Morphology and Notes on Morphogenesis during Cell Division of Deviata
polycirrata n. sp. and of Deviata bacilliformis (Gelei, 1954) Eigner, 1995

(Ciliophora: Kahliellidae) from Argentina

GABRIELA C. KÜPPERS and MARÍA C. CLAPS

Instituto de Limnologı́a Dr. R. A. Ringuelet (CCT LA PLATA CONICET), Avenida Calchaquı́ km 23.5, Florencio Varela 1888, Buenos Aires

Province, Argentina

ABSTRACT. Described herein are the morphology and certain morphogenetic stages of a new freshwater ciliate species, Deviata
polycirrata n. sp., and of Deviata bacilliformis recorded in the soil of a dried temporary pond from Argentina. Ciliates were studied alive
and after silver impregnation with Protargol. Deviata polycirrata n. sp. measures 130–180 � 45–70 mm in vivo. The species possesses 8–9
long cirral rows on the right and 9–13 on the left of the oral zone, and 3 dorsal rows of dikinetids. The adoral zone is composed of 39–48
membranelles. There are four macronuclear nodules and usually two micronuclei. A single contractile vacuole is located equatorially on
the left body margin. This new species mainly differs from its congeners in having a higher number of cirral rows, the three long dorsal
rows of dikinetids (vs. usually one to two dorsal rows of dikinetids), and a higher number of adoral membranelles. The other species
reported here, D. bacilliformis, is recorded for the first time in Argentina. Unlike previous observations on this species, on the dorsal
surface there are cirral rows that are preceded by cilia (combined cirral rows), and stomatogenesis begins with the proliferation of non-
ciliferous basal bodies some distance posterior to the buccal vertex.

Key Words. Buenos Aires Province, freshwater, kahliellid species, soil, Stichotrichida.

THE family Kahliellidae was erected by Tuffrau (1979) for
those genera that lack transverse cirri, have frontoventral so-

matic rows arranged longitudinally, have more or less distinctive
marginal rows, and well developed frontal cirri. Tuffrau (1979)
recognized 12 genera within the Kahliellidae but several of them
are now placed within different families (Lynn 2008). Eigner
(1995) redefined the kahliellids on the basis of morphological and
ontogenetic data as having more than one longitudinal cirral row
on the right side of the body and neokinetal anlagen developing
during morphogenesis. Eigner (1995) created two new genera,
Deviata and Neogeneia, and recognized a total of nine genera and
species within the family. In a subsequent paper, Eigner (1997)
transferred Deviata to the Oxytrichidae and considered the Kahli-
ellidae as a junior synonym of this family. On the contrary, the
family Kahliellidae is maintained by Lynn (2008) for organisms
with ventral cirral files that may be preserved through a variable
number of cell divisions before being resorbed and replaced
through additional new ( 5 neokinetal) anlagen. Since Eigner
(1995), two new Deviata species were discovered: Deviata este-
vesi (Paiva and Silva-Neto 2005) and Deviata rositae (Küppers,
Lopretto, and Claps 2007), although their division morphogenesis
has not been investigated. A recently published paper by Siqueira-
Castro, Paiva, and Silva-Neto (2009) proposes the transference of
D. estevesi to the genus Parastrongylidium Fleury and Fryd-Ver-
savel, 1984, as well as the description of a new Deviata species.

The aim of the present study is to describe the morphology and
some morphogenetic stages during the cell division of a new
Deviata species and of an Argentinian population of Deviata ba-
cilliformis (Gelei, 1954) Eigner, 1995, both by means of live ob-
servation and Protargol staining.

MATERIALS AND METHODS

Water samples were taken during February 2004 from a tem-
porary pond located near the Provincial Route 63, close to Dolores
city in the Buenos Aires Province, Argentina (3611805500S,
5713201200W) (Fig. 1). The following physical and chemical vari-
ables were measured with a multiparameter sensor (Horiba U21,

Kyoto, Japan): water temperature, electrical conductivity, dis-
solved-oxygen concentration, total dissolved solids, and pH.
Deviata bacilliformis was isolated from rewetted soil samples
from the dried bed of a temporary pond located near the city of
Poblet, Buenos Aires Province, collected during the summers of
2004 and 2005 (see Küppers et al. 2007 for the detailed descrip-
tion of the sampling site and sample treatment). Ciliates were
grown in the laboratory in raw cultures that were made by adding
table water and a wheat grain to the water sample and by resus-
pending the soil samples in Petri dishes. Observations on living
cells were performed under a stereomicroscope and a bright-field
microscope. Ciliates were picked from the cultures with micropi-
pettes and fixed in Bouin’s solution for treatment by the Protargol
technique according to Wilbert (1975). Living and silver-impreg-
nated cells were observed and measured under a bright-field mi-
croscope with a calibrated ocular micrometer at final magni-
fications of 125X, 425X, and 1,250X. The drawings of the organ-
isms in vivo are freehand sketches, and Fig. 2 was rendered by
scratching an ink-painted high-impact paper. Details of the silver-
impregnated organisms were traced with the aid of a drawing tube
under a bright-field microscope at a final magnification of 1,000X.
Photographs were also taken by bright-field microscopy. The ter-
minology used is according to Lynn (2008) and Eigner (1995).
The cirral rows on the right of the adoral membranelles in the in-
terphase organisms are numbered from left to right (i.e. R1, R2,
etc.), while the cirral rows on the left of the adoral zone are num-

Fig. 1. Type locality of Deviata polycirrta n. sp. near Dolores city in
the Buenos Aires Province, Argentina (star). NR, National route; PR, Pro-
vincial route.

