H y d r o b i o l o g i a vol. 50, 2. pag.
i o i - i [ 1, 1976
TINTINNIDA FROM COASTAL WATERS OF THE S.W.-NETHERLANDS I.
THE GENUS TINTINNOPSIS STEIN.
C. B A K K E R & W . J . P H A F F
D e lta In s titu te H y d ro b io lo g ic a l R e se a rc h , Y erseke, T h e N e th e rla n d s , c o m m u n ic a tio n no. 127
Received February 1, 1975
Keywords: lorica characteristics, phenotypic variability, environm ental conditions.
Introduction
M o st a u th o r s h a v e b a se d th e ir in v e s tig a tio n s o f T in tin n id
sy ste m a tic s o n th e w ell k n o w n ‘C o n s p e c tu s ’ o f K o fo id &
C a m p b e ll (1929). In th is w o rk all c h a ra c te ris tic s u se d fo r
s e p a ra tio n o f th e ta x a a re d e riv e d fro m th e lo ric a , th is
s tru c tu re sh o w in g a re m a rk a b le a d h e re n c e to ty p e ( K o ­
foid & C a m p b e ll, 1939).
H o w ev e r, B ra n d t (1906) a lre a d y a d m itte d th a t a c la s­
sific a tio n o n ly b a se d o n lo ric a sh a p e a n d s tru c tu re w as in ­
c o m p le te a n d d e m o n s tra te d a rtific ia l fe a tu re s e sp e cially
in re la tio n to th e h ig h ly v a ria b le lo ric a e o f n e ritic species.
C o n se q u e n tly , L a a c k m a n n (1908) a n d M e rk le (1909)
stu d ie d th e p r o to z o a n cells o f se v e ra l T in tin n id species as
w ell a n d in c lu d e d re p ro d u c tio n p h e n o m e n a . P a rtic u la rly
H o fk e r (1931) stressed th e sig n ific a n ce o f cell stu d ie s . H is
in v e stig a tio n s e n a b le d h im to c o m p o s e a k e y b a se d on
lo ric a as w ell as o n cell s tru c tu re s . H e m a d e im p o rta n t
o b s e rv a tio n s in re la tio n to th e d iffe re n t w ay s o f b u ild in g
u p the lo ric a by T in tin n o p s is c a m p a n u la . T h is species
sh o w ed a larg e v a ria b ility in o n e a n d th e sa m e sa m p le .
H o fk e r a rg u e d th a t lo ric a e m ay be c o n s tru c te d b y cells o f
th re e fo ld o rig in : 1. a f te r b in a r y fissio n th e u p p e r d a u g h ­
terc ell, k e ep in g th e old p e ris to m e , h a s to b u ild u p a new
bow l in itia lly c o n s tru c te d a s a c irc u la r stru c tu re . 2. so m e ­
tim es a cell is lo o sin g its o ld lo ric a a n d is b o u n d to m a k e
a new one: in th is c ase, H o fk e r s ta te d , th e lo ric a m a y be
c o n s tru c te d a lo n g th e w h o le la te ra l su rfa c e o f th e a n im a l.
3. fin a lly , new cells m ay d e v e lo p fr o m re stin g sta g es a n d
a re th e re fo re fo rc e d to b u ild new lo ric a e to o . M o re o v e r,
e n v iro n m e n ta l c o n d itio n s m a y d iffe r d u r in g lo ric a c o n -
l)i . H'. J u n k h.v. Publishers - The Hague. The N etherlands
s tru c tio n . A fte r b in a ry fissio n th e old lo ric a is m a in ­
ta in e d : c o n se q u e n tly , a n u m b e r o f lo ric a e o f d iffe re n t age
is p re se n t in c o u rse o f tim e , d e v e lo p e d d u rin g d iffe re n t
c o n d itio n s o f w a te r te m p e ra tu re , sa lin ity a n d w a te r
m o v e m e n t. C h a n g e s in sa lin ity e .g ., p r o v o k in g o s m o ­
re g u la to ry a ctiv ities o f th e cells, m ig h t lead to te m p o ra ry
c h a n g e s in b o d y fo rm , re su ltin g in a b e r r a n t sh a p e o f the
new lo ric a. N o ta b ly th e a b o r a l e n d o f T in tin n o p s is b e ro i­
dea a n d T. c a m p a n u la lo ric ae is v a ry in g s tro n g ly in re la ­
tio n to its sh a p e . T h e re fo re it is n o t s u rp ris in g th a t all
tr a n s itio n s h ave b e e n fo u n d fro m s h a rp ly p o in te d to
ev en ly ro u n d e d a b o ra l p o rtio n s o f T. b e ro id e a (fig. 2-3).
In th e sa m e w ay th e d e v e lo p m e n t o f th e b u tsc h lii- a n d
c y a t h u s - fo rm s o f T. c a m p a n u la (fig. 9) is to be u n d e r­
sto o d : H o fk e r o b se rv ed th a t sw a rm in g T. c a m p a n u la
w ith o u t lo ric a w ere ro u n d e d a n d b u ilt new b u tsc h lii- o r
c y a t h u s - l t k e b o w ls. T h e u p p e r d a u g h te rc e ll d e v e lo p in g
a fte r b in a ry fissio n , o n th e o th e r h a n d , is a b o ra lly p o in te d
o rig in a lly , fo rm in g a ty p ic a l c a m p a n u la - like lo ric a w ith
pedicel.
N o t o n ly the sh a p e o f th e a b o r a l p a r t, b u t a lso the
lo ric a le n g th is e x tre m e ly v a ria b le . G o ld (1969), c u ltu rin g
F avella, d e m o n s tra te d th a t in a g e in g c u ltu re s le n g th d e ­
c re a se d s tro n g ly in c o m p a ris o n to t h a t o f sp e c im en s
fro m n a tu r a l p la n k to n sa m p le s. O ra l d ia m e te r, o n th e
o th e r h a n d , a p p e a re d n e a rly c o n s ta n t. H a lm e & L u k k a -
rin en (i9 6 0 ) m e a su re d h u n d re d s o f lo ric ae o f T in tin n o p ­
sis lo b ia n c o i D a d a v a n d a ls o fo u n d a g re a t c o n s ta n c y o f
o ra l d ia m e te r.
W e h a d th e o p p o rtu n ity to s tu d y a larg e m a te ria l fro m
sev eral s ta tio n s o f th e sea a rm s , e stu a rie s , la rg e r a n d
sm a lle r b ra c k is h in la n d w a te rs o f th e S .W .-N e th e rla n d s
d u rin g a p p r o x . 5 y e ars (see M a p ). T h e larg e v a ria b ility o f
lo ric a sh a p e , d im e n sio n s a n d s tru c tu re , stre sse d the
to i

ANTWERP
M ap o f the area investigated with sam pling localities.
n e ce ssity n e v e r to id e n tify species e x a m in in g o n ly a few
sp e c im en s, b u t to stu d y in e a c h sa m p le a s m a n y sp e c i­
m en s a s p o ssib le . D u rin g p e rio d s o f p e a k a b u n d a n c e
d e n se p o p u la tio n s w ere sa m p le d : su c h sa m p le s a p p e a re d
p a rtic u la rly su ita b le to e s ta b lis h specific lim its o f v a ri­
a b ility . E x te n siv e stu d ie s o f larg e n u m b e rs o f lo ric ae
b e fo re d e sc rib in g a species w ere a ls o m a d e b y R o ss o lim o
(1926) a n d H a lm e & L u k k a rin e n (i9 6 0 ). In c o a s ta l w a te rs
o f th e te m p e ra te z o n e species o f th e fa m ilie s C o d o n e lli­
d a e (g e n u s T in tin n o p s is ) a n d C o d o n e llo p s id a e (genus
S te n o s e m e lla ) p r e d o m in a te . In th is first p a p e r w e h ave
tre a te d o n ly th e T in tin n o p s is species. A ll fig u re s h ave
b e e n d r a w n a f te r m e a su rin g th e o ra l d ia m e te r, len g th ,
m a x im u m d ia m e te r a .s.o. D im e n s io n s w ere p lo tte d a n d
lo ric a o u tlin e s w ere sk e tc h e d ; fin ally fig u res w ere c o m ­
p lete d w ith a ll d e ta ils (incl. e n c ru s ta tio n ).
