VOL. IV
D e p a r t m e n t of S y s t e m a t i c Z o o l o g y , U n i v e r s i t y of Łódź, Łódź, N a r u t o w i c z a 68, P o l a n d
Maria
WOLSKA
Study on the family Blepharocorythidae
Hsiung. III. Raabena
bella gen. n., sp. n. from the intestine of the Indian elephant
Badania
nad
rodziną Blepharocorythidae
H s i u n g . III. Raabena
g e n . n., s p . n . z j e l i t a s ł o n i a i n d y j s k i e g o
bella
S c a r c e i n f o r m a t i o n h a s b e e n r e p o r t e d c o n c e r n i n g t h e ciliates — p a r a s i t e s of
t h e e l e p h a n t intestine. A f e w species described belong to t h e families Cycloposthiidae
a n d Polydiniidae
(Entodiniomorpha).
Buisson
1923 described
Prototapirella
elephantis
f r o m t h e intestine of t h e A f r i c a n e l e p h a n t . K o f o i d
1935 described Polydinium
mysoreum
a n d Elephantophilus
zeta f r o m the int e s t i n e of t h e I n d i a n elephant. M o r e recently, L a 11 e u r 1958 gave t h e description of Thoracodinium
vorax f r o m t h e intestine of t h e I n d i a n e l e p h a n t f r o m t h e
zoological g a r d e n in A n v e r s .
W h e n e x a m i n i n g t h e faeces of t h e I n d i a n e l e p h a n t , I h a d o p p o r t u n i t y of
f i n d i n g t h e p r e s e n c e of a n e w ciliate. On account of its g e n e r a l m o r p h o l o g i c a l
c h a r a c t e r s as w e l l as of its morphogenesis, t h i s f o r m should be included into
t h e f a m i l y Blepharocorythidae
Hsiung. S o m e of its c h a r a c t e r s h o w e v e r evoke
the necessity to establish a n e w s e p a r a t e genus f o r it and t h e generic n a m e
Raabena is p r o p o s e d 1 .
T h e s t u d y of this n e w r e p r e s e n t a t i v e of Blepharocorythidae,
the family
w h i c h h a s b e e n t h e o b j e c t of m y i n t e r e s t ( W o l s k a 1966 a, 1966 b), m a y
i n t r o d u c e some f u r t h e r d a t a to our k n o w l e d g e of the p h y l o g e n e t i c relations in
this f a m i l y .
In the m a t e r i a l which becomes quicly decomposed, t h e ciliate r e m a i n s
s u f f i c i e n t l y w e l l p r e s e r v e d — t h e c i l i a t u r e is u n i m p a i r e d — a n d is s u i t a b l e f o r
description.
Material
and
methods
M a t e r i a l w a s t a k e n f r o m a f e m a l e I n d i a n e l e p h a n t , Elephas maximus (8 y e a r s
old) f r o m t h e Zoological g a r d e n of Ł ó d ź 2 . T h e s a m p l e s w e r e f i x e d in 10%
f o r m a l i o n . A f t e r f i l t r a t i o n a n d r i n s i n g in w a t e r , t h e ciliates w e r e e x a m i n e d
d i r e c t l y u n d e r t h e s e conditions. T h e s t u d y h o w e v e r is based m o s t l y on silver
i m p r e g n a t e d m a t e r i a l t r e a t e d w i t h t h e m e t h o d described p r e v i o u s l y ( W o l s k a
1966 a), or o n p r e p a r a t i o n s in w i c h t h e silver w a s r e p l a c e d b y gold.
1
The generic name Raabena has been chosen in honor of Prof. dr. Zdzisław
Raabe the Director of the Zoological Institute of the University of Warszawa,
Member of the Polish Academy of Sciences.
2
I wish to express my thanks to Direction of Zoological garden in Łódź for
enabling me investigation.
286
Mj W O L S K A
Results
Raabena
bella gen. n., sp. n.