Corresponding Author: G. Küppers, Instituto de Limnologı́a ‘‘Dr. R.
A. Ringuelet’’ (CCT LA PLATA CONICET), Avenida Calchaquı́ km
23.5, Florencio Varela 1888, Buenos Aires Province, Argentina—Tele-
phone number: 100 5411 4275 7799; FAX number: 100 5411 4275
8564; e-mail: gkuppers@fcnym.unlp.edu.ar

273

J. Eukaryot. Microbiol., 57(3), 2010 pp. 273–284
r 2010 The Author(s)
Journal compilation r 2010 by the International Society of Protistologists
DOI: 10.1111/j.1550-7408.2010.00474.x

Published by
the International 
Society of 
ProtistologistsEukaryotic Microbiology

The Journal of

mailto:gkuppers@fcnym.unlp.edu.ar


Fig. 2. Morphology of Deviata polycirrta n. sp. in vivo. CV, contractile vacuole.

274 J. EUKARYOT. MICROBIOL., 57, NO. 3, MAY–JUNE 2010



bered from right to left (i.e. L1, L2, etc.) following Paiva and Sil-
va-Neto (2005). Anlagen are indicated with Roman numerals.
Voucher slides of Deviata polycirrata n. sp. (accession numbers
MLP63 and USNM1135042) and of D. bacilliformis (MLP64)
were deposited in the Colección de Invertebrados of the Museo de
La Plata, Argentina, and the Smithsonian Institution, Washington,
USA.

RESULTS

Description of Deviata polycirrata n. sp. (Table 1 and Fig. 2–
4, 9, 14–19). The body size in vivo is length 130–180mm and
width 45–70 mm. The body shape is variable; sometimes elliptical
and slender, with both ends being rounded, but sometimes glob-
ular with the anterior end being pointed and flattened and the
posterior rounded. The body is very flexible. The cytoplasm is
colorless but appears dark gray at low magnification (o10X)
because of the presence of densely packed refractive globules
� 10 mm in width and ingested wheat starch from the cultures.

Cortical granules are absent. The contractile vacuole is equatorial
and on the left margin of the body and lacks collecting canals
(Fig. 2). At times a defecation vacuole was observed at the pos-
terior end. There are always four macronuclear nodules, with the
two anterior and two posterior connected by a thin strand, some-
times faintly stained. The macronuclear nodules are variable in
shape and usually are bilobed or ellipsoid. There are usually two

micronuclei, each one often located between a pair of macronu-
clear nodules. The adoral zone of membranelles represents 30% of
total body length (calculated on the average values measured on
silver-impregnated cells). This adoral zone is composed of 39–48
membranelles of four basal-body rows each. Paroral and endoral
membranes consist of dikinetids and the former is shorter at its
proximal end and slightly curved. In several specimens these
membranes intersected optically due to a preparation artifact, be-
cause the cells became very inflated after the treatment with Pro-
targol (Fig. 14–19). The buccal cavity is small and flattened, with
cytopharyngeal fibers extending posteriorly from the cytostome.
In the buccal field there are three strong frontal cirri and one bu-
ccal cirrus next to the paroral membrane. Behind the middle fron-
tal cirrus there are two cirri aligned with the buccal cirrus (R2).
Behind the rightmost frontal cirrus there is a row of 6–11 cirri
(R3) that extends up to the buccal vertex. There are 17–21 long
rows of cirri (i.e. rows that extend beyond the buccal vertex), of
which 8–9 are located on the right of the adoral zone of mem-
branelles (R4–R12) and 9–13 on the left of it (L1–L13) (Fig. 3, 4).
The first long cirral row from the right (R4), usually shorter than
the others, terminates at the posterior third of the body, without
reaching the posterior end. Sometimes this row is barely shorter
posteriorly; and in 10 individuals from a total of 57 organisms
observed, it reached the posterior end of the body. The other long
cirral rows from the right terminate at the posterior end of body.
The row R5 is slightly shorter anteriorly than its neighboring

Fig. 3, 4. Morphology of Deviata polycirrata n. sp. after Protargol impregnation. 3. Ventral view. 4. Dorsal view. AZM, adoral zone of mem-
branelles; B, buccal cirrus; CF, cytopharyngeal fibers; D1–D3, dorsal kineties one to three; L1–L9, left rows of cirri one to nine; Ma, macronuclear
nodule; Mi, micronucleus; P, paroral membrane; R1–R12, right rows of cirri 1–12. Scale bar 5 50 mm.

275KÜPPERS & CLAPS—NEW KAHLIELLIDAE FROM ARGENTINA



rows. The long cirral rows from the left abut anteriorly on the
adoral zone of membranelles and extend up to the posterior end of
the body. The anterior cirri from each row possess nine, six, or
four basal bodies; while the subsequent cirri are composed only of
four or two basal bodies. The rows of cirri are spiraled and those
that are located laterally commence ventrally at the anterior end,
and terminate dorsally at the posterior end. There are three long
dorsal rows of dikinetids (D1–D3) that are bounded by the long
rows of cirri on the dorsal surface and extend between the anterior
and posterior ends of the dorsalized cirral rows (Fig. 4). The dorsal
kinety D1 is slightly longer in the anterior portion. The dikinetids
of dorsal kinety D2 are densely packed within the row and are
therefore more numerous than those in the other two dorsal rows;

D2 also sometimes extends further posteriorly. The dorsal kinety
D3 is one to two basal body pairs shorter anteriorly than the other
two rows of bristles. According to Eigner (1995), retained parental
cirral rows in the interphase organism have widely spaced cirri
and half the cirri of a neighboring row. These retained parental
rows seem to be absent in the species described here, as well as
caudal cirri (see notes on morphogenesis below).

