PROCEEDINGS
OF THE
INDIAN ACADEMY OF SCIENCES
VOL. X
SECTION B
BANGALORE CITY :
PRINTED AT THE BANGALORE PRESS, MYSORE ROAD
1940 '
CONTENTS
SECTION B VOL. X
No. 1 July, 1939
PA UK
Competition in Fungi. I. A Study of the Growth Reactions of
Non-Parasitic Fungi in Associated Culture . . T. S. SADASIVAN 1
Effectiveness of Chemical Fertilisers on the Growth and Water
Requirement of Wheat .... B. N". SINGH AND J. B. SINGH 27
Some Aspects of the Anatomy of Anura (Amphibia,) A Review
L. 8. 11 A MAS w AMI 41
Oxidation of Thiols and Ascorbic Acid in the Latex of Papaya .
0. V. GANAPATH.Y AND B. N. SASTRI 81
On the Developmental Morphology of Androgynous Receptacles in
Marchantia palmata Nees K. S. SRINIVASAN 88
Bryozoa from the Bagh Beds .--. G. W. CHIPLONICER 98
No. 2August, 1939
Studies in HelminthologyTrernatode Parasites of Birds
JVlAKUNi) BJ-CHARI LAL 111
Little Leaf A Transmissible Disease of Brinjal
K. M. THOMAS AND C. S. KRISHNASWAMI 201
Testicular Ova in Uraeotyphlus narayani Seshachar
B. B. SBSIIAOHAB 213
Two New Additions to the List of the Indian Aspergilli ....
* I. FROILANO DE MELLO 21.8
No. 3 September, 1939
Compounds of Phosphorus in Milk I .
B. N". ACFIARYA AND S, 0. DEVADATTA 221
Phosphorus, Calcium and Magnesium in Milk II
1>. N. AOHARYA AND S. C. DBVADATTA 229
IV
No. 4 October, 1939
PAGK
Studies in Sorghum Sudanense, Stapf The Sudan Grass ....
. G. N. BANGASWAMI AY Y AN GAR AND B. W. X. PONNAIYA 237
Lameliibranchs from the Bagh Beds . . . . G. W. CHIPLONKEB, 255
On Some Stone Implements from Hoshangabad (Central Provinces)
MANOHAH LAL MTSRA 275
Noe 5 November, 1939
On Some Nematode Parasites from Afghanistan . , 8. A. AKHTAK. 287
No. 6 December, 1939
A Systematic Account of Some South Indian Diatoms
G. VENKATARAMAN 293
Study in Diseases of Fish: Fin-Rot A Bacterial Disease of Fins of
Fish HAMIDKHAN 369
COMPETITION IN FUNGI
I. A Study of the Growth Reactions of Non-Parasitic Fungi
in Associated Culture
BY T. S. SADASIVAN
(From the Department of Botany, Lucknow University}
Received November 14, 1938
(Communicated by Dr. S. N. Das Gupta, Ph.D., D.I.C.)
Introduction
THE earlier works on mixed culture are mostly studies on growth reactions
of fully developed mycelia in a given medium. The present work was under-
taken to study the influence of one fungus upon another in paired and asso-
ciated cultures in media variously modified, with a view to gaining an in-
sight into the nature of competition in fungi.
In nature, competition occurs whenever more than one fungi associate
together, and the success or failure of a fungus depends upon the chemical
and physical nature of the habitat and upon the environment. It must
also depend, to some extent, upon the effect the fungi exert upon one another.
A study of the latter aspect is the main objective of this paper.
Growth reactions in artificial culture are not necessarily true indications
of what may occur on host in nature (Machacek, 1928), but nevertheless, it
was thought that studies in simpler media are likely to throw more light on
general problem of competition, particularly with regard to saprophytes, than
studies using complex host tissues.
This paper which is the first of a series deals with the growth reactions
primarily of two non-parasitic fungi, Fusarium and Dendryphiella, in asso-
ciated cultures. The detailed investigation was restricted to these two fungi
only since it was found that a larger number of combinations would make
the work unwieldy. Similar experiments with parasitic organisms will follow
in subsequent papers of the series.
Material and Method
Six fungi were utilized for preliminary observation. The fungi differed
both in the microscopic and macroscopic characters, and these afforded an
1
Bl p l
2 T. S. Sadasivan
easv means of distinguishing the competing strains in mixed growths. Out
of these six only two fungi were chosen for a detailed investigation. The
names of the fungi together with a very short description of their macro-
scopic characters are given below.
1. Fttsarium sp. - - Mycelium white, moderately fluffy, non-zoning,
submerged mycelium very little.
2. Dendrvphiella sp. . . Mycelium dark, fluffy, non-zoning, submerged
mycelium abundant.
3. Phoma sp. . . Mycelium white, fluffy, zoning with distinct
dark-red rings. Visible as light band at the
top very clearly in the substratum.
4. Helminthosporium sp. . . Mycelium greyish-white, slightly fluffy, zoning,
submerged mycelium very little.
5. Monilia sp. . . Mycelium dark-grey, moderately fluffy, zoning
with rings of darker grey colour. Submerged
mycelium very little.
6. Gibberella sp. . . Mycelium whitish-grey, mostly superficial, thin
growth, non- zoning.
The actual combinations in which the fungi were paired are given
along with the experimental details.
The method of investigation consisted in inoculating a given plate of
nutritive medium with two fungi (members of a pair) and comparing the
growth rates and the relative area occupied by the competing fungi with
those of the controls.
The strains were inoculated at different degrees of proximity, viz., 2 cm.
apart, adjacent (two inocula touching each other) and mixed, with individual
controls.
For purposes of getting the average rates of growth and also for verifica-
tion of results replicates of three petri-dishes were used in each kind of in-
oculation with two controls for each of the strains.
The growth rates of the competing fungi and of the controls were noted
every twenty-four bours together with other features of interest. The
growths arising from mixed inocula were examined under binocular micro-
scope whenever required.
The standard synthetic medium used throughout the experiment as
basal medium, except where otherwise stated, had the composition : glucose
Competition in F^tng^ / 3
2-Ogtn., potassium nitrate 2-0 gm., magnesium sulphate 75 gm., potas-
sium phosphate 1-25 gm., potato starch 10-0 gm., shredded agar 15-0 gm.
and distilled water 1 litre.
The detailed work on Fusarium and Dendryphiella was carried out in
the standard medium variously modified by the addition of N/10 hydro-
chloric acid and malic acid in acid series, sodium carbonate and sodium
hydroxide in alkali series, in different percentages ; the details of which are
given in appropriate places.
Petri-dishes 2 cm. deep and 11 cm. in diameter were used in all cases.
The different media were poured 1 - 5 cm. deep to enable to make necessary
observations in the substratum as well. The usual methods of sterilizing
petri-dishes and media -were employed.
At the commencement of the work the purity of the various fungi was
assured by taking monohyphal cultures and maintaining stock cultures in
tubes of standard synthetic medium.
For purposes of inoculations equal amount of inocula were used from the
peripheral regions of tlie cultures of four or five days growth in standard
media and for mixed inocula cultures these were thoroughly mixed by means
of a scalpel and then inoculated.
All operations were carried out in aseptic condition.
The work was done at temperatures ranging between 18-20 C.
Preliminary Observations
As has been already stated six strains were utilized for the experiment.
The monohyphal cultures cf all these six strains were grown in standard
synthetic medium in order to compare their growth rates and their morpho-
logical characters. A short account of the macroscopical characters of each
of the strains has already been given ; the rate of growth for different strains
is given graphically in Text-Fig. L
4
T. S. Sadasivan
80
03 45 67
Number of days
8
Monilia sp.
Fusanum sp.
Dendryphiella sp.
Phoma sp.
Gibberella sp.
Helminthosporium sp.
TEST-FIG. 1. Graphs showing the radial spread in mm. of the fungi Monilia sp.,
Gibberella sp., Dendryphiella sp., Htdminthosforium sp., Phoma sp.
and Fusarlum sp. in standard synthetic medium.
It will be seen from Text-Fig. 1 that Monilia has the fastest growth rate
followed by Gibberella, Helminthosporium, Fusarium, Dendryphiella and
Phoww in descending order.
For experiment on competition the strains were paired in such a way
that each member of a pair had mycelium of different colour, to enable easy
detection of hyphse in mixed growth. The combinations employed were
as follows :
1. Fusarium
and
Dendryphiella
2. Fusarium
and
Gibberella
. . white, non-zoning,
dark, non-zoning,
white, non-zoning.
. . greyish-white, non-zoning.
Competition in Fungi / 5
3. Phoma . . white, zoning.
and
Helminthosporium . . greyish-white, zoning.
4. Phoma . . white, zoning.
and
Monilia . . dark-grey, zoning.
5. Phoma . . white, zoning.
and
Dendryphiella . . dark, non-zoning.
These six strains in the above five combinations were inoculated 2 cm.
apart, adjacent and mixed, in standard synthetic medium plates. Controls
were kept.
The results show that in the combinations employed the growth rates
of the associated cultures were not much different from those of the control
strains.
In adjacent and mixed cultures the faster strain usually enveloped the
slow growing strain and the later appeared as sectors. For example in
adjacent and mixed cultures of Dendryphiella and Phoma, Dendryphiella
which is slightly fast growing occupied the major portion of the growth.
Mixed and adjacent cultures of Dendryphiella and Phoma were, however,
exceptions. Although the former was decidedly fast-growing, it was the slow
growing Phoma that dominated, and Helminthosporium appeared as small
sectors in a major growth of Phoma.
The appearance of sectors as found in the above mixed cultures in
isolated areas is interesting. Obviously these had their origin in the original
inoculum, but their continuity was difficult to establish. Among the other
features of interest were the behaviour in regard to the formation of colour
and zonation in the mixed growths.
Colour. In the region where the young hyphse of Monilia and Phoma
arising from inocula planted 2 cm. apart came in contact with each other,
there a few millimetres inside the growing edge of Phoma culture, was formed
a single dark-red band very clearly visible in the substratum. As the faster
Monilia colony started encircling the Phoma, the dark-red band appeared at
the region of fresh hyphal contact, while the colour became fainter at th,e
first place of appearance. The red band gradually moved further up where
the younger hyphse of Monilia had come in contact with younger hyph;e of
Phoma.
Bl a F
6 T, S, Sadasivan
When the entire Phoma colony was enveloped by Monilia and the
younger hyphse of the latter occupied a position directly opposite to the
first place of contact with Phoma the band moved to the new place of contact,
the colour at the original place having disappeared.
The formation of colour at the line of contact of two colonies is well
known but the subsequent disappearance as noted here is peculiar.
The fresh, younger hyphse of Monilia induces the formation of the dark-
red band in Phoma colony. This must be due to the interaction of the
staling products of the two colonies. The subsequent disappearance of the
colour band may be attributed to the excess of a chemical substance due to
its accumulation or to the formation of a new chemical by the older
mycelium.
Zoning. In normal cultures Phoma and Helminthosporium produce
zonation although Dendryphiella does not. When Phoma and Helmintho-
sporium were inoculated adjacently to form a mixed growth Helmintho-
sporium appeared as sectors. In the mixed growth the zonation of both
Phoma and Helminthosporium persisted ; the red concentric rings of the former
forming almost an unbroken ring with dark bands of the latter. When the
growth arose from the mixture of inocula, the important point observed was
the complete absence of zonation in Phoma and the persistence of the same
in Helminthosporium. The identical results were obtained with Phoma
combined with Dendryphiella. In adjacent cultures Phoma zoned very well
as in controls but in mixture culture zonations in Phoma were singularly
absent.
Detailed Observations
With a view to making a detailed investigation it was decided to
concentrate on two strains, that is one pair of strains only, instead of the five
pairs used for preliminary observation. In order to find out the best reacting
strains all the six strains were inoculated individually in 0-1, 0-25, 0-5, 1-0
and 2-0 per cent, malic acid in standard synthetic medium. Dendryphiella
and Fusarium having proved more sensitive to the acids, were chosen for the
investigation.
As has already been seen Fusarium and Dendryphiella are also easily
distinguishable microscopically by the striking difference in the mycelial
colour -.Fusarium being white and Dendryphiella dark-grey. Microscopically
the difference lay in the hyphal characters and branching. Further profuse
sporulation occurred in Dendryphiella where its growth met that of Fusarium.
This last character particularly facilitated the detection of the boundary of
Competition in Fungi /
the Dendryphiella growth, especially in the case of partial and complete
suppression. For these experiments the fungi were inoculated 2 cm. apart
and adjacently.
Glucose series. The combined effect of various salts in the standard
medium having failed to yield much result, individual salts like glucose and
potassium nitrate were tried in different concentrations using agar as the
basal medium.
Glucose was added to 1-5 per cent, agar in the strengths 0-1, 0-2, 0-4,
0-5, 0-6, 0-75, 1-0, 1-5, 2-0, 4-0, 6-0, 8-0, 10-0 and 12-0 per cent, steamed
instead of autoclaved for sterilization and plated. Fusarium and Dendry-
phiella were inoculated 2 cm. apart and adjacent controls were kept.
The results are given in Text-Figs. 2 and 3 where the diameters of six
days growths are plotted against the glucose concentration for 2 cm. apart
and adjacent cultures respectively.
60
50
1 30
20
10
01 02
" 075 ..... 1 ; """15 20 4t
Glucose per cent.
6Cf~8 r O ...... 100 1 ~12 :
Fusarium (Control)
,, (2 cm. apart)
Dendrypliiclla (Control)
,, (2 cm. apart)
TEXT- FIG. 2. Graphs showing the behaviour of F-usar'nim sp. and. Dcudryphiclla sp.
in, associated culture and in control in glucose series. Inocula
placed 2 cm. apart.
T. S. Sadasivan
1 30
fe 20
^
\A A
^
i
* ' B ^ / %
f *io * \ o m
.5 10
i
t
100-20304W06 075 10 15 20 40
Glucose per cent.
&0 BO 10-0 12-0
Fusarhim
Dendryphiella
(Control)
(Adjacent)
(Control)
(Adjacent)
. 3. Graphs showing the. behaviour of the two fungi F-usarium sp. and Dendry-
phlella sp. in associated culture and in control in glucose series.
Inocula placed adjacently.
It will be seen from Text-Fig. 2 that in the cultures where the fungi had
been inoculated 2 cm. apart :
1. Fusanum and Dendryphiella are inhibited in the presence of each
other as compared to the controls. Of these, however, Fusarium shows a
greater inhibition than Dendryphiella.
2. At lower concentrations of glucose 0-10 -6 per cent. Fusarium
has a lower growth rate than Dendryphiella both in control and in paired
cultures. At 0-75 per cent. Fusarium accelerates and throughout the higher
concentrations employed, the one in control remains by far the fast
growing.
3. In completing cultures at concentrations 0-3 to 4-0 per cent.
Dendryphiella has a faster growth rate than Fusarium, but beyond 4 it is
the other way round.
4. At concentrations 0-1 to 0-6 per cent. Fusarium and Dendryphiella
colonies do not meet when inoculated 2 cm. apart and the mycelia of the two
growths thin out as they approach each other.
It will be seen from Text-Fig. 3 that in adjacent culture :
1. Throughout the series Fusarium and Dendryphiella are inhibited in
the presence of each other as compared with the controls except in the case
Competition in fungi /
of Fusarium at 0-3 per cent. Of these however, Dendryphiella is very
markedly inhibited by Fusarium.
2. Fusarium dominated throughout. Dendryphiella in addition to
being retarded has three points of complete inhibition at 0-1, 0-2 and 0-4
per cent. There is a slight growth at 3 per cent, which probably is due to
the associated Fusarium being slightly less active at that concentration.
The almost complete inhibition at concentrations 0-1-0 -4 is especially
interesting since about that region Dendryphiella in the control has a faster
growth than Fusarium.
3. As in the case of 2 cm.-apart-culture from 0-75 per cent, and
beyond Fusarium outgrows Dendryphiella.
4. In competing cultures Fusarium always has a decidedly faster
growth rate.
In Text-Fig. 4 is illustrated diagrammatically the dominance of Fusarium
over Dendryphiella in adjacent cultures where Fig. 4 (A) indicates the complete
A.
TEXT-.FIG. 4 AF. 'Diagrammatic representation of the relative area occupied by
Fusarium. sp. and Dendryphiella sp. in various concentrations of
glucose. Inocula adjacent.
Shaded area represents Fusarium.
Unshaded area represents Dendryphiella.
Dots indicate sporulation of Dendryphiella.
Fusarium and. Dendryphiella inoculated adjacent in 0-1, 0-2 and 0*4%
glucose showing complete suppression of Dendryphiella.
Same strains in 0-5% glucose showing the appearance of Dendryphiella.
Same strains in 1-5% glucose showing an acceleration of Dendryphiella
growth over that in 0-5%.
Same strains in 2 -0 % glucose showing an acceleration.
Same strains in 1-0% glucose showing a retardation of Dendryphiella.
Dendryphiella showing an acceleration over 4*0% with, the addition of
8-0%.
D.
E.
F.
10
T, S* Sadaslvan
inhibition of Dendryphiella as seen in -1, 2, -4% glucose ; the fluctua-
tions in the relative area occupied by the two strains in other concentra-
tions are shown in Fig. 4 (A-F).
Potassium nitrate series. Potassium nitrate was next tried with 1-5
per cent, agar in strengths 0-1, 0-25, 0-5, 0-75, 1 -0, 1-5, 2-0, 4-0, 6-0,
8-0, 10-0 and 12-0 per cent. Fusarium and Dendryphiella were inoculated
2 cm. apart and adjacently. Controls were kept.
The results are given graphically in Text-Figs. 5 and 6 where the dia-
meters of six days growths are plotted against the potassium nitrate concen-
trations for 2 cm. apart and adjacent cultures respectively.
10025
: 5 ~ 075^ 10 K'SCT^ 6 :
Potassium nitrate per cent.
