The taxonomy and nomenclature of certain palmelloid, planktonic

459
J . a n . Soc. (Bot.),56, 388, p . 459
Printed in && B&in
The taxonomy and nomenclature of certain palmelloid,
planktonic, green algae described previously
BY J. W. G . LUND, F.L.S.
Freshwater Biological Association, Ambleside, Westmorland
-
Since the description (Lund, 1956) of certain planktonic algae I have received, through
the kindness of Professors A. M. Matvienko of Kharkov and J. V. Roll of Kiev, an
article and a book by Korshikov (1941, 1953) which were unknown to me. In these
Korshikov describes and figures most of them under different names. I n addition,
Bourrelly (1959) and Ettl (Komarek & Ettl, 1958) have considered Qemellicysth
Teiling and Cecidochloris Skuja. The following is a revision of the nomenclature and taxonomy of these algae in relation t o these works and some additional observations.
(1) Coe7tococcu.s planctonieus Komhikov, 1953 (Radiowccwr planktonicua Lund, 1956:
694)
Although I have often seen this alga in the plankton of Windermere in the last four
years and used clonal cultum for experimental purposes no features additional to those
already described have been detected.
In placing this alga in Radiococcus, I followed Teiling (1946) in not considering the
fine, radially striated mucilage of the envelope a character of generic importance,
although it is present in the species previously described. Komhikov (1953) describes
a number of azoosporic algae which form microscopic, mucilaginous colonies which only
differ in the shape and arrangement of the cells within. For these he erects three new
genera, Coenocystis, Coenochloti8 and Coenococcus, which lie near to 8phaerocystis Chodat
pro parte and Radiococcus Schmidle.
Coenochloris differs from Coenococcus in the grouping of the cells (autospores) closely
tbgether in the centre of the colony. In Coenococcw they are dispersed more or less
irregularly and the centre of the colony is often free of them (Korshikov, 1953, figs. 295,
296a, Radiococcw planktonicw Lund, 1956, fig. 1A, C). From Radiococcwr both genera
difFer in the absence of radially striated mucilage. The cells of young colonies of Coeno&is
(e.g. Korshikov, 1953, fig. 296b, 297a) are in the very centre of the colony, but
this feature is less marked in older colonies. Goenmyatis also has structureless mucilage,
the cells are elongate and not restricted to the central area of the colony.
The position of Sphaerocystis is not clarified by Komhikov, He never saw zoospores
(of. Lund, 1952) and agrees that it has nothing to do with CTlOeococcus Braun (Lund, 1952,
1967). He doea not figure it, in marked contrast to almost every other species in his
mcbgnificent monograph. It seems, then, that although he says it is a common alga he
bas not studied it in detail. This does not resolve the problem as to what are the limits
af this residue of Chodat’s (1897) genus, 8. echrozteri would appear to come close to
Coenococcw. S. polycocca of Komhikov (1953: 327, fig. 301) ia so different that it should
be placed in another genus. It resembles some of the stages of Qbeocystis planctonicu
(Naeg.) Lemm. in Skuja (1948, Taf. XII, figs. 2, 4).
In these genera there are some differences in the fate of the wall of the mother cell
duripg reproduction, but it is not clear that these are significant taxonomically.
Komhikov says that the cells of C o e 7 b o c o ~planktonicus are liberated by the mucihghizetion of the wall of the mother cell, this being the only respect in which his account
of Coemmccwplanktwicus differsfrommine of Radiococcus planktonim. In my material
460
J. W. G.LWND
however, pieces of this wall may sometimes be seen faihtly after liberation of the autospores (Lund, 1956, one such piece shown in fig. 1C). Therefore this does not seem to be
a real point of difference on which a separation might be based.
It may be questioned whether the differences between Radiococcwr, Co~nococcue,
Coenocyitia and Coenochloris are of generic value. (I exclude Sphrrocyatia from this
consideration since the identity of its type species, S . schro-teri, is too uncertain for the
name to be used). However, it is impossible to anewer this question for we know nothing
of the cytogenetica of these algae. The distinctions are in accord with present practice
in the Chlorococcales where a large part of the taxonomy is based on convenience for
identification. The retention of these genera, also avoids changes in nomenclature which
are no more likely to be final than the present arrangement.
Since their cells are not centrally located in a wide, radially striated mucilage investment, Radwcoccw planktonicma Lund mdR.pelqicu Teil, are therefore best removed
from that genus and placed in Coelzococcwr. The correct name for the ibst is then C .
phnctonicue Komh. and for the second C. pelagicus (Teiling) comb.nov. (Radiococcus
pclagicus Teiling, 1946).