Some variability in the cirral pattern was observed. In one
specimen, R4 is very short and does not surpass the buccal vertex,
while R5 reaches the posterior end of the body (Fig. 19). Another
specimen possessed a short row of five narrowly spaced cirri be-
tween R3 and R4. In still another specimen this extra row pre-
sented 13 cirri, R4 ended subequatorially, and there was another

Table 1. Mophometric data on Deviata polycirrata n. sp.

Character �X M Min. Max. SD n

Body length in vivo 152 150 130 180 16.8 10
Body width in vivo 61.5 65 45 70 10 10
Body length 181.2 172.5 155 220 20.9 20
Body width 130.2 127.5 100 180 21.2 20
AZM, length 54.7 55 50 70 5.2 20
Cytopharyngeal fibers, length 48.8 49.8 41.5 53.9 5.2 5
Macronuclear nodules, number 4 4 4 4 0 20
Macronuclear nodules, length 14.3 14.5 7.5 23.2 3.7 20
Macronuclear nodules, width 8.5 8.3 5.8 12.4 1.2 20
Distance between macronuclear nodules 23.7 21.6 14.9 37.3 6.6 20
Micronuclei, number 1.7 2 1 2 0.4 20
Micronuclei, length 5.4 5.8 4.1 6.6 0.6 20
Micronuclei, width 4.3 4.1 3.3 5 0.5 20
Adoral membranelles, number 43.3 43 39 48 2.2 20
Frontal cirri, number 3 3 3 3 0 20
Buccal cirri, number 1 1 1 1 0 20
Cirri behind middle frontal cirrus, number 2 2 2 2 0 20
Cirri behind rightmost frontal cirrus, number 8.9 9 6 11 1.1 20
Long cirral rows, total number 19 19 17 21 1.3 20
Right long cirral rows, number 8.4 8 8 9 0.5 20
Left long cirral rows, number 10.6 11 9 13 1.3 20
Cirri in R4, number 34.1 34.5 30 38 2.5 10
Cirri in R5, number 47.6 47 43 52 2.9 10
Cirri in R6, number 55.9 55.5 51 63 4 10
Cirri in R7, number 58 54.5 52 85 9.8 10
Cirri in R8, number 55.9 55 52 62 3.3 10
Cirri in R9, number 54.3 54 49 60 3.1 10
Cirri in R10, number 47.9 48 43 52 2.5 10
Cirri in R11, number 42.8 42.5 37 48 3.1 10
Cirri in R12, number 40.3 41 38 42 2.1 3
Cirri in L1, number 35.2 35.5 34 37 1.1 10
Cirri in L2, number 37.2 38 32 40 2.7 10
Cirri in L3, number 36.8 37 32 40 2.1 10
Cirri in L4, number 37.6 38.5 32 43 3.7 10
Cirri in L5, number 37.1 36.5 34 41 2.5 10
Cirri in L6, number 37.7 37 33 44 3.4 10
Cirri in L7, number 40.8 40 36 46 3.3 10
Cirri in L8, number 42.7 42 39 47 2.4 10
Cirri in L9, number 42.3 43 37 47 3.2 10
Cirri in L10, number 42.4 41.5 41 46 1.8 10
Cirri in L11, number 40.4 39 36 45 3.5 7
Cirri in L12, number 42 42 42 42 0 2
Cirri in L13, number 45 45 45 45 0 1
Dorsal kineties, number 3 3 3 3 0 20
Dorsal bristles, length 2.1 2.1 1.7 2.5 0.3 10
Dikinetids in D1, number 28.2 28 22 32 3 10
Dikinetids in D2, number 33.5 34 30 36 2.4 10
Dikinetids in D3, number 22.5 22.5 20 25 1.6 10

Measurements are in micrometers and unless indicated, taken on Protargol stained specimens.
AZM, adoral zone of membranelles; D1–D3, dorsal rows of dikinetids 1–3; L1–L13, long rows of cirri on the left of the adoral zone of membranelles;

M, median; max., maximum value; min., minimum value; n, number of observations; R4–R12, long rows of cirri on the right of the adoral zone of
membranelles; SD, standard deviation; �X, arithmetic mean.

276 J. EUKARYOT. MICROBIOL., 57, NO. 3, MAY–JUNE 2010



Fig. 5–8. Morphogenesis during cell division of Deviata polycirrata n. sp. after Protargol impregnation. 5. Middle divider in ventral view. 6. Middle
divider in dorsal view. Primordia of dorsal kineties (arrowheads). Parental dikinetids among new proliferating basal bodies (�). 7. Late divider in ventral
view. Additional rows of cirri (arrowheads). Connection of anterior fragment of parental R6 with the new row generated by anlage VI of the proter (�). 8.
Late divider in dorsal view. Dark stained zones (?). Ma, fused macronuclear nodules; Mi, micronucleus; R4–R6, parental rows four to six; VI, anlage six
of the proter; I0–VI0, anlagen one to six of the opisthe. Scale bars 5 50mm.

277KÜPPERS & CLAPS—NEW KAHLIELLIDAE FROM ARGENTINA



Fig. 9–19. Micrographs of Deviata polycirrata n. sp. in interphase (9, 14–19) and during morphogenesis (10–13) after Protargol impregnation. 9. In-
terphase organism in ventral view. Cirral row that ends subequatorially (arrowhead). 10. Middle divider in ventral view. Field of basal bodies (arrowhead).
Posterior fragment of parental row R5 (�). 11. Magnification of field of basal bodies between anlagen VI and VI0. 12. Magnification of the right side of oral
primordium of the opisthe. 13. Late divider. Connection of anterior fragment of parental R6 with the new row generated by anlage VI of the proter (ar-
rowhead). 14–19. Disposition of undulating membranes. Arrowheads point to additional cirri (18, 19). Ma, fused macronuclear nodules. Scale bars 5 50mm.