Fusarium (Control)
80 10-0 12-0
o ^a- Dendryphiella
(2 cm. apart)
(Control)
TEXT-FIG. 5.-
mmmwmmmotmautmm (2 cm. apart)
-Graphs showing the behaviour of Fusarium sp. and Dendrijphivlla, sp.
in associated culture and in control in KNO 3 series. Iiicoula placed
2 cm. apart.
It will be seen from Text-Fig. 5 that in cultures 2 cm. apart, there was
virtually no difference in the growth rates shown by Dendryphiella, Fusarium
and their controls, up to the concentrations of potassium nitrate 0-1-1-5
per cent. Beyond this concentration Fusarium definitely takes upper hand,
but nevertheless the fungi in associated cultures have almost similar growth
rates to those of the controls, showing that the interaction of the two fungi
has resulted into inhibition of growth rates only to an insignificant extent.
Competition in Fungi* /
11
In adjacent culture (Text-Fig. 6) in media up to the concentration of
0-75 per cent, the growth rates of associated cultures and the controls are
01 025 05 075 "10 1-5 20 40 60 80
Potassium nitrate per cent.
Fusarium
---- o ---- Dendryphiella
(Control)
(Adjacent)
(Control)
(Adjacent)
TEXT-FIG. 6.-
-Graphs showing the behaviour of: Fusariu.m sp. and Dendryphiella sp.
in associated culture and in control in I\NO :l series. Inocula placed
ad j ace ally.
almost the same except for that of Fusarium which shows a definite retarda-
tion. Beyond 0-75 per cent. Fusarium, both in the control and in associated
culture, is much faster than Dendryphiella. In concentrations 0-1 to 0-5
per cent. Dendryphiella dominates Fusarium, they are almost equal at
0-75 per cent, but beyond that concentration Fusarium completely domi-
nates Dendryphiella as is shown by the difference in the growth rate between
the two. At 4: per cent. Dendryphiella shows very slight growth which is
completely inhibited at 10 per cent, although the control maintains a
steady rate of growth. Dendryphiella seems to exert no influence up on
Fusarium at relatively higher concentrations as judged from its rate of
growth.
The domination of Fusarium by Dendryphiella in lower concentrations
of potassium nitrate (0-1-0 -5) and the domination of Dendryphiella by
Fusarium in higher concentrations (1-0-10-0) and complete inhibition of
the former by Fusarium in still higher concentrations are illustrated dia-
grammatically in Text-Fig. 7 (A-I).
12
T, S* Sadaslvan
iG. 7 ..4-7. Diagrammatic representation of relative area occupied by F-usariwn.
and Dendryphiella in various concentrations of KNO 3 . Inocula
adjacent.
Shaded area represents Fusaruim.
Unshaded area represents Dendryphiella.
Dots indicate sporulation of Dendryphiella.
A. Fusarium recessive with a sectorial growth in 0-1 % ;
B. Fusarium accelerates with addition of .25% ;
C. Fusarium further accelerates with addition of 0-5 % ;
D. Fusarium growth is equal to that of Dendryphiella growth in 0-75% ;
E. Dendryphiella retards and occupies a sector in Fusarium growth in
1-5%;
F. Dendryphiella further retards with addition of 2-0% ;
(r. Dendryphiella occupies a still smaller part in 4*0% ;
H. Dendryphiella occupies a very small area in 6 .0 % and 8-0 % ;
/. Entire growth is that of Fusarhtm and there is no growth of Dendry-
phiella with addition of 10-0 and 12-0% KNO 8 (Diagrammatic).
It is seen from Fig. 7 that Fusarium starts as a small sector in 1 per cent
potassium nitrate (7 A), increases in size with the increase in concentration
(7B-7C), attaining the same size as Dendryphiella becomes reduced to a very
small sector in 4 per cent. (7G), still smaller in 6 per cent, and 8 per cent.
(7H) till there is no growth of Dendryphiella at 1.0 per cent, and 12 per cent,
(71, and Plate I, Figs. 16-20).
Competition in Fungi /
13
Acid series. Malic acid was added to standard synthetic medium in
strengths 0-025, 0-05, 0-1, 0-2, 0-3, 0-4, 0-6, 0-75, 1-0, 1-5, and 2-0 per
cent, and plated. Fusarium and Dendryphiella were inoculated adjacent
and 2 cm. apart. Controls were kept.
The results are shown graphically in Text-Figs. 8 and 9 where the dia-
meters of six days growths are plotted against acid concentrations for
adjacent and 2 cm.-apart- cultures respectively.
It will be seen from Text-Fig. 8 that in adjacent culture Dendryphiella
shows a marked increase in growth in 1 per cent, to 1 - 75 per cent, over
Fusarium although iri lower concentrations, it is either being dominated by
0050102 04 06075 10
Malic acid per cent.
Fusarium
15
20
(Control)
(Adjacent)
Dendryphiella (Control)
mmmttmamommmmimmz ^ (Adjacent)
TEXT-FIG. 8. Graphs showing the two fungi Fusaruim sp. and Dendryphiella sp.
in associated culture and in. control in Malic acid series. Inocula
placed adjacently.
Fusarium or has a similar growth rate. In 2 cm,.-apart-culture (Text Fig. 9),
throughout the series the two fungi run closely parallel to each other and
to the controls.
Far interesting is the fact that the controls of both the strains, Dendry-
phiella and Fusarium, stop growth at about 1-5 per cent. acid. But when
they are inoculated 2 cm,, apart and adjacent, the point of total inhibition
is raised from 1-25 to 1-75 per cent. The result demonstrates a marked
increase in the tolerance of acid due to the association of the strains.
14
T. S. Sadasivan
60
-i
| 50
'^40]
O
1 30'
fe 20'
1 10
00-050102 04 0-6 075 10
Malic acid per cent.
1-5
2-0
Fusarium
(Control)
,, (2 cm. apart)
Dendryphie lla ( Control )
,, (2 cm. apart)
TEXT- FIG. 9. Graphs showing the behaviour of Fufiaritnn sp. and 'Dendruph'lcUa sp.
in associated culture and in control in Malic acid scries. Inocula
placed 2 cm. apart.
Alkali series. The effect of alkali on the competing strains was next
tried by using sodium hydroxide and sodium carbonate.
Sodium hydroxide. This was added to standard synthetic medium in
the following strengths and plated 0-01, 0-025, 0-05, 0-075, 0-1, 0-25 and
0-5 per cent. Inoculations were made 2 cm. apart and adjacently. Controls
were kept.
The results are shown graphically in Text-Figs. 10 and 11 where dia-
meters of six days growth are plotted against concentrations of sodium
hydroxide for 2 cm. apart and adjacent cultures.
The behaviour of the strains in the series was very interesting. It will
be seen from Text-Fig. 10 that in cultures 2 cm. apart Dendryphiella dominat-
ed in percentages 0-01, 0-025, 0-075 and 0-1, but all the time both growing
slower than in controls. At 0-25 per cent, the Fusarium so far slower than
Dendryphiella becomes faster both in 2 cm.-apart-culture and in the
control, due to the sudden drop in the growth rate of the latter. At - 5 per
cent, when the controls have ceased to grow both the strains still continue with
Fusarium somewhat dominating.
In adjacent culture the reactions of the fungi are very similar to that
for cultures 2 cm. apart as already explained. In the latter case, however,
Competition in Fungi /
15
0-010025 0-05 0-075 01 025
Sodium hydroxide per cent.
Fusarium
(Control)
(2 cm. apart)
**^** > " - **""~"* Dendryphiella (Control)
MHiiMoniMM (2 cm. apart)
EXT-PlG. 10. Graphs showing the behaviour of Fusarium sp. and DendrypJnella sp.
in associated cultures and in control in NaOBE series. Inocula placed
2 cm. apart.
"001 0025 0-05 0-075 01 0-25
Sodium hydroxide per cent.
Fusarium (Control )
(Adjacent)
o Dendryphiella (Control)
iHiOM ,, (Adjacent)
. 11. Graphs showing the behaviour of Fusarium sp. and Dendryphiella sp.
in associated culture and in control in NaOH series. Inocula placed
adjacently.
16
S* Sadaslvan
Fusariwm shows peculiar fluctuations,^., acceleration at one strength and
retardation at the next.
It will be seen from Text-Fig. 12 (A--G) that in a medium containing
0-01 per cent, sodium hydroxide, Fusarium is confined to a small sector-like
growth in a major growth of Dendryphiella [Text-Fig. 1.2 (A)]. In 0-25 per
TKXT-FIG. 12 A-G. Diagrammatic representation of the relative area occupied by
Fusariam sp. and Dendrypkiella sp. in various concentrations of
NaOH. Inocula adjacent.
Shaded area represents Fusarium.
Unshaded area represents Dendryphiella.
Dots indicate sporulation of Dendryphiella.
A. Fusarium occupies a small sector in. a major growth of Dendryphiella
with the addition 0-01% NaOH to the standard medium;
J5. Fusarium accelerates with the addition of 0-025% ;
C. Fusarium retards again with the addition of 0-05% ;
D. Fusarium accelerates with 0-075% ;
E. Fusarium again retards with addition of 0-1 %;
F. Fusarium growth is more than that of Dendrypliiella with 0*25% ;
G. Fit-sariuM dominates the growth and Dendrypkiella occupies a sector
at 5 % NaOH (Diagrammatic) .
cent, sodium hydroxide the two growths occupy equal area. In 0-05 and
1-0 per cent. Fusarium is restricted to a small sector, while in concentra-
tion 0-075 per cent. Fusarium outgrows Dendryphiella and occupies half
the area of the entire growth and at 0-5 per cent, dominates Dendryphiella
completely restricting it to a very small sector.
A comparison of Text-Figs. 11 and 12 will show that in this case there
is no correspondence between the relative growth of the two fungi and the
Competition in Fungi /
17
area occupied by them in adjacent cultures. It is seen from Text-Fig. 11
that in all concentrations up to 0-1 per cent. Dendryphiella greatly dominates
the Fusarium, having almost twice the growth rate, in spite of that Fusamm
at 0-025 and 0-075 per cent, of NaOH occupies area equal to that occupied
by Dendryphiella ; these are the concentrations when Fusarium shows
accelerated growth.
It is interesting to note that in these series as in the others already
mentioned, the associated fungi are capable of growth in higher alkali
in adjacent culture and in culture 2 cm. apart while they have ceased to grow
in the control plates.
Sodium carbonate. This was tried in the following percentages with
standard synthetic medium 0-01, 0-025, 0-075, 0-1, 0-2, 0-4 and 0-75.
The two strains Fusarium and Dendryphiella were inoculated 2 cm,, apart
and adjacent. The results are shown graphically in Text-Figs. 13 and 14
where diameters for six days growth of the competing strains and of the
controls are plotted against various concentrations of sodium carbonate for
2 cm. apart and adjacent cultures respectively.
. 50
3
^ 0-010-025 007501 02 04
Sodium carbonate per cent.
Fusarium (Control)
,, (2 cm. apart)
o Dendryphiella (Control)
HHOBMMOHBWHBM ,, (2 cm. apart)
13. Graphs showing the behaviour oC Fusarium sp. and Dendrijphic.Ua sp.
in associated culture and in control in NaoCO^ series with inocula
placed 2 cm. apai't.
18
T. S. Sadasivan
0-010-025 0-07501 0-2 04
Sodium carbonate per cent.
Fus a rium ( Control )
(Adjacent)
----- c --- Dendryphiella (Control)
wmamvmsmQmvmmmi ,, (Adjacent)
TEXT-FIG. 1.4. Graphs showing the behaviour of Fusarium sp. and Dendryphiella sp.
in associated culture and in control in Na 2 C0 3 series. Inocula placed
adjacently.
It will be seen from Text-Fig. 13 that in the concentrations of sodium
carbonate employed Fusarium and Dendryphiella growing 2 cm. apart do not
show significant difference from the controls upto 0-4 per cent. At 0-75
per cent, however, both the fungi continue to grow while there are no growths
in the controls.
Particularly interesting results are seen in adjacent culture (Text-Fig. 14).
In media containing 0-01 per cent, and 0-025 per cent, sodium carbonate
Fusarium has the same growth rate as in the control ; at 0-075 per cent.
there is a sudden drop. At 0-1 per cent, of sodium carbonate the growth
rate shows a marked increase and at 2 per cent.it is again virtually the
same as that of the control. At 0-4 per cent, the competing Fusarium has
a slightly higher growth rate and beyond that at 0-75 per cent, where the
growth of the controls has completely ceased, it continues to grow.
A comparison of Dendryphiella in control and in adjacent culture with
Fusarium is still more interesting. In adjacent culture Dendryphiella has a
much slower growth rate. A slight addition of sodium carbonate 0-01
per cent, brings down the rate considerably and at 0-025 the growth of the
fungus is inhibited. At 075 per cent. Dendryphiella shows definite accele-
ration. Dendryphiella is finally inhibited at a concentration beyond 1
Competition in Fungi /
19
per cent, sodium carbonate although growth in the control continues till
75 per cent.
The results are of great importance. They show a definite inhibitory
influence exerted by Fusarium on Dendryphiella. In sodium carbonate
concentration where Fusarium is active, Dendryphiella is considerably
inhibited and with the decrease in the activity of Fusarium, Dendryphiella
shows a corresponding increase. Beyond 0-1 per cent, however, Fusarium
completely checks the growth of the fungus.
A B
C.
D.
C D
TEXT- Fro. 15 ,4 -D.~ Diagrammatic representation of the relative area occupied by
Fusariuni and Dendryphiella in various concentrations of NaoCO 3 .
Inocula adjacent.
Shaded area represents Fusar'nun.
Unshaded area represents Dendryp/iiell.a.
Dots indicate sporulation of Dendryphiella.
A. Dendryphiella, forms a sector in an entire growth of Pufiarium at 0-01 % ;
#. Very little growth of Dendryphiella. seen with the addition of 0-025% ;
Dendryphiella accelerates with the addition of 0-075% ;
There is no growth of Dendri/phiella, the dominant strain being Fut*urium
in percentages 0-1. to 0*75 of NaoOO.v
Text-Fig. 15 (A-D) will show diagramtnatically the inhibitory influence
exerted by Fusarium on Dendryphiella in sodium carbonate series. At
0-01 per cent, and 0-075 per cent. Dendryphiella forms a small sector, at
0-025 per cent, it is restricted to a very small area near the centre, at 0-075
per cent, it occupies a much bigger area and is completely inhibited at 0-1
per cent.
Discussion
A certain amount of work has been done on the growth of fungi in mixed
culture. Harder (191.1), Cook (1924), Porter (1924), Machacek (1928),
20 T 8 S- Sadasivan
Vanin and Vladimirsky (1982), Endo vSigeru (1932, 1933) and others have
worked on different aspects of the problem.
Harder has given a detailed account of changes in the rate of growth,
in hyphal structures, of the formation of colouring matter, phenomenon of
aversion, etc., in mixed cultures of a number of strains. He has also demon-
strated that one mycelium does not kill another. Vanin and Vladimirsky
(1932) have shown that in the mixed cultures of Memlius lacrymans and
Coniophom cerebella, at an early stage, the mycelium of each grows normally
without interfering with each other. I v ater the mycelium of Merulim
lacrymans outgrows that of Coniophora cerebella causing a considerable retar-
dation in the growth of the latter. Endo Sigem (1932) has shown that the
presence of micro-organisms is one of the factors controlling the sclerotial
formation in Sclerotium oryza-sativcz. The inhibition and retarding action
of one fungus on another in mixed culture have been worked out in detail
by Porter (1924). Cook (1924) has made an interesting study on the succes-
sion of fungi in artificial culture and Machacek (1928) on the association of
phytopathogens.
On the applied side of the problem Endo Sigeru (1933) has found that
the antagonistic action of several organisms prevented appearance of a
disease due to Hypochnus Sasaki-i Shirai. Vasudeva (1930) has demon-
strated that the parasitic activity of a fungus is greatly retarded by the
presence of a non-parasitic fungus and Asthana (1936) has found some
interesting results on the effect of various fungi on the parasitic vigour and
other characteristics of Botrytis cinerea.
The problem is approached from altogether a different view-point in this
investigation. An attempt has been made here to find out how two fungi
will behave in different growth conditions in associated cultures when they
are grown contiguously or a little distance apart and how will they influence
each other's growth.
It is evident from the various experimental data given that fungi are
influenced not only by the change in the composition of the medium but also
by the association of one strain with the other. A profound alteration in
growth reactions occur when these are grown adjacently and 2 cm. apart in
altered condition of the medium.
Where a fast-growing strain was paired with a relatively slow-growing
one, it was usually the faster strain that outgrew and enveloped the slower
one, but reverse condition was also found where the slow-growing Phoma
strain in mixed and adjacent cultures had completely dominated the faster
Helminthosporium restricting it to small sectors.
Competition in Fungi / 21
It is in the detailed investigation of the behaviour of Fusarium and
Dendryphiella, however, that more interesting results were obtained.
The domination of one strain over another as seen from the extent of area
occupied by the competing strains of Fusarium and Dendryphiella in a mixed
growth arising from inoculation either put adjacently or 2 cm. apart, in
various concentrations of the chemicals employed is very instructive.
Fusarium is only slightly faster than Dendryphiella in standard medium.
Nevertheless in almost all the series so far employed, glucose, potassium
nitrate, sodium carbonate and sodium hydroxide as the concentration was
increased Fusarium gradually dominated Dendryphiella in adjacent culture.
In the highest concentration of potassium nitrate and sodium carbonate
employed the growth was all Fusarium to the total inhibition of Dendry-
phiella.
The details of reaction in different series, however, differed to a large
extent. In glucose, for example, there were three inhibition points at
relatively low concentrations of 0- 1, 0-2 and 0-4 per cent, where Dendry-
phiella was totally suppressed. But at 0-2 per cent., however, there was a
certain amount of growth of Dendryphiella, which probably is due to the
associated Fusarium being slightly less active at that concentration.
