, go down with the
tide and do the uppermost parts on your way back.
Collecting can be done by species (a single species and label per bag: numerous bags
will be needed, but sorting out will be much easier); note the field identification of each
number. Sometimes preference is given to collecting by biotope (a pool, a rock wall,
a phorophyte: several species in a single bag, but with a single label). In species-rich
areas or time shortage the latter method is being used. Always add ample seawater
in the bags as to avoid decay by temperature rise or desiccation. Also add a label,
which corresponds with a number in your note book (plexiglass plate) where you
add: the detailed ecology of the collecting site (air-exposed/submerged at low tide;

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 pool: vertical/overhanging/sloping wall/(sand-covered) bottom; epilithic/epiphytic (on
what?); the level relative to the tides (above high tide level (supralittoral); between high
and low tide (intertidal (high -, mid -, low -)); under low water mark (subtidal)). Make
notes on morphological characters which will be lost after processing the specimens,
such as growth form (isolated plants, individual tufts, gregarious, forming intricated
cushions); growth direction (erect, ascendant, prostrate, pendulous); in situ colour:
some seaweeds are iridescent when alive; some seaweeds change colour upon
drying; consistency: membranous, gelatinous, cartilaginous, stiff, brittle; eventually,
presence of reproductive structures.
ALWAYS collect several specimens as to illustrate morphological variability and to
be able to look for fertile specimens. ALWAYS collect complete specimens, including
the holdfast as this might be a character needed for identification: presence of a disc,
haptera, rhizoids, a bulbous structure.
While collecting, be aware of possible danger: even with a seemingly smooth sea, a
sudden big wave can emerge (Fig. 46C).
9.1.4. Coming back from the field
If the laboratory is far from the sea and not provided with seawater, collect a (plastic)
drum with seawater for sorting out the specimens, or … sort out in the field! (Figs 46F,
G).
9.1.5. Sorting out the specimens
If the species have been collected individually, put them in separate trays (vials) and
add the field number. If the collecting was made by collecting site, put the collection of
one bag and its label in a large tray and sort out the different species in smaller trays
(vials) giving them each a subnumber (e.g. collection from site 3: species 3a, 3b, 3c,
3d) (Fig. 47A).
9.1.6. Finally numbering and labelling the species
Copy the data from your field notebook or from the plexiglass plate on the computer
or in the final notebook: date, place, general description of the site.
Each species gets a final serial number, preferably preceded by the collector’s initials
(e.g. HEC = Herbarium Eric Coppejans). Start with 0001 and go on all of your life:
e.g. day 1: HEC 0001-0024, day 2 HEC 0025 – 0056 and so on). Add the detailed
ecological data from the field as well as the morphological data (eventually add
observations carried out in the laboratory). A HERBARIUM SPECIMEN WITHOUT A
(complete) LABEL IS SCIENTIFICALLY (almost) USELESS!
Individual labels are printed out and added to the herbarium specimens. All these label
data are introduced in a database. This way data can be retrieved by: collector, place,
period, genus or species level (over different regions, oceans); herbarium, formalin
preserved, silicagel, culture specimens.

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 The final label
Number: HEC 16128 (eventually +F, +S, +L; see further)
Name: Caulerpa racemosa (Forsskål) J. Agardh
Locality: Sri Lanka, Galle, in front of the lighthouse
Collection date: 15 August 2008
Ecology: on the sand-covered bottom of a low intertidal pool
Morphology: thallus fleshy, dark green, with starlike, slightly iridescent light green
stripes; well attached to the substratum by numerous rhizoids; stolons prostrate,
intricated; uprights with short rachis and densely set vesicular branchlets,
resulting in a grape-like appearance
Collector: Eric Coppejans
Identification: W.F. Prud’homme van Reine (+ date of identification)

9.1.7. Preparation of a herbarium specimen

- Take a tray and fill it with clean SEAwater;
- Put a (cork)floater in the water (or an inclined smooth surface; Fig. 47B);
take a bristol card (or strong drawing paper) of the size adapted to that of the specimen
that you want to prepare;
- Write the serial number IN PENCIL in the right down corner;
- Put the bristol paper and the seaweed in the tray, on the floater (Fig. 47C);
- Choose (a) nice, complete specimen(s) (with holdfast; eventually fertile);
- Arrange the specimen(s) in an optimal way, by pushing the floater under water (Figs
47D, E); filamentous, supple specimens can be spread by a small brush;
- Take the floater, bristol card and specimen slightly inclined out of the water and let
the surplus of water run off (Fig. 47F);
- Put the bristol card + specimen on a newspaper on a horizontal surface and let it
air-dry somewhat (Fig. 47G); don’t leave it in the sun and don’t wait too long: the
specimens should not shrivel!
- Put the air-dried specimens between newspapers, covered by a fleece (Figs 48A-E),
preferably regularly alternated by corrugated cardboard (for aeration);
- Close the plantpress with belts or put weights on them (Fig. 48F);
- Keeping the plant press in the sun or adding a ventilator directed on the press
increases the drying speed, avoiding molding of the specimens; NEVER put the plant
press in an oven (unless it is a drying oven with ventilation)!
- Change the newspapers (not the flieces) daily until the specimens are dry;