Description of the species
a. T. b a ltic a B ra n d t, (fig. ia -d ).
T h e c h a ra c te ris tic sh a p e o f a ty p ic a l T. b a ltic a is p re ­
se n te d in fig. i a. T h e lo ric a sh o w s a fla rin g m o u th , n u c h a l
c o n s tric tio n , in fla te d b o w l a n d c o n ic a l p o in te d a b o r a l
e n d , th e la tte r so m e tim e s b e in g slig h tly c o n v e x a t o n e
side a n d slig h tly c o n c a v e a t th e o th e r side. T h e o r a l rim
m a y b e d e n tic u la te .
T h e v a ria b ility o f th is species is larg e . M a n y lo ric ae
h a v e th e a p p e a ra n c e o f fig. I b a n d c, sh o w in g th e e n la rg e ­
m e n t o f th e b o w l c lo s e r to th e a b o r a l e n d t h a n th e sp eci­
m e n o f fig. i a . S o m e tim e s in d iv id u a ls a re fo u n d as
1 02
p ic tu re d in fig. i d , la c k in g a n y e n la rg e m e n t o f th e h o u se
su b o ra lly a n d g ra d u a lly n a rro w in g in a b o r a l d ire c tio n .
T h e sp e c im en s o f fig. i b a n d c so m e tim e s sh o w a n n u la ­
tio n .
D im e n sio n s: o ra l d ia m e te r: 36-48/1, m a x im u m w id th
o f th e b o w l 36-50/t; le n g th 55-106/11. T h ese d a ta a g re e w ith
th o se giv en b y B ra n d t (1906) a n d S ilv a (1950). B alech
(1948) stu d ie d m a te ria l fro m A tla n tic O c e a n w a te rs o ff
U ru g u a y a n d fo u n d sm a lle r d im e n sio n s (o ra l d ia m e te r:
30-40/i; m a x im u m d ia m e te r o f th e b o w l: 2 8 -35/i; len g th
4 2 -7 2 /i), a p p a re n tly o w in g to th e p re v a ilin g h ig h e r w a te r
te m p e ra tu re in th is s u b tro p ic a l reg io n .
T h e ra n g e o f v a ria b ility d e m o n s tra te d in fig. ia - d w as
a lre a d y e sta b lish e d b y e a rlie r a u th o r s . F ig . i a c o rre ­
s p o n d s v e ry w ell w ith B ra n d t’s (1906) fig. 6, pi. 15; w ith
B a le ch ’s (1948) fig. 22, pi. 3 a n d fig. 44, pi. 4; a n d w ith
S ilv a ’s (1950) fig. 2, pi. I. T h e m o st c o m m o n f o rm o f th e
species, fig. i b a n d c, is sh o w n to o b y B r a n d t’s (1906) fig.
9, pi. 15 a n d b y S ilv a ’s (1950) fig. 5, pi. 1. F in a lly , the
ra th e r a b e r r a n t sh a p e o f fig. i d c o rre s p o n d s w ith B ra n d t’s
(1906) fig. 15, pi. 15. W e d id n o t o b se rv e B r a n d t’s (1905)
sp e c im en s o f T. b a ltic a w ith s tro n g ly e lo n g a te d c o lla r (his
fig. 4, pi. 16).
H o fk e r (1931) w ro n g ly c o n sid e re d 7 . b a ltic a an
e lo n g a te d T. fim b r ia ta . W e w ere n o t a b le to d isc o v e r a n y
tr a n s itio n a l stag e b e tw ee n T. fim b r ia ta a n d T. b a ltic a in
th e W e s te rn S c h e ld t e s tu a ry w h ere d e n se , p a r tly o v e r­
la p p in g p o p u la tio n s o c c u r. T. b a ltic a is a p o ly h a lo b e sp e ­
cies e x te n d in g in th e m e s o h a lin e z o n e , w h e re a s T. f i m ­
b ria ta is a tr u e b ra c k is h w a te r species, c h a ra c te riz in g the
m e s o h a lin e p a r t o f th e e s tu a ry a n d e x te n d in g in to the
lo w e r re a c h e s o f th e p o ly h a lin e zo n e. T. b a ltic a B ra n d t
a n d T. fim b r ia ta M e u n ie r a p p e a r to d iffe r m a rk e d ly in
sh a p e (c o m p a re figs. 1 w ith 7), d im e n sio n s a n d sa lin ity
p re fe re n ce .
J ö rg e n s e n (1927) s u p p o s e d T. v a sc u lu m M e u n ie r, T.
strig o sa M e u n ie r a n d T. tu r b o M e u n ie r to b e id e n tic a l
w ith T. b altica. T h is m ay b e ju s tifie d , in o u r o p in io n , in
la b c d
Fig. I . Tintinnopsis baltica Brandt (ia , b, c from the Western
Scheldt, loc. 3, 5; id from the Eastern Scheldt, loc. 9).
re la tio n to T. v a sc u lu m a n d T. strig o sa . T. v a sc u lu m
h o w e v e r lac k s a fla rin g m o u th , is e n la rg e d in its u p p e r
h a lf a n d is a b o ra lly ro u n d e d . T. strig o sa , o n th e o th e r
h a n d , d o e s in d e e d re sem b le T. b altica. T h e se tw o species
( T . v a sc u lu m a n d strig o sa ) a re d e sc rib e d b y M e u n ie r
(1919) fro m a few sp e c im en s o n ly ; th e re fo re th e y a re to o
in su ffic ie n tly k n o w n to b e m a in ta in e d as d is tin c t species.
T. tu rb o , o n th e c o n tra ry , w e c o n sid e r a v a lid species (see
Pg- 12).
F in a lly th e p o s itio n o f T. n itid a B ra n d t a n d T. sin u a ta
B ra n d t h a s to b e d isc u sse d in re la tio n to T. b a ltica . M a r ­
sh a ll’s (1969) fig u re 43, pi. 2 re p re s e n tin g T. n itid a , n o
d o u b t b e lo n g s to T. baltica. B ra n d t’s (1906) o rig in a l
fig u re s 5 a n d io , p i. 15, h o w e v e r d iffe r m a rk e d ly fro m
M a rs h a ll’s o n e . H is T. n itid a th e re fo re m a y be a v alid
species. T. s in u a ta B ra n d t is la rg e r, m o re e lo n g a te d (v ase­
sh a p e d ) a n d sh o w s in fla tio n o f th e b o w l a b o v e th e m id d le
(B ra n d t, 1906, fig. 3, 4, pi. 15). A c c o rd in g to J ö rg e n s e n
(1905) T. n itid a v a rie s s tro n g ly a n d sh o w s tr a n s itio n s to
T. s in u a ta a s in d ic a te d b y his fig. 116 pi. 18. O u r p re lim ­
in a ry c o n c lu sio n is th a t T. n itid a a n d T. s in u a ta h a v e to be
u n ite d . In c lu s io n h o w e v e r o f th is species in to T. b a ltic a
seem s n o t yet ju stifie d .
b. T in tin n o p s is b e ro id e a S te in (fig. 2-8)
T h e d iffic u lties o f th e e x a c t d e lim in a tio n o f sy ste m a tic
e n titie s w ith in th e g e n u s T in tin n o p s is a re w ell illu stra te d
b y th e figs. 2 a -h a n d 3a-f. T h e lo ric a e o f b o th series h ave
been fo u n d in sm a ll p o ly h a lin e in la n d w a te rs. T h e sp e c i­
m en s o f fig. 2 w ere sa m p le d in w in te r (8-2-’72) th o se o f
fig. 3 in su m m e r (22-8-’72). T h e y d iffe r m a in ly in w id th
o f o ra l d ia m e te r: 50-60/2 (a v era g e 53,5/1) in th e w in te r
series, 40-42,5/2 in th e su m m e r series. D im e n s io n s o f th e
Fig. 2. Tintinnopsis beroidea Stein (from Inlaag 1953, loc. 13).
lo ric a m a y p a rtly d e p e n d o n w a te r te m p e ra tu re . M o stly
species a re la rg e r in th e c o ld e r p a rts o f th e ir ra n g e ( M a r ­
sh a ll, 1969). E q u a lly a species o c c u rrin g th r o u g h o u t th e
y e a r m a y re a c h la rg e r d im e n sio n s in w in te r th a n d u rin g
su m m e r. A s u s u a l le n g th s o f lo ric a e o f b o th series v aried
m u c h m o re th a n o ra l d ia m e te rs: th e le n g th ra n g e o f
series 1 w as 60-135M, ° f series 2: 50-1 io/u.
