Trachelomonas Mg2

Observations on Some Species of Trachelomonas Grown in Culture
Author(s): E. G. Pringsheim
Source: New Phytologist, Vol. 52, No. 3 (Sep., 1953), pp. 238-266
Published by: Wiley on behalf of the New Phytologist Trust
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[ 238 ]
OBSERVATIONS ON SOME SPECIES OF

TRACHELOMONAS GROWN IN CULTURE
BY E. G. PRINGSHEIM
(Continuedfromp. I I3)
TAXONOMY
(i) GeneralConsiderations
It is not proposed to followthe historicaldevelopmentof Trachelomonas taxonomyin
detail. That would only show how slow the progresswas, in spite of the amassingof
descriptionsand names. Even establishedknowledgeof the cell body was lost again.
Cohn (I852) recognizedthe similarity ofthe cell bodyto thatofEuglena. Klebs (I883)
knew almostas much of the organizationof Trachelomonas as can be foundin modern
books, although, followingin the main Ehrenberg, he described only six species.
Dangeard's contributions(I889, I90I) do not give much more. Chodat (I909) finds
the principlesof differentiation quite uncertain. Lemmermann(I9IO) gives a reliable
surveyof the morphology,thoughhe overlookssome older papers, and describesforty
taxonomic forms. His contributionin the Suisswasserflora(1913) is similar. Conrad
(I9I6) complainsagain of the insecurityregardingthe validityof species, which could
onlybe overcomeby 'mesurationsmethodiqueset surtoutdes culturespures', because
'la forme,la taille,les ornementationsde la coque sontextremement variableset probable-
mentsujets a de profondesfluctuations'.He feltthatnot even the beginningof a satis-
factoryinsighthad been reached.The vaguenessof conceptsas to whichfeaturesare of
importance,and how theyare influencedby conditionswas in a remarkablecontrastto
the attractiveclearnessofthe naturalobjectsto be systematized:the envelopeswhichcan
so easilybe collectedand preserved.
To Conrad (I9I6, p. 195) is due the arrangement of the species of Trachelomonas into
fourgroups: 'coque spheriqueou subspherique;coque ovale, plus ou moins allongeeou
cylindrique;coque ovale, plus large que longue; coque en formed'cEufa base tournee
vers le haut.' This systemis too artificialto indicatetrue relationshipwhich mightbe
corroboratedwith the help of other characters,where such are available. Conrad's
systembecame the basis of Deflandre'selaboratekeys.
None of the existingattemptsat a systemof taxonomywas, it seems, meant to be
natural;theirpurposewas to assignto everyenvelopea place whereit could be foundfor
identificationpurposes. As the discrepancybetweenthesetwo aims has, in othergroups,
oftenbeen emphasized as unsurmountable,it is as well to say that apparentlysystems
whichareunnaturalare generallyalso shortlived.It is doubtfulwhetherin Trachelomonas
a systemreflecting can be achieved at presentbut, if at all, it can only be
true affinity
attainedon the groundof new observations.
A naturalbiological systemmust primarilyrest on a numberof confinedgroups of
formsheld togetherby as manycommoncharacteristics as possible,so thattherecannot

Observations
on somespeciesofTrachelomonasgrownin culture 239
be any doubt as to theiraffinity. These groups have then to be arrangedin a pattern
showingtheirplaces nearertogetheror fartherapart. Forms which cannotbe put into
one of the primarygroupsreceivea place betweenthemor at the fringesof the arrange-
ment. Taxonomic systemsseem always to originatethis way. Groupings on single
characteristics,as well conceived as theymay be, have somethingartificialabout them
and, althoughtheymay be usefulforsome time,do not last, because new featuresturn
up whichindicateanotherconnexionbetweenspecies and make it more difficult to allot
the rightplace to new formsto be inserted.
In Trachelomonas thefollowingcharacteristics offerthemselvesfortaxonomicpurposes:
shape, size, colour, and ornamentationof the envelopes, number of chromatophores,
formationof pyrenoids.These are distinctivefeatureswithinthe genus,whileflagellum,
eye-spot,nucleus, and reservoirsystemdo not seem to furnishuseful charactersof
differentiation,because theyare almostidenticalin all the species.
The strainsinvestigatedcan be arrangedin groupsthe typespecies of whichmay be
the following:(i) T. hispida, (2) T. volvocina,(3) T. bernardinensis,(4) T. conspersa (syn.
Strombomonas), (5) T. deflandrei, (6) T. bulla.
To group(i) belongstrains3, 6, I2, I6, 23, 24, 29 and3I representing T. hispidatyp.,
althoughtheyare not quite identicalwithone another,strains4 and I I: var. acuminata,
and strain5: var. coronata. It is not easy in a groupofverysimilarformsto differentiate
betweenhereditaryand induced variations,when these are slight. Envelope formation
is so easilyinfluencedby conditionswhich are neitherquite constantnor homogeneous
in soil-watercultures.Thus, partof the typicalstrainswere at firstbelievedto belongto
varietypunctataLemm., but turnedout to produce spines in old cultures.Those here
describedas varietiesacuminataand coronataare not in everyenvelope distinguishable
fromthe type,among the eightstrainsof which I have, however,never observedtail
conesor longspinesaroundtheporerespectively. Middelhoek (I95I, p. 232), whoalso
foundcountlesssmall intermediateforms,believes: 'une etude du protoplasteest indis-
pensablepour obtenirune determination plus ou moinssatisfaisante'but does notappear
to have undertakensuch a study.
Characteristics of T. hispidaare: double sheathedpyrenoids,relativelylargenumberof
chromatophores (I0-I2), pronouncedcolorationofenvelope. Collar,ifat all, short,often
raggedor spiny. Envelope ovoid, ellipsoidalor slightlycylindrical.
To group(i) shallalsobe assignedstrains7, I0, I3, I5, I7, 25, 32 and37, all ofwhich
are alike and have been identifiedas T. zorensis,the commonestof the species foundin
the largervicinityof Cambridge.The shape ofthe envelopeand thetwo or threechroma-
tophoresare reminiscentof T. oblonga,fromwhichthe species is different in its greater
size, the ornamentationand colour of the envelope,and its double-sheathedpyrenoids.
By the last this species shows a relationto T. hispida,fromwhich it is different, in its
smallersize, i.e. 20: 8 , comparedwith 30; 20 1 in the latter,which may explain the
small numberof chromatophores. The mostadvanced envelopesof T. zorensisresemble
in
T. hispida theirspinyappearance.
Group (2) contains:strainsI9, 26, 28 and 43, T. volvocina; strain8, T. volvocinopsis;
strain2I, T. volvocinopsis var. spiralis;strains9, i8, 26 and 36, T. oblonga;strain22,
T. oblongavar. punctata; and strain27, T. similis. Common to these formsare: inner
pyrenoids;two layeredenvelopes; two (or three) chromatophores;ring around pore;
quick,irregularmovement;no conspicuousornamentation, althoughtinydotsor delicate

240 E. G. PRINGSHEIM
spiralsmaybe present.The ringmay be supplementedby a cylinder,reachingoutward
or into the interior.
Group (3) consistsof: strainsI, 33, 34 and 40 representing T. bernardinensis;strain30,
T. pertyi;strains20, 4V, 42, T. lefevrei.In spite of a certainsimilarity withmembersof
the firstgroup,group (3) is herekeptapartbecause of the different pyrenoidtype,but it
is not yetquite satisfactorily defined.The appearanceof the oval envelopeis commonto
the strainshere included. T. bernardinensis has collar and tail cone, T. lefevreionlythe
collar,and T. pertyinone of the two.
Group (4), of which only strain2, Strombomonas was available,is well defined.The
clone appears,by its extraordinary variability,to representin itselfa numberof forms
which have been given names by previous authors.This formhas the same kind of
pyrenoidsas groups(2) and (3), but is different in its envelopesand certainfeaturesof
the protoplast.
Group (5) is representedby strains 35 and 44 with 'naked' pyrenoidsdevoid of
paramyloncaps, and with a location of the nucleus different fromthe majorityof the
strains. It is a new species.
Group (6), formedby species withoutpyrenoids,comprisesstrain38, T. varians,and
strain39, T. bulla. These species are otherwisenot particularlysimilar,except in the
great number of small, lens-like chromatophores.If there is a correlationbetween
dimensionsofthe body and numberof chromatophores, thisis not strict.T. varianswith
a diameterof 22-25 / has numerouschromatophores, comparedwithabout ten chroma-
tophoresin T. volvocinopsis witha diameterof i4-i6 /; but T. volvocinaand T. oblonga,
withabout the same dimensionsas the latter,have onlytwo and rarelythreechromato-
phores.There seems, however,to be a correlationbetweensize of chromatophoresand
pyrenoidformation, as is also recognizablein the genus Euglena. The largestchromato-
phoresusually have pyrenoids with paramylonsheaths,the smallesthave no pyrenoids,
and T. deflandrei withnaked pyrenoidsis intermediatein this respect.
It is clearthatall thesegroupstogetherembraceonlya smallpercentageof the species
of Trachelomonascompared with the multitudewhich have been described and the
probablystillgreaternumberthatexist.
(2) Descriptionof species