In subsequent concentrations, Dendryphiella occupied a relatively small
sectorial area throughout and was never again completely suppressed even
in the highest concentration employed. In sodium carbonate series too the
major growth of Fusarium contained a sector of Dendriphiella whose area
fluctuated in higher concentrations. Dendryphiella was completely inhibited
in concentrations 0-15-0 -75 per cent.
In potassium nitrate series it is the Fusarium that appeared as a small
sector, which gradually became larger as the concentration of potassium
nitrate increased finally suppressing the entire growth of Dendryphiella.
The result in sodium hydroxide series, however, was more complicated.
Fusarium appeared as a small sector at the lowest concentration. But in-
stead of a steady increase in the area with the increase in the concentration of
sodium hydroxide there was alternate rise and fall. A semi-circular area in
one concentration was followed by a small sector in the next till at - 5 per
cent. Fusarium dominated the growth and restricted Dendryphiella to only
a small sector.
A comparison of the results indicates that on the whole, the reaction of
one strain against another is more pronounced in adjacent than in 2 cm,-
apart-culture.
-72 T, So Sadasivan
The influence of Fusarium and Dendryphiella upon each other Is well mani-
fested In the difference between the growth rates of the strains in associated
culture on one hand and their controls on the other. The growth rate of a
fungus depends upon the chemicals constituting the media and their concen-
trations, but the reaction of the competing strains to the media is also pro-
foundly modified by the association of one fungus with the other. The modi-
fication in growth rate results in either retardation 01 acceleration.
The acceleration of growth rates of the competing fungi in associated
cultures was primarily observed in higher concentration of chemicals. For
example, both Fusarium and Dendryphiella showed acceleration over their
controls beyond 0-4 per cent, sodium hydroxide in adjacent culture, and
in the same concentration of sodium carbonate in cultures 2 cm. apart. In
malic acid series It was found only in concentration as high as 0-75 per cent.
and over. Comparatively lower concentration may also bring out the same
result. Thus Fusarium showed faster growth rate in adjacent culture beyond
0-1 per cent, sodium carbonate.
The acceleration of growth raises the final inhibition point of the strains.
The strains in these cases maintained their growths in concentrations where
the controls had ceased to grow. It was found in all the series except that
of glucose. For example, in malic acid series while the controls failed to
grow at about 1-25 per cent, both Fusarium and Dendryphiella continued
to grow at 1-5 per cent, and had respectively diameters of 3-0 mm. and 7-0
mm. in adjacent culture and 2 -5 mm. and 5-0 mm. in 2 cm. -apart- culture.
Similarly in sodium hydroxide series at 0-5 per cent, while control had
failed to grow altogether, Fusarium and Dendryphiella had diameters 3 mm.
and 2-0 mm. respectively in cultures 2 cm. apart, and 5-0 mm. and 3-0 mm.
respectively in adjacent culture. In sodium carbonate again at 0-75 per
cent, in 2 cm.-apart-culture Fusarium and Dendryphiella had diameters of
7-0 mm. and 4-0 mm. respectively while controls showed no growth. At
the same concentration, in adjacent culture diameter of Fusarium proved
to be 9^0 mm. against no growth of control, Dendryphiella had stopped growth
at 0-15 per cent, sodium carbonate.
The retardation of growth rates of the competing fungi was usually observed.
In the lower concentrations and sometimes throughout the series.
In malic acid there was retardation for both the strains in concentrations
between 0-01-0.75 per cent. In sodium hydroxide, the strains growing
2 cm. apart had a slightly slower growth rate in lower concentrations in
adjacent culture only Fusarium was strongly retarded below 0-4 per cent
In sodium carbonate in 2 cm.-apart-culture, Fusarium and Dendryphiella
Competition in Fungi / 23
were both retarded below 4 per cent, while the limit for the same in adjacent
culture was 0- 1 per cent. The degree of retardation, however, varied. This
was evident in all concentrations of potassium nitrate. In glucose series
too, in adjacent culture, there was a slight retardation throughout. Dendry-
phiella on the other hand, show r ed slight fall in growth rate in 2 cni.-apart-
culture which became very marked in adjacent culture with inhibition points
at 0-1, 0-2 and 0-4 per cent, concentrations.
The retardation culminating in the complete suppression of Dendry-
phiella by Fusarium occur at 0-15 per cent, sodium carbonate while the
diameter of the control is 25-0 mm. There is a similar and equally striking
inhibition of Dendryphiella at 10 per cent, potassium nitrate when the control
fungus shows a diameter of 20-0 mm.
It is possible that the media in the neighbourhood of the growths become
modified, either to the advantage or to the disadvantage of the strains by the
staling substances produced by them and the observed differences in the
growth rates are due to the differential reaction of the strains to thes e
modified media. Or as it has been suggested by Machacek (1928) the dif-
ferences may be due to unequal assimilation of available food from the media
causing the starvation of one organism.
That these factors play an important role in the growth reactions of
fungi is well known, but it is not improbable that actual contact (Das Gupta,
1934) or the mere presence of hyphse may, to a certain extent, be responsible
for some of the observed results.
At particular strengths of sodium carbonate (0-2,0-4 and 75 per cent.)
in adjacent cultures the retarding influence of Fusarium was so great that
Dendryphiella, in spite of having a favourable growth rate in the controls, is
totally suppressed by Fusarium. In such a condition where there was no
growth of Dendryphiella the accelerated growth of Fusarium over its control
could not possibly be due to diffusion of staling products from Dendryphiella.
It may be concluded therefore that at least in certain media even the mere
presence of an inoculum of Dendryphiella placed adjacently to that of
Fusarium has a striking effect on the acceleration of growth of the latter.
It is apparent from, the study of the results obtained from different
series of experiments that growth reaction of associated non-parasitic fungi
as represented by two strains under consideration, are extremely variable.
They show various degrees of tolerance and inhibition. Machacek (1928)
distinguished several types of association on the basis of mutual reaction,
viz (1) one organism causes complete inhibition of its associates ; (2) asso-
ciated organisms are mutually tolerant on each other and so on. Results here
>4 T. S 8 Sadasivan
show however, that no hard and fast rule can be laid down ; in fact, inhibi-
tion 'tolerance, etc., as shown by the fungi are but functions of the chemical
composition of the growth medium which are profoundly modified by the
Interaction of the organisms upon each other.
Summary
In order to ascertain the nature of growth reactions in associated cultures
of non-parasitic fungi six strains were used in five different combinations.
Fusarium sp. and Dendryphiella sp. ; Fusarium sp. and Gibberella sp. ;
Phoma sp. and Helminthosporium sp. ; Phoma sp. and Monilia sp. ; Phoma
sp. and Dendryphiella sp.
The paired strains were inoculated mixed, adjacent and 2 cm. apart in
media differing in composition of acid, alakli and nutritive chemicals and
their reactions were studied. Detailed work was done only with one pair
Fusarium sp. and Dendryphiella sp. Some of the more interesting results
are given below :
In the majority of cases usually the faster strain of a pair enveloped
the slower one and the latter appeared as sectors. But Phoma, a slow growing
strain, was found to dominate the faster strain of Helminthosporium.
Dendryphiella and Fusarium showed differential reaction to the various
media employed. There was also a pronounced influence of each fungus on
the other as judged from the relative area occupied by these in mixed growth
as well as from their growth rate.
The domination of one strain over another as indicated by the extent of
area occupied by the competing strains of Dendryphiella and Fusarium m
adjacent and 2 cm.-apart-culture varied with the composition of media
employed. Generally in lower concentrations Fusarium appeared as major
growth with Dendryphiella as sector. In some cases Fusarium appeared as
a small sector in the major growth of Dendryphiella. Ultimately, however
in all the series in relatively higher concentrations, it was always the Fusarium
that dominated the growth ; sometime to the complete inhibition of Dendrv-
phidla.
The accelerating influence of Fusarium and Dendryphiella upon each
other was manifest at comparatively higher concentrations and rarely at
lower, where Fusarium and Dendryphiella both accelerated over their controls
both in adjacent and 2 crn.-apart-culture. The immediate, important
result of the acceleration was the raising of the final inhibition point of the
strains in almost all the series. Here the individual strains in associated
Competition in Fungi /
25
cultures continued to grow in media wliere controls had ceased to grow
altogether.
The retarding influence of Fusarium and Dendryphiella upon each other
was evident in the lower concentrations. It was more pronounced in
adjacent culture. Dendryphiella was nore influenced by Fusarium than
Fusarium by the former. In glucose, in adjacent culture," Fusarium totally
inhibited the growth of Dendryphiella at three low concentrations (0-1, 0-2
and 0-4 per cent.). The most interesting was the complete inhibition of
Dendryphiella by Fusarium at 0-15 per cent, sodium carbonate and
10-0 per cent, potassium nitrate in adjacent cultures although in control
Dendryphiella showed diameter of 25-0 mm. and 20-0 mm. respectively.
There is a suggestion that the presence of a mere inoculum of one fungus
may also, in certain cases, modify the growth reaction of another.
I wish to express my grateful thanks to Dr. S. N. Das Gupta for suggest-
ing the problem and offering me ready guidance and criticism throughout.
Asthana, B. P.
Cook, M. T.
Das Gupta, S. N.
Endo Sigeru
Harder, R.
Machacek, J. E.
Porter, 0. L.
B2
LITERATURE CITED
" Antagonism in fungi as a me as '.ire of control in
1 Red-leg ' disease of Lettuce," Ann. Bot., 1936, 4,
201-07.
" Succession of fungi on culture media," Amer.
Journ. of Botany, 1924, 11, 91-99.
11 Studies in the genera Cytosporina^ Phomopsis and
Dlaporthe. VI. On the conversion of one strain of
Diaporthe perniciosa into another," Phil. Trans.
Roy. Soc. Loud., 1934, Ser. B, 223, Xo. 497. 121-61.
" Studies on the antagonism of micro-organisms. II.
Growth of Hypochnus Sasdkii Skirai, as influenced
by the antagonistic action of other organisms," Bull.
Miyasald College of Agri. and Forestry, 1932, 4,
133-S5.
" Studies on the antagonism of micro-organisms. IV.
Growth and pathogenicity of Sclerotiiun oryzm
sativce Sawada in the presence of other organisms,"
ibid., 1933, 5, 51-75.
" L r eber der Verhalten von Basidiomyceten und Asco-
myceten in Mischkulturen," Natunc. Zeitschr. /.
Forestu. Landw., 1911, Nos. 3-4.
" Studies on the association of certain phytopathogens,"
MacDonald College, McGill Univ. Tech. Bull., 1928,
No. 7.
" Concerning the characters of certain fungi as exhi-
bited by their growth in the presence of other fungi,"
Amer. Journ. of Bot., 1924, 11, 168-88.
F
26 T. S* Sadasivan
Vanin and Vlaclimirsky . . " On the biology of the fungus Mcrulius lacrijmans
and Coniophora cerebella," Bulletin of the Leningrad
Institute for Controlling Farm and Forest Pesls, 1932,
No. 3, 57-74.
Vasuveda R. Sahai " Studies in the physiology of parasitism. "XII. On the
effect of one organism in reducing the parasitic
activity of another," Ann. Bot., 19130, 44, 557 -ft 1.
DESCRIPTION OF PLATE FIGURES
Fros. KV-20. Photographs showing the behaviour of Fuftariutu sp. and Dendri/phiclla
sp. when grown adjacently in varioris cencentrations of KNO 3 . The
darker colony represents Dendrypliiella. X 3..
FIG. 16. -25% KNO 3 . Dendrypliiella occupies major area and Fusarium a small
sector.
FIG. 17. 0.75% KNO 3 . Dendryphiclla and Fusarium occupying equal area.
FIG. 18. 1% KNO 3 . Fusarium occupying major area.
FIG. 19. 2% KN0 3 . Fusarium. occupying still greater area and Dendryphiella
forming a sector.
FIG. 20. 6% KNO 3 . Fusarium completely dominating. Dendryphiella restricted
to a small growth near the inoculum (dark).
T. S. Sadasivan
Proc. hid. Acad. ScL, B, vol. X, PL I
19
EFFECTIVENESS OF CHEMICAL FERTILISERS ON
THE GROWTH AND WATER REQUIREMENT
OF WHEAT
BY B. N. SINGH AND J. R. SINGH
(From the Institute of Agricultural Research, Benares Hindu University)
Received November 22, 1938
Introduction
IN the previous contributions from this Experiment Station on the problem
of water requirement of a large number of crop plants 10 ' 11 besides the
specific and varietal differences and the critical periods, the influence of yield
and duration of life-cycle of the plants and also of the soil-moisture on their
water requirement were studied in detail. In this communication are pre-
sented the results of experiments on the effectiveness of certain chemical
fertilisers on the growth and water requirement of wheat.
With the increasing application of fertilisers in the successful production
of crops, the problem of water requirement is becoming more and more
complicated, this being particularly true when the varied after-effects of
dressings are studied in conjunction with the water relations of the plants.
Another problem of considerable importance is the requisite supply of water,
both as regards the amount and the opportune moments, when the fertilisers
are added so that the maximum growth and yield of the crop may be ob-
tained a problem on which no detailed information is yet available. In
an early investigation on the problem of water requirement Lawes 4 found
that the water requirement of certain plants, specially peas, clover and wheat
was diminished whereas that 'of others, e.g., barley and beans was considerably
increased in response to manuring with chemical fertilisers. Sachs 9 ob-
served a similar depression in transpiration rate when potassium nitrate
and ammonium, sulphate were added to soils. A distinct economising effect
on the water requirement of white mustard in response to the addition of
potash specially in soils containing smaller amounts of water was observed
by Maercker 7 while Hartwell and his co-workers 3 recorded an increase
in transpiration of wheat seedlings due to the addition of potassium chloride
(cf. Gardner 2 ). Reed, 8 on the other hand, noted that transpiration was
increased on the addition of lime and sodium phosphate whereas the reverse
27
28 B, N. Singh and J B R. Singh
elfect was observed in the case of potassium sulphate and sodium nitrate.
1 vim tlier 5 - <J working under Indian conditions, compared the water require-
ment of a number of crop plants supplied with and without manures
and found a decline of the same in the manured ones. His figures for all
the crops in general, are, however, much higher than those obtained at this
Station 10 apparently due to the unsuitability of his experimental technique
of estimating transpiration.
It is tints apparent that this problem of water requirement of crops in
relation to dressings with different artificial fertilisers awaits further detailed
and critical study since variations in the effect of the different fertilisers on
one plant and also of a single fertiliser on different plants are of common
occurrence. Investigations were therefore conducted using the improved
methods of experimentation to elucidate the effects of certain important
fertilisers, e.g., ammonium sulphate, ammonium phosphate, superphosphate,
potassium sulphate and potash mixture on the growth and water requirement
of wheat under the semiarid tropical conditions of this locality and are
reported in the present paper.
Procedure
Pure strain seeds of wheat var. Pusa 1.2 of as nearly uniform size, weight
atid relative density were grown in porous earthenware pots, 9 x 12 inches
iu size, to ensure proper soil aeration. Air-dry and well-sieved farm soil
was thoroughly mixed with each fertiliser separately at the rate of 1 -3 gm.
per 10 kgm. of soil. The weight of one acre foot of the farm soil was then
determined and the quantity of fertiliser added calculated in terms of Ibs.
of N a , PaO 5 or K 2 O added per acre. The amounts of nutrients added as
Ibs. per acre is shown in Table I. Before packing the pots, the mixed soil
was damped a bit with a little quantity of water. The pots containing these
soil-manure mixtures were then grouped into several sets according to the
nature of the manure added.
To ensure homogeneous growth under as natural conditions as possible
and also to obtain the best conditions of nutrition the potted plants were
buried upto the height of the pots in soil-fertiliser mixture of the same
proportion as that in the particular set of pots in shallow trenches lined
with, bricks side by side with the field plants.
At successive intervals six of the experimental pots selected at random
were taken out and sealed air-tight in metal containers with wax-petrolatum
mixture after the manner indicated in an earlier publication from this
Research Station. 10 Every morning the loss incurred through transpiration
Chemical Fertilisers on Growth & Water Requirement of Wheat 29
Influence of fertilisers upon the growth of wheat
Plant charactreistics
Control
Ammonium
phosphate
(NssPsa)*
Ammonium
sulphate
(Nss)
S o "^
al
3 "Hi 2
o g
P., <a~
P
|t
T
fS
^ *
* < T-l
-
-~^
^ *
.2 rn
Stf
Significant
difference
P = 0-05
Height per plant, cm.
65-83
108-52
108-82
80-54
82-04
89-21
8.52
Tiller number per plant
3
8
7
5
6
7
1.64
Dry weight per plant, gm.
14-34
23-71
22-83
16-99
18-18
19-36
L92
Yield of grain per plant, gm.
4-23
8-53
7-73
5-44
6-04
6.74
2-21
Yield of straw per plant, gm.
10-11
15-18
15-10
11-55
12.14
12-62
2-56
Length of head, cm.
5-43
12-24
11-79
6-01
9-27
10-17
2-60
Weight of 100 grains, gm.
4.43
6-98
6-61
5-74
5.41
6-03
1-42
Volume weight of grains per c.c.,
gm .
0-72
0-93
0.91
0-83
0-85
0-86
Leaf area per plant, sq. cm.
