(cf. Fig . 18 .).

Description

surface of the spirotri shows deciliated different surface sculp (aboral) surface of Eup single ciliated dikinet polygonal cortical alve perioral region of Para dikinetids of Colpoda ly ciliated dikinetids of um. Fig. 13. Doubly a heterotrich . Figs. 14-17. ciri of the hypotrich omposed of several Note: some kinetosomes pear as small stubs. Figs . ated somatic surface of the ptes showing deciliated surface-sculpturing of the g . 19 . Partially deciliated iliate Conchophthirus difference encountered in wo ciliates from the same chian Nyctotherus ova/is dikinetids and completely turing. Fig . 21 . Dorsal lotes eurystomus reveals ids or dorsal bristles and oli. Fig . 22. Deciliated I eft mecium . 386 CILIOPHORA, INTRO posterior to ciliated locomotor field . Figs. 24-26. Fig. 24. Oral region of Paramecium shows preoral and postoral sutures where kineties from left and right sides of body converge . Fig . 25 . Posterior suture of Paramecium. Fig. 26. Portion of anterior end of Conchophthirus curtus (cf . Fig . 19) shows presence of closely packed dikinetids of thigmotactic region , which attaches the ciliate to its bivalve host's gill-epithelium . arrangement and complexity of oral kinetids whose cilia create feeding currents and manipulate prey . There is generally a left- right asymmetry of oral structures , one side or the other being more prominent. DUCTION Figs. 27-37 Contractile vacuole pores of various ciliate genera. Fig . 27. Haptorian Didinium nasutum . Fig . 28 . Heterotrichian Nyctotherus ova/is. Figs. 29 , 30 . Peniculine Frontonia /eucas ; shows membrane flaps. Fig . 31. Peniculine Paramecium multimicronucleatum. Fig . 32 . Pen- iculine Neobursaridium gigas. Fig . 33. Hymeno- stomatid Tetrahymena pyriformis . Fig. 34 . Scuticociliatid Conchophthirus. Fig. 35.Suctorian Heliophrya. Fig . 36 . Suctorian Podophrya. Fig. 37. Posterior region of Paramecium multimicro- nucleatum shows cytoproct in posterior suture . Some postciliodesmatophoran karyorelictean ciliates have quite complex, not simple, oral ciliature (ct. Figs. 39-42; 66-68) . They are placed together because of presumably ancestral nuclear dualism, their similar somatic dikinetids (Lynn , 1981 ; Small and Lynn, 1981), and similarities in the sequence of their rRNA genes (Embley et al ., 1995; Hammerschmidt et al. , 1996) . The left border of the oral area may have paroral files of monokinetids (Fig .68), dikinetids (Figs . 66), or polykinetids (Figs. 67) , while the right border may have dikinetids (Fig.66) or polykinetids (Figs . 67, 6 8). Postciliodesmatophoran heterotrichs also have a ventral cytostome and oral area ; the somatic kinetid pattern resembles that of karyorelicteans, indicating close relationship of these two classes (Lynn, 1981, 1991 ). Almost all genera have many oral polykinetids CILIOPHORA, INTRODUCT a. TT (j!) b. f. extrusosomes of ciliates . (a) Mucocysts ready to discharge and discharging . (b) Rod-like mucocysts ready to discharge and discharging ; found in colpod eans , scuticocilatids , and nassophoreans. (c) Fibrous trichocyst of peniculine ready to discharge and discharging . (d) Three different kinds of toxicysts found in haptorians . All three may occur in the same species. (e) Haptocyst of su ctorian ready to discharge and discharged ; prey and prediitor joined . (f) Rhabdocysts of postciliodesmato- phoreans ready to discharge and discharging (after Hausmann , 1978) . often as a file , usually along the left border of the oral region (Figs. 72 , 73) . Oral dikinetids along the right border form paroral structures that are often precytostomal (Figs. 72 , 73) ; but they may be replaced or accompanied by right oral polykinetids. Protocruzia is exceptional : its simpler nuclear structure and somatic dikinetid suggest relationship with members of the Subphylum Postciliodesmatophora where Small & Lynn (1985) placed it. Figs . 