278 J. EUKARYOT. MICROBIOL., 57, NO. 3, MAY–JUNE 2010



extra short row of nine cirri posterior to the buccal vertex. In one
further specimen, R3 surpassed the buccal vertex, while R4 ended
at the center of the body.

Notes on morphogenesis of cell division in Deviata polycir-
rata n. sp. (Fig. 5–8, 10–13). A few well silver-impregnated
middle and late dividers were observed. Unfortunately, no early
dividers were found. In the middle divider (Fig. 5–6, 10–12), the
adoral zone of membranelles of the opisthe is almost completely
differentiated, although some basal bodies at its posterior end are
still not forming membranelles. The anlagen I0–IV0 of the opisthe
are lengthened and very likely developed from the oral prim-
ordium. The parental right row R6 has split earlier and formed a
long primary primordium anteriad, which developed parallel to
and on the right side of the anterior fragment of parental row R6.
This long primary primordium split into anterior and posterior
secondary primordia, becoming the Anlage VI of the proter and
Anlage VI0 of the opisthe. An irregular field of basal bodies re-
mains between both these anlagen (Fig. 5, 11). Anlage VI0 of the
opisthe lengthens posteriad by kinetosomal proliferation within its
length. Anlage V0 of the opisthe apparently developed from the
field of basal bodies (disaggregated cirri) between the Anlagen VI
and VI0, migrated posteriad, and is about to connect to parental
row R5. At this stage, the anterior fragment of parental row R6
remains unchanged. The undulating membranes (UM) of the
proter are in the process of dedifferentiation. The leftmost fron-
tal cirrus differentiates from the distal end of the UM primordium
and Anlage II from disaggregating cirri of parental row R2.
Anlage III of the proter is still not generated. In the right paren-

tal rows of cirri R7–R10 anlagen are developing at an equatorial
location within the rows. The left parental rows of cirri remain
unchanged. Dorsal kinety anlagen are also developing within the
rows at two levels in parental kineties D1 and D2, but only at the
posterior part of parental dorsal kinety D3. Parental dikinetids re-
main among the new basal bodies. The macronuclear nodules are
still not fused.

In the late divider (Fig. 7, 8, 13) there are supernumerary rows
and the new cirri are segregated. The new row generated by Anl-
age VI of the proter is about to connect with an anterior fragment
of row R6 (Fig. 7, 13). Apparently, the proters Anlage V devel-
oped within the anterior parental fragment of row R6. Anlage IV
of the proter could have developed within the parental row R5
posteriad. There is an extra row between Anlagen III and IV, pos-
sibly resulting from a complex neokinetal anlagen development.
The UM of the proter are reorganizing and have arisen from a
longitudinal split of Anlage I. The leftmost frontal cirrus differ-
entiated from the distal end of the UM primordium and the cirri of
Anlagen II and III are already segregated. The new cirri from
anlagen I0 to VI0 of the opisthe are also segregated and only a few
widely spaced parental cirri remain posteriorly. There is an extra
row on the right of anlage VI0, which seems to be resorbing and
could also be the result of a neokinetal anlage. The adoral zone of
the opisthe is completely differentiated and Anlage I0 split longi-
tudinally to form both UM. In the remaining right and left long
rows of cirri (i.e. R7–R9, L1–L11) two anlagen developed within
parental rows of cirri, one anteriorly and another posteriorly, from
which almost all new cirri are segregated. The dorsal kineties are

Table 2. Morphometric data on Deviata bacilliformis.

Character �X M Min. Max. SD n

Body length in vivo 105.7 105 65 154 28.5 10
Body width in vivo 44.8 42 35 70 10 10
Body length 111 112 98 133 12.9 15
Body width 56.5 56 42 70 8 15
AZM, length 24 24.5 21 28 2.2 15
Adoral membranelles, number 21 21 19 24 1.5 15
Cytopharyngeal fibers, length 37.2 39.2 28 42 5.4 5
Macronuclear nodules, number 2 2 2 2 0 15
Macronuclear nodules, length 20.2 19.6 14.7 30.1 5 15
Macronuclear nodules, width 7.2 7.7 5.6 8.4 1 15
Micronuclei, number 2.7 2 2 4 0.7 15
Micronucleus, length 2.5 2.4 2.1 3.1 0.3 15
Micronucleus, width 2.3 2.1 2.1 2.8 0.3 15
Frontal cirri, number 3 3 3 3 0 15
Cirri behind rightmost frontal cirrus, number 1 1 1 1 0 15
Buccal cirri, number 1 1 1 1 0 15
Long rows of cirri, total number 11 11 9 13 1 15
Right long rows of cirri, number 5.5 6 4 6 0.6 15
Left long rows of cirri, number 5.1 5 4 7 0.7 15
Cirri in R4, number 4 4 3 5 0.6 15
Cirri in R5, number 10.6 10.5 9 13 1.3 10
Cirri in R6, number 27.8 27 24 33 2.6 10
Cirri in R7, number 23.4 23 16 32 4.1 10
Cirri in R8 number 22.3 21.5 16 32 4.1 10
Cirri in R9, number 17.5 17.5 15 21 2 10
Cirri in L1, number 21.1 20.5 18 25 2.5 10
Cirri in L2, number 18.8 19 14 24 3 10
Cirri in L3, number 18.4 17 16 23 2.5 10
Cirri in L4, number 18.4 18 15 24 3.1 10
Cirri in L5, number 19.7 19 16 25 2.9 10
Cirri in L6, number 21 21 21 21 0 1
Cirri in L7, number 19 19 19 19 0 1
Dorsal kineties, number 1 1 1 1 0 15
Dikinetids in dorsal row, number 15.4 15.5 12 17 1.4 10

See abbreviations in Table 1.