W e a ssig n b o th series to th e sam e species as th e larg e st
p o r tio n o f sp e c im en s (60-70% ) w a s c h a ra c te riz e d b y th e
id e n tic a l a p p e a ra n c e o f fig. 2d a n d 3b. T h e c o n s ta n c y in
w id th o f o ra l d ia m e te r (n o ta b ly in series 2) e x c lu d e d th e
p o ss ib ility o f th e p re sen c e o f d iffe re n t species in b o th
series.
T h e g re a t v a ria b ility o f th e species in b o th series is
d e m o n s tra te d b y sev eral c h a ra c te ris tic s . A n n u la tio n w as
so m e tim e s c le a rly v isible in th e su m m e r fo rm s (fig. 3a),
b u t in m o st cases n o t o r h a rd ly d e m o n s tr a te d (fig. 3e). In
th e w in te r fo rm s a n n u la tio n w as n o t seen a t all. S lig h t
a n n u la tio n h o w e v e r m a y b e easily o v e rlo o k e d w h e n th e
e n c ru s ta tio n w ith p a rtic le s b e c o m e s d e n s e r (H a lm e a n d
L u k k a rin e n , i9 6 0 ), w h ic h w a s in d e e d th e c ase in th e
w in te r fo rm s (c o m p a re figs. 2 a n d 3). S o m e su m m e r
fo rm s sh o w e d g r a d u a l n a rro w in g (fig. 3 t) in o r a l d ire c ­
tio n , in so m e c ases fo llo w ed b y w id e n in g o f th e m o u th
(fig. 3d). O fte n th e a b o r a l p a r t o f th e lo ric a w a s sp h e ric a l
(30-40% , fig. íe , f, 3a, c, e ,) g iv in g th e sp e c ie s a T. lo b ia n ­
c o i a p p e a ra n c e . M o re o v e r th e sp ecies d e m o n s tra te d
se v e ra l tim e s a slig h t w id e n in g o f th e ro u n d e d a b o ra l p a rt
(fig. 2 a , e, 3a, d , e). T h e m o st c o m m o n a p p e a ra n c e h o w ­
e v er is sh o w n in figs. i d a n d 3b: lo ric a a b o ra lly b lu n tly
p o in te d , ty p ic a lly fo r T. b e ro id e a S te in .
O u r a ssig n e m e n t o f th e species to T. b e ro id e a S te in is
b a se d o n th e fo llo w in g c o n sid e ra tio n s:
Fig. 3. Tintinnopsis beroidea Stein (from Schelphoek, loc. 14).
103
 1 . T h e fa irly larg e a v e ra g e o ra l d ia m e te r (4 0 - 5 4 /2 ) is in
a g re e m e n t w ith th e d im e n sio n s giv en b y D a d a y (1 8 8 7 ) :
4 5 -5 0 /2 a n d E n tz S r. ( 1 8 8 4 ) : 5 0 -6 0 /2 . O fte n h o w e v e r sm a l­
ler w id th s a re g iven: 18-36/M (M a rs h a ll, 1 9 6 9 ) ; 3 0 - 4 2 / 2 ) J o r ­
g e n se n , 1 9 2 7 ). S m a lle r d ia m e te rs (ra n g e 2 0 -4 0 /2 ) w ere
g e n era lly fo u n d in th e fo rm s o c c u rin g in c o a s ta l se a w a te r
a n d th e la rg e r in la n d p o ly h a lin e w a te rs (L a k e V eere).
T h ese sp e c im en s a re p ic tu re d in fig. 4 .
2. J ö rg e n s e n (1927, fig. 5 ) d e sc rib e d T. b e ro id e a S te in
a c c o rd in g to E n tz S r. (1884). H e w as fo llo w e d b y K o fo id
& C a m p b e ll (1929). T h e ty p ic a l a p p e a ra n c e o f T. b e ro i­
d e a S te in em . E n tz S r. em . J ö rg e n s e n (c y lin d ric a l lo ric a
a b o v e c o n ic a l a b o r a l e n d ) is sh o w n b y 60-70% o f o u r
sp ecim en s.
3. A lso th e T. b e ro id ea fig u re s o f D a d a y (1887) a re im ­
p ressiv e b y th e ir v a ria b ility . B esides th e sp ecies th is
a u th o r d istin g u ish e d th e v a r i e tie s -a c u m in a ta ,- p la g io ­
s to m a a n d - c o m p re ssa . H is d e s c rip tio n o f species a n d
v a rie tie s strik in g ly a g re e s w ith o u r fig u res. D a d a y ’s sp e ­
cies (P I. 19, fig. 2, 14) c o v ers o u r a b o ra lly ro u n d e d sp e c i­
m ens: figs. 2b, 2c a n d 3c (K o fo id & C a m p b e ll, 1929: T.
ro tu n d a ta . Jö rg e n se n ). D a d a y ’s v a r. a c u m in a ta (P I. 19,
fig. 4 ) is id e n tic a l w ith th e ty p e (K o fo id & C a m p b e ll,
1929, p. 28). S o m e o f o u r sp e c im en s re sem b le D a d a y ’s
v a r. p la g io s to m a (P I. 19. fig. 3), th o u g h he is g iv in g a
sec o n d fig u re to o (P I. 19, fig. 6) sh o w in g a m u c h w id e r
m o u th . K o fo id & C a m p b e ll (1929) ra ise th is v a rie ty to
species, b u t n o tic e th a t th is f o rm is p e rh a p s o n ly a fla t­
te n e d T. b e ro id ea S te in . T h is fo rm w as a ls o fig u re d by
E n tz S r (1884, pi. 24, fig. 1). T h e v a r. c o m p re ss a D a d a y
fin ally (P I. 19, fig. 7-9), ra ised a ls o to species b y K o fo id &
C a m p b e ll (1929), is ro u n d e d a b o ra lly a n d sh o w s a slig h t
/ . a b e d
o K a ra S e a a ssig n e d to T. b ero id ea , th e o r a l d ia m e te r v a ry ­
4e f g h
Fig. 4. Tintinnopsis beroidea Stein (4a. b, c, d, f, g from Lake
Veere. loc. 6, 7; 4e from coastal seawater, loc. 15; 4h from Inlaag
1953, loc. 13).
104
5a
f b c
Fig. 5. Tintinnopsis beroidea Stein (5a from Lake Veere, loc. 7;
5b from the Eastern Scheldt, loc. 11:5c from the Western Scheldt,
loc. 3).
n u c h a l c o n s tric tio n , a c c o rd in g to o u r fig u re s 2e a n d 3d.
T h is la tte r f o r m re sem b les T. lo h m a n n i L a a c k m a n
(1908) w h ic h K o fo id & C a m p b e ll (1929) c o n sid e re d to be
b a se d o n in c o m p le te lo ric a e a n d w as sy n o n im iz e d by
th ese a u th o rs w ith T. su b a c u ta J ö rg e n s e n (= T. lo b ia n ­
c o i D a d a y , see p. 107).