39 , 40 . Oral region of 2 Karyorelictean protostomatid ciliates which exhibit mouthless phagotrophy and simple oral differentiation . Fig . 39 . Kentrophoros. Fig . 40 . Tracheloraphis . -9 Figs. 41 , 42 . Two Karyrelictean protostomatid ciliates with cytostomes and perioral dik inetids usually bearing longer cilia than those of somatic surface . Fig . 41 . Trachelocerca , the only karyorelictean genus with a prostom ial cytostome. Fig. 42 . Tracheloraphis patterns . Note the glabrous stripe (Fig . 42a.) . in which ingestion occurs by infolding of the stripe surface in the midregion (Fig . 42c) of the elongate vermiform cell. In another pattern , still considered Tracheloraphis , the anterior end appears to bear a cytostome-cytopharyngeal complex (Fig . 42b.) . It also bears a glabrous stripe but there is probably no ingestion by infolding in the midregion (Fig . 42d) . Figs. 43-49 . Genera of litstomate ciliates whose oral region is boun differentiated oral kinetids . Fig Monodium. Fig . 47. Phialina . Fig Figs. 50-58 . Genera of prostomate whose oral region is bounded by di oral dikinetids, within it small p make the "brosse ". Fig . 50 Dragesco et al. , 1974) . Fig . 51 (after Dragesco et al. , Apsiktrata. Fig . 53 . Holo 0/des/oeria. Fig . 55 . an undescr Spathidiopsis. CILIOPHORA, INTRODUCTION an haptorian ded by . 43 . Enchelydium . idium. Fig. 46. . 48 . Acineria . an ciliates fferentiated olykinetids that . Vasico/a (after . Pelatractus 1974) . Fig . 52 . phrya. Fig. 54 . ibed prorodontid . higma . Fig . 58. CILIOPHORA, :::·: .~.: .:.:.:....~.~·:· ·:.· .· :\~ Figs .59-62 . Genera of litostomatean trichostom- ateans showing increasing complexity of oral structures. Fig . 59. Oidesmis (after Grain 1966a) Paraisotricha (from Wolska 1964) . Fig . 62. Blepharocorys (after Wolska 1971 a). Figs . 63, 64 . Ciliates of the class Plagiopylea . Its oral structures (Fig . 69) and stomatogenesis relate it to the other ciliates. The sequence of the small subunit rRNA gene suggests that Protocruzia is related to the spirotrich ciliates (Hammerschmidt et al. , 1996), placing it in the Subphylum lntramacronucleata (Lynn , 1996) . The remaining ciliates are placed in the Subphylum lntramacronucleata Lynn, 1996, INTRODUCTION 389 haryngeal apparatus characteristic of litostom- atean haptorians . (A) Rhabdos of taxa having a temporary cytostome . Toxocysts (To) enclosed within fibrous annulus (FA) and ring of oral dikinetids. Transverse microtubular ribbons (T) support cytopharynx when it is formed. (B) Rhabdos of taxa having a permanent cytostome. Toxicysts, which are typically excluded from the oral region , are located either periorally or some distance from the oral region. Transverse microtubular ribbons (T) extend to support Cytopharynx. Nd-nematodesmata; P c- po stc iIi a ry microtubular ribbon; Ph-phagoplasm. oral structures . and share some very broad similarities in oral structures . Oral ciliature of phyllopharyngeans may be of pericytostomal and precytostomal kinetids (Figs. 80). 390 CILIOPHORA, IN Figs . 66-69. Ciliates with permanent cytostomes F that are ventral and never anterior. Figs . 6 6-6 8 . P Karyorelictean ciliates with complex ventral oral P structures . Fig. 67. Remanella . Fig . 