T h e ciliate is f l a t t e n e d laterally. T h e b o d y outline is m o r e or less ovoid, t h e
p o s t e r i o r end of t h e b o d y b e i n g m o r e n a r r o w t h a n t h e a n t e r i o r one (Fig. 1 A, PL
11—2). In t h e a n t e r i o r p a r t of t h e v e n t r a l side, p r o t r u d e s a vacuole of a n
u n k n o w n c h a r a c t e r , s i m i l a r l y as in t h e g e n u s Blepharocorys
(W o 1 s k a 1966 a).
Dimensions of t h e body: l e n g t h 44—67
w i d t h (or r a t h e r thickness) 27—36 ¡a.
Buccal o v e r t u r e is b r o a d l y open on t h e l e f t b o d y side a n d leads to a f u n n e l s h a p e d concavity w h i c h is slightly b e n t (Pl. 11, 2) a n d r e a c h e s a p p r o x i m a t e l y
as f a r as t h e half of t h e b o d y length. T h e deep groove .beneath t h e l e f t v e n t r a l
m a r g i n produces a sort of pocket in t h e a n t e r i o r p a r t of t h e b u c c a l f u n n e l .
A b o v e t h e buccal o v e r t u r e , on t h e l e f t — dorsal side p r o t r u d e s a s m a l l process
(the f r o n t a l process).
Fig. 1. Raabena bella gen. n.,
sp. n., A. General feature and
pattern of somatic ciliature (scheme). B. Ciliature of vestibulum.,
C. Course of the ciliary zone on
the frontal process and its transition to vestibulum. D. Sector of
the ciliary zone of the ventral lip
(right ventral side), delicate kineties on the territory of vacuole
T h e ciliature is composed of f o u r zones of long cilia located on t h e a n t e r i o r
a n d posterior p a r t s of t h e b o d y (Fig. 1 A, PL 11). T w o zones at t h e posterior pole
lie dorsally and v e n t r a l l y as r e l a t e d t o cytopyge. T h e v e n t r a l zone lies m a i n l y
on t h e l e f t b o d y side a n d only its s m a l l s e g m e n t passes over to t h e r i g h t side.
O n t h e c o n t r a r y , o n l y a s m a l l s e g m e n t of the dorsal zone e n c r o a c h e s u p o n t h e
l e f t side w h e r e a s its long s e g m e n t lies on t h e r i g h t side.
O n e of the a n t e r i o r zones r u n s along t h e r i g h t a n d v e n t r a l m a r g i n a n d p a r t ially along t h e l e f t one of t h e buccal a p e r t u r e . This c o r r e s p o n d s to t h e zone of
t h e v e n t r a l lip in t h e g e n u s Blepharocorys.
On t h e r i g h t - v e n t r a l side kineties
RAABENA
BELLA
GEN.
N.,
SP.
N.
287
of this zone are prolonged on the t e r r i t o r y of the vacuole. Here they are fained
and sometimes they lose the connection with the zone of their origin (Fig. 1 D).
The second anterior zone, similarly as in the genus Blepharocorys, begins at the
base of the f r o n t a l process on the right side, passes over to the left side of the
process, bends on its ventral m a r g i n and encroaches upon the right wall of the
anterior p a r t of buccal concavity (vestibulum?). The disposition of this zone on
the f r o n t a l process as well as its transition to the buccal concavity are illustrated
in Fig. 1 C. In its subsequent course this zone describes an arch on the right
anterior wall of the concavity (Fig. 1 A), it broadens and is prolonged as f a r
as t h e apex of the buccal funnel. In consequence, the right wall of the f u n n e l
becomes coated with longitudinal kineties (PI. IV l i b , 13) which densely cover
a t r i a n g u l a r field (Fig. 1 B). The buccal ciliary field is — in this w a y — a continuation of t h e somatic ciliary zone.