In the beginningit seemed, on accountof theirdiversity,to be almostimpossibleto
identifythe clones grownin culturewith species previouslydescribed.This difficulty
was graduallyovercomeas the technique improved. The more favourablethe condi-
tions, the more uniformare the individuals,and the identityof the strainswas then
more easilydetermined.It is thereforepossible to attributemost of the strainsisolated
to knownspecieswhich,in return,can now be betterdefined.This progressis due to the
greatnumberof cells of identicaloriginwhichcan be investigatedin a perfectstate. On
the other hand, populationsreared under less favourablecircumstancessupply some
clues for the understandingof aberrationsoccurringalso in nature. All the forms
appearing in one clone taken togethergive a picture of the inherentpossibilitiesof
a species. Deviations from the type, found under all conditionstested, are almost
certainlyhereditary,and strainsshowingthemcan rightlybe called varieties.
had to relyon luckychances of findingrich,homogeneous
As all earlierinvestigators
populations,it is not surprisingthat some new formswere found among the small

on somespeciesofTrachelomonasgrownin culture
Observations 24I
numberof strainsisolated in culture. Material which can easily be preservedwill be

useful in later work of identification,
furnishinga reliable stock of known origin.To
reformcurrenttaxonomy,experiencegatheredby investigatinghundreds of samples
would be necessary. I cannotundertakethisimmensetaskbut onlydescribethe strains
withwhich I became familiarin the course of my research.
Altogethersome fortyclones were isolated,some of whichprovedto be identical. In
groupingthesestrainsuse is made of the pyrenoidstructure.Featuresof the protoplast
should have priorityto those based on the envelope, and the shape and paramylon
depositionof the pyrenoidsis the onlyprotoplastcharacterin whichthe species clearly
differ.This groupingis tentativeonly,but it appears to bringrelatedspecies together.
The Trachelomonashispidagroup
There are two species with double-sheathedpyrenoids,one with up to a dozen
chromatophoresand ovoid envelopes,oftenwith a tendencyto produce a littlecollar:
T. hispida; and one with two to three chromatophores,and of smaller and narrower
figure:T. zorensis.
Members of T. hispidaare common: twelveof my strainsbelong to it. What is taken
as the type was found six times, and the varietiesacuminataand coronataeach once.
These formsare identicalin inner structure(Fig. 6) and do not differappreciablyin
dimensionswhichare generally29-3I : i8-20z j. The protoplasttendsto retirefromthe
envelope so that it becomes about cylindrical,oftenwith a littlewaist in the middle.
T. hispida(Perty)Stein emend. Defl. typ.was, forexample,isolatedfroma gardenpond
at Cambridgein JuneI940 and fromMadingleybrickpit near Cambridgein November
I942. When well developed it has a brown envelope with a low collar which is usually
ruggedat the edge, and withevenlydistributedspinesjust overtoppingthe outersurface.
In Deflandre'smonographT. hispidacomprisesthe type,fivevarietiesand two forms.
They are different fromthe main species in possessingspecial collar shapes, tail cones,
spines surroundingthe pore,havingspinesonlynearthefrontor back ends,or no spines
at all, respectively.Some of these charactersare additional,othersnegativecompared
withthe type. Some do not even fitintothe generaldiagnosis,forinstancethe absence
of spines or the possessionof a highcollar. It.is not statedwhyjust a formwithspines
but withouta conspicuouscollarand tailcone shouldbe thetype,as thelattercould quite
well be takento characterizethe morecompleteor advancedforms. If therewas a form
possessingboth tail cone and collar,thiswould be similarto T. bernardinensis, provided
thatprotoplasmiccharacteristics were neglected.
T. hispidavar. acuminataDefl. (Fig. 7A) was foundin June I940 in a ditchat Trum-
pington,Cambridge,fromwhichtwo strainswere isolated,different fromthe typeonly
in havinga small cone or thickeningof the posteriorend of the envelope,as indicatedin
Deflandre's fig. 222. Well-developed specimenshave short spines, and the same low
collar as the type. As usual, only a percentageshow ornamentations.T. hispidavar.
coronataLemm. (Fig. 7B) was isolated twice from a garden pond at Cambridge in
SeptemberI940, fromwhich the type had been taken earlierin the year. The crown,
i.e. spines decoratingthe edge of the collar,is distinctin manyindividuals.T. hispida
'var. punctata' Lemm. was collected at Fen Ditton, Cambridge, in April I945, at
Waterbeachfartherdown the River Cam in August I943, and froma pond at Debden
near SaffronWalden in September I945. Well-developedenvelopesfrommy cultures

provedto have,notonlya smallcollarlikethatofthetypebut also spines. Less advanced
envelopes may not of course have either. All these strainsbelong to T. hispida.The
appearanceof the envelopesand the innerstructureof the cells are typical.
A A
D D
Fig. 6. T. hispidaPerty. A, type: one cell with,anotherwithoutspines; chromatophores,and paramylon
around pyrenoids. B, type; variation in shape. C, type; small paramylon grains in characteristic
arrangementon chromatophores. D, naked cell, cf. also Fig. i, p. IOI.
In all theseclonesnakedcellswerealso observed(Figs. i, 6 D). They resembleto some

degreeEuglenagracilisin beingalmostcylindricalwitha bluntposteriortip,and in having
about the same number of discoid chromatophoreswith double-sheathedpyrenoids.

Observations
They differin the almostaxial insertionof the flagellum,in greatermetaboly,the longer
flagellum,and the much largereye-spot. All the membersof the group are apt to forgo
theirenvelopesunder unfavourablecircumstancesbut, like othernaked Trachelomonas
cells,cannotbe inducedto multiplyin thisstateso thatpopulationsentirelycomposedof
naked cells could not be obtained.
A B
Fig. 7. T. hispida. A, var. acuminataDefl., occasional occurrenceof tail-cone; protoplastbegins to leave

envelope; B, var. coronataLemm., various shapes of ornamentssurroundingpore.
T. zorensisis so farthe only otherrepresentative of the firstgroup. No varietiescan

be discernedamong the eightstrainsisolated, but for slightdifferences in size. It was
foundin the same gardenpond alreadymentioned,in JuneI940, in a ditchat Fen Ditton
in Apriland December I943, in a pond at Shelfordin April I943, in a ditchat Fen Ditton
in April I945, in a pond at Debden in SeptemberI945, in waterwithmud fromthe River
Cam in July I946, and on Coe Fen, Cambridge,in August I949. The last strainis
largerthan the others: 29-3I: I7-i8 /t,while most envelopes of the otherstrainsare
2I-23: I5-I7 /t.The uniformity withinthis species is alreadymentionedin Deflandre's
monograph(p. 92).
In the elongate shape, the pore and the small number of chromatophoresthere is
a certainsimilarity to T. oblongawhich,however,is smallerand has innerpyrenoids.The
two or three,sometimesfourpyrenoidsare situatedin a chromatophore each, which are
narrowas seen fromthe side, and roundish,oftenfringedor lobed, particularlyat their
posteriorends,as seen fromthefront.In comparisonwith T. hispidathesmallernumber
of chromatophorescould be explained by the inferiorsize of the protoplast. Well-
developedenvelopesareoftensimilarto thoseof T. hispidain ornamentation (Fig. 8 A-D).
Naked cells have,underunfavourableconditions,e.g. putrefaction, been seen occasion-
ally in all the strains.They are similarin shape to those of T. hispida,namelyalmost
cylindrical,withthe nucleus near the posteriorend, not distinctlystriated,moderately
metabolic,having two or three chromatophores, in a few cases more. Their lengthis
43-50[k,theirwidthabout ionu.
The envelope is eitherelongateoval or, more often,almostcylindricalin the middle
region.This deviationfroman ellipsoidshapeis notmentionedin Deflandre'smonograph,
but is a regularfeaturein all mystrains,and is also shownin twoofthefiguresreproduced
by Deflandre. Well-developed envelopes are golden brown and bear roughlyspiral,
irregularly band-likeornamentations, or wartswhichmay even become bluntprocesses,
and theenvelopeis thenrelatively thickas also pictured,thoughin a somewhatexaggerated
way, by Middelhoek (I95i). The thickeningby brown layersis not always regularly
New Phytol.52, 3 I'

distributedoverthewhole surface.The frontend is oftenalone or morestronglycovered
by thesedeposits(cf. Fig. 3). The pore is generallythickenedring-likefashion,muchlike
that of T. volvocinaor T. oblonga. In rareinstancesa low collarwas found.
C D
Fig. 8. T. zorensis Defl. A, appearanceof cells withenvelope,chromatophores and nucleus. B, toyp:
individual withtwo chromatophores;bottom: two views, high and deeper focusing,aftercell has been
turned by go'. C, three naked cells with different
arrangementof chromatophores. D, variationof
shapein a clone(IO).
TheTrachelomonas volvocinagroup
The T. volvocina groupis composedof formswitha naturalrelationship forthe
establishment ofwhichtheshapeoftheenvelopeis no suitablecharacter.If shapeis
takenas theprimary featureforgrouping, somepertinentforms areexcluded.Thereare,
inaddition,specieslikeT. v,olvocinopsis
and T. varianswhoseenvelopeswouldplacethem
as equal partnersintothisgroup,butforthegreater numberofsmallchromatophores
whichseemto indicateeithera looserrelation, or evenan onlysuperficial
similarityby
convergence. T. variansis,moreover,differentin havingno pyrenoids,following a rule
moreclearlyrepresented in Euglenawheresmalldiscoidchromatophores are always
devoidofpyrenoids. To embraceT. volvpocinopsisand T. 7varians
thegroupwouldhave
to be definedin a widersense.
Characteristics
commonto thenarrowergroupare: (I) two chromatophores withinner
pyrenoids;(2) eye-spotrelativelylarge; (3) nakedcells not spindle-shapedor cylindrical,