14-96
28-25
25-12
17-53
19-78
20-96
Time of blossoming, days after
sowing
58
63
63
60
60
58
Duration of floral development,
days
20
23
23
21
20
23
Time of fruiting, days after sowing
78
86
86
81
80
81
Time of harvest, days after sowing
103
111
111
106
105
106
* This indicates that nitrogen and P 2 5 are added at the rate of 33 Ibs. per acre. Similar
figures for the other treatments indicate the same in respect to the active constituents.
was recorded and the soil-moisture was brought to the original level by the
addition of the equivalent quantity of water. The moisture content of the
soil was determined at intervals of seven days. This was done by taking
representative samples of known weight from different pots and drying in
steam oven at 100 C. The loss incurred in drying was calculated in terms
of percentage loss on dry weight basis. It would appear from the values
of moisture content thus obtained (Table II) that the plants were growing
under 22% soil moisture content since the very beginning of the age-cycle,
thus ensuring optimum moisture conditions for studies on the water
requirement of cereals. 11
A full record of meteorological observations was maintained at this
Research Station. The evaporating power of the atmosphere was also
B. N. Singh and J. R. Singh
TABLE II
Moisture content of soil at successive stages of age-cycle
Moisture
Evapora-
Fresh
Dry
Moisture
Moisture -
content on
tion per
^ ,. ,. r .,T
(.
weight of
weight of
content per
holding
percentage
hour in
sJ.
~ " Date
: composite
! sample
composite
sample
100 gm.
of soil
capacity
of soil
water-holding
capacity
gm. per
1000
j
basis
sq. cm.
i am.
gm.
l-j
.. ; 15-11-1935
i 12.12
9-64
22-21
58-13
38-27
7-12
.. ; 22-11-1935
1 12-02
9-38
22-00
57-92
37-99
25-68
13-1S
..: 29-11-1935
i 12. IS
9-56
21-54
58-90
37.13
25-65
19-24
..; 0-12-1935
i 12-19
9-59
21-32
57-54
37-05
25-62
25-30
.J 13-12-1935
; 11-81
9-16
22-45
58-77
38-20
31-36
. .! 20-12-1935
12.35
9-62
22-07
58-29
37-87
24-52
37-42
..: 27-12-1935
12-01
9-35
22-13
58-08
38-11
43-48
. j 3-1-1936
! 12-32
9-63
21-97
57-03
38-00
49-54
. .. 10-1-1936
1 12-22
9-49
22-37
58-55
38-21
..
55-60
..I 17-1-1936
j 11-85
9-23
22-12
58-03
38-12
24-50
61-66
.. 24-1-1936
12-20
9-48
22-43
57-63
38-92
67-72
I
.J 31-1-1936
I 1L93
9-33
21.78
57.50
37.89
25-63
73-7S
7-2-1936
1 11-99
9-41
2L53
57-09
37-71
26-66
79-84
.. 14-2 -1 93C
\ 11-89
9-23
22-35
58-47
38.22
26-69
85-90
.. 21-2-1936
1 12.11
9-49
22-25
58-09
38-31
27-12
91-96
.. 28-2-1 93G
i 12-15
9-46
22-20
58-39
38-02
29-32
recorded from a shallow water pan and used for estimating how far transpira-
tion at successive stages of growth was related to the evaporating power of
the atmosphere. The seeds were sown on 15th November 1935 and the
crop harvested between 6th March to 14th March 1936.
As during the previous studies 10 - 11 the evaporation of water from
soil under similar conditions of atmospheric variables was also measured
during the period of experimentation. The total amount of water required
for a successful wheat crop in terms of gallons and acre inches was finally
calculated after taking into account the water transpired by the plant and
that evaporated from the soil during the life-cycle of the corp.
Chemical Fertilisers on Growth & Water Requirement of Wheat 3 1
Opportunities were also availed to study the response of the crop to
the addition of the different fertilisers. Thus, the variations in the morpho-
logical characteristics of the differently treated plants and also their dry
matter accumulation were noted at successive stages of their life-cycle and
were utilised to evaluate the influence of the manurial treatments both on
the growth and water requirement of the crop. The final yield of the plants
were also similarly noted. Statistical treatments consisted in analysing the
same by the Fisher's method of analysis of variance.
Experimental Results
Growth Characteristics
In consequence of the addition of the fertilisers distinct beneficial effects
on the growth of the plants are noted, prominent variations from, treatment
to treatment in the morphological characteristics appearing soon after the
establishment of the seedlings. Statistical analysis of the data of the various
characteristics collected at the final stage (Table I) indicate that in general
the treatments have given significantly better results than the control
series.
In all the treatments, the height of the plants increases significantly,
the best result being shown when N 33 P33* (ammonium phosphate) is added.
Nitrogen alone (N 33 ) given as ammonium sulphate gives better results than
P 66 (superphosphate), K 78 . 7 (potassium sulphate) and N 16 . 6 K 18 P 16 . 5 (potash
mixture) although this last treatment is significantly better than the other
two. Similarly, the tiller number is also significantly increased in response
to the addition of the fertilisers. N 33 P 33 (ammonium phosphate) again
shows the best result which is also better than K and P added alone. N 33
(ammonium sulphate) and N 16 . 5 K 18 P 16 . 5 (potash mixture) in this case show
similar results and are better than K applied alone.
The period when the blossoms appear for the first time is slightly delayed
in reponse to the different treatments as compared to the control plants
except when N 16 . 5 K 18 P 16 . 5 (potash mixture) is^applied. This delay is only
two days for K 78 . 7 (potassium sulphate) and P 66 (superphosphate) and
five days for N 33 (ammonium sulphate) and N 33 P 33 (ammonium phosphate)
series. Similarly, the manured plants require 1-3 days more for their floral
development except in the case of P 66 (superphosphate) applied alone.
The time when the fruits appear for the first time is also delayed in the
manured plants, a maximum delay of 10 days being noticed in the cases of
* This indicates that nitrogen and P 2 5 are added at the rate of 33 Ibs. per acre.
Similar figures have been used in the paper for the other treatments also.
! -u/ iUiL dnu -\ 33 i
E. X. Singh and J. R. Singh
. X 38 Ps8 (ammonium phosphate). Finally; the
cl':v f,r harvest 2-8 days later than the control plants,
' * "' ;L v*:r -cLssary for both the ammonium treated series.
- -^ -^ t^ tic addition of the fertilisers except when K is applied
^"VV .J^ nl srhhate, tLe length of the head is also increased signifi-
^ ._- ' :: 7\ aj;i!:-i,:niri pnosphate) shows the best result and is better
* ^ K / 1 T ul^ lied alone. The absolute weight of the grains is also signifi-
^ 7*V. -Ir^ci Wept again in the cases when K and P are applied alone,
*1 r: ' ^r:^ ::r ^nitlc^t differences between the N 33 (ammonium sulphate),
X-J. Jr^urhim pnosph^te) and N 16 . 5 K 18 P 16 . 5 (potash mixture). The
v-l::r:tr .vrlzLt of grains also bhows similarly better results with the N and
XP series.
The yield of the grains and straw per plant also varies considerably with
the different fertiliser treatments. K (potassium sulphate) and P (super-
phosphate) applied alone do not however show any significant increase in
the yield of the grain while these two treatments along with NKP (potash
mixture I also indicate the same when the straw yield is considered. N and
X P /ammonium sulphate and phosphate) give best yield of grain ; while the
former is better than K (potassium sulphate) the latter one shows higher
yield than both K (potassium sulphate) and P (superphosphate). The NKP.
combination (potash mixture), however, missed the level of significance in.
the case of straw yield by a very narrow margin.
Although the growth of the crop is best in N (ammonium sulphate) and
XP (ammonium phosphate) series in all respects, the chief disadvantage of
these series lies in the fact that the plants are often prone to lodging
while this is minimised in the K 78 . 7 (potassium sulphate) series and
characteristically absent in the other two series.
The dry weight of plants when plotted against age gives the usual
sigiaoid type of curve (Fig. 1). The curves in general rise gradually to a
maximum becoming more marked with the different treatments after the
day and show later the maintainance of the definite gradation.
A statistical analysis of the final dry weight per plant of the differently
treated sets shows a significant increase with all the treatments, the best
shown by N SS P 33 (ammonium phosphate) and N 33
f ammonium sulphate) which are also better than K, P and NKP treatments.
24
23
22
21
20
19
18
17
16
15
* 14
1 13
112
ill
^
3 10
9
8
7
6
5
4
3
2
1
Chemical Fertilisers on Growth & Water Requirement of IV heat 33
NP
Control
.,- ^ . Ammonitim sulphate
- Potash sulphate
P *-. Superphosphate
NKP ^n , K. * Potash mixture
NP M i n i. . * Amraonium phosphate
14 21 28 35 42 49 56 6^ 70 77 84 91 98 103
Age in days
FIG. 1
Growth, of wheat as influenced by fertiliser treatments
34 B. N. Singh and J. R. Singh
When the relative growth rates at successive stages of the life-cycle of
the plant are considered (Fig. 2) three characteristic peaks are noticed in the
treated and also in the control plants. Of the three maxima the first one is
14 21 28 35 42 49 56 63 70 77 84 91 98 103
Age in days
FIG. 2
Relative growth, rate of wheat under fertiliser treatments
Control
X - Ammonium sulphate
Potash sulphate
XKP * * * Potash mixture
XP Ammonium phosphate
p _.. Superphosphate
Chemical Fertilisers on Growth & Water Requirement of Wheat 35
observed at the early vegetative phase (21-28 days), the second one at the
pre-flowering period (42 days) while the final one is noticed at the flowering
stage (63 days). Plants grown with the different fertilisers exhibit a marked
increase in their rate of growth as compared to the control plants, specially
during the vegetative and the flowering stages when the need for nutrition is
most felt. A perusal of the curve further indicates that the intensity in the
rate of growth seems to be partially correlated with the nature of manure
added. Thus, in the seedling stage the N 33 P 33 (ammonium phosphate)
treated plants exhibit the maximum growth rate while in the flowering stage
this is shifted to N 33 (ammonium sulphate) series. The significance of these
results would be discussed in a subsequent contribution.
Water Requirements. When transpiration per unit dry weight is plotted
against the age of the plant four characteristic phases become evident
(Fig. 3). The seedling stage (14th day) is characterised by a high transpira-
tion activity which goes on declining till a low efficiency is indicated during
the pre-fiowering stage (49th day). There is again a sharp rise and a high
transpiration rate is observed during the flowering stage (63rd day). Soon
after this, there is again a fall and the lowest transpiration rate is obtained
during the senile stage. The curves of the plants for both the control and the
treated series show these four characteristic phases, variations being noted
in each case only in the amount of water transpired for the plants grown
under different fertiliser treatments. One of the chief characteristics of the
treated plants is the lower value of transpiration per unit dry weight as
compared to the control plants. It is interesting to observe that the plants
receiving K 78 . 7 (potassium sulphate) show the least amount of transpiration
while NP, N, NKP and P follow the decreasing order of efficiency. This order
does not remain constant since towards the latter stages of the growth,
the order reverses to NP, N, NKP and K. A study of the data presented
in Fig. 3 further shows that, as in the control plants, the plants of the other
series also exhibit the two critical periods of water requirement which are
at the seedling and the flowering stages of their life-cycle. The addition
of the fertilisers therefore does not bring about any drift in the critical
periods when the maximum supply of water should be available for the
growth of wheat plants.
The high transpiration efficiency of differently treated plants during the
very early seedling stage and the pre-flowering decline to a very low value,
and the more or less similar trend of the values for evaporating power of the
atmosphere (Table II) indicate that the ups and downs noted in the general
contour of the curves may partially be related to the rise and fall in the
1 X. Sir/en and J. R.
14 21 28 35 42 49 56 65 70 77 84 91 98 103
Age in
FIG. 3
per unit dry weight at successive life-stages
with different fertiliser treatments
C
Control
X Ammonium sulphate
p : Ammonium phosphate
p . Superphosphate
K Potash sulphate
-t Potash mixture
Chemical Fertilisers on Growth & Water Requirement of Wheat 37
evaporating power of atmosphere. It may further be remarked that similar
curves for transpiration/ dry weight ratio at successive stages of growth were
obtained with cotton, rice and tobacco 10 thus indicating that the pre-
flowering decline in water requirement is a characteristic feature of majority
of crop plants.
An examination of the data on the total water requirement of the
plants (Table III) indicates that it is considerably reduced, when the plants are
TABUS III
Influence of fertilisers on the water requirement of wheat
Water requirement
based on
Average
Treatments
transpi-
ration per
plant, gm.
Dry
matter
accumu-
Grain
yield
lation
Control
5841-91
407
1381
Ammonium phosphate (N 33 P 33 )
8186-92
345
960
Ammonium sulphate (N 33 )
8808-17
386
1139
Potassium sulphate (K 7 s , 7 )
5548
327
1019
Superphosphate (P 66 )
7251-25
399
1200
Potash mixture (N 16 . 5 K 8 P 16 . 15 )
7688-85
397
1141
Critical difference (P = -05)
163-26
11-16
519-8
manured. The plants grown without any dressing of fertiliser require more
water for their successful growth (unit dry matter production) than those
supplied with the artificial manures. A reference to the critical difference
even on the 5% level of significance (Table III) shows, however, that the
differences with the unmanured series are significant in all the treatments
except in the cases where P 66 (superphosphate) or N 16 . 5 E 18 P 16 . 5 (potash
mixture) are applied. The values obtained for .these two treatments are of
particular interest since the effects of different treatments are clearly shown.
A perusal of the data further indicates that amongst all the treatments
K 78 . 5 (potassium sulphate) reduces the water requirement to the maximum
extent. When the data of water requirements based on grain yield are
reviewed a similar decrease is noted although due to a high error no signi-
ficant difference is obtained. The water requirement of plants both on the
basis of dry matter accumulation and yield shows two distinct orders as are
K X. S::;:rh a:::i J. R. Singh
i;f K series (potassium sulphate) has
while for the latter XP (ammonium
TABLE IV
i.':. ..':,.-. -_. ...t*.._. ._' . iii^'.&cls
Water require-
_, + , ,, * * r - J Tatments
ment on grain
-. : .-:.t : ..^ >!
yield basisf
. ...,../. . . . . l.*A' AL.K' .::::: m phosphate
1-000
-"'*/ 1.0r~r> P.'^a'-iuni sulphate
1-061
, , ,- i '- 1 t * , .. I- ISO Ammonium sulphate ..
1-186
, , 4 .VT. . . .- I--I4 Potash mixture
1-188
*,-,;' -:!.!* .. . . I -219 ' Superphosphate
1-250
: , : ,. .. . . 1- 244 Control
1-438
V.. - v .- --L- 1 In tf/rnis of pot a-?! urn sulphate taken as 1-000.
V t". i' - - v --C 1 in icrn:-? of ammoniun: phosphate taken as 1-OCO.
\lirti tj.e question of absolute amount of water necessary for the crop
the above fertilisers are added is discussed, a computation of the primary
'latj. rfrr.v? that the quantity varies from 7-4 to 9-6 acre Inches (Table V)
IM: thr different scries. It Is Indeed interesting to note that the total
nuantitv of water necessary Is slightly less when the crop is manured either
by K 'potassium sulphite; or P ; superphosphate) alone. But when the
cror> is fertilised otherwise the total irrigation in acre inches is increased
beyond that required for the uninanured plants by about 5-12%
although such an extra expenditure Is more than heavily repaid by the high
Increase In yield obtained In all these cases. One of the greatest advantage lies
in the fact that no Increase In the number of Irrigation Is necessitated although
a slightly heavier demand of water Is made. It is therefore apparent that
the fertilisers may be advantageously used for obtaining higher yield at a
relatively low cost of irrigation.
Thus,, In addition to Inducing favourable variations in morphological
other plant characteristics, the benefit of the use of the fertilisers lie in
Inducing drought resistivity effecting thereby an economy in the water
balance of the plant system. From a purely economic point of view their
application the twofold advantage of greater yield accompanied by a
lower cost of irrigation.
Chemical Fertilisers on Growth & Water Requirement of Wheat 39
TABLE V
Influence of fertilisers on the water requirement of wheat in acre inches
and the yield of both straw and grains in Ibs. per acre
Total water
_ , i Yield per acre
Total ;
Treatments
required hy
crop per acre
in gallons*
water
in acre
inchest
Grain ' Straw
in Ibs. in Ibs.
Control
196548
8 '6 ! 1232 ! 2944
Ammonium phosphate
204508
9-0
1855
3301
Ammonium sulphate
217802
96
1681
3283
Superphosphate
193458
8-5
1473
2962
Photas mixture
214151
9-4
1644
3079
Potassium sulphate
166602
7-4
1374
2917
*' These values were found out by adding the water transpired by the crop and the water evapo-
rated from the soils in gallons per acre.
f These values have been calculated on the basis of amout of water required to irrigate one
acre of wheat.
Summary and Conclusions
A detailed study has been made to elucidate the after-effects of the
addition of certain fertilisers, e.g., ammonium sulphate, ammonium phos-
phate, potassium sulphate, superphosphate and potash mixture on the growth
and water requirement of wheat.
Treated plants in general exhibit decided improvement in their growth
than the untreated ones. While the height, tiller number and total dry
matter accumulation per plant are significantly better in all the fertilised
series than the unmanured ones, K 78 . 7 (potassium sulphate) and P 66 (super-
phosphate) applied alone do not show significantly higher increase in
the yield of grain and straw as also of the length of earhead or the absolute
weight of the grains.
The general effect of the fertilisers is to cut short the water requirement
of the treated plants to values much below that obtained for the control ones
although with P 66 (superphosphate) and N 16 . 5 K 38 P 16 .5 (potash mixture) the
differences are not statistically very significant. The total amount of water
necessary for raising a successful crop of wheat is therefore less in certain
cases, e.g., potash sulphate and superphosphate, while in other series it
increases beyond the values recorded for control apparently due to the
40
B. N. Singh and J. R. Singh
very high yield of grain obtained in tliese cases. Thus, the application of
fertilisers besides showing the higher yield of the crop has the added advan-
tage of minimising the cost of irrigation.
1. Fisher, E. A.
2. Gardner, P. D.
3. Hartwell, B. L.,
Wheeler, H. J., and
Pember, F. R.
4. Lawes, J. B.
5. Leather, J. W.
7. Maercker, M.
8. Reed, H. S.
9. Sachs, J.
10. Singh, B. N., Singh,
R. B., and Singh, K.
11. an( i Singh, B. R.