(2) Ceciduchbrie adnata (Korsh.) comb.nov. (Chhrophyaema adnata Korshikov, 1941;
Cecidochlol.Qeetichogloeacr Skuja, 1948: 100; see also Lund, 1966: 595)
Skuja and I were unaware that this alga had been described previously as Chlorophyaema
Korshikov (1941: 62).
Skuja (1948)found that a characteristic feature was the solution of the mucilage of the
host colony, Stichogloea doedarleinii (Schmidle) Wille. He also found the species occasionally on other muailaginous, colonial dgrte, but it is not clear whether he saw the same
solution of the mucilage in these hosts. Korshikov (1941) and I (Lund, 1956) did not
observe this; indeed we both observed it on algae which do not have mucilage sheaths
(Asterionelkc and D i m b y o n ) . It does not seem absolutely certain that what Skuja
observed W&B the destruotion of the mucilage of the host. When immersed in Staurastrum
anatinurn Cooke & Wilh (Lund, 1956, fig. 1D, F, c f , H) there is no sign of this. The alga
is completely surrounded by the mucilage of the host plant. Skuja (1948) says that it
leaves a clear pook mark. If, however, the developing Cecidochlorirr pushes aside the
surface mucilage of the host during the enlargement of its membrane, the result may look
the same. The hole left by it will fill up if the mucilage of the host alga is diffluent enough
to flow back or if new mucilage is produced.
Equally, if Skuja’s interpretation is correct, this solution of the host mucilage does
not seem to be a generic difference judging from his own accounts of Chlorophyaem
mdcrocyatidh Skuja (1966: 156-7, Taf. 22, figs. 31-33, Taf. 23, figs. 1, Z),an alga which
Ettl (1968) tramfere to Chlorangium because the cells do not lie in the inflated vesiole
developed from the wall of the mother cell. Skuja only distinguishes Cecidochloris from
this species by the absence of a b a d stalk (foot), the ability of the vegetative cells to
become motile, and the insertion of the daughter colonitxi within the enlarged membrane
of the mother cell (Einachachtelung). Whether the stalk of Chlorophyeema microcystidis
beoomes immersed in the host by dissolution of its mucilage is not stated.
“he direct metamorphosis of vegetative cells into swarmers is really only the normal
formation of zoospores with the development of motility delayed while growth continues.
This is the difference between volvocine and chlorococcoid (incl. palmelloid and dendroid
forme) green algae. In zoosporio coccoid forms assumption of motility is not possible
after the daughter cells have reaohed a certain stage in development, and hence only
zoospores can be produced. The rwson why there is no clear demarcation between these
group of motile and non-motile algae is that this ‘certain stage’ may be reached early
or late so that there are all trmmitions between a phase of motility and one without
motility. This may be Been in a series of species and genera, or in one and the same
Taxonomy and nomenclature of certain planktonic algae
461
lapeciea (e.g. palmelloid stages of terrestrial species of Chlumydomonae and species of
ChEorowccum).
Can the genus CecidochlorisSkuja (1948) be separated from Chlorophysemal The latter
wag erected (Pascher, 1927) to include species of Chlamydomonas in whose life-history
the non-motile phase is dominant. These algae (see also Pascher, 1940) are attached to
various substrata by a pad or stalk, which may be only a low cushion of iron-encrusted
material. They differ from allied genera (see Ettl, 1958) in the enlargement of the membrane so that the original cell and the following generations (usually one or two) lie in
an inflated vesicle, a condition only found in some of the other genera during the development of the zoospores (e.g. Pascher, 1927, fig. 440d, g: Chlorangium inhuerens (Bachm)
Ettl, 1958).
Ettl(l958) places Anachin's (1929-30) Chlorophysema sessilis in Cecidochloris since the
habit and cell structure are identical. Pascher (1940) had previously said that this
speoiea does not belong to Chlorophysema but gave no reaBons ; Korshikov (1953) retains
it in this genus. However, Ettl (1958 : 333) says Chlorophysema and CecidochEoris have
the same cell structure, the latter being characterized by its'habit, that is living in the
mucilage of other algae. This ignores the fact that Cecidochkwis stichogloea. Skuja
(Chkwophysema admta Korsh.) lives also on algae without mucilage (Korshikov, 1941,
1963; Lund, 1956). There does not seem to be any distinction on Ettl's grounds.