Observations
but pear-shaped;(4) envelopewithtwo layers,one colourless,one goldenbrown,smooth
or lightlyornamented,neverspiny.
This groupincludes: T. volvocina,T. oblonga,T. oblongavar.punctatan.var.T. similis,
among the strains investigated,and certainlymany more formswhich, for lack of
cytologicalinformation,cannotbe groupedhere withcertainty.
Fig. 9. T. volvocinaEhrbg. (I9). Three cells are given: one with two, one with three chromatophores,
and one with detached protoplast. A naked cell at half magnification.( x 500.)
T. volvocinatyp.was not commonin theplaces wherematerialwas collected,although

it is, along with T. hispida,the species most frequentlymentioned.The envelopesmay
oftenhave been confusedwith those of T. volvocinopsis or with formswhich are not
strictlyspherical,as T. volvocinais supposed to be. I foundit onlytwice,and even then
not everyspecimenwas 'exactementspherique'. The firstwas collectedat Fen Ditton,
Cambridge,in June I943, the second strainwas takenfroma pond witha greatvariety
of Euglenineaein thevicinityof Debden near SaffronWalden, wherein SeptemberI945
it was presentin various samples. The two inner pyrenoidsare generallywell seen
(Fig. 9). When growthis luxuriant,thereare sometimesthreeand even more chromato-
phores,each witha pyrenoid.The eye-spotis large,comparedwiththe size of the cell.
The hue of the envelope is given by Deflandreas variable,and in factdepends on the
amount of manganeseavailable. Since the coloured layeris verythin,the envelope is
neverdarkbrownas in otherspecies.
All the featuresof T. volvocinaare repeatedin T. oblongaLemm. (Fig. io), but for
a slightlysmhaller transversediameter,i.e. I2-I3 I as comparedwiththe length,and the
diameterof I4-I 5 ,u in T. volvocina.The twospeciesaretherefore closelyrelated,although
separatedby I4 pages in Deflandre'smonograph.The annularringand the structureand
colouroftheenvelopesare exactlythesame. The similarity goes so farthatevena proposal
totakethemas varietiesofone speciescould hardlybe opposed,particularly sincethereare
intermediate forms,and also varietiesof T. volvocinawitha shorterlongitudinalthantrans-
verseaxis. Whyshould a formnotbe a varietywhichdiffers fromthetypein theopposite
direction!Furthermore, thereare varietiesof both species possessingseveralumbilicate
,7-2

of the envelopewhichgive the impression
structures of multipleflagellar
openings.
These varieties,both named 'umbilicophora'by Deflandre,pointagain to a close
relationship.
Fig. io. T. oblongaLemm. (i8). Three individuals are represented:one with two chromatophores,one
with three, and one with detached cell body. Four naked cells are given at half magnification.
( x 500.)
Of the speciesof Trachelomonaswitha relatively long collar,onlyone was found

(strain27),collectedSeptemberI 945 from thesamepoolnearDebdenwhereT. volvocina,
T. zorensis,etc.,werealsotaken.It is hereincludedintheT. volvocinagroup,becauseof
thestructure ofthecellbodyandtheappearance oftheenvelope.A satisfactory identifi-
cationwasnotattained. Thereis probably a numberofsimilarforms.T. similisStokesis
punctated likemystrainbutlarger.(Cf. also Balech,I944, and Bourrelly, I947.)
Fig. iI. T. similis Stokes (?). Three cells with envelopes, with two, three, and four chromatophores
respectively,and naked cell. Details of collar (seen in the plane of the collar's bending) and pyrenoid
with paramyloncap.
The strainheredescribed(Fig. II) iS I9-20 H: i6 t, sometimes somewhat longer,the

neckis crooked,3,uwideand oftenslightly longer,therearemostly three,butoftentwo
orfourchromatophores, theflagellum is morethandoublethelengthofthebody. Naked
cellsare roundedin frontand conicalbackwards.
has envelopesso similarto thoseof T. volvocinathattheycannotbe
T. volvocinopsis
and sometimes
differentiated evenumbilicate ones. Deflandre(I926, p. 58) confesses:

on somespeciesofTrachelomonas grownin culture 247
Observations
'Cette espece ne differede T. volvocinaque par ces chromatophores.'There are said to
be tendevoidofpyrenoids.I am cultivating a form,congruentwithT. volvocinopsis Swir.,
and foundat Trumpington,Cambridge,in May I940, the chromatophores of which are
t(<~~~ 0U,'1'
0 \
B
Fig. I2. T. volvocinopsis Swir. A, type (8): shape and envelope structurevary. Two spherical,one of the
rarerellipsoid envelopes, and one with 'umbilicae', are represented.Three naked cells show the pear-
shaped body and the lips around the reservoiropening which are also seen in one of the encased
protoplastswhich has retractedfromthe pore. B, var. spiralis.This strain(2I) is described as a variety
althoughit shows slightdifferences
of T. volvocinopsis, fromthe type,apart fromstriationof envelope.
The size is the same, but naked cells are more elongateand metabolic. Envelope shows spiral striation
by lines of irregularwidth, sometimesfinerones alternatingwith broader striae.
each possessed of an innerpyrenoid(Fig. I2 A), and also a varietyT. volvocinopsis n.var.

spiralis(Fig. I2B), collectedin August I943 in a ditch near Waterbeach,Cambridge,
havinginnerpyrenoidstoo. Deflandreappearsto have seen the same form,which(p. 58
and fig.41) he givesas T. varians,var. spiralis. Accordingto the figureit has no tube, as
T. varians should have, and the appearance was evidentlyvery similarto my form.
Dimensions are not given.

Some doubt remainsin respectto T. variansDefl. Deflandre(1926, p. 58) places it
nextto T. volvocinopsis,whileearlierauthorsconsideredit to be a varietyQfT. volvocina.
Deflandredoes notseem to have seen it and appears to relyon Swirenkoforthe lack of
pyrenoids. In this he is right,as alreadyconfirmedby Lund (I937). Anotherfeatureof
T. variansis thetube ornamentingthe openingof the envelope. Similar structuresare,
however,picturedfor a numberof formsrelated to T. volvocina. They are given in
Middelhoek'sfigures(I951) of T. variansbut are notsufficiently
knownin theotherforms
to ascertainwhethertheyare of the same constructionthat,in T. varians,is character-
ized by the connexionbetweenthe tube and the ring(cf. Fig. i8, p. 258). In specimens
with a strongstainingby manganesethese ringsand tubes are likewisebrown.
The Trachelomonasbemardinensisgroup
Several strainsisolatedwere giventhe name T. bernardinensis, althoughtheywere not
quite in agreementwith the diagnoses given by Vischer (1915) and Deflandre(1926,
p. i i8). This was done because no otherspecies seems to be more similar,and because
the differences concernthe cell body, not the envelope,and may have been difficult to
observe in the originalmaterial.
Differencesfromthe descriptiongiven by Deflandreare: (i) the greaternumberof
chromatophores,namelymore than a dozen instead of 7-10; (2) T. bernardinensis is
supposed to have no pyrenoids,while all my strainshad them; (3) the flagellarlengthis
describedas being i-Ii timesthe lengthof the envelope,while my strainshad a much
longerone; (4) manyindividualsare more roundishthan those depictedby Deflandre
(I926, pl. 12). On the otherhand, some importantcharacteristics fitwell, forinstance
the lemon-shapedenvelope,the relativelyhigh,oftensomewhatirregularcollar,thesmall
cone at the posteriorend, and the sculptureof the envelope(Fig. 13).
As regardsthe differences the followingconsiderationsmaybe pertinent.The number
of chromatophores is best determinedby thatof the pyrenoidswhichwere not observed
by the earlierauthors.The flagellumis thickbut fragile. It can, in small hangingdrops,
be clearlyseen in its whole length,sometimeseven being straight. If, however,it is
treatedwith osmic vapour or iodine it breaks offat varyingplaces. Only by adding
strongosmium tetroxidesolutionto the preparationcan the flagellumbe fixed in its
entirelength. Althoughthe majorityof the individualsin a populationwere usuallynot
as long as previouslygiven, many had the shape recordedby Vischer and Deflandre.
Furthercharacteristics of T. bernardinensis
are: the eye-spotis largeand deep red; few
specimenswerefoundwithprotoplastsdetachedfromtheenvelopes;thechromatophores
are up to fifteen in number,discoid,sometimeslobed, the pyrenoidsprotrudingtowards
the interiorand covered by cup-shaped paramylonbodies. The tail-cone is always
present,althoughit maybe small. The hindmostpartof the envelopehas two layers,an
innerhyalineand thinone, coveredby the thicker,brownouterlayerwhich looks as if
brokenthroughby the innerone at the base of the cone (Fig. 12). This is also illustrated
by Deflandre'sfigs.657 and 674. Sometimesthe cone appears to be 'open' at the end,
and sometimesit is reducedto a low brownthickeningof the envelope. Dimensionsare
30-36: I8-21 It. The collar is 4-5t high and 5-6 t wide.
This speciesmightpossiblybe placed as a varietyto T. hispida.The latterhas normally
no cone, but var. acuminatais 'acuminee plus ou moins au pole posterieurou se forme
une esp&cede papille ou un leger cone mince', which were actuallyseen in two of the

Observations
strainscultivated(cf. p. 241), where it has exactlythe same appearance,withthe inner
layerbreakingthroughthe outer (cf. Deflandre,1926, figs.222a, b). Then thereis the
whichis wantingin thetypicalT. hispida,but is almost
distinctcollarof T. bernardinensis
as well developed in var. crenulaticollis (Maskell) Lemm. (Deflandre, fig. 223). No.
variety,it is true,has been describedshowingboth featurescombined,but judging by
@/.E X
-, .~~~~~~~~~.......
Fig. I 3. T. bernardinensis Vischer. Normal cells with detached protoplast and variations in shape, collar
and tail-cone. Cells with long, fine spines and naked ones, like those represented, are rare.
would not seem to be so widely separatedfrom

available diagnoses, T. bernardinensis
T. hispidaas its positionin Deflandre'ssystemseems to indicate,if it had not inner
pyrenoidswhich show that thereis no true relationshipto T. hispidawith its double-
sheathedpyrenoids.
The firststrain of T. bernardinensis was collected in a small pond not far from
Prague; similarones were foundin Britain(clay pond near Wray Castle, Windermere,
Wales,May I95I).
May I947, footprint, For goodgrowthno calciumcarbonate
must
be presentin thesubstrate.By addingstarchor cerealgrainsricherculturesare obtained.
These showwell the ornamentation of the envelopewhichis givenas denselyscrobulate,
and in theformastriataas delicatelypunctateand possessedofsomewhatspirallyarranged
striae Both kindsof ornamentsare foundin the same clone; theirdifference is therefore
of no taxonomicsignificance.