D a d a y rig h tly d e cid ed to a g re a t v a ria b ility o f T. b e ro i­
dea. H e h a d th e d is p o s a l o f a very ric h m a te ria l fro m th e
M e d ite rra n e a n (G u lf o f N a p e ls), sa m p le d d u rin g 8
m o n th s . P e rh a p s h e w as im p re ssed (as th e p re se n t
a u th o rs ) b y th e n e a rly c o n s ta n t o ra l d ia m e te r (45-50/2) o f
m o st o f his sp e c im en s (o n ly fo r h is species h e m e n tio n s
lo w e r v alues: 27-45/2). O u r sa m p le s to o c o n siste d o f dense
su s p e n s io n s o f T. b e ro id ea . S u c h sa m p le s a re v e ry sui­
ta b le to e sta b lish specific v a ria b ility ra n g es.
J ö rg e n s e n (1912) u se d B ra n d t’s (1906) fig. 5, pi. 16
(id en tifie d b y B ra n d t a s T. b e ro id e a S te in ) to d e sc rib e T.
p a rv u la . T h is fo rm w as sm a lle r a n d m o re sle n d e r th a n T.
b ero id ea . T h e m a in d iffe re n c e fro m th e la tte r w as the
slig h tly c o n s tric te d o ra l p a r t o f th e lo ric a . T h e b o w l is
w id est in its m id d le p a r t. T h e c o n ic a l a b o r a l p o rtio n is
b lu n tly p o in te d , w ith a n a n g le o f ± 6 o ° , see fig. 5.
D im e n sio n s: o r a l d ia m e te r: 18-26/2; m a x im u m w id th :
24-32/2; len g th : 49-65/2. B ra n d t (1906) m e a su re d a n o ra l
d ia m e te r o f 24/2, v. B re em en (1905): id e m 24-28/2; M e u ­
n ie r (1929, fig. 28, pi. 22, id e n tifie d a s T. b e ro id e a ) idem :
14-20/2; S c h u lz (1965): id em 21-22/2. K o fo id & C a m p b e ll
(1929) u n ite d T. f u s u s M e u n ie r w ith T. p a r v u la . T. p a r ­
vu la o c c u rs in th e B altic, a lo n g th e n o rw e g ia n c o a s t, in
th e s o u th e rn N o r th S e a a n d a d ja c e n t se a -a rm s a n d e s tu ­
arie s a n d in p o ly h a lin e in la n d w a te rs (L a k e V eere).
S o fa r, s e p a ra tio n o f T. p a r v u la fro m T. b e ro id ea
seem s ju stifie d . R o ss o lim o (1926) h o w e v e r, fig u red
tw e n ty so m e w h a t la rg e r sp e c im en s o f a species fro m the
ing fro m 31-38/2 a n d sh o w in g p a rtly T. p a r v u la sh a p e : see
fig. 6 (d e riv e d fro m R o sso lim o ). T h ese fig u re s d e m o n ­
s tra te c o n v in c in g ly a series o f tra n s itio n a l sta g es b etw ee n
T. p a r v u la a n d T. bero id ea . W e h a v e to c o n c lu d e , th e re ­
fo re , th a t T. p a r v u la c a n n o t be m a in ta in e d a s a d istin c t
species.
6 ^^^
Fig. 6. T. beroidea Stein, after R ossolim o (1926).
S o m e tim e s th e p a r v u la -fo n tis a re c le a rly ta p e rin g to
th e a b o r a l e n d (fig. 7a) a n d th e n a p p r o a c h T. rapa M e u ­
n ier. T h is la tte r fo rm , b e in g o n ly slig h tly n a rro w e r (fig.
7b ), th u s m a y fa ll w ith in th e b e ro id e a -p a r v u la series to o .
F in a lly , we p u t to g e th e r all v ery sm a ll c y lin d ric a l lo ri­
cae, h a v in g a n o ra l d ia m e te r o f ± 15p (H o fk e r , 1922, fig.
81; v a n B re em en , 1905, fig. 16; L o h m a n n , 1908, fig. 12,
PI. 17; M e u n ie r, 1919, fig. 21, PI. 22), a n d sh o w in g a
slig h tly p o in te d a b o r a l e n d , a s T. n a n a L o h m a n n . O u r
sm a lle st sp e c im e n even m e a su re d 8 p , o ra l d ia m e te r (fig.
8b). T. m in u ta W ailes (fig. 8a) d iffe rs fro m th is species
o n ly in re la tio n to le n g th a n d th e s h o rte r a n d m o re ro u n ­
d e d a b o r a l e n d . T h ese c h a ra c te rs , a g a in , a p p e a r very
v a ria b le a n d th e re fo re a re in su ffic ie n t to m a in ta in T.
m in u ta as a d is tin c t species. K o fo id & C a m p b e ll (1929)
a lre a d y id e n tifie d T. fis tu la r is M e u n ie r as T. n a n a . C le a r
se p a ra tio n h o w e v e r o f T. n a n a fro m la rg e r sp ecies, n o ta ­
bly T. b e ro id ea , is n o t p o ss ib le , b e c a u se n o t o n ly T. n a n a
h a s a - b e r o id e a sh a p e , b u t a ls o th e re is a g ra d u a l tr a n s i­
tio n of -n a n a - to - b e r o id e a - d \m e n s io n s . S m a lle st
b e ro id e a -lo ric a e m ay sh o w o ra l d ia m e te rs b e lo w 20p;
th u s , th e b o u n d a ry b e tw e e n T. b e ro id ea a n d T. n a n a is
q u ite a r b itra r y . W e n a m e d th e sp e c im e n o f fig. 8c T. n a n a
o w in g to its e lo n g a te d c o n ic a l a b o r a l e n d , b u t w e m ig h t
c lassify th is fo rm a s w ell a s T. b e ro id e a in view o f th e high
d e g ree o f v a ria b ility o f th e sh a p e o f th e a b o r a l e n d o f the
la tte r species.
O n th e o th e r h a n d it seem s re a s o n a b le to s u p p o s e th a t
th e m in u te fo rm s, a lth o u g h - b e r o i d e a - lik e in a p p e a r a n ­
ce, a re in fa c t n o t T. b e ro id ea . B ut th e e v id e n ce c a n n o t be
given o n th e basis o f d iffere n ce s in lo ric a s tr u c tu r e a n d -
d im e n sio n s. C u ltu re e x p e rim e n ts a n d su b s e q u e n t inves-
o s tra te d
-4-
7a b d u a lly (fig. 9a) o r a b r u p tly (fig. 9b ). P e d ic e l ru n s m o stly
Fig. 7. Tintinnopsis beroidea Stein (7a from the Eastern Scheldt,
loc. io , 7b from Lake Veere, loc. 7).
i 9
8a b c
Fig. 8. Tintinnopsis beroidea Stein (8a from Lake Veere, loc. 6,
8 b from S ch elp h oek ,loc. 14, 8c from the Eastern Scheldt, lo c .12).
tig a tio n o f th e p ro to z o a n cells m a y re v ea l e v e n tu a l re al
d iffere n ce s b e tw e e n n a n a - a n d b e ro id e a -fo rm s.
c. T. c a m p a n u la (E h rb g .) D a d a y (fig. Ça-g)
T h e v a ria b ility o f th is c o m m o n species is v e ry la rg e as
w as d e m o n s tra te d b y H o fk e r (1931). T h is a u th o r in ­
clu d e s th e fo llo w in g fo rm s in to T. c a m p a n u la : T. b u ts c h ­
lii D a d a y , T. c in c ta (C la p & L a c h m .) D a d a y , T. c y a th u s
D a d a y , T. e lo n g a ta D a d a y , T. in fu n d ib u lu m D a d a y , T.
lin d e n i D a d a y a n d T. u rn ig e r (E n tz S r) D a d a y . A ll these
species w ere c o lle cted to g e th e r in th e B ay o f N a p le s. A lso
J ö rg e n s e n (1899, 1927) id e n tifie d T. b u tsc h lii, T. c in c ta
a n d T. c y a th u s, o c c u rrin g in th e N o r th S e a a n d th e B altic,
w ith T. c a m p a n u la . A lre a d y v a n B reem en (1905) d e a lt
w ith a T. c a m p a n u la - c i n c t a - b u t s c h l i i - series. M a rs h a ll
(1969) s u p p o s e d th a t T. m a jo r M e u n ie r (A rc tic , N o r ­
w eg ian S e a , B altic) to o m ay b e lo n g to th is g r o u p . W e
a g ree w ith th ese a u th o r s a n d re fe r to H o fk e r (1931) fo r
a m p le a r g u m e n ta tio n . T. e c a u d a ta K o fo id & C a m p b e ll
(N o rw e g ia n S e a ) w as a lre a d y d e sc rib e d b y B ra n d t (1906)
a s T. c a m p a n u la v a r. a.