The anterior part of the buccal concavity, between the arched p a r t of the
ciliary zone a n d the m a r g i n of t h e overture, is lined with a delicate n e t w o r k
of fibers (PI. II 7). Pairs of short fibers r u n f r o m the arched p a r t of the zone
and bend backwards, their f r e e ends overlap tegularly. In this w a y they produce
a u n i f o r m f r a m i n g of t h e arched zone (PI. II 5, 6). Near t h e m a r g i n of the buccal
o v e r t u r e begin other, thicker fibers. These fibers split just near the right m a r g i n
and r u n to meet the f o r m e r f i b e r s almost touching them. On t h e left v e n t r a l
margin, w h e r e the groove called t h e pocket begins, the fibers penetrate into it
and r e m a i n undivided along a r a t h e r long distance (PI. I l l 9, IV 11 a). In t h e
deep p a r t of the pocket the fibers split and bend (PI. I l l 8, 9), pass over to the
right wall and then — as thin ones — reach the f r a m i n g of the arched zone
(partly seen also in t h e PI. II 5).
T h e deep n a r r o w e d p a r t of the funnel is strengthened b y thick semicircular
fibers which r u n along its dorsal and left side (PI. I 2, IV 11 a, 13). In many cases
it looks as if those fibers initiated of the last longest buccal kinety (the nearest
the dorsal side). So it looks in the PI. II 6. However possibly those fibers initiate
earlier on t h e right side. In PI. II 4 fibers on the right wall, a r e seen against the
background of buccal kineties. One fiber which runs above t h e anterior ends
of the buccal kineties is especially distinctly seen (the ends of kineties are t h e
anterior ones if we neglect the fact that the direction of kineties is here reversed). The semicircular fibers t e r m i n a t e on the left wall at the strongly impregnated longitudinal streak (PI. I l l 10, IV 11 a, 13). W h a t is the impregnated
s t r u c t u r e is difficult to decide. Possibly it is a bundle of fibers which is f o r m e d
of overlaping extermities of the semicircular fibers (PI. I l l 10). Impregnated
preparations fail to provide distinct images in this case. The streak seems to
p e n e t r a t e over the wall of the pocket but here the m u t u a l relation of the fibers
becomes complicated, the more so as a n e w fan-shaped fiber group appears
(PI. II 4 b). W h e r e these f i b e r s i n i t i a t e and w h a t is their relation to the l o n g i t u d i n a l streak a n d to t h e semicircular f i b e r s — this c a n n o t be elucidated p r e s e n t l y .
The elongated macronucleus lies usually at the middle of the body length,
n e a r e r t h e dorsal margin. Nevertheless it is sometimes shifted f o r w a r d s or
backwards. Its axis coincides w i t h t h e long body axis of the ciliate. The round
micronucleus lies in t h e concavity of the macronucleus at the half of its length
(Fig. 1 A).
One contractile (?) vacuole is present on the ventral side more or less at t h e
half of the body length. In cytoplasm n u m e r o u s spherical or ovoid shining
bodies are dispersed. They resemble those which are present in the " K o n k r e -
288
mentenvacuole" in Paraisotrichidae
and in
Buetschliidae.
The f e w dividing individuals observed indicate that the division occurs in
the same w a y as in t h e genus Blepharocorys
(W o 1 s k a 1966 a, 1966 b). The n e w
ciliature arises in the vacuoles. Each of the four zones arise separately. PI. I 3
and IV 12 show the f o r m e d fronto-buccal zone in its n e a r l y definitive form, and
the semicircular fibers.
Raabena gen. n., diagnosis
The ciliate flattened laterally. The extensive buccal o v e r t u r e on the left body
side. Four ciliary zones. One zone on the so called v e n t r a l lip, another one is
on the f r o n t a l process and penetrates into t h e buccal depression. Two other
zones are on t h e posterior pole. The contractile(?) vacuole is on the v e n t r a l body
side at the half of its lenght. Vacuole of u n k n o w n character lies in anterior
p a r t of the body.
Type — species: Raabena bella sp. n. P a r a s i t e of the Indian elephant intestine
{Elephas maximus L.).
Discussion
The genus Raabena gen. n. should be included into the family
Blepharocorythidae Hsiung because its body is elongated, flattened laterally, the ciliature is
reduced to several zones in the anterior and posterior parts of the body; it has
a f r o n t a l process, ciliated vestibulum, single contractile vacuole.
The type of morphogenesis is in Raabena the same as that which occurs
in the genus
Blepharocorys.