Three of mystrains,whichmaynot be completely identical,weredetermined as
T. lefevreiDefl.,although theinformation is notsufficient to be quitecertain.Theyare
derivedfrommaterial collectedat Shelford nearCambridge in AprilI943, froma cattle
footprintat the edge of a weedypond nearRadnorForestin Wales,AprilI95I, and
froma cattlepond (Leicestershire) neartheroadbetweenCorbeyand Uppinghamin
JuneI95I, respectively. Like thepreceding speciesit growsbeston one barleygrain
coveredwithsoiland water.
The envelopeis ellipsoidto ovaland sometimes relatively broad(Fig. I4). Whenwell
maturedit is reddishbrownand punctatedwithwhatappearto be fineholes. The
flagellar
openingis surrounded by a well-pronounced low collarwhichis generally
cylindricalbutmay,as an exception, be funnel-like or conicalwitha diameter smallerat
theopeningthanat thebase. In rarecasestheenvelopeis ornamented withirregular
zigzaglines.The posterior pole is eitherbroadlyrounded,or it bearsa brownknob,
a diminutive cone,or an irregular brownoutgrowth. No breaking through ofthetail-
cone,as described forT. bernardinensis,wasseen,butthisdifference mayonlybe a matter
of degree.
The cellbodycontains up totwenty discoidorpolygonal chromatophores withstrongly
protruding innerpyrenoids. The protoplast mayloosenits adherenceto theenvelope.
Naked,elongatedellipsoidalcells,and double-celled monsters wereobserved(Fig. 4).
This formis different fromT. bernardinensis in its smallerdimensions, particularlyin
length, and in theabsence,partlyresponsible fortheshorter shape,ofa well-developed
tail-cone;and fromT. hispidaby its conspicuouscollarand, of course,by its inner
pyrenoids. The relation ofT. lefevreiwithT. bernardinensis is notindicated inDeflandre's
monograph, wherethelatteris groupedamongthe' Caudatae',andtheformer amongthe
'Rotundatae',morespeciallythe'Ampulliformes', characterized by a collarofa height
seldominferior toitsdiameter, although inthediagnosis ofthespecies(p. I03) itis stated
to havea 'col peu eleve'. The differences do notappeartherefore to be clearenoughto
basemajortaxonomic groupsonthem.Whilethesimilarity betweenT. bernardinensis and
T. lefevreiis obvious,thereis a thirdspecieswhichI wouldliketo givea placenearto
them,although itsaffinity
is notbeyonddoubt.This formwasisolatedfroma pondnear
Debden (see p. 245) witha mixedflagellate. population,mainlyof Euglenineae,in
May I945 andcouldnotbe identified withanyknownspecies.Therefore itwillbe called
T.:pertyin.sp. It strikestheobserverby its broadlyellipsoidshapeand wideannular
thickening of 7-8p in diameteraroundthe flagellar opening,althoughmoreelongate
shapeswereoccasionally observed(Fig. I5). Dimensionsare 28-3I: 20-2I P. There
may be some relationto Playfair'sinsufficiently known T. granulosa(Deflandre,p. 95)
which,however,is smallerand does not seem to have such a wide ring. Otherwisethere
seem to be similaritiesas faras descriptionsand figuresgo. Concerningthe 'grana' which
gave the speciesits name,Deflandreasks 'ou perforeesfinement?'This is the impression
of the envelope of T. pertyilike that of T. lefrevrei:it seems to have denselysituated
perforations (Fig. I4). In addition,thereis a fineoblique striationwhich,however,may
belongto the periplast.This could not be decided because the protoplastalwaysadhered
to the envelope,and veryfew naked cells were seen. T. pertyimightbe confusedwith
T. hispidaforits generalshape and dimensionsand the envelope devoid of collar and
tail-cone,but forits innerpyrenoidsattachedto about ten to twelve large,deep green
chromatophores which are oftenso close togetheras almostto touch each other.

on somespeciesofTrachelomonas
Observations grownin culture 251
0~~~~~~~~
0 0
Fig. 14. T. lefevreiDefl. A, one individual with thick envelope and detached protoplast,one with thin
envelope, one with funnel-shaped collar, and one naked cell. B, various shapes, showing the great
diversityin outlines,collar and tail-cone.

If we take T. lefevreias having,comparedwith T. bernardinensis,a reducedcollarand
cone, T. pertyicould be consideredto go fartherin the same direction,althoughthis
notion shall not representan evolutionaryline which might have proceeded in the
oppositedirection.There mayalso be a relationof T. manginiDefl. to the presentgroup,
particularlyto T. lefevrei,providedthatits protoplasmicstructureis the same as thatof
only in its curvedcollar.
T. similis(see p. 246) whichis different
r ' f =0uNCS
0 130
o0 O 0
Fig. I5. T. pertyin.sp. Figures showing envelopes with wide pore,

chromatophores,pyrenoidsand naked cells.
The Strombomonasgroup
There are a number of Trachelomonas-like organisms,the 'Saccatae' according to
Deflandre's(I926) nomenclature, whichdifferfromthe restin havingcollarsand some-
titmeshollow tail-conesnot sharplydefinedbut with a gradual transitioninto the main
body of the envelope. Gordienko(I929) investigateda richmaterialof theseformsand
found intermediatesteps connectingT. conspersaPascher with T. zmiewikaSwirenko,
whichhad untilthenbeen supposed to be taxonomicallydifferent, mainlyin dimensions
and tail-cone. He proposed,therefore, to includeseven species into one. Lackey (I940)
uses fora group of severalsupposedlydifferent species similarto thosejust mentioned
the name T. urceolata,because his materialcomprised a large intergradingseries of
forms,afterPlayfair(I9I5) had alreadytried to make a taxonomicgroup of them.
Deflandre(I930) introducedfor the whole of his previous Saccatae the genericname
Strombomonas, holdingthemto be sufficiently different to be separatedfromTrachelo-
monas,in having a pliable, not brittleenvelope withoutspines or other well-defined
ornamentation, and a wide frontalpore.
Only one strainof this group was available formy investigations.It was isolated in
August I943 from a pool near Cambridge where it was accompanied by species of

Observations
Chlamydomonas, Chlorogonium, Cryptomonas and a colourlessPeridinieacean. It grows
abundantlyin cultureswith soil and water,especiallywhen enrichedwith ammonium-
magnesium-phosphate. Well developed envelopes are not readilyobtained.With addi-
tional organicfood, such as cereal grains,covered withsoil, it tends to live in a naked
state. Brown envelopes are mainlyfound emptyon the bottomor with sluggishcells.
They are pliable,generallycoveredwithdarkbrownparticlesofa manganesecompound,
and otherwisealmostcolourless.They containlittleiron.
Since pure cultureswerenotobtained,a deeperinsightintothe conditionsof envelope
formation was notgained.Whilea percentageoftheenvelopesundervariedcircumstances
regardingpH, humus-content,illuminationand temperature,are almost uniformin
shape, theirdiversitymay be extraordinary (Fig. i6A). It was not, however,possible
to obtain a definiteappearance at will, nor to establishunder which conditionsone or
the other prevailed. At least eightforms,which mighteasily be mistakenforspecies,
have been foundin subculturesof one clone. The correspondingfiguresin Deflandre's
maonograph are givenin brackets.The commonestformin my culturesis thatdescribed
by Pascher (I924) as T. conspersa(6i6). It was found in soil-watercultures:with soil
alone, withpeat and soil, calciumcarbonateand soil, ammonium-magnesium-phosphate
and soil, a grainof barleyand soil, peat and clay, and barleyand clay. This shows the
wide rangeof conditionsin whichthisstrainis able to multiplyand to producethetypical
shape, if thereis no competitionby otheralgae.
The T. conspersa form,generallyfoundin themajorityof specimens,is takenas charac-
teristic,and this name adopted for the strainwhich is thereforecalled Strombomonas
(Trachelomonas)conspersa(Pascher) Deflandre. It is irregularlysack- to flask-shaped,
withshortneck,not set offfromthe body of the envelope. Posteriorlyit is almostflator
slightlyrounded.The colour is a clear brownto reddishbrownor even deep sepia and
mainly(or solely?)caused by littledarkwarts.The otherformsfoundin the same clone
(Fig. i6 B) resembleT. zmiewikaSwir.(763-765) witha hollowtail-cone,T. acuminatavar.
verrucosaTeodoresco (768), T. affinis Lemm. (785), T. maskelliiDrez. (532), T. sowerbii
Skvortz.(605), and two formswhich could not be identified,one longish,shaped like
a French wine-bottle,one with a shallowwaist in the middle.
These differences are caused by the shape which the naked cell body assumesbefore
exuding the delicate primordialcase. If, for instance,the posteriorend is not fully
retracted,the envelopewill have a posteriorcone, ifit is withdrawn,the back end will be
flattenedor hemispherical.If thesides remainnearto cylindrical, the sac or bottleshapes
will originate,if theytoo are rounded,the envelopewill be egg shaped.
In cultureswithmuch organicmattertherewere greatnumbersof naked individuals,
completelydevoid of envelopes. As in otherspecies of Trachelomonas, thisphenomenon
is not caused by deficiencyin ironor manganesewhichonlypreventsthe envelopesfrom
acquiringthe brown hue, but not the thin case frombeing produced. Naked cells are
foundunderunfavourablebut nottoo harmfulconditions,and are theresultoftheproto-
plasts leavingtheirenvelopes.
The nakedcells of Strombomonas (Fig. I6 C) are different
fromthoseof mostmembers
of Trachelomonas properin being elongatespindle-shapedwitha long tail-likeposterior
end, pronouncedmetaboly,surpassingthat of any memberof Euglena, and a marked
striationalreadyobservedin T. conspersaby Deflandre(1926, p. 114). It is surprising
thattheydo not seemto have been describedas a species ofEuglena,thoughtheyare also