E x c e p t th e c h a r a c te r is tic - c a m /> a « « /a - f o r m s (fig. 9a-
c), w e f o u n d - m a j o r - (fig. 9d -e) a n d - b u t s c h l i i - (fig. 9!)
-s p e c im e n s . F ig . 9g re p re se n ts a d e v e lo p m e n ta l sta g e o f
a c h a ra c te ris tic c a m p a n u la . T h e lo ric a o f th e ty p ic a l
fo rm is c a m p a n u la te a n d p ro v id e d w ith a w idely fla rin g
m o u th a n d a s to u t ped icel (fig. 9 a-c). E v en th is c h a r a c ­
te ristic fo rm sh o w s a larg e v a ria b ility , a s d e m o n s tr a te d
b y a series o f fig u res fro m B ra n d t (1906). T h e b o w l is e n ­
la rg in g g ra d u a lly to th e o r a l p a rt. T h e o r a l re g io n m ay
sh o w a m o re o r less c le a r a n n u la tio n (fig. 9d). T h e e n ­
c r u s ta tio n w ith p a rtic le s m ay be d e n se a n d r a th e r re g u la r
(fig. 9a) o r m ain ly lim ite d to th e a b o ra l p a r t w ith pedicel
(fig. 9b, c, g). M a jo r - a n d b ü ts c h li i - ïo r m s dem on­
sp a rse a n d irre g u la r e n c ru s ta tio n (fig. 9d-f).
T ra n s itio n b e tw e e n b o w l a n d p e d ic el m ay p ro c e e d g r a ­
s tra ig h t b u t so m e tim e s sh o w s to r s io n ( B r a n d t, 1906;
M e u n ie r, 1919). S p e c im e n s w ith o u t p e d ic el m a y be
105
 d. T. c ylin d rica D a d a y (fig. i o )
A c c o rd in g to K o fo id & C a m p b e ll (1929) w e b a se th e
id e n tific a tio n o f th is species o n th e o rig in a l d e s c rip tio n o f
D a d a y ’s (1887) T. d a v id o ffi var. c y lin d ric a (D a d a y ), pi.
19, fig. 24).
T. c ylin d rica is c h a ra c te riz e d b y a n e lo n g a te d lo ric a ,
th e a n te r io r p a r t b e in g c y lin d ric a l f o r 75-80% o f th e to ta l
len g th . T h e a b o r a l p a r t n a rro w s to a s to u t p e d ic el, slihgtly
9a b c v a ry in g in le n g th a n d w id th . T r a n s itio n o f th e c y lin d ric a l
p a r t o f th e lo ric a in to th e p e d ic e l p ro c e e d s o fte n g ra d u a lly
b u t so m e tim e s a b ru p tly . E n c r u s ta tio n w ith c o a rse p a r ti­
cles is e sp e cially seen in th e a b o ra l p a r t a n d p ed icel. In
L a k e V eere th e g re a te r p a r t o f th e lo ric a is c o v e re d w ith
very sm a ll p a rtic le s (clay p a rtic le s a n d /¿-cells). T h e o ra l
d ia m e te r o f o u r sp e c im en s is strik in g ly c o n s ta n t in tim e
a n d sp a c e, v a ry in g o n ly b e tw e e n 35-42, m o st lo ric a e m e a ­
9d e f g su re d 36 d ia m . L e n g th v a rie d fr o m 120-223 U-
Fig. 9. Tintinnopsis cam panula (Ehrbg) D aday (9a, e, f, g from T h e species o c c u rs in E a s te rn S c h e ld t, W e s te rn S c h e ld t
the W estern Scheldt, loc. 4, 2, i; 9b, c, d, from the Eastern
a n d L a k e V eere, p re fe rin g sa lin itie s h ig h e r th a n io % o C 1’.
Scheldt, loc. 8 ,9 ).
H ig h e st n u m b e rs a re re a c h e d in su m m er.

evenly ro u n d ed a b o ra lly ( b u t s c h li i - fo rm s, fig. 9 f; e. T. fim b r ia ta M e u n ie r (fig. 11 a-c)

c y a th u s -fo rm s) o r a n g u la r (m a jo r -io r m s , fig. 9d, e; T. fim b r ia ta is a v ery c h a ra c te ris tic species, sh o w in g a
c in c ta -fo rm s). B u ts c h lii-fo rm s have a m o re fla rin g c o m p a ra tiv e ly slig h t v a ria b ility . B ra n d t (1906) fig u re d a n
m o u th th a n c y a th u s -fo rm s; tw ia -fo rm s a re n e a rly u n d e sc rib e d species (figs. 5, 7, PI. 17 a n d fig. io , PI. 18),
c y lin d ric a l th ro u g h o u t th e ir len g th . For fig u res of d o u b tle ss id e n tic a l w ith T. fim b r ia ta M e u n ie r (H o fk e r,
c y a th u s- a n d c in c ta -fo rm s: see M a r s h a ll (1969). T h e re ­ 1931). K o fo id & C a m b e ll (1929) w ro n g ly d e sc rib e d th is
fo re d ia g n o s is o f T. c a m p a n u la c a n n o t b e b a se d o n p re ­ fo rm as a n ew species: T. m e u n ieri.
sence o f a p e d ic el. H o fk e r (1931) gave a s a tis fa c to ry T h e c a m p a n u la te lo ric a n a rro w s in to a s tr o n g n u c h a l
e x p la n a tio n fo r th e o rig in o f fo rm s w ith o u t p e d ic el (see c o n s tric tio n . T h e m o u th is fla rin g to n e a rly th e d ia m e te r
in tro d u c tio n ). o f th e b o w l. T h e o ra l rim is c h a ra c te ris tic a lly ra g g ed .
D im e n sio n s: M a x im u m o ra l d ia m e te r: 70-88/1; d ia ­ T h e a b o r a l e n d is alw ay s p o in te d (fig. n a ) , so m e tim es
m e te r ± h a lfw a y th e lo ric a: 40-56/1; le n g th in c lu d in g very s to u tly (fig. 11b), so m e tim e s o n ly slig h ty (fig. n c ) .
p edicel: 135-172/1; le n g th w ith o u t p ed icel: 110-130/1. L o ric a d e n se ly e n c ru ste d w ith c o a rse p a rtic le s, a ty p ic a l
O c c u rra n c e : N o r th S e a , B altic, M e d ite r a n e a n S e a ,
A tla n tic O c e a n (Jö rg e n s e n , 1927; H o fk e r, 1931 ; M a r ­
sh a ll, 1969). In th e W e s te rn S c h e ld t e s tu a ry th e species is
c o m m o n ly fo u n d th r o u g h o u t th e p o ly h a lin e a n d w ith in
th e a - m e s o h a li n e z o n e, d e c re a s in g ra p id ly in th e
b ra c k is h w a te r. T h ese d a ta a re in a g re e m e n t w ith th o s e o f
H o fk e r (1922, 1931) a n d v a n B re em en (1905), tr a c in g th e
species in th e n o r th e rn , m o re sa lin e p a r ts o f th e fo rm e r
Z u id e rz e e o n ly (n o t b e lo w 8% c, C I ’) a n d w ith th o s e o f
S c h u lz (1965) o f th e E lb e e s tu a ry . T. c a m p a n u la is a su m ­
m e r fo rm : m a x im u m a b u n d a n c e o c c u rrin g d u rin g th e
p e rio d o f J u ly to O c to b e r (3 0 -4 0 /I, sa lin ity i4 ,3 -i7 ,2 % o
C l’). W e c o u ld n o t y e t d e m o n s tra te th e species in L ak e 10a b
V eere, th o u g h c h lo rin ity in su m m e r m a y rise fro m io to Fig. io. Tintinnopsis cylindrica D aday (10a from the Eastern
i4 % o . Scheldt, loc. io; 10b from Lake Veere, loc. 7).