279KÜPPERS & CLAPS—NEW KAHLIELLIDAE FROM ARGENTINA



lengthened by development within the rows. Dorsomarginal kine-
ties and caudal cirri are not formed. Extra rows of cirri are later
resorbed because parental cirral rows are absent in the interphase
organisms. Macronuclear nodules appear condensed in a single
unified structure, and the micronucleus is dividing.

Description of an Argentinian population of Deviata bacil-
liformis (Gelei, 1954) Eigner, 1995 (Table 2 and Fig. 20–27).
The body size in vivo is of length 65–154 mm and width 35–
70 mm. The body shape is variable; sometimes it is vermiform,
other times it is slightly pyriform and wider, round in cross section
but dorsoventrally flattened anteriorly. The cytoplasm is grayish–
blackish at low magnification (o10X) due to inclusions. Cortical
granules are absent. The contractile vacuole is located on the left
in midbody and have anterior and posterior collecting canals (Fig.
20). The macronucleus is composed of two nodules that are vari-
able in shape. Some specimens have two ellipsoidal or reniform

nodules, while in other specimens the anterior-most nodule is
bilobed. There are two to four spherical or ellipsoidal micronuclei.
The adoral zone is composed of 19–24 membranelles and repre-
sents approximately 22% of the total body length (calculated on
the average values measured on silver-impregnated cells). In the
frontal field, there are nine cirri, which are termed R1–R4. The R1
has one cirrus—the leftmost frontal cirrus; R2 has two cirri—the
middle frontal cirrus and the buccal cirrus; R3 has also two cirri—
the rightmost frontal cirrus and one cirrus behind it; R4 begins on
the right of the cirrus behind the rightmost frontal cirrus and is
composed of three to five cirri (Fig. 21). The anterior-most frontal
cirri have nine basal bodies each, while the cirrus behind
the rightmost frontal cirrus is composed of six basal bodies. There
are 9–13 long rows of cirri. Five (4–6) of these rows are located on
the right of the adoral zone of membranelles, while 5 (4–7) are on
the left of it. The first row that extends past the buccal vertex is

Fig. 20–22. Morphology of Deviata bacilliformis from Argentina. 20. Ventral view in vivo. 21. Ventral view after Protargol impregnation. Barren
basal bodies (arrowheads). 22. Dorsal view after Protargol impregnation. Combined cirral rows (arrowheads). CV, contractile vacuole; D, dorsal kinety;
Ma, macronuclear nodule; Mi, micronucleus; L1–L5, left rows of cirri one to five; R3–R9, right rows of cirri three to nine. Scale bar 5 20 mm.

280 J. EUKARYOT. MICROBIOL., 57, NO. 3, MAY–JUNE 2010



R5, which begins nearly at the same level as the last cirrus of R4
and extends just posterior to the equator of the cell. The anterior-
most cirri of long rows are composed mainly of four basal bodies
each, while the cirri behind them have only two basal bodies each,
and are consequently more delicate. There are combined cirral
rows on the dorsal surface: the two rows on the right of the dorsal
kinety are preceded by two pairs of dikinetids with the anterior
basal body bearing a cilium (Fig. 22). The single dorsal row of
dikinetids extends from the anterior to the posterior of the body,
and is parallel to the last left row of cirri, which is generally L5
(Fig. 22).

A few specimens were observed in the first stages of stomato-
genesis (Fig. 25–27). One to three pairs of non-ciliferous basal
bodies proliferated between the row of cirri that ends subequa-
torially (R5) and the first left long row of cirri (L1). These cirri
apparently were formed de novo and later a longitudinally elon-
gated anarchic field develops between R5 and L1. The contribu-

tion of the posterior cirri of the adjacent right row R5 to this oral
primordium was not observed.

Deviata bacilliformis was isolated from soil samples from a
dried temporary pond collected in January 2004 and rewetted in
December 2006, and from samples collected and rewetted in Jan-
uary 2005.

DISCUSSION

The new kahliellid species, D. polycirrata n. sp., was placed
within the genus Deviata on the basis of the following significant
features: it usually presents a long ventral row of cirri that termi-
nates subequatorially, typical parental cirral rows are absent; and
the mode of replication of some right ventral rows of cirri, which
develop multiple anlagen within the row during division morpho-
genesis (Eigner 1995). It is worth mentioning that D. polycirrata

Fig. 23–27. Micrographs of Deviata bacilliformis in interphase (23, 24) and during stomatogenesis (25–27) after Protargol impregnation. 23. Ventral
view. Long row that ends subequatorially (arrowhead). 24. Dorsal view. Dorsal kinety (arrowhead). 25, 26. Magnification of proliferating barren basal
bodies between R5 and L1 (arrowheads). 27. Anarchic field of basal bodies of the oral primordium (arrowhead). L1, left long row one; R5, right long row
five. Scale bars 5 20 mm.

281KÜPPERS & CLAPS—NEW KAHLIELLIDAE FROM ARGENTINA



n. sp. resembles Parastrongylidium because there are numerous
spiraled rows of cirri, but in Parastrongylidium most of these
rows replicate by means of anlagen generated within the parental
rows (Aescht and Foissner 1992; Fleury and Fryd-Versavel 1984).