254
0~~~4
Fig. i6. Strombomonas conspersaDefl. (2). A, two individualswithenvelopes differentin shape, one repre-
sentingformnot yet described,the other 'Trachelomonasconspersa'Pascher. B, envelopes of various
shapes, representing:firstrow T. zmiewikaSwir., T. acuminata var. verrucosaTeod., T. affinisLemm.
and T. Sowerbii Skvortz; second row, apparentlyundescribed forms. C, naked cells with strong
metaboly.

Observations
differentin havinginner pyrenoidsnot recordedfor that genus. Anotherdifferential
characteris the positionof the reservoiropeningwhichis median as in otherspecies of
Trachelomonas and of Lepocinclis,while it is one-sidedin Euglena.
The shape of the nakedprotoplastof Strombomonas in swimmingmaybe veryslender
witha pointedend and a flagellumabout Ii timesthe lengthof the body. Mostly,how-
ever,thereis a roundishanteriorand a repeatedlytaperingposteriorportionwithundu-
latingoutlines,as in otherEuglenineae(Pringsheim,1948). The roundingup can go so
far that only a small tail is leftwhich may become almost cylindricalwith a minute
finger-shaped process. In a stronglymetaboliccell new thickenings are formedwhilethe
old ones are vanishing.The facultyof almostball-likecontractionhas some bearingon
theshapingoftheenvelope(cf.p. 254). The nucleusis alwaysbehindthemiddleofthecell
and not much largerthanthe chromatophores.In specimenswithan envelopeit almost
touchesthe posteriorsurface. In naked cells it takespositionin the inflatedpartjust in
frontof the narrowerportion (Fig. i6 C). The chromatophoresare relativelysmall,
disk-shapedand sometimesslightlylobed. The pyrenoidswiththeircoveringparamylon
cap protrudeconsiderablyinto the interiorof the cell. Division takesplace in encased,
non-motileindividuals.
Althoughonlyone clone was investigated,it appears safe to say thatDeflandre'sable
descriptionof the genus Strombomonas has to be correctedin two respects. He denied
the existenceof pyrenoids(1930, p. 556),and he describedas species formswhichare at
least in some part modificationsof a small numberof species, caused by surrounding
conditions.
Strombomonas is differentfromTrachelomonas in the followingcharacteristics:enve-
lope not brittleowingto low contentof glass-likeferrichydroxideand withno proper
ornamentationbut covered with scatteredbrown particlescontainingmanganeseper-
oxide; in motilespecimensenvelopesare generallyalmostcolourless;the broad body of
the envelope shows a gradual transitioninto the neck; the protrusionof the pyrenoids
towardthe centreof the cell is pronounced;naked cells occur more often.
Euglenachlamydophora (Mainx, 1927, p. 342), which is likewisedescribedas lacking
pyrenoids,maybelongto Strombomonas, since nakedcells,similarto thoseof S. conspersa
and envelopesof a comparabletype (loc. cit. text-fig.8) have been observed.They are
somewhatsimilarto T. kufferathi (Deflandre,1926, fig.632).
Chlamydoblepharis France (I894), among the Volvocales, is an instance parallel to
Strombomonas forthe casualtyof envelopeformation.Under circumstancesas obtaining
in pollutedpools and liquid manure,cells ofPolytomauvellaare eithercoveredwithtiny
particleswhich do not seem to be bacteria(cf. Pascher, 1927, p. 379), or with a brown
coating containingiron and manganese. Such populationswere describedby France
undervariousspecificnamesas membersof Chlamydoblepharis. They owe theirenvelopes
to the oxidation on their surfaceof manganese and iron compounds presentin the
mediumin a reducedform.When grownin pure culturein an acetate+ peptonemedium,
like those used forPolytoma,theyare no longerdistinguishablefromthe lattergenus.
Trachelomonas
deflandrei
n.sp.
A species representedby two of my strains(35, 44) is particularlyinteresting
forthe
structureof its chromatophores.At firstno pyrenoidswere detectedin its numerous
chromatophores.In a youngerculturematerial,however,denser,lens-shapedportions

were detectedin the centreof each chromatophore whichbecame more pronouncedon
theapplicationofdiluteiodinesolution.They areidenticalin appearancewiththeso-called
naked pyrenoidsof Euglena deses(Dangeard, I90I, p. I89). In a few instancessmall,
cap-shaped paramylongranuleswere foundattachedto the middle of the inner,convex
side of the chromatophores (Fig. 17), but it was not clearwhethertheyhad a relationto
the pyrenoidsunderneath.It seems remarkablethat,althoughmostof the pyrenoidsin
Trachelomonas are of the same 'inner' type,irresponsiveof the othercharacteristics of
theparticulartaxonomicform,therearethreemoretypesofchromatophores, namelysuch
with double-sheathed,naked, and with no pyrenoids(cf. next paragraph),so that the
diversityis almostas greatas in Euglena.
A B
0
00
Qo
Fig. 17. T. deflandrein.sp. A, three individuals showing chromatophoreswith 'naked' pyrenoids,para-

mylon grain next to eye-spot and relativelyshortflagellum;shape and mutual position of chromato-
phores; rare instance of paramylongrain attached to chromatophore.B, seven envelopes of diverse
shape. C, position of nucleus.
Attemptsto identifythis species with the help of Deflandre'smonographwere not

successful. The majorityof the individualsin soil-waterculturesare cylindricalwith
hemisphericalends, slightlyless rounded in frontthanbehind,withan ill-definedring
aroundthe opening,and finelypunctated.The nearestapproach,as regardstheenvelope,
appears to be T. lacustrisDrez. which,however,accordingto Deflandre(1926, p. 75),
possessestruepyrenoids,no doubtwithparamyloncaps. T. cylindrica Ehrbg.shouldalso
resembleit but has a smooth surface. T. abrupta,though similarand with the right
numberof chromatophores withoutpyrenoidsand the same dimensions,is ellipsoidal-
cylindric,i.e. more rounded than my more cylindricalform,and not punctated but
scrobiculated.Identificationbecomesstillmoreuncertainbytheoccurrence,in myclone,
of individualswhich,thoughcylindricalin the main,show a tendencytowardsa conical
shape of the posteriorend. Since, therefore, the species does not seem to be identical
with any previouslydescribed,it shall be called T. defiandrei.Most of the individuals