106

_____ _ 3,
<' CD
(N
Il a b c
Fig. i l . Tintinnopsis fimbriata M eunier (from the Western
Scheldt, loc. i , 2, 3).
p h e n o m e n o n f o r T in tin n o p s is - s p e c ie s o c c u rrin g in t u r ­
b u le n t e s tu a rin e tid a l w a te rs w ith h ig h s u s p e n d e d m a tte r
c o n te n ts.
D im e n sio n s: o ra l d ia m e te r: 44-68/u; m a x im u m w id th
o f th e bow l: 48-68/u; le n g th 56-88/u. A v e rag e v a lu e s show
th e te n d e n c y o f d e c re a s in g d im e n sio n s w h e n w a te r te m ­
p e ra tu re is rising. D u rin g J u ly 1974, w a te r te m p . 17,7-
19,2 ° C , a v e ra g e d im e n sio n s w ere (36 sp e c im en s): o ra l
d ia m e te r: 55/2; m a x im u m w id th : 61/2; len g th : 74^2. A fte r
a m o n th , A u g u s t 1974, w a te r te m p . 19,5-20,4° C , a v era g e
v a lu e s w ere (57 sp ecim en s): o r a l d ia m e te r: 52,5/2: m a x i­
m u m w id th : 58/2; len g th : 71.5/2.
O b s e rv a tio n s o f H o fk e r (1931), in d ic a tin g th a t sp e c i­
m en s g rew lo n g e r w h e n sa lin ity in c re a s e d w ere n o t c o n ­
firm e d b y o u r d a ta . M e a s u re m e n ts o f 57 sp e c im en s
fro m 3 s ta tio n s w ith d iffe re n t c h lo rin itie s (7,6; 10,6;
12,2% o) re su lte d in a n e a rly c o n s ta n t ra tio o f len g th to
m a x im u m w id th (1,25; 1,26; 1,23 resp ectiv ely ).
H o fk e r (1931 ) sta te d th a t T. b a ltic a B ra n d t m a y be
id e n tic a l w ith T. fim b r ia ta . A fte r o u r o p in io n th is is n o t
ju stifie d : see p. 102.
M e u n ie r (1919) d e sc rib e d th e species fro m b ra c k is h in ­
lan d w a te rs in B elgium , o c c u rrin g a b u n d a n tly th e re . In
c o a s ta l w a te rs o n th e o th e r h a n d h e fo u n d th e species
sp a rse ly , in a g re e m e n t w ith H o fk e r (1922) a n d w ith o u r
o b se rv a tio n s. W e fo u n d in c id e n ta l T. fim b r ia ta in th e
p o ly h a lin e z o n e o f th e W e s te rn S c h e ld t e stu a ry . W e d id
n o t o b se rv e th e species in th e e u -p o ly h a lin e E a s te rn
S c h e ld t. B ra n d t (1906) fo u n d th e species in th e K iel B ay
a n d th e K a ise r W ilh e lm C a n a l. T. f im b r i a ta p ro v e s to be
a c h a ra c te r species o f th e (m ix o ) m e s o h a lin e z o n e o f th e
W e s te rn S c h e ld t e s tu a ry d u rin g su m m e r (B a k k e r & D e
P a u w , 1975). T h e species o c c u rre d a b u n d a n tly to o in th e
s o u th w e ste rn (m e so h a lin e ) p a r t o f th e fo rm e r Z u id e rze e
(H o fk e r, 1922; 1931). A c c o rd in g to S c h u lz (1961; 1965)
th e species lives in th e E lb e e s tu a ry to o , b u t n o t in g re a t
n u m b e rs .
f. T. lo b ia n c o i D a d a y (figs. 12 a-d, I 3)
A c c o rd in g to th e o rig in a l d e s c rip tio n o f D a d a y (1887),
T. lo b ia n c o i is a c y lin d ric a l, e lo n g a te d sp ecies o f larg e
le n g th (270 ju) a b o ra lly ev en ly ro u n d e d a n d sh o w in g a n
o ra l d ia m e te r sim ila r to th a t o f th e lo ric a: 45/2. M o s t d a ta
a g ree in re la tio n to th e w id th o f th e o ra l d ia m e te r. A larg e
m a te ria l fro m th e B altic (G u lf o f F in la n d ) w as th o r o u g h ­
ly in v e stig a te d b y H a lm e & L u k k a rin e n (i9 6 0 ). T h ey
e sta b lish e d a ra n g e o f 30-55 ß o ra l d ia m e te r, 90% o f th e
lo ric ae h o w e v e r fa llin g w ith in a ra n g e o f 35-45 p . S o m e
o th e r d a ta : B r a n d t (1906): 40-50 p ; B ie rn a c k a (1948): 46-
50 72; v a n B re em en (1905): 45 ju; M e u n ie r (1910): 36-40 p;
o w n m e a su re m e n ts; 37-47 ß . L e n g th v a rie s fro m 30-300
¿2, (H a lm e & L u k k a rin e n ); 220-390 ß (B ra n d t); 156-170 ß
(v a n B reem en ); 100-140 ß (M e u n ie r); o w n m e a s u re ­
m en ts: 144-210 ß .
T h e species sh o w s a larg e v a ria b ility . J ö rg e n s e n (1927)
c o n sid e re d T. lo b ia n c o i D a d a y a n d T. tu b u lo s a L e v a n d e r
d is tin c t species, T. tu b u lo s a d iffe rin g fr o m T. lo b ia n c o i
by a n n u la tio n a n d a b o r a l w id e n in g o f th e lo ric a . M e rk le
(1909) h o w e v e r, g ave a fig u re o f T. lo b ia n c o i w ith very
c le a r a n n u la tio n . N o ta b ly H a lm e & L u k k a rin e n d e m o n ­
s tra te d c o n v in c in g ly th a t tu b u lo s a - and lo b ia n c o i-
a p p e a ra n c e o c c u rre d in th e sam e m a te ria l, sh o w in g all
p o ssib le in te rg ra d a tio n s . In th e ir sa m p le s a b o u t a th ird o f
a ll sp e c im en s sh o w ed lobiancoi-appea.ra.nce, th e re m a in ­
d e r d e m o n s tra tin g a ll a m o re o r less e n la rg e d a b o r a l p a rt,
o fte n slig h tly p o in te d , re se m b lin g th e tu b u lo s a fo rm .
O u r m a te ria l (fig. 12 a -d ) w as n e v er tu b u lo s a -\ik e . S o fa r
a s w e e n c o u n te re d sp e c im en s w ith w id e n ed a b o r a l e n d o f
th e lo ric a , we h a d to assig n th ese fo rm s to T. b e ro id ea
S te in (see fig. 2a, e; 3a, d , e). W e n e v e r fo u n d T. lo b ia n c o i
in g re a t a b u n d a n c e a n d so we c o u ld n o t c o m p o s e a series
o f th is species c o m p a ra b le w ith t h a t o f H a lm e a n d L u k ­
k a rin e n . In fig. 13, d e riv e d fro m th e se a u th o r s , w e c h o o se
13 sp e c im en s o f th e ir series o f 100 re p re s e n tin g several
12a b e d
Fig. 12. Tintinnopsis lobiancoi D aday (12a from the Western
Scheldt, loc. 5; 12b from the Eastern Scheldt, loc. 11; i2 c ,d fr o m
coastal seawater, loc. 15).
1 07
 In o u r a re a th e species o c cu rs in h ig h -sa lin ity w a te rs
a n d in th e p o ly h a lin e e s tu a rin e z o n e. P e n e tr a tio n in to th e
m e so h a lin e zo n e, h o w e v e r, is p o ssib le .