LITERATURE CITED
Statistical Methods for Research Workers, 5th edition,
1934, Oliver and Boyd, Edinburgh, Tweeddale
Court, London, 33 Paternoster Row, E.G.
" Fertility of soils as affected by manures," Bull. 48,
Bureau of Soils, U.S. Dept. of Agric., 1908.
" The effect of the addition of sodium to deficient
amounts of potassium upon the growth of plants
in both water and sand cultures," Ann. Rept. R.I.
Acjric. Expt. Sta. 9 1908, 20, 299.
" An experimental investigation into the amount of
water given off by plants during their growth,"
Jour. Hort. Soc. London, 1850, 5, 38.
" Water requirements of crops of India," Mem. Dept.
Agri., Ind. Chem. Ser. 9 1910, 1, 8.
" Water requirements of crops in India," ibid., 1911,
, 10.
" Versuche uber die Beeinflussung des Wasserver-
brauche cler Pflanzen durch die Kalirohsalze,"
Jahrb. AtjriJc. Cliem. Versuchsst. Halle, 1895, 15 ;
also Arbeit Landw. Gesells., 1896, Heft 20.
" The effect of certain chemical agents upon the trans-
piration and growth of wheat seedlings," Bot. Gaz.,
1910, 49, 2, 81-109.
" Ueber den Einfluss der chemischen und physikalis-
chen Beschaffenheit des Bodens auf die Transpi-
ration der Pflanzen," Landw Versuclisst., 1859,
1, 203.
" Investigations into the water requirement of crop
plants," Proc. Ind. A cad. Sri., 1935, 1, 9.
" Growth and water requirement of crop plants in
relation to soil moisture," ibid., 1936, 4, 5.
ASPECTS OF THE ANATOMY OF ANURA
(AMPHIBIA) A REVIEW*
BY I,. S, RAMASWAMI
(From the Department of Zoology, University of Mysore, Central College, Bangalore)
Received July 15, 1939
(Communicated by Prof, A. Subba Rau, D.SC., F.R.M.S.)
CONTENTS
PAGE
I . Introduction . . . , . . . , 42
II. Literature Selected . . . , . , , , 45
III. Review :
(a) Narial Region . . . . . . . . . . 45
(b) Prechoanal Sac . . . . . . . . . . 47
( c) Septomaxilla . . . . . . . . . . 48
(d) Erninentia Olfactoria . . . . . . . . 49
(e) Sphenethmoid . . . . . . . . . . 50
(/) Subethmoidal Cartilage . . . . . . . . 50
(g) Maxillae . . . . . . . . . . 51
(h) Prevomer . . . . . . . . . . 52
(i) Palatine Bone . . . . . . . . . . 56
(j) The Nasal, Frontoparietal, Squarnosal, Pterygoid,
Quadratojugal and Parasplienoid Bones . . 58
IV. The Middle Ear Region . . . . . . . . 60
V. The Pterygoquadrate and its Attachments, and the Arteria
Carotis Interim . . . . , . . . 63
VI. Bursa Angularis Oris (Fuchs) or the Mundwinkeldriise . . 65
VII. The Lower Jaw . , . . . . . . . . 66
VIII. The Hyolaryngeal Apparatus of Microhylidse and Pelobatidae 66
IX. The Vertebral Column . . . . . . . , . . 68
X. Summary and Conclusions . . . . . . . . 71
XI. Acknowledgement . . . . . . . . 76
XII. Bibliography . . . . . . . . . . 77
XIII. Errata I^ist . . . . . . . . 80
* Based on the published papers of the author and accepted for the Doctor of
Science Degree of the University of Madras, March 1938,
41
42 L* S, Ramaswami
I. Introduction
AT the outset, it must be pointed out that tinder the title of anatomical
studies of Indian and some extrapeninsular Anura, I have examined the
cranium and larynx by the method of sections and also by gross study, and
the morphological features of the vertebral column of some Indian anuran
species by the latter method. It may be remarked here, that the study of
cranial morphology by the method of sections revived at the incentive,
given by Dr. de Villiers of South Africa, is still young. The various internal
anatomical characters exhibited by the anuran species have to be examined
comparatively and then only we can embark upon any generalizations. As
we go on working, the only procedure can be, therefore, to correlate and
compare such features as become manifest in the several families and sub-
families and then try to assess their usefulness from the view-point of taxo-
nomy. In trying to correlate these characters, we have to bear in mind that
the environmental conditions may have a profound influence upon the
organization of individuals, and that secondary modifications in response
to changed surroundings are often largely developed.
I have examined the cranial morphology of adult specimens of the
following species and in many cases more than one specimen has been
utilised for this purpose (author, 1932, 1932 a, 1934, 1935, 1935 a, 1937) :
1. Microhylida.
Microhyla ornata Dutn. & Bibr.
K&loula pulchra Giinth. part (K. pulchra taprobanica Parker, 1934)
Uperodon systoma Schneid (Cacopus systoma Schneid).
Glyphoglossus molossus Giinth.
2. Ranidce.
Rana hexadactyla I/ess.
Rana cyanophlyctis Schneid.
Rand curtipes Jerdon.
3. RhacophoridcB ( Polypedatidse).
Rhacophorus maculatus Blgr.
Rhacophorus microtympanum Giinth.
Philautus petersi Blgr.
Philautus chalazodes Giinth.
Philautus oxyrhynchus Giinth.
Some Aspects of the Anatomy of Anura (Amphibia) A Review 43
4. Bufonidce.
Bufo melanostictus Sclineid.
Bufo parietalis Blgr.
Bufo hololiits Gtinth.
Bufo beddomii Gtinth.
Necto-phryne misera Mocq.
5. Pelobatida.
Megophrys major Blgr. (M. gigas Jerd. = major Blgr.).
Scaphiopus holobrookii Harlan.
The descriptions of the sections of the larynx of Uperodon systoma
Schneid, Kaloula pulchra Giinth. part (K. pulchra taprobanica Parker) and
Microhyla ornata "Dum. and Bibr. (author, 19326) are given; and
morphological accounts of the larynx of Megophrys major Blgr. and
Scaph^op^ls hammondii (author, 1935 b) and of the hyoid of these and of the
South Indian Microhylid forms named above are dscribed.
The morphology of the adult vertebral column in the following species
is described (author, 1933) :
1. Rhacophorida (Polypedatidse).
Rhacophorus maculatus Blgr.
Rh. eques Gunth.
Rh. microtympanum Gunth.
Philautus chalazodes Giinth.
Ph. sylvaticus Rao.f
Ph. nasutus Giinth.
Ph. oxyrhynchus Gunth.
Ph. sp. (marked B in the Central College Museum collection).
2. Ranidcc.
Micrixalus saxicola Jerd.
Micrixalus sp. (marked A in the Museum collection).
Nyctibatrachus major Blgr.
N. pygmccus Blgr.
N. sanctipalustris Rao (1920).
Nannobatrachus kempholeyensis Rao (1937).
Ranct' beddomii Giinth.
R. bhagmandalensis Rao (1922).
t This form has been since discovered to be a new species of Nyctibatrachus by Rao
(1937), and is therefore, treated as Nycttoatrachus- sylvaticus Rao.
B3a F
44 L. S. Ramaswami
R. breviceps Sclineid.
R. brevipalmata Peters.
R. crassa Jerd. ( R. tigrina var. crassa Jerd.)
R. cyanophlyctis Sclineid.
R. curtipes Jerd.
7?. diplosticta Blgr.
R. gracilis Gravh.
R. intermedium Rao (1937).
R. leithii Blgr.
2?. leptodactyla Giinth.
R. limnocharis Wiegm.
R. malabarica Tsch.
R. pantherina (R. tigrina var. pantherina Fitz.).
JR. pavambiculamana Rao (1937).
R. semipalmata Blgr.
R. tenuilingua Rao (1937).
R. sauriceps Rao (1937).
The occurrence of a Bursa angularis oris or Mundwinkeldriise is
reported by me (1933 a] in Glyphoglossus molossus Giinth., Uperodon systoma
Schtieid, Microhyla ornataQum. andBibr. Kaloula pulchra Giinth (K. pulchra
taprobanica Parker) and in Rhacophorus maculatus Blgr. The Apodan
genera Ichthyophis and Ur&otyphlus were also investigated in this
connection.
With regard to the Microhylid species investigated by me, the genus
Kaloida Gray has been subsequently split into two, Kaloula Gray and Rama-
nella Rao by Parker (1934) and this splitting is based on the difference in
the nature of the prevomer. Rao (1925) described a new genus of Micro-
hylid frog from South India which he called Ramanella symbiotica, and he
makes no reference to the exact nature of the prevomer in this species.
This form, however, was later discovered to be none other than a species
of Kaloula, K. variegata Stoliczk. But Parker (1934) retains the generic
name Ramanella for accommodating five species which were originally
included under Kaloula Gray and according to him the difference. between
Kaloula Gray and Ramanella Rao is that in the latter, the prevomer is reduced
and broken up. My descriptions of K. pulchra were drawn up from a
specimen of the race which Parker subsequently described as K. pulchra
taprobanica (1.934).
As regards the other genera, I included Rhacophorus (Polypedates)
under the family Ranidse following the observations of Gadow (1901) and
Some Aspects of ike Anatomy of Anura (Amphibia) A Review 45
Boulenger (1.882 and 1890). But Noble (1927 and 1931) has dissociated
Rhacophorus from the Ranidse and erected a new family Polypedatidse, in
which he includes Rhacophorus (Polypedates], Philautus and some other
genera. Moreover, the genus Rana itself has been split into nine subgenera
by Boulenger (1920), viz., Rana, Tomopterna, Hildebrandtia, Ptychadena,
Aubria, Hylorana, Discodeles, Nanovana and Pyxicephahis. The two
species R. hexadactyla and R. cyanophlyctts come under the subgenus Rana
s. str., while R. curtipes is included under the subgenus Hylarana. This
splitting is very well supported by my anatomical observaions.
In describing the Bufonid head (author, 1937), I have remarked in the
introduction that Notaden is a member of the family Bufonida\ I have
followed Gadow (1901) and Noble (1931) in this, and Noble (1931) correctly
points out that neither Notaden nor Myobatmchns shows any " affinity to
bufonids found to-day outside Australia " and insists that they do not belong
to the subfamily Bufoninse which according to Parker (in litt.) is approxi-
mately equal to Bufonidse of Gadow and Boulenger. With regard to the
other genus Nectes, it was placed as a synonym of Pseudobufo by van
Kampen (1923) and this nomenclature is followed by vSmitli (1930). Further
the Bufonid genus Cophophryne of Boulenger is treated by Noble (1931)
under the family Pelobatidse.
In connection with the vertebral column, I have examined members
belonging to the two families Ranidac and Rliacophoridoe (Polypedatidse).
Subsequent to the publication of a note on the precocious nature of the 8th
and 9th vertebrae in Rhacophorus maximus by Mookerjee (1.932), it occurred
to me that " if it could be shown that the vertebrae arc uniformly precocious
in this genus Rhacophorus, then its inclusion under the family Ranidsc,
becomes a questionable procedure ". I have examined four species of
Rhacophorus of which one (R. maculatus) confirms the observations of
Nicholls (1915-16) in being diplasiocoelous.
II. Literature Selected
In referring to previous work I have largely restricted myself to recent
workers who have studied the skull of anuran species by the method of
sections. This does not necessarily mean that I have ignored the morpho-
logical descriptions of other authors, and in some cases I have not been able
to secure the necessary literature, and therefore, my bibliographical list may
not be exhaustive.
///, Review
(a) Narial region. The very first structure that we meet with in
the rostral end of the anuran examples studied is the prenasal cartilage
46 L S* Ramaswami
associated with the premaxilla. In the Microhylid examples studied, it is
noticed that Uperodon and Glyphoglossus possess both the prenasal cartilages
(prenasalis superior and inferior) while their congeners Microhyla ornata
and Kaloula pulchra (K. p. taprobanica) possess only the superior cartilage.
On p. 2 of my paper on the Glyphoglossus head (author, 1932 a) I have said,
" At the outset it must be pointed out that Glyphoglossus stands apart from
the other members of the group Engystomatidse in very many features.
Both the prenasal cartilages, the superior and inferior, are present and
support the premaxilla ". This gives an idea that the Microhylid
(Engystomatid) examples examined possess only one prenasal cartilage ;
the sentence ought to read " Glyphoglossiis possesses both the prenasal
cartilages and in this feature it resembles Cacopus, but in many other
characters it differs from the other members of the same family with which it
has been compared". In the Ranid forms (JR. hexadactyla, R. cyanophlyctis,
R. curtipes) and the Rhacophpridse (Polypedatidee), (Rh. wwculatus and
JR. microtympanum, Philautus petersi, P. chalazodes, P. oxyrhynchus) and
the Bufonid examples (B. melanostictus, B. parittalis, B. hololius, B. beddomii
and Nectophryne misera) both the prenasal cartilages are present.
Previous work. In the forms examined by de Villiers (Phrynomerus
1930^, Cacosternum 1931., Anhydrophryne 1,931 c, Ascaphus 1.934,
Rhombophryne 1934- a, Microbatrachella 1934 b) the prenasal cartilages
are double, while in Breviceps fuscus (1931^), the superior is reduced,
in Hemisus (1931 6), the inferior is absent. In Bufo (Schoonees, 1.930),
Phrynobatrachus (G. du Toit, 1933), Liopelma (Wagner, 1934), Rana
and Crinia (C. duToit, 1933 ; 1.934) and Spelceophryne (de Vos, 1935)
the prenasal cartilages are normally disposed.
The cartilaginous tectum of the nasal capsule gives rise anteriorly to the
cartilago alaris and cartilago obliqua. The alary cartilage gives attach-
ment to the superior prenasal cartilage, while from the cartilago obliqua,
depends into the nasal chamber, a plica. In Rana the plica (Bruner, 1902) is
described as a connective tissue projection into the nasal chamber depending
from the tectal cartilage. But in the forms examined by me (Microhylidse,
viz., Uperodon systoma, K. pulchra (K. p. taprobanica) and Microhyla ornata
(1932), Glyphoglossus molossus (1932 a} } Rani das (1935), Pelobatidae (19350)
and Bufonidse (1.937), the plica depends from the cartilago obliqua and not
from the tectum. In Rhacophoridae (Polypedatidae) (author, 1934) the plica
may depend from the tectum (examined species of Philautus) or from the
cartilago obliqua (Rhacophorus maculatus and R. micro tympanum) .
Some Aspects of the Anatomy of Anura (Amphibia] A Review 47
Previous work. According to de Villiers the obliquous suspension is
noticed in Phrynomems (1.9300), Cacosternum (1.931), Anhydrophryne
(1931c), Rhombophryne (1934 a), Microbatrachella (1934 6), and by
C. du Toit in Rana grayi (1933). Similarly in Phrynobatrachus
(G. du Toit, 1933), Bufo (Schoonees 1930) and. Spelaophryne (de Vos,
1935). However in lyiopelmidse (Wagner, 1934), Crinia (C. du Toit,
1934) and Breviceps fuscus (de Villiers, 1931 d), the plica depends from
the tecturn.
The nasal chamber referred to above is disposed in the form of three
sacs- the cavum principale, cavum medium and cavum inferius. It is
noticed that posteriorly, the cavum principale opens into the buccal cavity
by means of the. choana. The cavum medium gives rise to the ductus
nasolacrimalis on its external aspect which opens below the eye normally
by a single opening. In Scaphiopus holbrookii, the cavum medium is com-
paratively diminished in size. The cavum inferius, it may be noted in
passing, is differentiated into a recessus medialis towards the septum where
it is surrounded by glands in all the forms examined. Now, this recessus
medialis is considered to represent the Organ of Jacobson so commonly
met with in Reptiles and other examples. I^apage (1928) homologises the
cavurn inferius with the Organ of Jacobson and Howes (1891) is inclined to
believe that the supposed Organ of Jacobson in Amphibia is a maxillary
sinus. Gaupp, however, notes the recessus medialis as Jacobson' s Organ.
I have followed the descriptions of Gaupp and labelled the thickened internal
part of the cavum inferius as recessus medialis.
( 6) Prechoanal sac. Another important feature one meets with in the
sectional views of the narial region of Uperodon systoma, Microhyla ornata
and Kaloula pulchra (K. p. taprobanica} is the occurrence of a prechoanal
sac. In Uperodon systoma anterior to the choanal opening, there appears
a vestigial sac in the roof of the mouth the prechoanal sac. In M. ornata
and K. pulchra (K. p. taprobanica} the cavum principale opens on either- side
into a prechoanal sac and these two sacs open into the buccal cavity.
However, in the larval forms of one of these examples (M. ornata} no precho-
anal sac is noticed, though in the case of U. systoma, the choanse of the
tadpole enter into a spacious prechoanal sac which opens into the buccal
cavity posteriorly. I have not examined the larval forms of Kaloula since
I was not able to secure the tadpoles. Now, with regard to the morpho-
logical significance of the prechoanal sac found in the adult of M. ornata
or K. pulchra (K. p. taprobanica}, I noticed that a similar state of affairs
is met with in Phrynomerus (de Villiers, 19300). He notes that " it is
48 L e S. Ramaswami
more than probable that the sacs referred to above are vestiges of the Organ
of Jacobson". Then if this view is accepted, is it that the narial part of the
Jacobson's Organ is represented by the recesstis inedialis of the cavtun
inferius and the buccal division in Micro hyla and Kaloula by the prechoanal
sacs ? In the other examples investigated by me, a prechoanal sac is
absent from Gluphoglossus (author, 1932 a) t Rhacophorus miwotympanum,
Philautus peter si, P. chalazodes, P. oxyrhynchtts (author, 1934), Rana
hexadactyla, R. cyanopJilyctis, R. curtipes (author, 1935), Scaphiopus holbrokii
(author, 1935fl), Bufo parietalis, B. hololius, B. beddomii, Nectophryne miser a
(author, 1937) while in Rhacophor us maculatus (author, 1934) and Megophrys
major (author, 1935 a), it is present.