The only possible distinction seems to be the formation of some kind of attachment
organ in Chlorophysema. Astalk is formed in C. inertis (Korsh.) Pasch., C. apiocystifomne
(Artari) Pasch., C. wntractum Pasch. and C. ampliata Skuja. Transitions to a ferruginous
pad or solely this are seen in C. inertis p.p., C. ellipsoideum Pasch., and C. ovalis Skuja.
Figures and details of these species may be found in Pascher (1927, 1940), Korshikov
(1953), Skuja (1966), and Ettl (1958). Korshikov (1953) says on p. 70 th6t the cells are
attached to the substratum by a beak, but in his key on p. 71 separates C. sessilis and
C.adnuta from the other species by the absence of a beak.
As Chlorophysema sessilis Anachin and C . adnata Korsh. agree in the absence of any
definite attaching organ or basal iron incrustation they are best transferred to Cecidochloris as C. sessilis (Anachin) Ettl and C. adnuta (Korsh.) comb.nov. ( C . stichogloeae
Skuja), with the change that the grounds for generic separation are not those of Ettl
(1958) or Skuja (1950), but this absence of an-attaching organ. This is in agreement with
the taxonomic importance generally placed on the presence or absence of this feature in
the Chlorococcales and Chlorangiales. It also avoids further changes in nomenclature.
(3) QemeUicystis imperfecta (Korsh.) comb.nov. (Tetraspora imperfecta Korshikov, 1941 :
60; Gemllicystis neglecta Teiling, 1946: 67, emend. Skuja, 1948: 110; 828 also Lund,
1956: 598, Bourrelly 1959: 40, incl. G. lundii Bourrelly, 1959).
All the accounts cited above are more or less a t variance with one another. The essential
generic features on which there is now agreement are as follows: the cells are arranged in
pairs, apex t o apex. When young they are apically flattened, but before the next division
are generally globose. These pairs are further arranged in loose, irregular groups of four
or more cells dispersed with& a wide mucilage envelope which may itself split into two
parts. The massive, basically cup-shaped, chromatophore contains a large b a d pyrenoid.
Two anterior contractile vacuoles are present.
The points of dieagreement are the presence or absence of pseudocilia and flagella,
8 stigma in the non-motile stage, and the detailed structure of the ohromatophore and
the starch sheath to the pyrenoid. In addition to these accounts, algae showing certain
etrong resemblances to Q. neglecta have been described by Woronichin (1931) and Korshikov (1941)aa Pseudospp7raerocyotisplanctonica Woronich. and Tetrmpwa imperfectu Korsh.
Tbiling (1946) depicted as pseudocilia short threads which are undoubtedly two
flagella (Skuja, 1948; Lund, 1966), a feature which I have checked on several occasions
462
J. W. G.LUND
in the last two years. Pseudocilia were believed to be absent (Skuja, 1948; Lund, 1956),
though curious interrupted thread-like structures were visible in colonies stained in
Lugol’s solution (Lund, 1958, fig. 3G). Them were thought to be artifacts, but I have
since observed them with the aid of cotton blue and under phase contrast. They are not
always detectable, but when present there are never more than two to each cell. They
arise from the front end and almost a t once bend backwards sharply. They may reach the
periphery of the mucilage, but I have never seen them outside it.
The reason why Teiling (1946)did not see contractile vacuoles or a stigma seems to be
that hc examined preserved material. Thus in his figure 2 the mucilage envelope is
strongly contracted as is normal when formalin or other flxativea and stains are added.
The British specimens were said to have a stigma (Lund, 1966) and its occasional
apparent absence could have been an oversight because of its minute size. Skuja (1948)
found that the Swedish ones did not have a stigma but this did appear when vegetative
cells assumed motility. Bourrelly (1969)say13a stigma is often present. Further examination has shown that these accounts are not a8 divergent as originally appeared. A stigma
is not always present in the vegetative cells, but in the British specimens very commonly
is (of. Bourrelly, 1959). Skuja (1948) is therefore wrong in denying its presence but is
probably right in relating the development of a stigma to the assumption, or potential
assumption, of motility. A stigma is, however, often present in non-motile ceUs lacking
flagella.