Observations
are 28 t long and I4t wide. The envelopeis lightbrownand thin.The chromatophores
are 5-6 t in diameter.There is oftena conspicuousparamylongrain near the eye-spot
(Fig. i6), in a way similarto thatof certainspecies of Euglena,forinstanceE. variabilis.
The species is also peculiarin the positionof the nucleus whichis not close to the rear
end, as usual, but withitsfrontedge about in themiddleofthe cell,and on thewhole not
in the medianbut attachedto one side. No otherspecies withnakedpyrenoidshas been
found.
Groupof specieswithout
pyrenoids
My previousopinion(Pringsheim,1948, pp. 51-2) thateveryspeciesof Trachelomonas
has a pyrenoidin each chromatophorehad eventuallyto be droppedwhen two of my
strainswere establishedto be devoid of them. Althoughin quite a numberof species
previousauthorshad failedto findpyrenoids,theymay only have looked forthe para-
myloncaps coveringthem,and these are oftenhidden by an excess of otherparamylon
grainsin a naturalmaterial.
Only Deflandre's (1926, p. 58) statementfor his T. varians, for which the name
T. cerviculaStokes (I890) mightperhaps be appropriate,and Lund's (I937) detailed
descriptionof the same species seem sufficiently reliable,althoughthe latter'smaterial
was at firstsuspectedas objectionablebecause it showedan interconnexion ofthechroma-
tophoresto forma network,and thisis generallyattributableto a poor stateof the cells.
These doubts were removedwhen,aftersome search,the species was found in Wales
in April 1951 in a cattlefootprintnear a pond (cf. T. bernardinensis,p. 62) and proved
to grow well in soil-watercultureswith a little peat. Lund found the envelopes
22z5-26 t in diameterbut oftenlongerthan broad. My strainis 23-26 : 20-21 p and
confirmsDeflandre'sstatementthatsphericalcells are not the rule. Like Deflandreand
Lund I found the numerouschromatophoresnear together,oftenslightlypolygonal,
thoughnottouchingone another. Lund's observationthatthe nucleusis notmedianbut
laterallysituatedwas also confirmed.He doubts whetherthe innercontinuationof the
collar is typicalof the species; I foundit always,and I am able to give a more detailed
drawingof its shape (Fig. i8), whichshowsthatit is based on a ringlike thatof T. vol-
vocina. In soil-watercultureswithclay soil and a littlestarch,whichin thisspecies as in
othersproduces the best specimens,the envelopes are particularlydark,and the tube
around the openingis a deeper brown than the rest of the envelope. Division stages
showed the two daughtercells one behind the other,so that the plane of divisionwas
normalto the longitudinalaxis, as generallyin Trachelomonas.
T. bulla Stein is the otherspecies in whichthe authorityof Deflandre(I926, p. III0),
who appearsto have seen it repeatedly,was confirmed byobservationson a materialfrom
the Station d'HydrobiologieLe Paraclet (Somme, France) where it was foundin May
1951, together withotherspecies. A mat of decayingSpirogyrain a ditchseems to have
favouredtheirgrowth. Chemical reduction,indicatedby the occurrenceof hydrogen
sulphideand sulphurbacteria,had evidentlyconditionedthe presenceof iron and man-
ganesein theloweroxidationstateand thussupportedthemultiplication of Trachelomonas.
T. bullagrowswell in soil-watercultures,particularly whensupplementedwitha grain
of barleyunderneaththe soil, in a healthy,normalstate. No pyrenoidscould be found,
not even on applicationof iodine. This species holds an isolatedpositionin Deflandre's
monograph.There is no reasonto assume a relationto T. varians: not onlythe shape of

the envelope,but also the structureof the collar,are quite different.Shape and size of
T. bulla are remarkablyconstantand in accordancewithStein's and Deflandre'sfigures.
The envelopeis egg-shaped,usuallypunctatedand at the same timefinelystriated. It is
00
00
Fig. I8. T. varians Defl. One slightlyelongate, one spherical, and one curiously aberrant individual;
arrangementofchromatophores,naked cells and, at highermagnification( x 2000), variousappearances
of pore and tube.
Fig. I9. T. bulla Stein. Left: the commonestshape; right:one of the rare individualswithhigh collar and
tail-cone,showingalso pore to be much smallerthan basis of collar; detail: pore in optical section.
stainedby neutralred and methyleneblue which is not, or not so markedly,the case in

otherspecies.The collaris.different fromthatofmostmembersofthegenusin appearing
to be the directcontinuation of the oval main partof the envelope,thoughmuch thinner
(Fig. I9). It does notin factsurroundtheflagellaropeningbut formsa frustumofa cone,
sometimesnot quite symmetrical.Its innerand widerend is closed bya diaphragmwith
a small pore forthe flagellumthroughwhich no doubt the daughtercells escape after
division.The chromatophores are much likethoseof T. varians;the nucleus is here also
not quite at the posteriorend and slightlylateral.The flagellumis shorterthan in any
other species known to me. The species seems indeed to occupy an isolated position
withinthe genus.

grownin culture 259
Observations
The twopyrenoid-free speciesare puzzling. Are they'primitive'in theirprotoplasmic
structure,or have theylost theirpyrenoidsalong withthe reductionof chromatophore
size? The chromatophores are comparableto the small discoid ones withoutpyrenoids
foundin at leasttwo independentgroupsofthegenusEuglena,thosewithrigidand those
withpliable periplasts,represented, forexample,by E. acus and E. proximarespectively.
The similarityin their chromatophoresbetween two, otherwiseseeminglynot near
related,speciesof Trachelomonas makesthe groupingoftheformswithinthegenusmore
difficult.
(3) Taxonomicconclusions
The recognitionof Trachelomonas as a memberof the Euglenaceae is based on the
structureof the protoplast.This has, however,forvarious reasons,played no role in
the identification,differentiationand groupingof the species withinthe genus: (a) the
envelopes are easier to investigatethan the delicate innerformations,especiallywhen
the cells are inadequatelypreservedor full of paramylon;(b) in well-developeddark
brownenvelopesthe cell structuremaybe obscured,whiletranslucentspecimenscannot
readilybe referredto a taxonomicunit; (c) the protoplastof the majorityof the species
seems to be uniform.A thoroughstudyof manystrainsin cultureonly can revealthe
reliabilityof protoplasmicfeaturesfordifferentiation and grouping.
All the authors dealing with the genus found their diagnoses merelyon envelope
features. Deflandre,forinstance(I926, p. 8), mentionscell structuresonly to dismiss
themas unpracticable.He makesin thisrespectno difference betweenchromatophores,
pyrenoidsand paramylon,althoughit is knownthat the amount and formof reserves
change accordingto nutritionalconditions,while chromatophoresand pyrenoidsare
permanentorganelles. Deflandredoes not seem to have realized that everychromato-
phorepossessesone pyrenoid. In statingthat'ils peuventdisparaitrecompletement'he
relies too much on casual observationwhich preventshim from apprehendingthe
taxonomicvalue of this structurein the diagnosisof the genus and the differentiation of
the species. Pyrenoidscan be concealed,but theypersist. In observingthem,the fact
is valuablethattheirparamylonsheathsare oftenmorereadilyseen thantheythemselves
are, and also easierto see thanthe outlinesof the chromatophores, so thatthe numbers
of the lattercan be foundby countingthe paramyloncaps.
In estimatingthe cell featuressuitablefortaxonomicuse, those recommendablefor
identificationofspeciesmust be distinguished fromthoseapplicableonlyforconstructing
a systemofnaturalrelationshipalthough,afterits establishment, taxonomicunitsmaybe
hoped to be recognizedby easier means. The shapes of naked cells, for instance,are
entirelyuselessfortheworkerin thefield,althoughtheymaythrowlighton relationship.
The same would be true of featuresderivedfromspecialitiesof nucleardivision,while
chromatophores and theirpyrenoidsare morereadilyobserved. In practicetwo features,
derived fromthe numberof chromatophoresand the formationof pyrenoids,proved
helpful. It seemed clear from the beginningthat the shape of the body, whether
sphericalor ovoid or more cylindrical,could not have a deeper taxonomicsignificance
in Trachelomonas any more than it has, for example, in Chlamydomonas.The same
appliesto the size and distribution of the spines or the heightof the collar,since com-
parable structuresvarymuch in otherorganismsand are also subject to environmental
influences.
Assistancefromprotoplasmicfeatureswould therefore be verywelcomein an attempt
New Phytol. 52, 3 I8

at naturalclassification.No usefulfeaturescould, however,be foundin any of the cell
organs,exceptin the number,size and shape of the chromatophores and the formation
of theirpyrenoids. So long as the greatmajorityof the species has not in this respect
been investigated,naturalarrangement must in Trachelomonas remainincomplete.The
followinggroups can tentatively be discerned:(i) large,shield-shapedchromatophores
with double-sheathedpyrenoids; (2) similar chromatophoreswith inner pyrenoids;
(3) discoid chromatophoreswith 'naked' pyrenoids; (4) small lenticularchromato-
phoreswithoutpyrenoids.There seems to be a correlationbetween size and pyrenoid
formation.
These four groups are clear enough. Every species investigatedcould, without
vacillation,be referredto one of them; but the marginsbetweenthemcut across those
marked by envelope features.This difficulty in the taxonomic use of protoplasmic
structuresis seriousin T. varianswhich,in otherrespectsis similarto T. volvocinopsis,
and in the doubtfulaffinity of the formerto T. bulla in the same group(4). A problem
is also the affinity of the Strombomonas group to the species of Trachelomonas, part of
which have the same chromatophores, whileotherwisethereare remarkabledifferences.
The similarity ofthe cell bodyof Trachelomonas withthatofEuglenais easyto observe.
It becomesstillmoreobviouswhennaked Trachelomonas cells are envisaged.Their form,
metabolyand mode of swimmingrecallimmediatelycertainspecies of Euglena. Never-
theless,thereare characteristics of differentiation.
Klebs (I883, p. 87) comparesthe protoplasmicstructureof Trachelomonas to that of
Euglenagracilis,withwhichit has actuallymuchin commonin such speciesas T. hispida,
especiallyin the shape and in the pyrenoidsof the chromatophores.Dangeard (I90I,
p. 223) emphasizes with equal justificationthe similaritybetween the cell bodies of
Trachelomonas and that of Euglena velata. E. polymorphabears, by its great flagellar
length,the large nucleus and the size of the eye-spot,which is small in E. gracilis,still
moreresemblanceto Trachelomonas, but it is different
in its superiorsize, in theendosom
whichis readilyseen afterfixation,contraryto Trachelomonas, and in its stronglystriated
conspicuousperiplast.
For defining theaffinitybetweenTrachelomonas and theothergeneraoftheEuglenaceae
the innerstructureof the cells is most valuable. Like Mainx (I927, p. 308) and Chu
(1947, p. 78) I considerthe chromatophores, so divergentamong the membersof the
family,as of special importancefortaxonomicpurposesin the Euglenaceae (Pringsheim,
I948, p. 5I), although,owingto incompleteknowledgesomemistakeshad to be corrected.
The chromatophores seem to be all of the same typein the species of Phacus,Lepocinclis
and Colaciumrespectively,while there are differences in Trachelomonas regardingthe
pyrenoidsand they differwidely in Euglena. If only featuresof chromatophoresand
pyrenoidswereused,Euglenapisciformis, E. anabaena,E. gracilis,and otherswould appear
to be nearerto Trachelomonas hispidathanto Euglenavariabilisor E. proxima,the former
having large shield-likechromatophoreswith double-sheathedpyrenoids,the latter
smalllens-shapedones withoutpyrenoidsas theyare also foundin T. variansand T. bulla.
T. deflandrei has nakedpyrenoidsin discoidchromatophores of mediumsize likeEuglena
desesand E. mutabilis, and the innerpyrenoidsof most species of Trachelomonas, which
have not been observedin Euglena,withthe possibleexceptionof Mainx's (I926, p. 157)
E. mucifera, is a warningagainstpronouncinga closer relationshipbetween Trachelo-
monasas a whole withany group withinthe genus Euglena.