Previous work. I will only refer to those forms where a prechoanal sac
is noticed. Phrynoments (de Villiers, 19300) has a paired prechoanal
sac. Cacosternum (de Villiers, 1931 a) has an unpaired one ; similarly
in Breviceps and Probreviceps (de Villiers, 1932 a), Hemisus (de Villiers,
1931 6), Spelaophryne (de Vos, 1935) and Rana grayi (C. du Toit,
1933).
(c) Septomaxilla. Associated with the narial cartilages, viz., lamina
superior cristse intermediae and lamina inferior cristae intermedise, the
septomaxilla (internasal of Gaupp) makes its appearance. This bone,
possessing posteriorly diverticula or limbs, is noticed to embrace the
recessus sacciformis or the infundibulum (the passage between the cavum
principale and cavum medium). In the case of Rana (author, 1935) the
posterior portion of the septomaxilla is triradiate ; this arrangement is not
seen in the case of Scaphiopus holbrookii (author, 1935 a). It is noticed in
the latter example that after the fusion of the two limbs (the superior laminal
and inferior laminal) of the septomaxilla as in Rana, the bone again appears
as two investments of the planum and continues to be so till it finally disap-
pears, thus differing from the Ranid type where the bone is noticed to delimit
the anterior extremity of the planum. Now, in the Bufonidae (author, 1937)
the bone follows the common plan in the examined species of Bufo while in
Nectophryne misera the disposition is slightly different. At any rate, there
is a common feature between the two genera ; a limb of the bone appears
below the plica in Bufo and slightly behind the plica in Nectophryne misera
a feature not met with in the Ranidae and Rhacophoridae (Polypedatidse). In
the latter family (author, 1934) the arrangement of the bone is Ranid in nature.
The minor variations noticed are not of great significance but it is to be noted,
however, that the nature of the bone itself has been interpreted in two ways.
According to I^apage (1928) the bone is considered as " originally a cartilage
Some Aspects of the Anatomy of Anura (Amphibia) A Review 49
bo lie " and one of the criteria in determining this is "in certain places for
example, where the septomaxillary is in contact with the lamina superior of the
crista intermedia, these cartilages seem to be continuous with the septomaxil-
lary a fact which further supports the view that the septomaxillary arises in
the cartilage " (p. 413). Now in the case of the examples studied by me, in
certain regions of the laminal cartilage of Uperodon systoma, Kaloula
ptdchra (K. p. taprobanica) and Microhyla ornata, the bone is actually in
contact with the cartilage thereby supporting I^apage's theory. This, in
fact, is really very pronounced in K. pulchra (K. p. taprobanica). I, there-
fore, concluded at the time that in these examples the bone was of carti-
laginous origin which, of course, was not supported by ernbryological
evidence. Subsequent study has convinced me that in a large majority
of forms like Rana, Rhacophorus (Polypedates) , Philautus, Bufo, Necto-
phryne, Megophrys and Scaphiopus, the bone is separated from the cartilage
by connective tissue ; and, more than that, " the attachemnet and apparent
continuity of one end of the bone with that part of the cartilage of the nasal
capsule does not mean that the bone was preformed in cartilage " (de Beer,
1937). A study of the development of the bone will finally settle the matter
and till then it may be treated as a membrane bone.
Previous work. W. K. Parker (1881) describes a large number of forms
without a septomaxilla, and it has been shown that his observations
are inaccurate in some cases. De Villiers and his school describe it
as a membrane bone.
(d) Eminentia olfactoria. Both anteriorly and posteriorly to the
choana, the solum may give rise to an elevated cartilage into the cavum
principale which is called the eminentia olfactoria. Such an elevation is
remarkably well developed in the fossorial Uperodon systoma, Microhyla
ornata and Kaloula pulchra (K. p. taprobanica). Uperodon is found to live
several feet below the earth, while Microhyla lives under loose sand and
Kaloula under rocks and the bark of trees. Further, in the case of Scaphiopus
holbrookii (author, 19350), Bufo (author, 1937), Glyphoglossus (author, 1932 a),
Rana hexadactyla and 72. fyanophlyctis and (author, 1935), the eminentia is
elevated. In the other examples investigated, viz., Rhacophorus (Poly-
pedates), Philautus, Rana curtipes, Megophrys major and the arboreal
Nectophryne misera, the eminentia is flat and not elevated. The African
School of anatomists headed by de Villiers put forward a theory that the
elevation of the eminentia is closely correlated with the fossorial mode of
existence of these animals. De Villiers (1932 a) significantly remarks in
this connection that " increase in the area of the eminentia olfactoria
SO L 8 S* Ramaswami
represents a purely physiological phenomenon, associated with adaptati
to terrestrial life and has been independently evolved in Bufonidse, Rani
and Brevicipitidse". This view fitted in extremely well when I worked (
the Microhylid examples but when I investigated the Ranid forms 1:
I?, hexadactyla and R. cyanophlyctis and R. curtipes, I was rather struck
the variation noticed in the nature of the eminentia. It is elevated
R. hexadactyla and .R. cyanophlyctis while in R. curtipes it is fiat.
is very well known that the former two examples are completely aqua
forms, and therefore, the occurrence of an elevated eminentia does not v<
well lit in with the theory that the elevated eminentia is closely correlal
with a fossorial mode of existence even though it may have been evoh
independently. If the two Ranid genera were terrestrial, the independ*
evolution of the elevated eminentia w r ould have amply borne out but sii
they are aquatic, I pointed out (1935, p. 6) that " the elevation of the ei
nentia has probably nothing to do with terrestrial adaptations of the Anu
It may, however, be said that the structure increases in area purely
response to the sensory requirements of the individual". Probably exp(
mental zoology may be able to clear this difficulty.
Previous work. The occurrence of an elevated eminentia is noticed
Phrynomerus (de Villiers, 1930 a), Breviceps (de Villiers, 1931 rf), JF
breviceps (de Villiers, 1932 a) and Spelceophryne (de Vos, 1935) and
reference is made to it in Rhombophryne (de Villiers, 19340).
Bufo angusticeps, Schoonees (1930) records an elevated eminem
(e) The sphenethmoid bone (Os en Ceinture Cuvier}. This b<
occurring in the ethmoid region is known as the girdle-shaped bone sinci
forms a girdle round the brain. It appears in the anterior sections on
lateral aspects of the brain and posteriorly extends ventrally also and
some examples a piece of trabecular cartilage is left unossified midventra!
when the bone is said to be distinctly paired. Such a feature is noticed
Uperodon systoma, Microhyla ornata, Kaloula pulchra (K. p. taprobanic
Rh. microtympanum and Philautus chalazodes. In the last two species :
sphenethmoid is feebly developed. However, in Mana hexadactyla, R. cya,
phlyctis, Megophrys major, Scaphiopus holbrookii, Philautus oxyrhynchus a
P. peter si, there is no ventral trabecular piece and the bone is one compL
girdle. In P. petersi the bone originates far anteriorly in the septum ns
In Rhacophorus microtympanum a feeble sphenethmoid is seen and in Ra
curtipes in the sphenethmoid region, no ossification is noticed.
(/) Subethmoidal cartilage. There is a remarkable feature noticed in 1
case of Rana cyanophlyctis in the ethmoid region. In the anterior region
Some Aspects of the Anatomy of Anura (Amphibia) A Review 51
the sphenethmoid, disposed between it and the parasphenoid, is a cartilage
which has been called the subethmoidal cartilage, also noticed in Rana gmyi
(C. du Toit, 1.933). What exactly is the significance of this cartilage, I am
unable to say.
Previous work.Vf. K. Parker's monograph (1.881) on the Anuran
skulls gives us an account of the sphenethrnoid but unfortunately
the double nature is not disclosed in many forms. The paired nature
of the bone is described by de Villiers in Phrynomerus (1930 a),
Rhombophryne (1934 a), by C. du Toit in Crinia (1934), by G. du Toit
and de Villiers in Hyperolius (1932), by Wagner in Liopelma (1934).
In the other forms whose cranial anatomy has been studied variations
are met with. In Cacosternum, the sphenethrnoid is noticed
posteriorly (de Villiers, 1931 ; 1931 a), the bone is completely girdle-
shaped in Phrynobatrachus (G. du Toit, 1933) and in Rana grayi
(C. duToit, 1933) and in Bufo angusticeps (Schoonees, 1930) portions
of cartilage are discovered in the bone. In Breviceps and Pro-
breviceps (de Villiers, 1932 a) and in Spelceophryne (de Vos, 1935), the
bone is wanting. In Ascaphus (de Villiers, 1934) an orbitosphenoid is
described and in ' I^iopelmidse ' .Wagner (1934) describes the sphen-
ethmoid as paired. Peculiarly in Hemisus (de Villiers, 1.931 b) the
bone has fused with the nasals above. Representatives of 58 genera
have been examined by Parker (1934) and the normal condition in
these is that the sphenethmoid is single ; however, variations in few
cases were also met with.
(g) Maxilla?. The premaxillse and maxillse do not show many varia-
tions. However, it may be remarked that in the Microhylid species
examined by me, viz., U. systoma, M. omata and K. pulchra (K. p. tapro-
banica), the maxillse (and also the prevomers) are edentulous. Now it has
been long established that no stress need be laid on the dentigerous or eden-
tulous nature of the bones, for in the same family, members possessing teeth
(e.g., Microhylid Dyscophinse) and also without them are found. Again
in Bufo and Nectophryne (author, 1937) the maxillse and prevomers are
edentulous while in Ranidse (the author 1935) and Rhacophoridse (Poly-
pedatidse) (author, 1934), the maxillse are dentigerous. Therefore, this
character cannot be utilised for systematic purposes especially with reference
to the Microhylid family.
Previous work.W. K. Parker (1881) describes a large number of forms
with and without teeth on the maxillae. In describing the maxillary
bones, de Villiers (1931) also notes that "Noble has repeatedly
52 L 9 S* Ramaswami
maintained that no great systematic value can be attached to the
absence or presence of teeth ". In describing the cranial morphology
of Cacosternum (1931, 1931 a) he notes that the species of this genus
behave rather differently ; C. bottgeri and C- capense possess pre-
maxillary teeth while C. namaquense does not but the maxilla are
dentigerous. Peculiarly, the Microhylid genera, Breviceps and Pro-
breviceps (de Villiers, 1932 a), Rhombophryne (de Villiers, 1934 a) and
Spelaophryne (de Vos, 1935) possess edentulous maxillae. Phryno-
merus (de Villiers, 1930 a) which is treated as a genus under the family
Phrynomeridse by Parker (1934) also possesses, like the Microhylid
genera referred to above, edentulous maxillse.
(h) Prevomer (Broom). The topographical disposition and the nature
of this bone is largely used in Anuran taxonomy. In the species that I
have examined of the Microhylid family, viz., U. systoma, M. ornata,
K. pulcJira (K. p. taprobanica) and Glyphoglossus molossus, the disposition
of the prevomer is based on a common plan. In U. systoma, the prevomer
appears divided anterior to the choana, is edentulous and there is a small
bone posterior to the choana, which I have called the degenerate palatine
in consonance with the nomenclature adopted in labelling the bone occurring
in this region in other anuran genera on purely topographical evidence.
In the second species of Uperodon, U. globulosum, the prevomer embraces
the choana (Parker, 1934) and a palatine is wanting. Now, since the palatine
is absent in this species, it is thought that in other species U. systoma also, a
true palatine is absent and the degenerate bone that is present postchoanally
[palatine according to me (1932) and Devanesan (1922)] may be the posterior
portion of the prevomer. If, however, it is proved that 8 7. systoma is more
primitive and possesses a prechoanal prevomer and a degenerate palatine,
then the condition noticed in U. globulosum should be interpreted as one in
which a fused prevomeropalatine bone is noticed. My slides of the head
of U. systoma do not provide any clue to this problem. In K. pulchra
(K. p. taprobanica) the prevomer is large and edentulous and in the alizarin
preparations that I have examined of K. pulchra (K. p. taprobanica), the
prevomer is well developed and the postchoanal portion is associated with
the well- developed palatine. I have already remarked that according to
Parker (1934), the genus Kaloula Gray is split into two genera Kaloula Gray
and Ramanella Rao, based on the nature of the prevomer, and since I have
not described any species belonging to Ramanella in these two papers (author,
1932 ; 1932 a), I shall not discuss the prevomerine condition in the latter
genus. On p. 11 of my paper (author, 1932), I have said that "The
Some Aspects of the Anatomy of Anura (Amphibia) A Review 53
vonier does not form a vomeropalatine and in Kaloula the vomer is super-
imposed by the palatine ". It ought to read that the palatine is super-
imposed by the vomer. Now in M. ornata, the prevomer is poorly developed
and a limb of it extends intrachoanally as in Phrynomerus (de Villiers, 19300)
but this does not embrace the choana posteriorly. I have clearly stated
this point in my paper on Rhacophoridae (Polypedatidse) (author, 1934,
p. 84) : " The occurrence of this intranasal prolongation of the vomer
(prevomer) is also noticed in one of the South Indian Engystomatid examples
Microhyla and in a large number of foreign forms [see Phrynobatrachus
(G. du Toit, 1933) and Phrynomerus (de Villiers, 1930)]". In my specimen
of Glyphoglossus molossus (author, 1932), the prevomer is small and there is
no intrachoanal prolongation as much as we see in M. ornata. But in a
figure drawn by Parker (1934, p. 72) of the ventral aspect of the cranium of
G. molossus, a prevomer which embraces the choana is shown. I am unable
to make out the palatine as an independent bone in the figure and in the
description we read " Prevomer undivided, the postchoanal portion over-
lying the palatine region and bearing, mesially, one or two knob-like promi-
nences (Fig. 30)". The figure suggests that a prevomeropalatine arrange-
ment is present. My description of the prevomer in Glyphoglossus (author,
1932 a) is that " at the base of the solum nasi the prevomer makes its appear-
ance and unlike the Brevicipiticbe and Microhyla, no extension of the pre-
vomer seems to embrace the choanse in Glyphoglossus". What I mean by
" embrace " is an intranasal prolongation of the prevomer as in Microhyla,
and not a post-choanal extension as seen in U. globulosum. As already
noted above, I have made this point clear in my paper on Rhacophoridse
(Polypedatidse) (author, 1934, p. 84, vide supra). Further my slides do not
disclose ' the postchoanal portion overlying the palatine region ' in Glypho-
glossus as described by Parker (1934, p. 72). This is due to the fact that
my observations were made 011 an immature specimen. In this, the bone
investing the ventral aspect of the antorbital cartilage which I called the
palatine is separate, and there is a prechoanal prevomer. When the adult
condition is reached, the prevomer and the palatine (if it is a palatine)
obviously fuse together (Parker in litt.) and a prevomeropalatine arrange-
ment is noticed ; or it may be that the palatine really represents the post-
choanal portion of the prevomer. Then, the figure drawn by Parker (1934,
p. 72), represents a prevomer with a postchoanal portion or it may be a
prevomeropalatine bone. Further, Parker points out that this genus
Glyphoglossus " ought, perhaps, to be united with Uperodon. Uperodon
globulosum is almost exactly intermediate between the type species of
Uperodon and Glyphoglossus". In adult U. systoma, then, there is a pre-
54 L. S. Ramaswami
chcanal prevomer and a postchoanal prevomer or palatine ; in G. niolossus,
the juvenile specimens show a prevomer and a postchoanal prevomer or
palatine and in the adult, the prechoanal and postchoanal portions fuse
representing either a prevomer or a prevomeropalatine. Similarly iu
U. globules-urn, the bone may be a prevomeropalatine or a true prevomer
with a postchoanal limb. Now, in the Rhacophorid (Polypedatid) family
examined (author, 1934), the prevomer is rather interesting. According
to Boulenger (1890), the difference between Rhacophorus (Polypedates} and
Phila-utus (Ixalus) consists in the presence of prevomerine teeth in the former
and their absence from the latter. I have noted in that paper (author,
1934, p. 82) that "This may not be so very safe and stable criterion of enough
systematic importance since Boulenger himself is doubtful about the exis-
tence of vomerine teeth in Rhacophorus dubius". Noble (1927, 1931) at any
rate, distinguishes Rhacophorus (Polypedates} from Philautus by the absence
of vomerine teeth in the latter and he points out that the genus is derived from
Rhacophorus by a ' loss of the vomerine teeth '. However this may be, the
family Rhacophoridae (Polypedatidse) according to the same author (1931),
is a natural group possessing cylindrical sacral diapop} r ses, intercalary carti-
lages and a diplasioccelous vertebral column, and further confirms his
statement by saying that " the anatomical evidence at present available
points toward the Polypedatidse as being a natural group 5 ' (p. 525). Within
the family, since Philautus has arisen from Polypedates by the loss of vomerine
teeth, Noble is inclined to believe that Philaittus is not a natural genus but
a polyphyletic one. The sectional views of the prevomer reveal that the
bone is dentigerous in Rhacophorus maculatus, Philautus chalazodes and
P. oxyrhynchus while in P. peter si and Rhacophorus microtympanum, it is'
edentulous. Thus, both among Rhacophorus and Philautus, there are
species with and without prevomerine teeth and it becomes apparent how
futile it is to utilise this character for purposes of further classification of
these two genera. If we accept Noble's dictum that the genus Philautus
represents a polyphyletic assemblage, then we are not justified in retaining
it as a single distinct genus ; we may have to merge the several species of
Philautus under those of Rhacophorus. This view is very well supported
by my observation of the prevomerine teeth for, both genera Rhacophorus
and Philautus possess species with and without prevomerine teeth. But the
otker aspects of cranial anatomy as reported by me (author, 1934) do not
warrant this, and therefore, it is very unlikely tjiat Philautus is, as Noble
thinks, polyphyletic, unless it be that the other characters, too, have been
developed independently on more than one occasion (Parker in litt.). In
the Ranid examples studied, viz., R, hexadactyla, R. cyanophlyctis and
Some Aspects of the Anatomy of Anura (Amphibia] A Revieiv 55
R. curtipes, the prevomer is normally disposed with no postchoanal portion,
is dentigerous and resembles that in R. temporaria (Gaupp, 1904).