68 . Ge/eei/a. but now considered more closely related to the spirotrichs . F p ( :=:' : ,.,.. \ ·,·1 \ ..I ': j : ,· " ; " \ \' l\\ \. Figs. 70-73. ~ \ i, L ,, : Genera of heterotrichean \j J and spirotri chean ciliates with compl ex ventral oral organ ellar comple xes . Fig. 70. The spirotrichean hypotrich Euplotes. Fig .71. The spirotrichean stichotrich Paraurosty/a . Figs 72 , 73 . Heteroyricheans Parablepharisma (Fig . 72 .) and F ( Eometopus) . c Usually, oral dikinetids border the right of the oral area (Figs. 74, 75, 79, 82 , 83) . 01 P the left, or posterior left, is a variable S Number of oral polykinetids (Figs. 70 , 7 4, n 75 , 77-79 , 82, 83). In the Classes o Colpodea, Nassophorea, and Oligohymeno- b phorea there are usually 3 oral polykinetids , c but like the spirotrich (Figs . 70 , 71 ), some e genera, such as Nassu/a (Fig. 77) and TRODUCTION \ ', ~ 0. ·... \ . .. '.. ·. ·. • •. ·. igs. 74-76 . Genera of colpodean ciliates. Fig . 74 . uytoraciella (from Nj ine , 1978) . Fig . 75 . latyophrya (from Dragesco et al. , 1977) . Fig . 6. Colpoda . 79./ !: ~· : igs. 77-79 . Genera of nassophorean and eniculine ciliates . The nassophoreans, Nassula 9 . Peniculine , Frontonia . ig. 80. A phyllopharyngean ciliate with ventral ytostome, Trithigmostoma. latyophrya (Fig . 75) , have many more. ome genera of the Subphylum lntramacro- ucleata have large nematodesmata, often rganized as a complex cyrtos ("pharyngeal asket") (Fig. 84). The nematodesmata of the yrtos (Fig . 84) originate at the proximal nds of kinetosomes and may or may not CILIOPHO ? " · ,?: ·.• ! ~:.·\ .. . ·."

Type species

Figures

Raw text

surface of the spirotri
shows deciliated
different surface sculp
(aboral) surface of Eup
single ciliated dikinet
polygonal cortical alve
perioral region of Para
dikinetids of Colpoda
ly ciliated dikinetids of
um. Fig. 13. Doubly
a heterotrich . Figs. 14-17.
ciri of the hypotrich
omposed of several
Note: some kinetosomes
pear as small stubs. Figs .
ated somatic surface of the
ptes showing deciliated
surface-sculpturing of the
g . 19 . Partially deciliated
iliate Conchophthirus
difference encountered in
wo ciliates from the same
chian Nyctotherus ova/is
dikinetids and completely
turing. Fig . 21 . Dorsal
lotes eurystomus reveals
ids or dorsal bristles and
oli. Fig . 22. Deciliated I eft
mecium .
386 CILIOPHORA, INTRO
posterior to ciliated locomotor field .
Figs. 24-26. Fig. 24. Oral region of Paramecium
shows preoral and postoral sutures where
kineties from left and right sides of body
converge . Fig . 25 . Posterior suture of
Paramecium. Fig. 26. Portion of anterior end of
Conchophthirus curtus (cf . Fig . 19) shows
presence of closely packed dikinetids of
thigmotactic region , which attaches the ciliate to
its bivalve host's gill-epithelium .
arrangement and complexity of oral kinetids
whose cilia create feeding currents and
manipulate prey . There is generally a left-
right asymmetry of oral structures , one side
or the other being more prominent.
DUCTION
Figs. 27-37 Contractile vacuole pores of various
ciliate genera. Fig . 27. Haptorian Didinium
nasutum . Fig . 28 . Heterotrichian Nyctotherus
ova/is. Figs. 29 , 30 . Peniculine Frontonia /eucas ;
shows membrane flaps. Fig . 31. Peniculine
Paramecium multimicronucleatum. Fig . 32 . Pen-
iculine Neobursaridium gigas. Fig . 33. Hymeno-
stomatid Tetrahymena pyriformis . Fig. 34 .
Scuticociliatid Conchophthirus. Fig. 35.Suctorian
Heliophrya. Fig . 36 . Suctorian Podophrya. Fig. 37.
Posterior region of Paramecium multimicro-
nucleatum shows cytoproct in posterior suture .