The f a m i l y Blepharocorythidae
was established by H s i u n g 1929 for the
genera Blepharocorys
Bundle and Charon J a m e s o n (presently Charonina
Strand). D o g i e 1 1934 considered this genus as identical with
Blapharocorys
Bundle, W o 1 s k a 1966 b reestablished genus Charonina Strand. C h a v a r i a
1933 added h e r e t h e n e w g e n u s Ochoterenaia w i t h one species O.
appendiculata.
S t r e l k o v 1939 a f t e r all included this species to the genus Blepharocorys
and
created a n e w genus Charonnautes for Ch. equi (Hsiung, 1930).
present only an insignificant
The genera Ochoterenaia
and Charonnautes
component of t h e f a m i l y a n d have not been studied with t h e method of silver
impregnation. In describing the family Blepharocorythidae
the attention should
b e in first place concentrated on the genus Blepharocorys
which constitutes the
bulk of the f a m i l y and on the genus Charonina. The genus Blepharocorys
with
its n u m e r o u s species which live in the intestine of Equidae (mostly in horse)
and also in some rodents, has not been studied in a sufficient degree either.
The extensive research w i t h application of t h e electron microscope, silver impregnation and other methods h a v e been initiated by G r a i n 1966. T h e study
of a m o r e r e s t r i c t e d scope, c o n c e r n i n g mostly t h e i n f r a c i l i a t u r e , have been b e g u n
b y me (W o 1 s k a 1966 a, 1966 b). Our data achieved as yet seem to be sufficient
to compare (mostly concidering the i n f r a c i l i a t u r e ) Raabena bella gen. n., sp. n.
w i t h the species of the genus Blepharocorys
Bundle and Charonina Strand.
Raabena bella gen. n., sp. n. has a more complet ciliature t h a n the species of
the genus Blepharocorys.
On its posterior end exist two r a t h e r large ciliary
zones. It should be recognized for t h a t reason that R. bella presents a more
primitive c h a r a c t e r of ciliature t h a n do the species of the genus
Blepharocorys,
and is in this respect n e a r e r Charonina ventriculi in which two ciliary zones
exist as well (W o 1 s k a 1966 b), although they are smaller and more reduced.
RAABENA
BELLA
GEN.
N.t
SP.
N.
289
In t h e r e p r e s e n t a t i v e s of t h e genus Blepharocorys
a n d Charonina t h e ciliat u r e of v e s t i b u l u m h a s no connection w i t h t h e somatic ciliature. This w a s s t a t e d by
G r a i n 1966 a n d W o l s k a 1966 a. In division the c i l i a t u r e of v e s t i b u l u m
arises i n d e p e n d e n t l y of t h e somatic one ( W o l s k a 1966 a, 1966 b). In R. bella
t h e c i l i a t u r e of v e s t i b u l u m is t h e continuation of t h e s o m a t i c one a n d it arises
in m o r p h o g e n e s i s as c o n t i n u a t i o n of the somatic zone. T h e buccal ciliature of
R. bella h a s t h e n c h a r a c t e r of v e s t i b u l a r ciliature w h i c h is c h a r a c t e r i s t i c of
Trichostomata.
It is m o r e p r i m i t i v e t h a n in t h e g e n u s Blepharocorys
a n d Charonina in w h i c h the buccal c i l i a t u r e becomes i n d e p e n d e n t of t h e s o m a t i c one a n d
arises i n d e p e n d e n t l y of it in ontogenesis.
W h e n c o m p a r i n g t h e b u c c a l ciliature of t h e r e p r e s e n t a t i v e s of t h e genus
Blepharocorys,
Charonina ventriculi
a n d R. bella, it b e c o m e s e v i d e n t t h a t t h e
m o s t a d v a n c e d is t h e evolution of ciliature in t h e species of the g e n u s Blepharocorys w h e r e it is not only i n d e p e n d e n t of t h e s o m a t i c one b u t is d i f f e r e n t i a t e d
into t w o s e p a r a t e groups. It is m o r e p r i m i t i v e in Charonina ventriculi
being
i n d e p e n d e n t of the somatic o n e h o w e v e r not d i f f e r e n t i a t e d . T h e m o s t p r i m i t i v e
is t h e buccal ciliature in R. bella b e i n g a b u n d a n t , u n d i f f e r e n t i a t e d a n d p r e s e n t ing the continuation of t h e s o m a t i c one.