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 - Mount the herbarium specimen on standard dimension sheets (eventually stick the
loose plants with glued paper strips, never directly with glue; certainly don’t plastify
them!!!) together with the label (Figs 49A, G);
- Store in a dry room, sheltered from direct sunlight (Figs 49B-F).

Fig. 47. Preparing herbarium specimens. A. Sorting out specimens in trays filled with
seawater; B. The cork floater in the tray filled with seawater; C. The numbered bristol card
on the floater; D. Arranging the specimens of the bristol card; E. Specimens arranged on the
bristol card, on the floater, still in the tray with seawater; F. Taking the floater and the bristol
card with specimens out of the water, letting drip off most of the water; G. The bristol card with
specimens is (shortly) air dried.

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 Fig. 48. Preparing herbarium specimens. A. In the plant press a corrugated cardboard and dry
newspaper is placed; B. Placing the bristol card with specimens on the newspaper; C. Putting
a fleece on the specimens; D. Adding a newspaper on the fleece; E. Adding a corrugated
cardboard on top; F. Closing the plant press.

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 Fig. 49. Storing specimens. A. Example of a seaweed herbarium specimen (Grateloupia
lithophila) and label in the ring binder at the GENT herbarium; B. The National Herbarium
of Sri Lanka in Peradeniya; C. The inside of the National Herbarium of Sri Lanka; D.
The cupboards where the specimens are kept in the National Herbarium of Sri Lanka;
E. Cupboard with the large herbarium specimens at the GENT herbarium; F. Cupboard
with smaller herbarium specimens classified in ring binders at the GENT herbarium; G. A
mounted specimen of Acrosorium ciliolatum (Harvey) Kylin, with field identification on the
full label and final identification on the ‘Determinavit-label’.

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 9.1.8. Formalin-preserved specimens
Most herbarium specimens can be resoaked for anatomical analysis, but most of the
time cells remain shrivelled and cytological details (e.g. plasts) difficult to observe.
Therefore it is better to keep (part of) a specimen in 4% formaldehyde (pure formalin
= 40%, so 1 part of formalin + 9 parts of SEAwater; the concentration is not critical and
even half the above will usually give good preservation). Add the same label (number)
as the serial number of the herbarium specimen and add ‘+F’ on the herbarium label
and in your data set as to indicate that there is a formalin-preserved specimen.
Formalin is a strong irritant and carcinogenic and therefore should be handled with
care, avoiding inhalation or direct contact with the skin. Store the formalin preserved
specimens in hermetically closed vials, out of the light, in a (preferably cool) room
with ventilation and NEVER in a room where persons are working on a regular basis
(separate store room)!!!
9.1.9. Silica-preserved specimens
Fragments of most herbarium specimens can be used for molecular analysis (in as
far as they have not been previously stored in formalin), but most of the time results
are (much) better when fragments are immediately dried in silicagel. Therefore,
Eppendorfs are being used, (almost) filled with fine-grained silicagel. The Eppendorfs
should be kept closed at all times, otherwise the silicagel would attract air humidity.
Only a small fragment (a few mm only) of an apical part of the specimen should be cut
off and cleaned and dried with a paper tissue. The young apices are less epiphytized
but still have to be cleaned as to remove the eventual single-celled epiphytes (e.g.
diatoms). The fragment is put in the Eppendorf and a tiny label with the same serial
number as the herbarium specimen is added (Fig. 50A). The Eppendorf should be
closed immediately (Fig. 50B) and somewhat shaked, as to completely surround
the fragment by silicagel: the quicker the drying process, the better the molecular
extraction will proceed. Some scientists prefer to dry two fragments for the case that
the DNA-extraction on the first fragment didn’t succeed. It is useful to indicate on the
top of the Eppendorf that it has been used (Fig. 50C). On the herbarium label and in
the data set ‘+S’ should be added as to indicate that there is a silicagel-preserved
portion. Of course this should be deleted from the data set as soon as the fragment(s)
have been used.

Fig. 50. Silicagel dried specimens. A. Putting a specimen in a labeled Eppendorf; B. Closing
the Eppendorf; C. Indicating that the Eppendorf has been used.

Molecular techniques are outside the scope of this field guide. For details we refer to
Hillis & Moritz (1996).

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