Bourrelly (1969) has divided the G. negkcta of the authors into two species. Q. lundii
has en undissected chromatophorn unlike that described by Skuja (1948). The starch
sheath to the pyrenoid consists of two caps, one lining the anterior and the other the
posterior part of the pyrenoid, with a small gap between them. It is perhaps s i e c a n t
that he says that this gap L ‘signe probable de croissence’, if by this he refem to cell
development;. Q. neglecta Teil. has an apically dissected chromatophore (Skuja, 1948,
Taf. XI, figs. 14, 16-21) and has, according to Skuja, a granular starch sheath to the
pyrenoid. Whether Bourrelly is correct in applying this name to Skuja’s description
seems uncertain. The type description (Teiling, 1946: 67) lhentions neither of these
features. Bourrelly (1969) presumably considers that as both Telling and Skuja found
i t commonly in Sweden they must have been observing the same alga. But Bourrellg
(1959) and Lund (1966) failed to find the characteristic chromatophore and Teiling
recorded his alga from Italy and Germany. It seems unlikely that only one speciem
exists in Sweden and another in various parts of Europe (see also Jaag & Nipkow (1951)).
Dr H. M. Canter has also found specimens in formalin material from Finland which agree
with Teiling’s (1946) description of Gr. neglecta. However, further examination of British
material, from the same lakes as before, leads to the view that both these algae belong
t o the same species which can therefore be called 0.neglectu Teil. em. Skuja (syn.
Q. Zundii Bourr.). Sometimes in British specimens the chromatophore is not unbroken,
though I have never seen such dissected specimens a8 some of those depicted by Skuja.
Indeed it appears that not all the Swedish speaimens have dissected chromatophorea
(e.g. Skuja, 1948, Taf. XI, fig. 15), while it seems that this feature may easily be overlooked (of. Skuja’s drawings a t lower magnitication, 1948, Taf. XI, figs. 10-13). Skuja
says ‘von aussen f.median zerschlitzt ’ and ‘interdum extus longitudinalites irregularitusve striato’ (Skuja, 1948: 112, 311). I n British material, when the cells are observed
from above, the upper margin of the‘chromatophore may sometimes be seen to be
somewhat undulate, and from the side t o have one or two slits. These features are easily
overlooked because the chromatophore is so massive that the undulations or slits are
masked by the parts beneath, only becoming visible by careful focusing with a n
immersion lens.
Pseu~?o8phaerocystieplanctonica Woronichin (1931) (see Korshikov, 1953: 74) is so like
Gemdicystia neglecta that despite the sketchy deacription the only significent difference
seems to be the occtlsional presence of a papilla (compare the two cells in fig. 176,
Taxonomy and nomenclature of certain planktonic algae
463
Korshikov, 1953). This papilla has not been seen in the rich material examined by myself
or Skuja (1948) and so Pseudosphaerocystis plumtonica cannot be considered as synonymous with Qemellicystis neglecta. On the other hand, it seems certain that it does belong
t o the latter genus and is,therefore, better named Qemellicystisplunchica (Woronichin)
comb.nov.
Tetrmpora imperfecta Korshikov (1941: 60) is also so like (
I
neglecta
. that the only
question concerning its identity with it is over the pseudocilia. The apparent absence
of a stigma, flagella or dissected chromatophores is clearly not significgnt from what has
been said already. Korshikov’s statement concerning the pseudocilia is in translation
as follows ;
As regards the pseudocilia, in the vast majority of cases they are completely absent. Only
by very careful investigation, with the help of immersion objectives, c a n one sometimes see
on the anterior end of the cells faintly visible muciiaginous lumps or threads, which form
probably because of the destruction of the pseudocilia in thevery beginning of their formation.
In certain cases I was able t o see on the front end of the cell pairs of tiny mucilaginous globules,
resembling those which get formed from the aggregation of threads (fig. 3). Thus in fact the
pseudocilia are here as if reduced and this waa the cause of my naming the organism T.
imperfectu. However, careful examination of a large number of colonies showed that in some
catm the cells have a supply of real pseudocilia and this permits me to include the given organiam in the genus Tetraspora, despite a series of peculiarities unusual for this genus. In
figure 4 two c e h with pseudocilia are depicted. The latter were so thin that only with immersion lenses and strained attention did I succeed in tracing them from the beginning to the
end and in depicting them with the help of a drawing apparatus. The contour of the mucilage
is invisible by normal investigation and was drawn after application of indian ink.The length
of the pseudocilia was about 60p. Immediately from their base they bend in the opposite
direction backwards to the periphery, coming forward out of the mucilage into the surrounding
water for approximately half their length. They stain with extraordinary difEculty and this
greatly impedes their being found. In material from the river Niva the cells apparently
always had pseudocilia.
I n order to emure, so far as possible, that Korshikov’s exact meaning is not altered,
I have translated this passage in a literal manner.