Is really this t y p e of v e s t i b u l u m c h a r a c t e r i s t i c of Trichostomata?
Are
Blapharocorythidae
to be considered as Trichostomata?
G r a i n 1966 also p u t
t h i s question, leaving it w i t h no a n s w e r till m o r p h o g e n e s i s is s t u d i e d .
T h e t y p e of m o r p h o g e n e s i s w h i c h occurs in Blepharocorythidae
is not c h a r a c t e r i s t i c of Trichostomata
(W o 1 s k a 1966 a, 1966 b) b u t of t h e h i g h e r g r o u p s
of ciliates. T h e d e v e l o p m e n t of this e v o l u t i o n a r y b r a n c h —
Blepharocorythidae— h a d f o l l o w e d a q u i t e specific t r e n d . H a d t h e y b r a n c h e d off f r o m Trichostomatai? Or is it p e r h a p s a specific evolution of parasitic Gymnostomata?
Little
a l t e r a t i o n is needed to m a k e t h e ciliature of t h e posterior end of Didesmis —
w h i c h is composed of t w o a d h e r i n g zones — q u i t e s i m i l a r to t h a t of Raabena.
T h e f l a t t e n i n g of t h e b o d y exists a l r e a d y in Didesmis. G r a i n 1966 stressed
t h i s c h a r a c t e r as e x c e p t i o n a l in Buetschliidae.
T h e only contractile vacuole in
Didesmis
has a s i m i l a r position as in Raabena. A s i m i l a r position h a s t h e
K o n k r e m e n t e n v a c u o l e in Didesmis as t h e vacuole w i t h kinetosomes in Raabena.
T h a t vacuole is p r e s u m a b l y (W o 1 s k a 1966 a) an a t r o p h i e d K o n k r e m e n t e n vacuole. T h e a n t e r i o r ciliary z o n e in Didesmis is r e a l l y composed of t w o ones,
w h i c h a r e f o r m e d s e p a r a t e l y in ontogenesis. Each of those zones m i g h t develop
in a d i f f e r e n t m a n n e r in Blepharocorythidae.
One of t h e m r e m a i n e d on t h e
s u r f a c e , t h e second o n e w r a p p e d r o u n d t h e f r o n t a l process (new s t r u c t u r e ) ,
dived p a r t l y inside p r o d u c i n g at f i r s t t h e v e s t i b u l a r ciliature of Raabena; in t h e
s u b s e q u e n t d e v e l o p m e n t it d i f f e r e n t i a t e s into t h e s o m a t i c a n d buccal p a r t in t h e
genus Charonina a n d Blepharocorys.
It schould b e p o s t u l a t e d t h a t a leap in
t h e d e v e l o p m e n t o c c u r r e d — p e r h a p s in connection w i t h t h e a t r o p h y of non —
ciliated kinetosomes on t h e b o d y wich exist in Didesmis — a n d caused t h a t
m o r p h o g e n e s i s a s s u m e d t h e f e a t u r e of r e s t o r i n g t h e c i l i a t u r e e n t i r e l y de novo
w i t h o u t transmission of a n y r u d i m e n t s of t h e p a r e n t a l ciliature to t h e opisthe.
Summary
A n e w ciliate genus: Raabena bella gen. n., sp. n. h a s b e e n described f r o m
t h e intestine of t h e I n d i a n e l e p h a n t Elephas maximus L. This ciliate should b e
included into the f a m i l y Blepharocorythidae
Hsiung. T h e specific d e v e l o p m e n t
of this f a m i l y is discussed.
290
Mj W O L S K A
STRESZCZENIE
Autorka opisała nowy gatunek orzęska Raabena bella gen. n., sp. n. z jelita słonia
indyjskiego Elephans maximus L. Orzęsek ten powinien być włączony do rodziny
Blepharocorythidae
Hsiung. Autorka omawia swoisty rozwój tej rodziny.