Comparison of Deviata polycirrata n. sp. with congeners. At
the moment, there are four species described within the genus
Deviata. These species can be distinguished by the number and
disposition of dorsal rows of dikinetids and/or the number of
macronuclear nodules (Berger and Foissner 1987). Deviata
polycirrata n. sp. differs from the type species, Deviata abbreves-
cence Eigner, 1995, mainly in the number of dorsal kineties (3 vs.
2), the number of cirral rows (17–21 vs. 7–8), and the number of
adoral membranelles (39–48 vs. 19–26, respectively; Eigner
1995). Compared with D. bacilliformis, this newly isolated spe-
cies not only possesses a higher number of dorsal kineties (3 vs.
1), more cirral rows (17–21 vs. 9–10), and more adoral mem-
branelles (39–48 vs. 18–21, respectively), but also its contractile
vacuoles are different because there are collecting canals only in
D. bacilliformis (Berger and Foissner 1987; Eigner 1995; Gelei
1954). Moreover, the species described by Fleury and Fryd-Ver-
savel (1984) as D. bacilliformis (formerly Kahliella bacilliformis)
has 3 dorsal kineties, but unlike D. polycirrata n. sp. there are
7–10 long rows of cirri. As was suggested by Eigner (1995), the
isolate of Fleury and Fryd-Versavel (1984) is possibly a different
species. In contrast to D. polycirrata n. sp., the type population of
D. estevesi has 2 dorsal kineties, 10 rows of cirri, 28–33 adoral
membranelles, and the contractile vacuole is also located in mid
body, away from the margins. Siqueira-Castro et al. (2009) trans-
ferred D. estevesi to the genus Parastrongylidium after studying
its morphogenesis. The new kahliellid described here differs from
D. rositae in the number of macronuclear nodules (4 vs. 7–14), the
number of dorsal kineties (3 vs. 2), the number of rows of cirri
(17–21 vs. 6), and the number of adoral membranelles (39–48 vs.
14–18, respectively) (Küppers et al. 2007). Finally, like D.
polycirrata n. sp., D. brasiliensis Siqueira-Castro, Paiva & Silva
Neto, 2009 usually has 4 macronuclear nodules, but these species
differ in the number of long rows of cirri (17–21 vs. 7–11), the
number of dorsal kineties (3 vs. 2), and the number of adoral
membranelles (39–48 vs. 18–31, respectively).

Within the genus Kahliella Corliss, 1960 there are some species
with four macronuclear nodules that should also be compared with
the new isolate reported here. Kahliella spirostoma Alekperov,
1988 was found among putrefying vegetation from a freshwater
reservoir in Azerbaijan (Alekperov 1988) and, like D. polycirrata
n. sp., has a high number of adoral membranelles (45–50 vs. 39–
48) and three dorsal rows of bristles, but the latter differ in the
number of long rows of cirri (3 rows on the right of the adoral
zone of membranelles (AZM), 2 rows on the left of it vs. 8–9 on
the right of the AZM and 9–13 on the left of it). Another species
with four macronuclear nodules is Kahliella quadrinucleata Drag-
esco, 2003; but relative to that species, the new isolate described
here presents also a higher number of long rows of cirri (17–21 vs.
10–12), of dorsal kineties (3 vs. 2), and more adoral membranelles
(on average 43 vs. 22). Unlike the species described in the genus
Deviata, in both K. spirostoma and K. quadrinucleata, the first
right long row of cirri on the ventral side terminates at the pos-
terior end of the body, as can be seen from the illustrations in
Alekperov (1988) and in Dragesco (2003). In Deviata this row
ends equatorially, and this characteristic is considered diagnostic
for the genus (Eigner 1995). Further morphological and ontoge-
netic studies on K. spirostoma and K. quadrinucleata are needed
to determine whether these species should be transferred to the
genus Deviata.

Notes on morphogenesis during cell division of Deviata
polycirrata n. sp. We observed some well silver-impregnated
morphogenetic stages that should be compared with the morpho-

genesis of D. abbrevescens studied by Eigner (1995), and with
that of D. brasiliensis studied by Siqueira-Castro et al. (2009). In
the new isolate, Anlagen VI for the proter and for the opisthe (VI0)
seem to develop from a split long primary primordium. Anlage V0

of the opisthe is generated in the field of basal bodies (disaggre-
gated cirri) between the fragmented long primary primordium
(Anlagen VI and VI0) and later deviates to and develops within
the posterior fragment of parental row R5, as occurs in
D. abbrevescence and possibly in D. brasiliensis. On the con-
trary, the rows R4 and R5 of the proter seem to be generated in a
different way in the three species. In D. polycirrata n. sp., Anl-
agen IV and V of the proter possibly develop from R5 and R6,
respectively; while in D. abbrevescence Anlage IV originates in
R4 (I4 in Eigner 1995) and further extends into R5 (I5), with
Anlage V originating from the anterior region of R6 (I6). In D.
brasiliensis, Anlagen IV and V are formed in the anterior regions
of R4 and R5, respectively. In addition, in the present new species
some morphogenetic events occur at a different time compared
with D. abbrevescens. For instance, at the same time the long
primary primordium splits and the Anlage V0 is connected to the
posterior parental fragment of row R5, Anlagen IV and V of the
proter are still not generated. A comparison of Eigner’s Fig. 12
would indicate that these phenomena occur at the same time in D.
abbrevescens. In the late divider, when Anlage VI of the proter is
about to connect with the old parental anterior fragment of row R6
in D. polycirrata n. sp., there are extra rows between Anlagen III
and IV of the proter and also an extra row on the right of Anlage
VI0 of the opisthe. As has been stated by Eigner (1995), these
extra rows could result from the complex neokinetal anlagen
development and are later resorbed, because extra rows are
absent in the interphase organism. The field of barren basal
bodies between Anlagen VI of the proter and Anlage VI0 of the
opisthe was not mentioned by either Eigner (1995), or Siqueira-
Castro et al. (2009). These basal bodies persisted in the late
divider as well.