Some postciliodesmatophoran karyorelictean
ciliates have quite complex, not simple, oral
ciliature (ct. Figs. 39-42; 66-68) . They
are placed together because of presumably
ancestral nuclear dualism, their similar
somatic dikinetids (Lynn , 1981 ; Small and
Lynn, 1981), and similarities in the
sequence of their rRNA genes (Embley et al .,
1995; Hammerschmidt et al. , 1996) . The
left border of the oral area may have paroral
files of monokinetids (Fig .68), dikinetids
(Figs . 66), or polykinetids (Figs. 67) , while
the right border may have dikinetids
(Fig.66) or polykinetids (Figs . 67, 6 8).
Postciliodesmatophoran heterotrichs also
have a ventral cytostome and oral area ; the
somatic kinetid pattern resembles that of
karyorelicteans, indicating close relationship
of these two classes (Lynn, 1981, 1991 ).
Almost all genera have many oral polykinetids
CILIOPHORA, INTRODUCT
a. TT (j!) b.
f.
extrusosomes of ciliates . (a) Mucocysts ready to
discharge and discharging . (b) Rod-like
mucocysts ready to discharge and discharging ;
found in colpod eans , scuticocilatids , and
nassophoreans. (c) Fibrous trichocyst of
peniculine ready to discharge and discharging . (d)
Three different kinds of toxicysts found in
haptorians . All three may occur in the same
species. (e) Haptocyst of su ctorian ready to
discharge and discharged ; prey and prediitor
joined . (f) Rhabdocysts of postciliodesmato-
phoreans ready to discharge and discharging
(after Hausmann , 1978) .
often as a file , usually along the left border of
the oral region (Figs. 72 , 73) . Oral
dikinetids along the right border form
paroral structures that are often
precytostomal (Figs. 72 , 73) ; but they may
be replaced or accompanied by right oral
polykinetids.
Protocruzia is exceptional : its simpler
nuclear structure and somatic dikinetid
suggest relationship with members of the
Subphylum Postciliodesmatophora where
Small & Lynn (1985) placed it.
Figs . 39 , 40 . Oral region of 2 Karyorelictean
protostomatid ciliates which exhibit mouthless
phagotrophy and simple oral differentiation . Fig .
39 . Kentrophoros. Fig . 40 . Tracheloraphis .
-9
Figs. 41 , 42 . Two Karyrelictean protostomatid
ciliates with cytostomes and perioral dik inetids
usually bearing longer cilia than those of somatic
surface . Fig . 41 . Trachelocerca , the only
karyorelictean genus with a prostom ial
cytostome. Fig. 42 . Tracheloraphis patterns .
Note the glabrous stripe (Fig . 42a.) . in which
ingestion occurs by infolding of the stripe surface
in the midregion (Fig . 42c) of the elongate
vermiform cell. In another pattern , still
considered Tracheloraphis , the anterior end
appears to bear a cytostome-cytopharyngeal
complex (Fig . 42b.) . It also bears a glabrous
stripe but there is probably no ingestion by
infolding in the midregion (Fig . 42d) .
Figs. 43-49 . Genera of litstomate
ciliates whose oral region is boun
differentiated oral kinetids . Fig
Monodium. Fig . 47. Phialina . Fig
Figs. 50-58 . Genera of prostomate
whose oral region is bounded by di
oral dikinetids, within it small p
make the "brosse ". Fig . 50
Dragesco et al. , 1974) . Fig . 51
(after Dragesco et al. ,
Apsiktrata. Fig . 53 . Holo
0/des/oeria. Fig . 55 . an undescr
Spathidiopsis.
CILIOPHORA, INTRODUCTION
an haptorian
ded by
. 43 . Enchelydium .
idium. Fig. 46.
. 48 . Acineria .
an ciliates
fferentiated
olykinetids that
. Vasico/a (after
. Pelatractus
1974) . Fig . 52 .
phrya. Fig. 54 .
ibed prorodontid .
higma . Fig . 58.