g. T. tu rb o M e u n ie r (fig. 14 a -f)
13
T h e lo ric a sh o w s m a x im u m w id th in its m id d le o r u p p e r
p a r t a n d n a rro w s (so m e tim e s a b ru p tly : fig. i o a , b) to th e
o ra l p o r tio n c o n sistin g o f a s h o rt c y lin d ric a l c o lla r. T h e
c o n ic a l a b o r a l e n d is ro u n d e d o r b lu n tly p o in te d . T h e
h o u se , is s tro n g ly e n c ru ste d w ith larg e p a rtic le s.
13
T h e o rig in a l d e s c rip tio n o f M e u n ie r ( 1 9 19) is r a th e r in ­
c o m p le te a n d h a s b e e n b a se d o n a few sp e c im en s. T h e
sh a rp ly p o in te d a b o ra l e n d in his fig u re (fig. 27, pi. 22) h a s
n o t b e e n o b se rv e d b y a n y a u th o r . B ra n d t ( 1906) d e sc rib e d
a n d fig u re d (fig. 3 pi. 16) a f o rm a s T. n u c u la F o l?, w hich
w as id en tifie d b y K o fo id & C a m p b e ll (1929) a s T. tu rb o .
o d..;-*
T h is fo rm sh o w s a so m e w h a t m o re sw o lle n a p p e a ra n c e
th a n M e u n ie r’s species. S c h u lz ’s (1965) m ic r o p h o to g ra p h
o f T. tu rb o (fig. 4, pi. 2) is c h a ra c te riz e d b y a c o n ic al
13 a b o r a l p a r t w ith a lm o s t s tra ig h t c o n to u rs re se m b lin g o u r
fig. 9 f. S c h u lz h a d th e o p p o rtu n ity , to s tu d y a large
m a te ria l a s w ell a s w e h a d .
D im e n s io n s : o ra l d ia m e te r: 29-40 p., a v era g e 35 p \
m a x im u m w id th : 45-54 p , a v e ra g e 48 p ; le n g th 64-81 p ,
a v e ra g e 69 p . S c h u lz (1965) m e a su re d a n a v e ra g e o ra l

Fig. 13. Tintinnopsis lobiancoi D aday, after Halm e . Lukka- d ia m e te r o f 33 p , m a x im u m w id th o f 51 p , a n d le n g th of

rinen (i960). 66 p.
H o fk e r (1922) id e n tifie d a m u c h la rg e r sp ecies fro m
d e v e lo p m e n ta l a n d a d u lt sta g es o f T. lo b ia n c o i. T h ey th e fo rm e r b ra c k is h Z u id e rz e e (o ra l d ia m e te r: 55-60 p',
c o n sid e r th e fo llo w in g species b e lo n g in g to th e lo b ia n c o i-
series: T. tu b u lo s a L e v a n d e r, T. k a ra ja c e n sis B ra n d t, T.
s u b a c u ta J ö rg e n s e n , T. tu b u lo s o id e s M e u n ie r, T. p is til­
lu m K o fo id & C a m p b e ll, T. r o tu n d a ta J o rg e n s e n , T. lata
M e u n ie r, T. strig o sa M e u n ie r, T. tu r b o M e u n ie r, T. b a lti­
ca B ra n d t. A fte r o u r o p in io n T. tu r b o a n d T. b a ltic a a re
w ell d e fin e d species (see p . 108 a n d 102) n o t to be c o n fu se d
w ith d e v e lo p m e n ta l stà g es o f T. lo b ia n c o i, a lth o u g h re ­
se m b lin g th ese stag es. T. strig o sa m ay be id e n tic a l w ith
T. b a ltic a (J o rg e n s e n , 1927; see p. 103). A s to th e re m a in ­
d e r o f th e a b o v e m e n tio n e d species, w e a g re e w ith H a lm e
& L u k k a rin e n . B r a n d t (1906) a lre a d y s ta te d th a t h is T. k a ­
ra ja cen sis o n ly d iffe re d fro m T. lo b ia n c o i in r e la to n to its
sm a lle r d im e n sio n s. T. m a c ro p u s M e u n ie r a n d T. lo h ­
m a n n i L a a c k m a n a p p e a r to b e lo n g to th is series to o :
K o fo id & C a m p b e ll (1929) ra n g e d th ese fo rm s a lre a d y
u n d e r T. s u b a c u ta J o rg e n s e n . F in a lly , th e series h a s to
b e classified a s T. lo b ia n c o i D a d a y , th is sp ecies n a m e
h a v in g p rio rity , a n d n o t a s T. tu b u lo s a L e v a n d e r a s H a l­ Fig. 14. Tintinnopsis turbo Meunier (from the Western Scheldt,
m e & L u k k a rin e n in c o rre c tly d id . loc. 3).

108
m a x im u m w id th : 70-78 /r; len g th : 100-115 m) as T. tu rb o
a n d la te r (i9 3 1 ) a s T. lo h m a n n i L a a c k m a n n (sy n o n im i­
zed b y K o fo id & C a m p b e ll, 1929, w ith T. su b a c u ta J ö r ­
g en se n = T. lo b ia n c o i D a d a y a c c o rd in g to H a lm e & L u k ­
k a rin e n , i9 6 0 ). T h e sh a p e o f th e lo ric a re p re s e n te d by
H o fk e r’s ( 1931) fig. 12 b , h o w e v e r, a p p r o a c h e s M e u n ie r’s
(1919) d e s c rip tio n a n d n o t L a a c k m a n ’s (1906).
Fig. 12a, o n th e o th e r h a n d , sh o w s a n e lo n g a te d o ra l
p a r t w ith so m e w h a t in c re a s in g d ia m e te r o f th e o ra l rim
a n d c a n n o t be c la ssified , in o u r o p in io n , a s T. tu rb o . H o f­
k e r’s ( 19 3 1) fig. 13, fin ally , re sem b les to so m e e x te n t o u r
fig. 9 f a n d S c h u lz ’ (1965) fig. 4, pi. 2.
W e n o te d a lre a d y (p. 11) th a t H a lm e & L u k k a rin e n
(i9 6 0 ) sy n o n im iz e d n u m e ro u s species w ith T. lo b ia n c o i
D a d a y , a .o . T. tu rb o .
E x a m in in g th e ir fig u res w e feei t h a t th e sm a lle st sp eci­
m en s m a y lo o k tu r b o - like (see o u r fig. 13), b u t in fa c t a re
d e v e lo p m e n ta l sta g es w ith in c o m p le te lo ric a e a n d th u s
T. tu rb o m u st n o t be id en tifie d w ith th ese fo rm s.
T. tu r b o is a c h a r a c te r species o f th e o lig o h a lin e
(S c h u lz, 1965), a n d th e m e s o h a lin e z o n e ( B a k k e r & D e
P a u w , 1975) o f th e e stu a rie s o f E lb e a n d W e s te rn S c h e ld t
resp . T h e species o c c u rs th r o u g h o u t th e y e a r, d e m o n ­
s tr a tin g p e a k s in s p rin g a n d su m m e r.