The presence of extra rows of cirri in four interphase specimens
from the present study is not considered as retention of parental
cirri, because these were densely packed within the rows and the
rows did not have half the cirri of a neighboring row, as was sug-
gested by Eigner (1995) to recognize a retained parental row in
interphase. These specimens could also be interpreted as postdi-
viders with parental or neokinetally generated rows still unre-
sorbed completely.

Further studies are needed to make a complete description of
the morphogenesis during cell division of D. polycirrata n. sp.,
although the observations on the few dividers found support the
placement of the new isolate within the genus Deviata. As stated
in preceding paragraphs, D. polycirrata n. sp. typically differs
from all its congeners by having three dorsal rows of dikinetids, a
higher number of cirral rows, and a higher number of adoral
membranelles. Unfortunately, gene sequence data were not sur-
veyed at the time of preparing the slides in 2004. Nowadays, these
data are increasingly being considered necessary for the accurate
description of new species of protists, especially in those forms
that are not as rich in morphological features (Lynn and Simpson
2009). Nevertheless, morphological traits make this new isolate
clearly different from the other species within the genus Deviata
and diagnosis of the ciliate as a new species is warranted.

Subclass Stichotrichia Small & Lynn
Family Kahliellidae Tuffrau
Deviata polycirrata n. sp.

Diagnosis. Body size in vivo, 130–180 � 45–70 mm, with 8–9
long cirral rows on the right of the adoral zone membranelles and
9–13 on the left of the latter, plus 3 long dorsal rows of dikinetids.
Adoral zone with 39–48 membranelles. With four macronuclear

282 J. EUKARYOT. MICROBIOL., 57, NO. 3, MAY–JUNE 2010



nodules and one to two micronuclei. Single contractile vacuole
located on the left body margin.

Etymology. The term polycirrata is a composite from the
Greek poly- (many), and the Latin cirratus (cirrated, curly-
haired), and refers to the presence of numerous rows of cirri.

Type locality. Near Dolores city, Buenos Aires Province,
Argentina, near the Provincial Route No. 63 (3611805500S,
5713201200W).

Type habitat. Temporary pond covered by the floating mac-
rophytes Lemna sp., Spirodella sp., and Limnobioum spongiae.
The new isolate was found under the following physical–chemical
conditions, which were measured at the sampling site: pH 7; elec-
trical conductivity, 242mS/cm; water temperature, 15.5 1C; total
dissolved solids, 0.16 g/l; depth, 40–55 cm.

Type material. One Protargol-stained hapantotype slide is de-
posited in the Colección de Invertebrados from Museo de La
Plata, Buenos Aires Province, Argentina (accession number MLP
63), and a paratype slide is registered in the Ciliate Type Slide
Collection of the Smithsonian Institution, Washington, USA
(accession number USNM1135042).

Deviata bacilliformis was discovered in Hungary (Gelei 1954)
and later recorded by other authors in Israel (Berger and Foissner
1987). Fleury and Fryd-Versavel (1984) found it in France,
although it was possibly misidentified (Eigner 1995). In Africa,
Dragesco (2003) recorded D. bacilliformis in moss samples from
Rwanda, and Foissner, Agatha, and Berger (2002) found this spe-
cies in terrestrial habitats from Namibia. For Argentina and the
rest of South America, it represents a new finding. The morpho-
logical characters of D. bacilliformis in this study differ from the
observations of Berger and Foissner (1987) in the number of rows
of cirri (9–11 or occasionally 13 vs. 10–11), the number of mac-
ronuclear nodules (invariably 2 vs. 2–4), and in the presence of
combined cirral rows on the dorsal surface. The most constant
trait in the Argentinian population, comparable to the European
isolates, is the pattern of cirri in the frontal field. In contrast, the
African population of D. bacilliformis described by Dragesco
(2003) presents three to five macronuclear nodules, and as seen
from the illustrations (fig. 33–37 in Dragesco 2003), there are two
cirri behind the rightmost frontal cirrus (vs. one cirrus in the pres-
ent study and in Berger and Foissner 1987), and the short row of
cirri in the frontal field is lacking (R4 with three to five cirri in the
present study; row five with three to six cirri in Berger and
Foissner 1987).

The formation of the oral primordium during the morphogen-
esis in D. bacilliformis begins apparently de novo, from the barren
basal bodies between R5 and L1. These basal bodies proliferate
and an anarchic longitudinal field develops between R5 and L1. In
D. abbrevescens stomatogenesis begins with the proliferation of
basal bodies by disaggregation of the fifth and sixth posterior cirri
of the row that ends at the center of the ventral surface (I4 sensu
Eigner 1995), eventually forming two conspicuous fields that later
fuse. Then the posteriormost cirri of I4 contribute to the formation
of triangle-shaped primordium that differentiates the membrane-
lles. Whether posteriormost cirri of R5 contribute to the formation
of the oral primordium in D. bacilliformis could not be determined
because individuals in later stages of stomatogenesis were not
found. The presence of non-ciliferous basal bodies between the
right row that ends at the subequatorial region of the cell and the
first left row of cirri was also observed in D. rositae, although
specimens in subsequent morphogenesis stages were also not
found by Küppers et al. (2007). Siqueira-Castro et al. (2009) also
reported similar structures in one specimen of D. brasiliensis, but
with no apparent relationship to the formation of the oral prim-
ordium. It is worth mentioning that stomatogenesis is apokinetal
in Parastrogylidium martini Fleury and Fryd-Versavel, 1984 and

Aescht and Foissner (1992) stated that the morphogenesis of Pa-
rastrongylidium oswaldi Aescht and Foissner, 1992 occurs as in P.
martini. Further ontogenetic studies of species within the genus
Deviata are necessary to ascertain the stomatogenetic mode, with
the phylogenetic consequences that these would imply, and also to
clarify the complex neokinetal anlage development in some ven-
tral cirral rows.