Observations grownin culture 26i
AlthoughLepocinclishas some featuresin commonwith Trachelomonas, forinstance
the long flagellum,the large eye-spot,and the whole appearanceof the individuals,its
small, lens-shapedchromatophoresshow the similarityto be due to evolutionarycon-
vergenceratherthan to near relationship.In fact,Lepocinclisis nearerto Phacus, with
which it has in commonthe rigidperiplast,the type of chromatophoresand the small
number,mostlytwo, of special, conspicuous paramylonbodies. These charactersalso
betraya relationshipof boththesegenerato thatgroupor subgenusofEuglena,to which
E. spirogyra, E. tripteris,
E. oxyuris,and othersbelong.
The genus Trachelomonas, judged by its protoplasmicfeatures,is not as homogeneous
as Lepocinclisand Phacus seem to be, althougha thoroughinvestigationmay reveal
a greaterdiversitytheretoo, as the Phacuspyrumgroup makes probable. The variety
within Trachelomonas is, however,verymuch less than in the genus Euglena, so that,
if featuresderivedfromthe envelopeswere not available,or onlynakedcells were com-
pared, so that one had to relysolely on protoplasmiccharacters,not more than a few
species of Trachelomonas would be discernible. Envelope formationappears to react
stronglyto minuteconstitutional differences,generallymuch morethan to alterationin
surroundingconditions,so that shape and ornamentation of the enveloperevealdelicate
hereditaryvariationswhich,in othergenera,if recognizedat all, would be considered
to be of small importance.Only minutetaxonomicgroupswould be foundedon them,
perhaps 'formae' in Euglenaviridisand E. deses,or physiologicalvarietiesin E. gracilis.
While the genus Trachelomonas as a whole is well established,the positionregarding
taxonomywithinit is not satisfactory. The groupingof species solelyby the shape and
sculptureof theirenvelopesleads to much overlappingand confusion.The rejectionof
protoplasmicfeaturesto be used fordiscerningsubgroupsand species is not convincing.
The usefulnessof the envelope alone for taxonomyis reduced by the occurrenceof
populationscomposed of individualswidelyvaryingin appearanceand oftenconnected
by intermediatesteps. Questions connected with this difficulty will be discussed
singly.
(i) It is difficult
to decide whichoftheformsin a taxonomically homogeneousor clone
populationshall be consideredas 'the type', i.e. the nearestexpressionof the inherent
morphological ofthetaxonomic
tendencies unit..Lemmermann
(I9IO, p. 519), Conrad
(I9I6, p. i) and Deflandre(I926, p. 3) recommendpure culturesas the remedy,but
not one seems to realize that withoutdifferentiation betweenhereditaryand unstable
charactersthereis no reliablebasis fordescribingspecies at all. Even successfulcultures
oftenaffordonlya limitedporportionof well-developedenvelopeswithall the characters,
of the species definingthe type,althougheach individualwill, under equal conditions,
develop into a populationof the same mixed appearance.
(2) It is discouraging,whentryingto identifya givenmaterial,to findthatformscan
be put in two or more different places in the existingsystems,for instancethe most
perfectof all, thatby Deflandre. Actuallyone and the same species can be foundunder
two different names at places farfromone another.T. ovata Stokes cannotbe separated
fromT. hispida,to whichT. manginiand T. lefevrei mayalso be referred.This uncertainty
is due to neglectofthe principlethatonlywell-developedspecimensshouldbe used, and
to the predominantrelianceon shape.
(3) The developmentof the individualis not sufficiently known,so that stages in
maturingcannot be distinguishedfrom hereditaryvarietiesnor from modifications.
x8-2

z6z E. G. PRINGSHEIM
Althoughsinglecells could not be watcheddevelopingunderthe microscope,experience
gained frommanycultureexperimentspermitscertainconclusions.
(4) There is no definitionof taxonomicterms in Deflandre's monograph: species,
varietiesand formsare enumeratedwithoutstatingthe meaningof theseterms. In the
presentpaper a species is to be a group of individualsidenticalin essentialfeaturesand
reproducingtrue to form.A varietyis different fromthe type in one or a few inter-
connectedminor,thoughhereditary, details. A modificationis different by responding
to outerconditions.
It is notyetpossibleto setup subgenera,but nucleiofaffiliation are alreadydiscernible.
Because of the insufficient number of species investigated,the significanceof many
featurescannot be established; but one conclusioncan safelybe drawn: numbersof
similarformsmay be grouped togetherto constitutecollectivespecies which,in their
turn,when closelyrelated,formsubgenera.The numberof the latterwill be small,the
numberof minortaxonomicunitsverylarge.
Well-developedformsare readilycharacterized.T. hispidatyp. and T. volvocinatyp.
are such formswhichare mostfrequently mentioned.The characteristics of the former
are: cylindro-ellipsoidal
to ovoid envelopewithoutor withlow collar,devoidoftail-cone;
punctateto spiniferous;8-I2 chromatophores withdouble-sheathedpyrenoids.Higher
collars,unevenlydistributedspines,tail-cones,and longerspines around the pore, are
hereditary features,characterizing otherspecies,and varietiesof T. hispida.T. volvocina
is definedby sphericalor slightlyelongateenvelope; ring-shapedthickeningaroundthe
pore; envelopetwo-layered,plain or delicatelypunctate;two (or three)chromatophores
with single-sheathed,inner pyrenoids; while cylindricalstructuresaround the pore,
spiral striation,envelopeswith a largeror witha shortermain axis, as well as a greater
number of chromatophores or lack of pyrenoids,are hereditaryand exclude the form
fromthe type.
There is no confirmation oftheview of Klebs (I883, p. 88) and Deflandre(I926, p. 35)
thatenvelopesofcertainspeciescannotbe distinguished fromincompleteones ofanother
species. These statementsseem to be based on samples which did not containtypical
envelopesof eitherspecies (cf. p. 255). If indeed specimens,defectivein ornamentation,
are takenas representing thetype,thentheymaynot be distinguishable fromincomplete
specimensofanotherspecies.The mistakeis due, partlyto the difficulty of distinguishing
modificatory fromhereditaryfeatures,partlyto the notion that organismsfromthe
habitatarealwaysin a perfect
condition
(cf.,however,
Fritsch,I935, pp. 732-4).
DISCUSSION
The extraordinary diversityin theappearanceoftheenvelopesof Trachelomonas in regard
to size, shape, colour, ornamentation, collar and tail-conecan, with the help of clone
cultures,be shownto be caused, partlyby hereditary variation,partlyby the influenceof
surroundingconditions. For the establishmentof a taxonomicsystemthesetwo sets of
factorshave to be separated,and to makethe systemmorenatural,the featuresexhibited
by the protoplasthave to be takeninto consideration.When Deflandre(I926, p. 46), in
respectto these,means: 'Les particularites...sont ou tresvariables(pyre-noides, para-
mylon)ou d'une etude delicate(chloroplastes,noyaux)', one can onlyinsiston tryingto
overcomethese difficulties.