Discussion
E v id e n tly th e w ay o f d e v e lo p m e n t o f th e lo ric a m a y d if­
fe r stro n g ly , a c c o rd in g to th e d e v e lo p m e n ta l p h a se
d u rin g w h ic h th e a n im a l h a s b u ilt its lo ric a a n d a lso
a c c o rd in g to th e p re v a ilin g e n v iro n m e n ta l c o n d itio n s
(see in tro d u c tio n ). In th is w ay th e la rg e p h e n o ty p ic v a ri­
a b ility o f th e lo ric a o f o n e species in th e sa m e sa m p le is
e sta b lish e d . T he im p o rta n t o b s e rv a tio n s of H o fk e r
( 1931) h a v e s tro n g ly c o n tr ib u te d to a b e tte r u n d e r­
sta n d in g o f th e p o ssib le c a u se s o f lo ric a v a ria b ility . It is
re a so n a b le to s u p p o s e th a t p a rtic u la rly c o a s ta l se a w a te r
a n d b ra c k ish in la n d w a te rs o ffe r g o o d o p p o rtu n itie s fo r
th e d e v e lo p m e n t o f g re a t v a ria tio n in T in tin n id species,
o w in g to th e s tro n g flu c tu a tio n s o c c u rrin g in th e c h e m i­
c al a n d p h y sic a l e n v iro n m e n t a n d to th e p re se n c e o f
e stu a rin e g ra d ie n t situ a tio n s . T h e re fo re it is n o t ju stifie d
to use th e te rm ‘F o rm e n k re is ’ in th is c ase. A ‘F o rm e n -
k re is ’ is d e fin e d (M a y r, 1965) as a n a g g re g a te o f g e o ­
g ra p h ic a lly re p re se n ta tiv e (a llo p a tric ) sp ecies a n d s u b ­
species. In his d isc u ssio n a b o u t p o ly ty p ic sp ecies (= F o r ­
m e n k re ise ) M a y r, c itin g D ö d e rle in (1902) a n d T o rto n e s e
(1950) a rg u e s th a t m o st m a rin e a n im a ls in d e e d sh o u ld be
u n ite d in to c o m p re h e n s iv e p o ly ty p ic sp ecies, b u t th a t
so m e tim e s th e su b sp e c ie s a re m ere ly in d iv id u a l v a ri­
a n ts . B ra n d t (1896) c o m p a re d T in tin n o p s is lo ric a e fro m
d iffe re n t g e o g ra p h ic a l re g io n s, d e sc rib in g re la te d fo rm s
a s d is tin c t species b u t u n itin g th ese sp ecies in to ‘F o rm e n -
k re ise ’. W ith o u t e x c lu d in g th e p o ss ib ility o f g e o g ra p h ic
v a ria b ility in T in tin n o p s is species, we s h o u ld like to a t ­
ta c h th e g re a te s t im p o rta n c e to in d iv id u a l v a ria tio n here
to o .
O f c o u rse we h a v e to c o n sid e r th e p o ssib le o c c u rre n c e
o f lo ric a v a ria b ility o n a g e n etic b ase. E n v iro n m e n ta l
c h a n g e s d u rin g la te sta g es o f c o n ju g a tio n c a u s e d p e rm a ­
n e n t m o r p h o lo g ic a l c h a n g e s o f P a ra m a e c iu m spec. (Jo l-
los, fro m K u d o , 1966). R e c o m b in a tio n o f n u c le a r m a te ­
rials m ig h t be e ffected in su c h a w ay th a t th e g e n o ty p e is
a lte re d (K u d o ).
F in a lly we m a y m e n tio n e v e n tu a l h y b rid iz a tio n as a
cau se o f p h e n o ty p ic v a ria b ility . W e m a d e a single o b s e r­
v a tio n o f a p o ssib le c o n ju g a tio n o f T in tin n o p s is c y lin ­
d ric a a n d T. b e ro id ea (fig. 15). W e c o u ld n o t d e m o n ­
s tra te , h o w e v e r, th a t c o n ju g a tio n re ally h a d ta k e n place,
a n d , if so, th a t th e h y b rid e s m ig h t p ro v e v iab le . K u d o
(1966), d isc u ssin g e x p e rim e n ts to realize c o n ju g a tio n b e ­
tw e en d iffe re n t species o f P a ra m a e ciu m , s ta te s th a t sp e ­
cies o f th is g e n u s a re in c o m p a tib le w ith o n e o th e r.
T h u s in d iv id u a l p h e n o ty p ic v a ria b ility , g e o g ra p h ic
v a ria b ility a n d g e n o ty p ic c h a n g e s o f th e p ro to z o a n cell
m ay a ll be re sp o n sib le fo r th e larg e v a ria b ility in lo ric a
sh a p e . H o w ev e r, d iffere n ce s in lo ric a sh a p e sh o u ld n o t
p rim a rily be re la te d to (su b ) specific (g e n o ty p ic ) d iffe r­
e n ces b e tw ee n th e T in tin n id cells. In d iv id u a l p h e n o ty p ic
v a ria tio n e v id e n tly p lay s a n all im p o r ta n t ro le in local
p o p u la tio n s . T h e re fo re it is n o t ju stifie d in o u r o p in io n ,
to d e sc rib e new species fro m a m a te ria l c o n s is tin g o f o nly
o n e o r a few sp e c im en s, a s M e u n ie r (1910, 1919) d id .
K o fo id & C a m p b e ll (1929) ig n o re th is a s p e c t in th e ir c o n ­
15
Fig. 15. Tintinnopsis cylindrica D aday and T. beroidea Stein:
conjugation? (from the Western Scheldt, loc. 4).
109
sp e c tu s a n d c o n se q u e n tly h a v e n o t e sc a p e d to u n a c c e p ­
ta b le sp littin g o f species; m a n y in a d e q u a te ly k n o w n
fo rm s o r d iffe re n t fo rm s fro m a c o h e re n t g ro u p h a v e
b e e n ra ised to species.
S tu d y in g s e p a ra te sp e c im en s f ro m o u r (figs. 2 a n d 3)
and R o s s o lim o ’s (fig. 6) b e ro id e a m a te ria l, o n e m ay
id e n tify th e se fo rm s easily w ith d iffe re n t d e s c rib e d sp e ­
cies. T h e a s p e c t o f in d iv id u a l v a ria tio n in b u ild in g o f th e
lo ric a de se rv e s fu ll a tte n tio n o f a ll w o rk e rs o n T in tin n id
sy ste m atics. C la s sific a tio n h a s to b e b a se d o n in v e s tig a ­
tio n o f a larg e m a te ria l. U ltim a te ly , p ro b le m s o f lo ric a
d e v e lo p m e n t a n d sh a p e m a y d e fin ite ly b e so lv ed b y c u l­
tu rin g . C o m b in a tio n o f field in v e s tig a tio n s a n d la b o r a ­
to ry e x p e rim e n ts w o u ld a p p e a r m o s t fru itfu ll. R e n ew ed
a tte n tio n to fu ll life cycles (C o rliss, 1961) m ig h t lea d to a
b a se fo r re v isio n o f p re s e n t-d a y c la ssific a tio n s.
Summary
T h e species o f th e T in tin n id g e n u s T in tin n o p s is S te in ,
o c c u rrin g in th e c o a s ta l w a te rs o f th e S W .-N e th e rla n d s ,
a re fig u re d a n d d e sc rib e d . W e d e m o n s tr a te d a larg e v a ri­
a b ility o f se v e ra l sp ecies in re la tio n to th e sh a p e o f th e
lo ric a. N e a rly a ll lo ric a c h a ra c te rs a p p e a re d to v a ry c o n ­
sid e ra b ly , p a rtic u la rly th e sh a p e o f th e a b o r a l p o r tio n o f
so m e species (T . b e ro id e a S te in ). T h e o ra l d ia m e te r o f
th e lo ric a , o n th e o th e r h a n d , v a rie d m u c h less. T h e
m a n y fo rm s w ere cla ssified in to 7 species on ly : T. b a ltic a
B ra n d t, T. b e ro id e a S te in , T. c a m p a n u la (E h rb g ) D a d a y ,
T. c y lin d ric a D a d a y , T. fim b r ia ta M e u n ie r, T. lo b ia n c o i
D aday, T. tu r b o M e u n ie r. L o ric a e o f so m e species
d e m o n s tra te d e x tre m e v a ria b ility , n o ta b ly th o s e o f T.
b e ro id ea . O th e rs o n th e c o n tr a ry , a p p e a re d r a th e r c o n ­
s ta n t: T. cylin d rica , T. fim b r ia ta . V ery s tro n g in d iv id u a l
v a ria tio n w a s d e m o n s tra te d in lo ca l p o p u la tio n s (fig. 2
a n d 3). T h e re fo re it is n o t ju s tifie d to d e sc rib e n ew species
fro m sp a rse m a te ria l. F o r d e sc rip tiv e w o rk , e sp e cially o n
n e ritic T in tin n id s , in v e s tig a tio n s o f a g re a t n u m b e r o f
sp e c im en s is a d v o c a te d .
Acknowledgements
W e th a n k m r. J . v. d . E n d e f o r his a ssista n c e in th e re p ro ­
d u c tio n o f th e fig u res.
IIO
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