In the Pelobatid examples studied Megophrys major and Scaphiopus hoi-
brookii (author, 1935*2), the prevomer is single on either side and possesses
two limbs posteriorly. In M. major (author, 1935, p. 72, Fig. 5) " on the
ventral aspect of the eminentia the prevomer is noticed and posteriorly,
it is divided into two between which the Rachendrii.se are present (Fig. 5 a)
and more posteriorly the internal limb (Fig. 5 6) is present and this is the in-
trachoanal portion of the prevomer," which, however, disappears posteriorly.
In S. holbrookii, 'Disposed at the tip of the bony eminentia is the large
vomerine bone' (p. 72). "The vomer (prevomer) (Figs. 4 and 6 v) in
Scaphiopus underlies the large bony eminentia and, as in Megophrys, in pos-
terior regions the bone is present in the form of two parts with loose connec-
tive tissue and the Rachendruse in between " (pp. 73 and 74). Posteriorly,
the maxillary limb of the prevomer alone persists and this also disappears
after cutting through some or more sections posteriorly. In Bufo and
Nectophryne misera (author, 1937), the prevomer is small and edentulous.
It does not extend postchoanally. In my introductory chapter on the
Morphology of the Bufonid head, I have remarked about ' the absence of
the teeth from the jaws (except Notaden) . .' and this implies that Notaden
possesses dentigerous jaws. The implication was not intended ; Notaden also
lacks teeth in the jaws, but prevomerine teeth are present. In passing, it
may be noted that the systematic position of Notaden is changed now, and
I arn not in possession of the literature referring to it (see, however,
Noble, 1931," p. 498).
Previous work. W. K. Parker (1881) depicts the prevomer in a large
number of forms and he draws figures of the skulls of two species of
Microhyla under the name of Diplopelma and also that of Calhtla
pulchra. H. W. Parker (1934) describes, amongst other Micro-
hylidse, the disposition of this investing bone in Uperodon, Kaloula,
Ramanella, Microhyla and Glyphoglossus. In Uperodon, ' Prevomer
entire or divided (Fig. 31 and 32), the postchoanal portion overlying
the palatine region and sometimes bearing a raised knob at its mesial
end ' ; in Kaloula ' prevomer undivided, the postchoanal portion
overlying the palatine region and raised into a strong, sometimes
crenulate ridge (Fig. 33) '. However, Smith (1930) draws a palatal
aspect of K. pulchra (Fig. 10, p. 121) and in the description he says
' vomer forming a sharp transverse ridge behind the choana ; palatine
much reduced, its inner extremity underlying the vomer '. In
Phrynomerus (de Villiers, 1930 a), a vomeropalatine arrangement is
56 L. S* Ramaswami
seen; in Cacosternum (de Villiers, 1.931 a), vomer is edentulous and
large"'; in Breviceps (de Villiers, 1931 d), there is an intranasal prolonga-
tion which is absent from Probreviceps (de Villiers, 1932 a)\ in Hemisus
(de Villiers, 1931 J), the vomer is absent ; in Rhombophryne (Noble and
Parker, 1926 ; de Villiers, 1934), it is disposed in two parts ; in Asca-
phus (de Villiers, 1934), there is a toothed prevomer with the rostral
portion edentulous; in Spelaophryne (Parker, 193-1 ; de Vos, 1935),
the prevomer is edentulous; in Crinia (C. du Toit, 1934), a divided
prevomer is present ; in R. grayi (C. du Toit, 1933), it is simple and
undivided; in Phrynobatmchus (G. du Toit, 1933), the edentulous
prevomer fringes the choana and in Bufo (Schoonees, 1930), the bone
is edentulous. Also in Hyperolnts (0. du Toit and de Villiers, 1932)
the vomer is without teeth.
(/) Palatine bone. In the Anura, the palatine bone shows considerable
variation. Forms with and without, or with degenerate palatine
or with a prevomeropalatine are not uncommon. Among the
forms studied by me, in the Microhylid genera, ' the palatine observes
a sequence in reduction'. Kaloula has a comparatively well-developed
palatine ; Uperodon systoma a vestigial one (see below) while M. ornata
has lost it. Smith (1930) remarks that in the case of K. pulchra (K. p. tapro-
banica), there is a ' reduced palatine ' and when we compare this with the
other two South Indian Microhylid examples, it must be said that the
palatine is comparatively well developed. In the case of U. systoma, a
bone on the ventral aspect of the quadratoethmoidal commissure has been
labelled by Devanesan (1922) as palatine and I have also followed the same
nomenclature. Whether it is a true palatine or only a postchoanal portion
of the prevomer is rather difficult to say and the probabilities have been
discussed on pp. 53-54. With regard to Microhyla, I have noted that the
palatine is absent (author, 1932, p. 11) and that 'Parker (23) finds that
among the species of Microhyla an orthogenetic series could be established,
ranging from forms having a palatine to forms completely devoid of it J . As
early as 1881, W. K. Parker figured two species of Microhyla under the name
of Diplopelma where he has drawn a well-developed palatine. Obviously,
the genus contains species both with and without a palatine bone. H. W.
Parker (1928) as stated above, had studied all the species of the genus
Microhyla and had come to the same conclusion. Pie draws an interesting
correlation when he says that ' only rarely does it persist when the post-
choanal portion of the prevomer is absent'. In the Ranid and Rhacophorid
(Polypedatid) forms examined (author, 1934, 1935), the palatine is normally
disposed on the ventral aspect of the antorbital cartilage.
Some Aspects of the Anatomy of Anura (Amphibia) A Review 57
In Megophrys major a single specimen of which I secured from the
Indian Museum at the time I reported, I have noted (author, 1935#) that
the palatine is absent and investing the ventral aspect of the quadrato-
ethmoidal commissure is a projection of the maxilla. This projection
forms the posterior boundary of the choana. However, after examining a
large number of species of Megophrys, including M. major, Parker informs
me (in litt.) that a palatine is normally present and therefore, my specimen
does not represent the normal condition. Subsequently I examined another
specimen of M. major and also discovered the same feature noticed in the
first one with regard to the palatine. Mr. H. W. Parker of the British
Museum has examined my second specimen. It is to be noted that the bony
extension of the maxilla described by me (author, 1935 a) as forming the
posterior boundary of the choana which I propose to call maxillo-palatine,
is topographically identical with the palatine of other Anura and therefore,
Parker (in litt.) is inclined to label this merely as palatine bone. Further
he states that if it is to be regarded as a maxillo-palatine, one of the two
assumptions have to be made, viz., that it is a bone not foreshadowed in
any frog and appears for the first time here or that the processus frontalis
of the maxilla has grown ventrally to the antorbital cartilage. If, however,
following Parker, it is labelled as palatine, the union of the maxilla and
palatine in this region is lost sight of and probably this is the first anuran
where such a condition of fusion has appeared.
In S. holbrookii, the palatine is wanting. Further, in the same paper
(1935 <2, p. 75) it is noted that ' in all the examined species of Microhyla
(H. W. Parker, 1928) the palatine is wanting .... ' This is not correct, and
therefore, it should be re-stated that in the examined species of Microhyla,
the palatine may be present or absent. In Bufo (author, 1937), the palatine
is normally disposed while in Nectophryne misera, the bone is wanting.
On p. 1163, I have noted in my paper on the morphology of the Bufonid
Head (1937) that ' Peculiarly, however, in the arboreal Nectophryne, the
palatines are wanting. This, I am informed, gives an impression that
in all the species of the genus Nectophryne, the palatine is wanting which
of course is not what I mean ; my idea is that in the examined species of
Nectophryne, the bone is wanting, which I have clearly pointed out in the
summary of that paper.
Previous work. W. K. Parker (1881) draws the disposition of the
palatine in a large number of forms described by him like Callula
pulchra and two species of Microhyla, which he describes under the
.name of Diplopelma, D. ornatum vel rubntm and D. Berdmorei (?).
5g L* S. Ramaswami
The descripiton of Parker of these two species ate as follows : " In
D ornatwn vel nibrum, the lateral rudiments of the girdle bone are
less and run into their own ate partially ; yet an endosteal deposit,
with scarcely any perichondrial bone (ectosteal palatine) runs tip
to the cheek' " He further mentions the presence of a postpaktine
being separated from the pterygoid. In D. Berdmorei, the palatine
is feebly developed. There is no girdle bone and in both species
Parker has figured a prevomer. I have not been able to secure
H. W. Parker's paper (1931) where an account of the prevomer and
palatine of Uperodon is given. The same author (1934) also de-
scribes the palatal bones in a large number of Microlvylidrc. In
Phrynomenis (de Villiers, 1930 a), a prevomeropalatine is formed;
similarly in Liopelma (Wagner, 1934) ; in Cacoslernum (de Villiers,
1931), Crinia (C. du Toit, 1934) and R. grayi (C. du Toit, 1933), Bufo
(Schoonees, 1930), the bone is normal. In Hemisus (de Villiers, 1 931 6),
Breviceps and Probreviceps (1932 a), Ascaphus (de Villiers, 1934) and
Spelaophryne (de Vos, 1935), a palatine is wanting. In Rhombo-
phryne (Noble and Parker, 1926; de Villiers, 19340), the posterior
portion of the prevomer overlies the palatine. According to W. K.
Parker (1881) in B. melanost ictus, there is a lopsided variation in the
development of the palatine which, however, is not seen in my
specimens.
(;) The Nasal, Frontoparietal, Squamosal (paraquadrate Gaupp} t Ptery-
goid, Qiiadratojugal (quadratomaxillary Gaupp} and Parasphenoid.Eliv
variations noticed in these investing bones are minor and present no great
morphological significance. The nasals may be so situated that the
sphenethmoid is exposed between them and the frontoparietals, or as in
Nectophryne misera and Megophrys major, the nasals may be separated
anteriorly where the ossified tectum can be seen. The frontoparietals
extend as far as the nasals anteriorly and posteriorly they cover the synotic
region. I have not noticed any co-ossofication between the frontoparietals
and the exoccipitals, but in Philautus peter si t the frontoparietals are united
raesially and in P. oxyrhynchus, the frontoparietals of either side unite with
the pro-otic bone. Both in Megophrys major and Scaphiopiis holbrookii, the
nasals and frontoparietals are studded dorsally with bony asperites, while in
the other examples, the bones are smooth. With regard to the parasphenoid,
the anterior end may show a divided appearance in some examples
(Scaphiopus holbrookii and Philautus peter si}.
The squamosal (paraquadrate Gaupp) in Ran a is a triradiate bone
according to Gaupp with an anterior, posterior upper and lower arms. In
Some Aspects of the Anatomy of Anura (Amphibia) A Review 59
the Microhylid examples studied by me, viz., U. systoma, M. ornata and
K. pulchra (K. p. taprobanica) , a gradual reduction in the development of
the arms could be studied and the sequence of reduction being in the order,
Kaloula, Microhyla and Uperodon. The posterior arm is reduced in Kaloula
and Microhyla while it is absent from Uperodon (Devanesan, 1922). In
Glyphoglossus molossus, it is not reduced. In the Ranid, Polypedaticl,
Bufonid and Pelobatid genera examined by me, the bone is typically Ranid
in nature and these are, therefore, of no great significance.
In discussing the suspensorial region of U. systoma, K. pulchra (K. p.
taprobanica) and M. ornata, I have said (author, 1932, p. 67) that <f In
Kaloula the processus quadratus becomes one with the processus ptery-
goideus and thus all the three, quadratomaxillary, paraquadrate and
pterygoid invest this cartilage, while in Rana only the quadratomaxillary
invests it". In all the Microhylid species examined and also in Rana, it is
only the quadratojugal that invades the processus quadratus, while the
squamosal and pterygoid invest it.
Now, in the descriptions of the cranium of the several Anura studied.
by me, I have designated the squamosal bone as the paraquadrate bone
following Gaupp and de Villiers. According to the latter author (.1936), 'the
paraquadrate in the Anura is not an investing bone of the otic capsule and
as such cannot be a squamosar. Further, he points out that it is typically
a membrane bone of the palatoquadrate, and adduces embryological evidence
from the study of Urodela and describes a quadrato-maxillary in the apodan
Boulengerula (1930). A year prior to the publication of this paper, Brock
(1.935) having studied the temporal bones in lyizards, Birds and Mammals,
pointed out that, "There is no need for Gaupp 's term ' paraquadrate ' for
the reptiles and amphibia, since it is merely a synonym for squamosal, a
bone which may be defined as a membrane bone primarily associated with the
lateral surface of the quadrate ] with the migration of the quadrate (incus)
into the tympanic cavity, the squamosal of the mammal secondarily becomes a
bone of the otic capsule'. Probably, de Villiers was not aware of this work (?)
or at any rate, he does not refer to it. In the latest book on the develop-
ment of the vertebrate skull, de Beer (1937) has pointed out that since
Gaupp was not able to discover in Amphibia a squamosal of the type seen
in Birds and Mammals, he called it a 'paraquadrate'. This view becomes un-
tenable when we realise that 'the living Amphibia with their large quadrate
cartilages are highly specialised, and that with the reduction of the quadrate
to the incus which becomes protected by the tegmeii tympaiii, the squamosal
which covered the quadrate in the lower vertebrates, becomes a covering
60 L a S a Ramaswami
bone of the auditory capsule in the higher forms. . . . '. Thus, It becomes un-
necessary to introduce the term paraquadrate for describing the squamosal.
It may not be out of place here to remark that de Villiers (1936) has
also discussed Gaupp's denomination of the quadratojugal as the quadrate-
maxillary. He points out that this is not a sesamoid bone and is represented
by a quadrate squame in Gymnophiona where it is described as being absent.
According to him, (< Gaupp's nomenclature is followed in preference to what
might be called a purely topographic one". Brock (1935) does not make a
critical reference to this bone nor does de Beer (1937), though in describ-
ing the osteocranium of Amphibia, the latter author retains the term
quadratojugal and treats it as a synonym of Gaupp's quadratomaxillary.
Previous work. I am only making a brief reference to these investing
bones in the other forms that have been studied ; the frontoparietals,
nasals and the parasphenoid are not always normally disposed ; in
Hemisus (de Villiers, 1931 i), the sphenethmoid fuses with the nasals.
In Phrynomerus (de Villiers, 1.930<z), there is a large frontoparietal
fontanelle ; in Cacosternum (de Villiers, 1931 ; 19310), these and the
pterygoid and squamosal are Ranid in disposition, but the fronto-
parietals are poorly developed. In Hemisus (op. cit.) the pterygoid
is normal while the anterior process is absent from the squamosal.
The frontoparietals are fused together and is disposed over the nas-
ethmoid. In Breviceps (de Villiers, 1932 a), the frontoparietals are
separate while in Probreviceps the nasals and frontoparietals fuse
together. In all the three genera (Hemisus, Breviceps and Probreviceps)
the quadratojugal is absent, and probably on account of this, de
Villiers notes that the squamosal and pterygoid have fused together.
In Ascaphus (de Villiers, 1934), the nasals and frontoparietals are
separate ; similarly the pterygoid and squamosal. In Rhombophryne
(de Villiers, 19340), the disposition of these bones is Ranid. In
Spelczophryne (de Vos, 1935), the squamosal is reduced ; the quadrato-
jugal is absent and in the pterygoid and squamosal have fused and
the frontoparietals are narrow bones. In the other examples (Crinia,
Rana grayi, C. du Toit, 1934 and 1933 respectively) and Bufo
(Schoonees, 1.930), the bones are normally disposed.
IV. The Middle Ear Region
In sectional views the relationship of the pterygoid and squamosal
to the middle ear and to the sound conducting apparatus is clearly made
out. I shall now describe the sound conducting apparatus in brief. This
consists of columella and stapes. According to Gaupp, the columella is
Some Aspects of the Anatomy of Anura (Amphibia} A Review 61
composed of three parts ; a cartilaginous pars exterua plectri (extra-
stapedial of Parker) which abuts on the internal aspect of the tympanic
membrane ; a pars media plectri (mediostapedial of Parker) which is bony
and an internal pars interim plectri (interstapedial of Parker) which plugs
the anterior region of the f enestra where the cartilaginous operculum (stapes
of Parker) closes or fits into the foramen ovale. In the Ranid forms (Gattpp)
from the pars exterua plectri arises a dorsal process (commonly met with in
Reptilia) called the processus or pars ascendens plectri or laterohyal
(suprastapedial of Parker) which gains attachment with the crista parotica.
The tympanic membrane or tympanum, a modified region of the skin in
that region is held taut by means of an annulus tynipanicus cartilage. In
some examples, the tympanum is ' hidden ' being covered over by the
unmodified skin. In the Microhylid examples studied by me, the tympanum
is not externally visible or in other words the skin in this region is not
modified to cover the tympanum. By ' hidden' we mean, therefore, the
condition noted above. Giinther, as early as 1858, described in detail the
classification of Anurous Batrachians depending mainly on external and
readily ascertainable characters, in which the question of " hidden tym-
panum. " as described by Prashad (1.91.8) and Boulenger (1920) is also dis-
cussed. Mivart (1869) in a paper " On the classification of Anurous
Batrachia " utilises certain osteological characters some of which are drawn
from Cope's studies. With regard to the ear, Mivart points out that he
agrees with Giinther, in that " the Batrachians with imperfectly developed
ear would form together an unnatural group and would be separated too
far from other allied forms". He divides, therefore, Anura, with reference
to teeth, tongue and perfect or imperfect ear. Baini Prashad (1918) writing
on the middle ear of Anura, introduced a new terminology, and on looking
through his bibliography list, I find no reference to Gunther (1858) and
Mivart (1869). According to him, the ' tympanic area ' is the name given
"to the area of skin situated on the temporal patch on the side of the head.