Observations
Paramylon,of course, as a reservesubstance,depositedand consumed accordingto
needs (Dangeard, I9OI, p. I78), can only in certaincases become usefultaxonomically,
forinstancethe largebodies of paramylonin Euglenaspirogyraor Phacus longicauda,or
the caps on the pyrenoidsofmanyspeciesof Trachelomonas. The tediousnessof studying
protoplasmicstructuresshould not be used as an excuse to neglectthem. Where the
difficultyof definingtaxonomicunitsis great,as it is indeed in Trachelomonas (Conrad,
I943, p. 7), everyavailablecharacteristic is welcome. Only ifthisendeavourhas achieved
a certaindegreeof perfection,can the practicalaim of identifying the formsmeet with
success. Clone-cultureexperimentsshowedthatdoubts of variousauthors(cf. Conrad,
I9i6) regardingthe taxonomicvalidityof small differences in the appearance of the
envelopes,forinstancein the sculptureof the surfaceand the shapes of the collars,went
too far. Taxonomic formsbased on them may thereforebe valid, and the extreme
lack of constancythatpreviousworkerswereafraidof, does not seem to exist,exceptin
characterswhichare readilymodifiedand depend mainlyon the availabilityof iron and
manganese.
In spite of this advance,the diversityof Trachelomonqs specimens,foundtogetherat
the habitat,is not easily explained. One reason may be that a numberof species are
generallyassociated,no doubt because oftheirsimilarity in ecologicalrequirements, and
that near related formscannot always be distinguishedby mere inspection. Earlier
observersdid not realize how numerousare small hereditarydeviationsfromthe type.
The interpretation usually given, and based on differences in age, so that the most
elaborateand dark-colouredenvelopesbelongto the oldestindividuals,probablyhaving
been inhabitedby severalgenerationsofcellsin succession,and thosewiththin,colourless
envelopesto theyoungestspecimens,is notconvincing.It is derivedfromtheexpectation
thatat least one of the daughtercells mustproducea new,at firstverydelicateenvelope,
and that,in a naturalpopulation,these can readilybe recognized.
The beliefthat a definiteappearance is always connectedwith a certaintaxonomic
constitution is notsupportedby evidence,and the hypothesisbased on differences in age
is also not in conformity with observations:(i) thereare populationswhich are more
homogeneousthanthe hypothesiscould explain,and in whichno 'old' and 'young' cells
exist; (2) afterturningabout, a cell can hardlycontinueto elaboratean envelope,for
instanceto formspines,at the same and properplace wheretheyhad been begun by the
parentor grandparentcell; (3) thickness,ornamentation, and colourof envelopesdo not
always go parallel; (4) in cultures with a great varietyin appearance, many empty
envelopes are found,while in more homogeneouspopulationsalmost none are devoid
of livingprotoplasts.
These observationscan be explainedbest on the assumptionthatlifeand reproduction
of the cell are to some degreeindependentof the developmentof the envelope. Cells of
everyspecies testedmultipliedin soil-waterculturesto formhealthypopulationswhich
weregenerallycomposedof individualswithenvelopesin variousdegreesof accomplish-
ment. During the phase of quick divisionsthe protoplastsare healthywithlittlepara-
mylonand clear chromatophores.Few of the individualsshow brown envelopes.The
betterthe nutrition, forinstancebythe additionof ammonium-magnesium-phosphate to
soil-watercultures,the more pronouncedand long lasting is this stage. Only when
multiplication declines,do betterdevelopedenvelopesbeginto be morenumerous,while
the cells are less activeand containmore paramylon.

Soil-waterculturesare not much improvedby the additionof iron and manganese
which do not appear to be deficientunder these circumstances.Pure culturesin soil-
extract,and stillmore in hay-extract media, show a much improvedgrowthon addition
offerrousiron,and producenormallyshaped and colouredenvelopesonlyifa manganous
salt is also added. These resultsshed some lighton the causes of diversitybetweenthe
individualsin a population. Afterheavymultiplication onlypartof the individualsmay
be able to acquire sufficient ironand manganeseto produceperfectbrownenvelopes,so
thatmanyremainthin and faintlycoloured.
Deficiencyin ironand manganeseis not instrumental in the occurrenceof naked cells
whichwere occasionallyobservedin almosteverystrain.They are due to unfavourable
conditionsand can be producedforinstanceby too high a concentration of manganous
salts,or by putrefaction.
From formswithincompleteenvelopeswhichrevertto normalunder suitablecondi-
tions, must be distinguishedvarietieswhich reproducetrue to pattern. Expectations
derivedfromliteraturehave been confirmed:verysmall deviationsin collars and tail-
cones and in sculpture,thoughnot so much in the exactsize and shape of the envelopes,
are hereditary.These are particularlyvariable in the Strombomonas group, where the
degreeof contractionand roundingup of the protoplast,prior to the productionof
theprimordialhyalinecase, has a greatinfluenceon the shape oftheenvelope,so thatthe
formswhich have been given specificnames have to be re-investigated to clarifytheir
taxonomicvalidity. Moreover,manyof the all too numerousspecies of Trachelomonas
are in theirdiagnosesonly based on emptyenvelopeswithoutthe organismitselfbeing
knownat all.
The use of the shape ofthe envelopesas a primarycharacteristic has separatedrelated
forms,and the emphasison ornamentation, the degreeof whichvarieswithconditions,
adds to the difficulty of placing a format itsrightplace in the system,and therefore also
of its identification.In Deflandre'stables some formsmaybe allocatedto two or three
different positions. By furtherculture experimentsa considerablesimplificationand
luciditymaybe attained,withouteverysinglespecies being investigatedexperimentally.
Ecologicallyalso muchhas stillto be done. In Klebs's monographonlya fewremarks
give some relevantinformation.He states(I883, p. 88) thatthe biologicalpropertiesof
Trachelomonas are essentiallythe same as those of otherEuglenaceae, except that they
are moreeasilyharmedby a changein conditionswhichmayalso have the effectthatthe
protoplastsleave theirenvelopes withoutprevious division. Gaidukow (I905, p. 253)
characterizesthe habitatsof Trachelomonas as being richin iron and findsthe colour of
the envelopeto dependon the ironcontentofthewater.The formerpointis also stressed
by Lemmermann(I9I0, pp. 290, 5i8). Deflandre(I926, p. 33) finds,on the contrary,
thatthe colourof Trachelomonas in waterparticularlyrichin ironis notdifferent fromthe
normalappearance,whileMainx (I927, p. 348) differs fromKlebs in his observationthat
Trachelomonasshows little sensitivityto variationin surroundingconditions.These
differences in opinionneed notimplymistakeson the partofone or the otherauthor,but
illustratethe danger,of generalizationand of the lack of insightinto the conditions
essentialforgrowth,envelopeformationand coloration. Lackey(I940) findsTrachelo-
monasin the pollutedwaterof a river,the greatestnumberoccurringin the zone where
recoveryfromputrefaction is takingplace.
In fact,Trachelomonas in generalbelongsto the Euglenaassociation,but populations

Observations grownin culture 265
are oftencomposed almost entirelyof Trachelomonas, althougha numberof species is
represented,and it may also be foundwithCryptomonadsor Volvocales withoutother
Euglenaceae.There aretherefore ecologicalquestionswhichmaybe hopedto be answered
withthehelpofcultureexperiments and comparison.There is,however,no greatdiversity
in therequirements ofthestrainsinvestigated,
althougha few,forinstanceStrombomonas,
seem to be speciallyadapted,and theconditionsforelaboratingenvelopesmaybe slightly
The apparentuniformity
different. maybe somewhatmisleadingowingto the similarity
of the habitatswhere my materialwas taken, mainly ditches and ponds in lowland.
Species planktonicin largerwaters,or inhabitingbogs and swampsmay be different.
SUMMARY
The greatdiversityof shapes and ornamentations withinthe genus Trachelomonas not
only rendersidentification but reducesthe ecologicalvalue of theseformswhich
difficult
otherwise,by the good preservability of theirenvelopes,could be eminentlyhelpfulin
manyways. As it is, the numberof problemsincreaseswiththe labour devotedto the
group,a clear sign thatit has been neglected.
Fromvariousplaces,mostlyin thevicinityofCambridge,forty-four clonesof Trachelo-
monaswere grownin soil-watercultures,some of them also freefrombacteria.These
strainsbelongto thirteenspecies. By varyingthemediumand inspectingthepopulations
aftershortor prolongedmultiplication and underdifferent conditionsoftemperature and
illumination,hereditarydeviationscould be discerned from reversiblemodifications.
Many forms,whichhad been consideredas varieties,are actuallycaused by deficiencyin
iron and manganesesupply.The formerelementis responsiblechieflyforthe structure,
the latterfor the colorationof the envelopes, though both are indispensablefor the
organismsto develop. Multiplicationin Trachelomonas is to some part independentof
the elaborationof the envelope,and so is the periodicityof division. As in otheriron
organisms,variabilityin the appearance, particularlyof the inorganic deposits, is
considerable.
A usefultaxonomicsystemcannot be based on the envelopesalone which,in many
instances,are the only knownparts of the organisms.While most of the protoplasmic
structuresare uniformin thewholegenus,thenumberof chromatophores, and especially
the existenceand structureof the pyrenoidsvaryand seem to give a clue as to natural
relationship.
laya firmer
While it is hoped thattheseinvestigations foundationforour knowledgeof
the genus,the expectationthatit would be possible,by droppingmanynames,to reduce
thenumberof taxonomicforms,did notcometrue. Whilemodifications could be cut out,
new varietieswereadded to the knownones duringinspectionof a richmaterialgathered
in natureand grownin culture.The multitudeof formsis more obvious than in other
groups,because the susceptiblenatureof the inorganiclayersof the envelopes reveals
differences which,withoutthesedeposits,could onlybe recognizedby a thoroughstudy
of physiologicaland morphologicaldetails,or possiblynot at all.

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Observations on Some Species of Trachelomonas Grown in Culture

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OBSERVATIONS ON SOME SPECIES OF

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(2) Descriptionof species

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numberof strainsisolated in culture. Material which can easily be preservedwill be

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In all theseclonesnakedcellswerealso observed(Figs. i, 6 D). They resembleto some

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Fig. 7. T. hispida. A, var. acuminataDefl., occasional occurrenceof tail-cone; protoplastbegins to leave

T. zorensisis so farthe only otherrepresentative of the firstgroup. No varietiescan

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T. volvocinatyp.was not commonin theplaces wherematerialwas collected,although

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Of the speciesof Trachelomonaswitha relatively long collar,onlyone was found

The strainheredescribed(Fig. II) iS I9-20 H: i6 t, sometimes somewhat longer,the

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each possessed of an innerpyrenoid(Fig. I2 A), and also a varietyT. volvocinopsis n.var.

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would not seem to be so widely separatedfrom

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Fig. I5. T. pertyin.sp. Figures showing envelopes with wide pore,

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Fig. 17. T. deflandrein.sp. A, three individuals showing chromatophoreswith 'naked' pyrenoids,para-

Attemptsto identifythis species with the help of Deflandre'smonographwere not

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stainedby neutralred and methyleneblue which is not, or not so markedly,the case in

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