CILIOPHORA,
:::·: .~.: .:.:.:....~.~·:· ·:.· .·
:\~
Figs .59-62 . Genera of litostomatean trichostom-
ateans showing increasing complexity of oral
structures. Fig . 59. Oidesmis (after Grain 1966a)
Paraisotricha (from Wolska 1964) . Fig . 62.
Blepharocorys (after Wolska 1971 a).
Figs . 63, 64 . Ciliates of the class Plagiopylea .
Its oral structures (Fig . 69) and
stomatogenesis relate it to the other ciliates.
The sequence of the small subunit rRNA gene
suggests that Protocruzia is related to the
spirotrich ciliates (Hammerschmidt et al. ,
1996), placing it in the Subphylum
lntramacronucleata (Lynn , 1996) .
The remaining ciliates are placed in the
Subphylum lntramacronucleata Lynn, 1996,
INTRODUCTION 389
haryngeal apparatus characteristic of litostom-
atean haptorians . (A) Rhabdos of taxa having a
temporary cytostome . Toxocysts (To) enclosed
within fibrous annulus (FA) and ring of oral
dikinetids. Transverse microtubular ribbons (T)
support cytopharynx when it is formed. (B)
Rhabdos of taxa having a permanent cytostome.
Toxicysts, which are typically excluded from the
oral region , are located either periorally or some
distance from the oral region. Transverse
microtubular ribbons (T) extend to support
Cytopharynx. Nd-nematodesmata; P c- po stc iIi a ry
microtubular ribbon; Ph-phagoplasm. oral
structures .
and share some very broad similarities in
oral structures . Oral ciliature of
phyllopharyngeans may be of pericytostomal
and precytostomal kinetids (Figs. 80).
390 CILIOPHORA, IN
Figs . 66-69. Ciliates with permanent cytostomes
F
that are ventral and never anterior. Figs . 6 6-6 8 .
P
Karyorelictean ciliates with complex ventral oral
P
structures . Fig. 67. Remanella . Fig . 68 . Ge/eei/a.
but now considered more closely related to the
spirotrichs .
F
p
(
:=:' :
,.,.. \ ·,·1
\ ..I
': j
: ,· "
; "
\ \'
l\\ \.
Figs. 70-73.
~ \
i,
L
,, :
Genera of heterotrichean
\j
J
and
spirotri chean ciliates with compl ex ventral oral
organ ellar comple xes . Fig. 70. The spirotrichean
hypotrich Euplotes. Fig .71. The spirotrichean
stichotrich Paraurosty/a . Figs 72 , 73 .
Heteroyricheans Parablepharisma (Fig . 72 .) and
F
( Eometopus) .
c
Usually, oral dikinetids border the right of
the oral area (Figs. 74, 75, 79, 82 , 83) . 01 P
the left, or posterior left, is a variable
S
Number of oral polykinetids (Figs. 70 , 7 4,
n
75 , 77-79 , 82, 83). In the Classes
o
Colpodea, Nassophorea, and Oligohymeno-
b
phorea there are usually 3 oral polykinetids ,
c
but like the spirotrich (Figs . 70 , 71 ), some
e
genera, such as Nassu/a (Fig. 77) and
TRODUCTION
\ ',
~ 0.
·... \
. ..
'.. ·. ·.
• •. ·.
igs. 74-76 . Genera of colpodean ciliates. Fig . 74 .
uytoraciella (from Nj ine , 1978) . Fig . 75 .
latyophrya (from Dragesco et al. , 1977) . Fig .
6. Colpoda .
79./
!: ~· :
igs. 77-79 . Genera of nassophorean and
eniculine ciliates . The nassophoreans, Nassula
9 . Peniculine , Frontonia .
ig. 80. A phyllopharyngean ciliate with ventral
ytostome, Trithigmostoma.
latyophrya (Fig . 75) , have many more.
ome genera of the Subphylum lntramacro-
ucleata have large nematodesmata, often
rganized as a complex cyrtos ("pharyngeal
asket") (Fig. 84). The nematodesmata of the
yrtos (Fig . 84) originate at the proximal
nds of kinetosomes and may or may not
CILIOPHO
? " ·
,?: ·.• ! ~:.·\
.. . ·."