REFERENCES
B u i s s o n J. 1923: Les infusoires ciliés du tube digestif de l'homme et des mammifères. Trav. Lab. Parasit., Fac. Med., Paris 1—201.
C h a v a r r i a M. Ch. 1933: Estudios protistológicos II. Ochoterenaia
appendiculata
gen. nov., sp. nov., nuevo infusorio del intestino del caballo (Equus caballus
Linn.) de Mexico. An. Inst. Biol., Nac. Mexico, 4, 191—196.
D o g i e l V. A. 1934: Angaben über die Ophryoscolecidae des Wildschafes aus
Kamzschatka, des Elches und des Yaks, nebst deren zoogeographischen Verwertung. Arch. Protistenk. 82, 290—297.
G r a i n J. 1966: Etude cytologique du quelques ciliés Holotriches endocommensaux
des Ruminants et des Equidés. Centre National de la Recherche Scientifique.
H s i u n g T. S. 1929: A survey of the protozoan fauna of the large intestine of the
horse. (Abstr.) J. Parasit. 16, 99.
H s i u n g T. S. 1930: Some new ciliates from the large intestine of the horse. Trans.
Amer. micr. Soc. 49, 34—41.
K o f o i d C. A. 1935: On two remarkable ciliate protozoa from the caecum of the
indian Elephant. Proc. nat. Acad. Sei., Wash., 21, 501—506.
L a t t e u r B. 1958: Les ciliates Pylydiniinae.
Thoracodinium
vorax n.g., n. sp.,
Cellule, 59, 269—296.
S t r e l k o v A. 1939: Parasitićeskije infusorii iz kiśećnika neparnokopytnych sem.
Equidae. Ucen. Zap. leningr. gosud. pedagog. Inst. Gercena, 17, 1—262.
W o l s k a M. 1966 a: Study on the family Blepharocorythidae
Hsiung. I. Preliminary
remarks. Acta Protozool. 4, 97—103.
W o l s k a M. 1966 b: Study on the family Blepharocorythidae
Hsiung. II. Charoûina
ventriculi (Jameson). Acta Protozool. 4, 279—283.
EXPLANATION OF PLATES I—IV
Raabena bella gen. n., sp. n.
1: General view, ciliary groups are impregnated
2: Semicircular fibers and fibers of the "pocket"
3: Division, a) semicircular fibers of the opisthe, optical section, b) ciliature of
vestibulum of the opisthe another optical section
4: Vestibulum. a) view of the dorsal side, b) view of the ventral side, fan-shaped
fibers
5: Isolated fronto-buccal zone, the zone of ventral lip and fibers coating vestibulum
6: Isolated fronto-buccal zone, fibers running from the arched part of the zone
7: Fibers of the anterior wall of vestibulum
8: Anterior part of the ciliate body showing the inside of the pocket; splitting of
fibers is seen
9: View of the "pocket" fibers: single in the anterior segments and split in the
deeper part
10: Isolated kineties of vestibulum, semicircular fibers and the longitudinal streak
11: Left ventral margin and interior of vestibulum. a) kineties of the ventral lip,
fibers in the "pocket" and semicircular fibers are seen, b) vestibulum kineties on
the right wall (the same specimen as in a) another optic section)
12: Fronto-buccal zone of the opisthe and semicircular fibers
13: Isolated kineties of vestibulum, semicircular fibers and the longitudinal streak
Photomicrographs of impregnated preparations.
Magnification: 1—3 1000 X, 4—^ 2C00 X, 7—13 2600 X
A C T A PROTOZOOL. VOL. IV, 27
M. Wolska
P L A T E III
auctor phot.
ACTĄ P R O T O Z O O L . VOL. IV, 27
P L A T E IV
A C T A P R O T O Z O O L . V O L . IV, 27
M. Wolska
P L A T E III
auctor phot.
A C T Ą P R O T O Z O O L . VOL. IV, 27
M. Wolska
P L A T E IV
auctor phot.