ACKNOWLEDGMENTS

Dr. Bettina Gullo from Facultad de Ciencias Naturales y
Museo, Universidad Nacional de La Plata, Argentina, is greatly
acknowledged for bringing the samples where the new species was
found. We are also thankful to Prof. Dr. Norbert Wilbert for his
constant help and advice; to Santiago Nenda for his help with the
photographs; to Drs. Lı́a Lunaschi and Fabiana Drago for the use of
the drawing tube in their laboratory; and to Dr. Donald F. Haggerty,
a native English speaker, for editing the manuscript. Consejo Nac-
ional de Investigaciones Cientı́ficas y Técnicas (CONICET) is
acknowledged for financial support. This is contribution number
877 from the Instituto de Limnologı́a Dr. R. A. Ringuelet.

LITERATURE CITED

Aescht, E. & Foissner, W. 1992. Biology of a high-rate activated sludge
plant of a pharmaceutical company. Archiv. Hydrobiol., 90(Suppl.):
207–251.

Alekperov, I. K. 1988. Two new species of infusoria (Ciliophora, Hypot-
richida) from fresh waters of Azerbaijan. Zool. Zh., 67:777–780.

Berger, H. & Foissner, W. 1987. Morphology and biometry of some soil
hypotrichs (Protozoa: Ciliophora). Zool. Jb. Syst., 114:193–239.

Corliss, J. O. 1960. The problem of homonyms among generic names of
ciliated protozoa, with proposal of several new names. J. Protozool.,
7:269–278.

Dragesco, J. 2003. Infaciliature et morphometrie de vingt espèces de ciliés
hypotriches recoltés au Rwanda et Burundi, comprenant Kahliella
quadrinucleata n. sp. Pleurotricha multinucleata n. sp. et Laurentiella
bergeri n. sp. Trav. Mus. Natl. Hist. Nat. ‘‘Grigore Antipa’’, 45:7–59.

Eigner, P. 1995. Divisional morphogenesis in Deviata abbrevescens nov.
gen., nov. spec., Neogeneia hortualis nov. gen., nov. spec., and Kahli-
ella simplex (Horváth) Corliss and redefinition of the Kahliellidae
(Ciliophora, Hypotrichida). Eur. J. Protistol., 31:341–366.

Eigner, P. 1997. Evolution of morphogenetic processes in the Orthoamp-
hisiellidae n. fam., Oxytrichidae, and Parakahliellidae n. fam., and
their depiction using a computer method (Ciliophora, Hypotrichida).
J. Eukaryot. Microbiol., 44:553–373.

Fleury, A. & Fryd-Versavel, G. 1984. Unité et diversité chez les
hypotriches (Protozoaires Ciliés) I. – Approche morphogénétique
par l’ étude de quelques formes peu différenciées. Protistologica,
20:525–546.

Foissner, W., Agatha, S. & Berger, H. 2002. Soil ciliates (Protozoa,
Ciliophora) from Namibia (Southwest Africa), with emphasis on two
contrasting environments, the Etosha region and the Namib desert.
Denisia, 5:1–1063.

Gelei, J. von. 1954. Über die Lebensgemeinschaft einiger temporärer
Tümpel auf einer Bergwiese im Börzsönygebirge (Oberungarn). III.
Ciliaten. Acta Biol. Hung., 5:259–343.

Küppers, G. C., Lopretto, E. C. & Claps, M. C. 2007. Description of
Deviata rositae n. sp., a new ciliate species (Ciliophora, Stichotrichia)
from Argentina. J. Eukaryot. Microbiol., 54:443–447.

Lynn, D. H. 2008. The Ciliated Protozoa. Characterization, Classification,
and Guide to the Literature. 3rd ed. Springer, Dordrecht, the Nether-
lands. p. 15–61, 357–359, 361–362.

Lynn, D. H. & Simpson, A. G. B. 2009. From the editors: describing new
taxa of unicellular protists. J. Eukaryot. Microbiol., 56:403–405.

Paiva, T. S. & Silva-Neto, I. D. da. 2005. Deviata estevesi sp. n.
(Ciliophora: Spirotrichea), a new ciliate protist from a restinga lagoon
in Rio de Janeiro, Brazil. Acta Protozool., 44:351–362.

283KÜPPERS & CLAPS—NEW KAHLIELLIDAE FROM ARGENTINA



Siqueira-Castro, I. C. V., Paiva, T. S. & Silva-Neto, I. D. da. 2009. Mor-
phology of Parastrongylidium estevesi comb. nov. and Deviata brasi-
liensis sp. nov. (Ciliophora: Stichotrichia) from a sewage treatment
plant in Rio de Janeiro, Brazil. Zoologia, 26:774–786.

Tuffrau, M. 1979. Une nouvelle familie d’hypotriches, Kahliellidae n.
fam., et ses consequences dans la repartition des Stichotrichina. Trans.
Am. Micros. Soc., 98:521–528.

Wilbert, N. 1975. Eine verbesserte Technik der Protargolimprägnation für
Ciliaten. Mikrokosmos, 64:171–179.

Received: 11/15/09, 02/03/10; accepted: 02/05/10

284 J. EUKARYOT. MICROBIOL., 57, NO. 3, MAY–JUNE 2010