Korshikov’s and my observations agree in several respects. The pseudocilia are
always difficult to see. This may or may not mean that he is correct in saying that they
are often really absent. When seen their course is very difficult to follow. They bend
backwards in a very characteristic manner. His observations differ from mine in the
occasional presence of mucilaginous bubbles in the place where a t other times pseudocilia
are found and the passage of pseudocilia well beyond the mucilage investment. The
‘bubbles’ may be of W e r e n t character or related to the production of flagella. That the
pseudocilia did indeed project beyond the edge of the colony and were not extruded
during contraction of the mucilage in stains or fixatives is clear from Korshikov’s
use of indian ink. Korshikov (1941, Tab. 3, fig. 4) shows them as continuous lines in
contrast to my broken lines (Lund, 1956, fig. 3G) but this is probably because I have
drawn the parts of the protoplasm which stain and are not continuous in pseudocilia.
It is just such parts which are likely t o be stained by iodine and cotton blue and to stand
out in phase contrast.
There can then be no doubt that a t least part of the material included in Tetrmpora
imperfecta is identical with Gemellicystis neglecta. The only uncertain part of Korshikov’s
(1953) material is that found in the River Niva in which all the cells had psehdocilia.
Tetrmpora imperfecta Korsh. may therefore be typified by the organism on which the
major part of his description is based and the Niva material excluded as being either
aberrant or belonging to a different organism. Then T . imperfecta Korsh. as typified here
has priority over Teiling’s (1940) G e d i c y s t i s neglecta. ,The alga should be separated
from Tetrmpora because of the characteristic cell arrangement and structure, while
25
JOURN. L L ” . S0C.-BOTANY, VOL. LVI
464
4
J. W. a. LUND
even if the paeudocilia really are such, they are indeed imperfect and unlike thoee of
Tetraapwa. The name should then be Gemdicysti8 imprfecta (Korah.) comb.nov.
(4) Paulschdzia p 8 e ~ v o l t m x(Schulz) Skuja, 1948: 118 (Tetraepora p~eudovolvox
Schulz, 1923; T . eimplex Korahikov, 1941; Schulziellu pseudovolvox Teiling, 1942)
The range of structure deecribed and depicted for Tstraspora eimplix Korshikov (1941)
ia within that in Schulz (1923), Teiling (1942), Skuja (1948) and Lund (1956). Skuja’s
name has priority 88 it is based on T .psewlovolvox Schulz (1923, Schuhulzielh peeudovolvox
Teil.). The genus ie clearly diatinct from Tetraspa (Lund, 1966).
(6) Paulschulzia tenera (Korah.) comb.nov. (Tetrccspwa tenera Korahikov, 1941; Tetra8 p ~ ehgam
a
Womnichin, 1949; Paulschulzia elegans (Woron.) Fott, 1964; P.ekgam
(Womnich.) Lund, 1966)
Tetraqma tenera Korahikov (1941) is identical with P . ehgam. Since this name has
priority over Woronichin’a (1949) Tetraepora elegaw, the correct name is Paubchulzia
tenera (Korah.) comb.nov.
The author’s thanks &re due to Mr R. Row for kindly critioizing the manuscript and
eepecially for advice on nomenclature, and to his wife (DrH. M. Canter) for checking
the observations on the chromatophores of aemeUicy8tis mglecta.
BUMMdBY
On the bash of further investigation and literature previoudy unobtainable, the
correct names and synonymy of certain palmelloid planktonic green dgm is considered
to be aa follows:
(1) Coenowccwr pkc?tctoniowKorsh. (Radiococcwplanctonicw Lund.).
(2) Coerwwccw, pehgicua (Teil.) comb.nov. (Rudwwccwrpkqica Teil.).
(3) Cecidochlmie adnata (Korsh.) combmov. (~h.h?vphysema
adnata Korsh.; CecidochEoria
8tiCbghU4 Skuja),
(4) Gemellicystis neglecta Teil. non Bourr. (Tetraapora imperfectu Korsh. ; GemeElicy8tth
neglecta Ted. mnsu Bourrelly (1959); a. ludii Bourr.).
( 6 ) GemeUicy8tie phndoniea (Woronich.) comb.nov. (P8eud08phuerocystia planctonica
Woronich.).
(0) Pauhchu~ziap8eudovoltmx(Schulz)Skuja (Tetraeporap8eudoorolvoa:Schulz. ; T.8 i m p h
Korsh. ; Schulziellu pseudovolvox Teil. ; Paubchulzia p8tudovolvox (Schulz em.
Teiling)).
( 7 ) Paukchulzia tenera (Korsh.) comb.nov. (Tetraaporatenera Korsh. ; T.ekgans Womnich. ; Paubchulzia ekgans (Woronich.) Fott; P . elegailw (Woronich.) Lund.).
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