This area is continuous with the skin. . . . " Further, the tympanic membrane
"is quite a distinct structure lying immediately underneath the so-called
tympanic membrane of authors. It can be easily separated from the skin
covering it". It is obvious that it is unnecessary to use this fresh terminology,
for Boulenger (1897, 1920) after referring to the previous workers, has
remarked that, 'The tympanum, or drum of the ear, is absent in Bowibinator
and Pelobates. When present it may be concealed under the skin, as in
some specimens of Discoglossus, Pelodytes and Bufo vulgaris, or appear on
the temple behind the eye as a round or oval disc covered with thin skin '.
Rightly does de Villiers (1930 a, p. 689) point out in this connection that
B4
62 L. S, Raniaswami
" A ' hidden tympanum ,' is in any case a dangerous cliche, for it is never
anything but hidden, whether the superficial ectoderm is thin and trans-
parent or thick and undifferentiated. "
Now, with regard to the plectrum, it is noticed that in none of the
Microhylid examples studied is a suprastapedial process developed. Fur-
ther, the pars externa plectri expands into an oval cartilage on the inner
aspect of the skin within the annulus tympanicus and on account of its
expanded nature and its suspension from the end of the pars externa, it is
not like the Ranid pars externa plectri, where the pars media attaches
itself to the middle of the pars externa. This cartilage is therefore, composed
of pars externa plectri plus the extraplectral cartilage. Whatever this dif-
ference may be, the plectral apparatus of the Anura can be homologised with
the similar one noticed in Reptiles (Abel, 1929); if this view is not accepted
by all, then there is no point in calling the cartilage extrastapedial, for the
whole pectral apparatus is external to the operculum. Therefore, I have
followed the terminology used by de Villiers and called it the " extra-
plectral " cartilage. A brief reference may be made to the operculum.
According to Versluys (1924) this structure is developed in response to the
needs of a terrestrial life. In the terrestrial forms like U. systoma, M. ornata,
K. pulchra (K. p. taprobanica) and Glyphoglossus molossus, the operculum
is well developed and possesses a knob on the exterior aspect for the attach-
ment of an opercular muscle. An operculum is also noticed by me in
aquatic forms like Rana hexadactyla, R. tigrina, R. curtipes, and Rhaco-
phorid (Polypedatid) examples and in the terrestrial forms like the species
of Bufo and in the arboreal Nectophryne misera. It would, therefore, be
extremely interesting to make a comparative study of the development of
this structure in these forms and examine if the statement which Parker
(1934) makes with reference to the Microhylidas that ' the operculum is
normal in the family and retains its muscular connection with the scapula '
is applicable to the other families also.
Previous work. I have already referred to the classificatory importance
of the ear region as discussed by Mivart (1869). The occurrence
of a normal middle ear, eustachian passage and plectral apparatus
is described in Phrynomerus (de Villiers, 1930 a), Cacosternum (de
Villiers, 1.931), Anhydrophryne (de Villiers, 1931 c), Microbatrachella
(de Villiers, 1934 6), Rana grayi (C. du Toit, 1933), Crinia (C. du Toit,
1934) and Spel&ophryne (de Vos, 1935). In Hemisus (de Villiers,
1931 b) the middle ear, tympanic membrane, annulus tympanicus,
eustachian passage and plectral apparatus are absent. Parker (1934)
Some Aspects of the Anatomy of Anura (Amphibia) A Revieiv 63
also notices the same feature in Melanobatrachus, Hoplophryne (and
presumably Parhoplophryne also). Similarly in lyiopelmidse (Wag-
iier, 1934). While the plectrum and middle ear are developed in
Aglossa (de Villiers, 1932), the eustachian tubes enter the buccal
cavity by a median opening. The division of the tympanic cavity
in Probreviceps is complete while in Breviceps (de Villiers, 1 932 a) it
is incipient. Further, in the former species an opercular portion
of the M. levator scapulae superior is differentiated while the muscle
and processus opercularis are absent from Breviceps and Phrynomerus
(de Villiers, 1.931^). In this feature, these two examples differ consider-
ably from the Microhylid character enumerated above and it may be
noted here again that Parker (193-1) has created a separate family
for the accommodation of Phrynomerus. He also refers to the taxo-
nomic importance of the ear region.
V. (a] The Pterygoquadyate and Us Attachments
In the larval condition of the Anura, it is noticed that the pterygo-
quadrate gains cartilaginous attachment with the cranium in three regions
normally. The first is the quadratocranial commissure and in the majority
of cases a quadrato ethmoidal commissure is also formed and is anterior to
the quadratocranial one. The quadrato-etlmioidal commissure connects
the processus maxillaris posterior of the lamina-orbito-nasalis and the
processus pterygoideus of the quadrate. Next comes the processus
ascendens which gains attachment with the pila antotica (or the
orbital cartilage). Now, in some cases it is noticed that the pterygoquad-
rate may also give rise to a process in this region, which may articulate
with the ventral wall of the cranium ; if this palatobasal articulation
is anterior to the palatine nerve (VII), it is described as the true basal
articulation. In Anura generally, this process is posterior to the palatine
branch of the seventh cranial nerve, and therefore, it is called the
pseudobasal process and the articulation is similarly known as the pseudo-
basal articulation. InRana (de Beer, 1937), a larval pseudobasal connection
is formed and this is replaced by a pseudobasal articulation when the tadpole
metamorphoses into the adult. Now, the last connection in anuran larvae
is the processus oticus which unites the pterygoquadrate with the otic
capsule over the carnioquadrate passage. This process lies laterally to the
head vein and anterolaterally to the hyomandibular branch of facial nerve.
Exceptions to what has been described above with regard to the pseudobasal
process and the processus oticus are also found. When the tadpole under-
goes metamorphosis, the larval quadratocranial commissure is lost and is
B4A
64 L. S. Ramaswami
replaced by the quadratoethmoidal commissure ; the processus ascendent
disappears completely ; the adult oticus connection is formed by the rota-
tion of the processus muscularis and its subsequent fusion with the crista
parotica. A pseudobasal articulation is found in Rana (incompletely auto-
systylic) while in Bufo, it is completely autosystylic. With this adult arrange-
ment of the pterygoquadrate as the background, if we examine the Micro-
hylid, Rhacophorid (Polypedatid), Ranid, Bufoiiid and Pelobatid genera
studied, it is noticed that in all these, a pseudobasal process is developed.
This articulates with the subocular shelf (there being no basitrabccular
process) posteriorly to the palatine nerve and is ventral to the vena capitis
lateralis. It is obvious from a review of the previous literature on this topic,
that morphologists have followed Gaupp's nomenclature and called it the
processus basalts, and it was de Beer (1926) who pointed out the difference
between the true and pseudobasal processes. In all the forms examined by
me [Microhylidse, 1.932, 1932 a ; Rhacophoridse, 1934 (Polypdeatidae), Ranidiu
(1935), Pelobatidse (1935 a) except Bufonidse (author, 1937)] there is an
incomplete autosystyly ; however, in Rhacophonts maculafats (author, 1934,
Plate, IX, Fig. 5), there is a slight fusion of the internal end of the pseudo-
basal process with the ventral part of the otic capsule. Until the tadpoles
of this species are examined, the significance of this connection is difficult
to ascertain. At any rate, this is not like the one noticed in Bufonidie
where the entire pseudobasal process fuses with the ventral wall of the otic
capsule. An antorbital and an oticus connection are uniformly noticed
in the forms examined by me.
Previous work. A pseudobasal process is described in all the species
examined by de Villiers [except in Ascaphus (1934) where a true
basal process is present] and by C. and G. du Toit, de Vos and
Schoonees in the forms examined by them. The last author de-
scribes a complete autosystyly in Bufo angusticeps. In Hemisus
(de Villiers, 1931 6), the absence of a " basal " process is noted.
V. (b) Arteria Carotis Inter na
In describing the orbito-temporal and posterior region of the cranium,
a brief reference may be made to the disposition of the arteria carotis interim.
In the larval Anura (Gaupp, 1893; de Beer, 1937) it is noticed that the carotid
artery becomes intramural after entering through the carotid foramen.
Here it gives rise to the ophthalmica magna artery which gains exit through
the metoptic foramen and the anterior and posterior cerebral arteries. The
intracranial palatine artery which is also given off by the carotid gets out
through the craniopalatine foramen. Now, when the tadpole undergoes
Some Aspects of the Anatomy of Anura (Amphibia) A Review 65
metomorphosis, the trabecular portion separating the metoptic foramen
and the carotid artery breaks down, so much so that the vessels (ophthalmica
magna and internal carotid arteries) look as though they enter the cranium
through the oculomotor foramen. An anomalous condition, analogous to,
though different from this is also noticed in some fishes like Atniuvus, etc.
In the Ranid forms examined by me, a similar disposition is also seen and
I am examining the larvae of these and also of the other genera to note if
this phenomenon also occurs.
VI. Bursa angular is or is (Fuchs, 193.1) or the Mundwinkeldruse
The occurrence of this gland (?) was first described under the name of
' Mundwinkeldruse ' by de Villiers in Anhydrophryne (1931c). It was later
found to be present in some Amniote examples also and since the name
Mundwinkeldruse was preoccupied, Fuchs (1931) introduced the phrase
' Bursa angularis oris ' to describe it. This is a lymphocytic accumula-
tion occurring in the angle of the mouth of frogs between the maxillary
and pterygoid bones. In my study of this gland (1933 a), I have noticed that
it uniformly occurs in all the four Microhylid examples, U. systoma, M.
ornata, K. pitlchra (K. p. taprobanica) and Glyphoglossus molossus ; further,
it is also present in the Ranid and Rhacophorid (Polypedatid) genera
examined. It is, however, wanting in Pelobatid and Bufonid examples
studied by me. I have remarked that the exact function of the gland is
not yet known and Mtiller (1932) has gone to the extent of questioning the
glandular nature of it. The gland is ill-developed i-n Glyphoglossus molossus
(young specimen) but well-formed in Uperodon systomci, M. ornata, K. pulchra
(K. p. taprobanica) and Rhacophorus maculatus. At" the time I reported,
I suggested that the probable line of evolution may have proceeded with
Glyphoglossus as the starting point and the type seen in Rhacophorus as
having taken its origin and proceeded on one side while that seen in Kaloula>
Microhyla and Uperodon, on the other. This, according to me, is intended
mainly to give an idea of the interrelationship of the Microhylid examples
investigated by me. At the same time, I have cautioned that this is of no
phyletic significance and particularly when we know that Breviceps fuscus
lacks one and de Villiers (1933 a) himself is not quite sure if the other species
of the genus Breviceps are devoid of it. This should not cause surprise for
B. fuscus exhibits a series of peculiar features, as e.g., the reduced
superior prenasal cartilage, absence of palatine, sphenethmoid, qudrato-
jugal bones and of an opercular muscle ; the fusion of the cartilago obliqua
posteriorly with the septum, the fusion of the pterygoid and squamosal
bones and the absence of a " basal " process.
66 Lo S* Ramaswami
Previous work. The gland lias been described In Anhydrophryne (tic
Villiers, 1931 c), Probreviceps (1.932$), Rhombophryne (de Villiers,
1934 a), Spelaophryne (de Vos, 1935), Rana grayi (C. du Toit, 1933)
and Crinia (C. du Toit, 1934). In Phrynomerus, Caco sternum and
Hemisus, de Villiers does not mention about the gland while in
Breviceps fuscus, de Villiers (1.931 d) notes the absence of the gland
even though it is closely related to Probreviceps (de Villiers, 1932**).
Schoonees (1930) notes its absence in B. angusticeps.
VII. The Lower Jaw
The lower jaw is characterised uniformly in the Auura by the posses-
sion of the two membrane bones, viz., the angular and the dentary and a
single cartilage bone the mentomandibular. The membrane bones are
investments of Meckel's cartilage. The dentary is only met with in the
anterior sections of the lower jaw. In the Microhylid examples studied, viz.,
U. systoma, M. ornata, K. pulehra (K. p. taprobanica) and G. molosstts, there
is a Meckelian epiphysis which projects postero-internally (author, 1932 b)
parallel to the jaw on either side. The presence of this was first noticed by
Devanesan (1922) in U. systoma. In the foreign Phrynomeridae and Micro-
hylidse studied by de Villiers, its occurrence is also noticed (Phrynomerus
1930 a, Breviceps 1.931 d\ Probreviceps 1932 a, Rhombophryne 1934 a, and
by de Vos in Spelaophryne, 1935). In Caeosternum and Hemisus (de Villiers,
1931. and 1931 b respectively) it is absent and therefore, they do not show
this Microhylid affinity. It may be pointed out here that in the Ranid,
Rhacophorid (Polypedatid) and Bufonid examples studied by me, the
epiphysis is absent. Further, de Villiers (1934 b) discovered the absence of
it in Microbatrachella, and therefore, excluded it from the Microhylidse ; he
notes (p. 414) " I agree with Ramaswami (1932 b) that ' it is a lateral epi-
physis of Meckel's cartilage uniformly occurring in the Engystomatidrc '.
The absence of this process in Mierobatrachella definitely excludes the genus
from Brevicepitidse ". From the same point of view, Caeosternum and
Hemisus cannot be included under Microhylid^ and rightly does Parker
(1934) omit these two from the Microhylid family. Thus, this character
of the possession of Meckelian epiphysis will be very useful in discussing
the affinities of the Microhylid genera.
VIII. The Hyolaryngeal Apparatus of Microhylid^ and Pelobatidat
As already pointed out, I have studied the sectional views of the
larynges of both sexes of U. systoma, M. ornata and K. pulehra (K. p. tapro-
banica) and the gross anatomy of the larynges of various Ranid and Rhaco-
phorid (Polypedatid) species. With regard to the Microhylid hyolaryngeal
Some Aspects of the Anatomy of Anura (Amphibia) A Review 67
apparatus, I have recorded certain interesting features, everyone of which,
has been confirmed by the later worker Trewavas (1933) and in a personal
communication, she has stated that my paper (author, 1932 b) was unfortu-
nately not available to her at the time. I have also studied the gross ana-
tomy (1935 b) of the hyolaryngeal apparatus of Pelobatidse, and noted that
there is considerable variation with regard to the cricoid element. With
regard to the hyoid apparatus, I have shown that a demarcation line could
be drawn between the subfamilies Megophrynse and Pelobatinse. In the
case of the Megophryne hyoid apparatus, a lateral foramen is absent while
in the Pelobatinae, it is well developed. There is, however, one exception.
In the case of Megophrys fees (Beddard, 1911) the occurrence of a lateral
foramen is noted, and this escaped my attention when I wrote my paper,
Further, in the case of Pelobates fuscus, W. K. Parker (1881) does not depict
a lateral foramen, though according to the observation made above,
the hyoid of Pelobatinae must possess a lateral foramen. Probably, Parker's
delineation is incorrect, or it is an individual variation, for it is noticed that
in his paper on the development of the hyoid apparatus of Pelodytes, Ride-
wood (1897, Fig. 12) draws, for comparison, a figure of the hyoid of Pelobates
fuscus, where a lateral foramen is clearly shown. Thus, so far as is known
to me, the Pelobatine hyoid apparatus possesses a lateral foramen while
in the case of Megophrys (except M. fece] the lateral foramen is absent and
M. fe& may be treated as a connecting link between the two.
Now, with regard to the hyolaryngeal apparatus of the Microhylid
examples studied (author, 1932 b), the following important points are
noted :
(1) The hyoid plate has a cartilaginous or bony, beak-like portion
between the postero-medial processes. This feature was already
noticed by W. K. Parker (1881) in Callula pulchra and two species
of Microhyla which he described under the name of Diplopelma,
D. ornatum vel rubram, and D. Berdmorei (?). It is rather unfortu-
nate that neither Trewavas (1933) nor myself referred to Kaloula
and Microhyla, though Trewavas did refer to Gastrophryne of
Parker (1.881). With regard to Microhyla, the difference in nomen-
clature is responsible.
(2) An extra-hyal associated with the anterior cornu is well represented.
Parker (1881) does not draw the occurrence of an extra-hyal in his
specimen of Callula pulchra. Trewavas (1933) notes its presence.
In Microhyla okinavensis, an extra-hyal is absent according to
Frazier (1924).
68 L* S* Ramaswami
(3) A cartilage of Santorini (cartilago apicalis Gaupp) is absent.
(4) The cricoid annulus does not possess an oesophageal process (spina
oesophagea Gaupp). This process is commonly met with in the
female specimens of the Ranid species studied by me. However,
in only two examples of Ranids, viz., Rand tigrina and R. breviceps,
the male may also possess this process though this feature is not
uniformly noticed. Therefore, this is an erratic variation and no
importance need be attached to it. Thus, since the oesophageal
process was noticed in all the Ranid females examined by me, it
may well be used as a sex determiner.
(5) The broncheal processes are long and embrace the root of the lung
as expanded plates.
And to these characters Trewavas (1933) adds five more, four of which
are with regard to the musculature of the hyolaryiigeal apparatus and the
other with reference to the postero-medial processes of the hyoid apparatus.
These characters taken together exclusively distinguish the Microhylid
hyolaryiigeal apparatus from that of the others.
I have mentioned on p. 40 of my paper (1932 6) that Microhyla and
Kaloula are devoid of a vocal sac, an observation made on insufficient
material. I have since examined the male members and have discovered
the presence of a small vocal sac, and as early as 1882, Boulenger reported
the occurrence of a vocal sac in the South Indian Microhylid genera.
Parker (1934), who has described these forms, also notices the presence of
vocal sac in Kaloula and Microhyla. In Uperodon systoma, Devanesan
(1922) also draws attention to the occurrence of a large vocal sac.
Previous work. Trewavas (1933) gives us an exhaustive account of
previous workers on this subject. However, she does not refer to
some of Blume's papers, and they are W. Blume (1931, 1932 and
1933).
IX. The Vertebral Column
The examination of the vertebral centrum in the Ranid and Rhaco-
phorid (Polypedatid) families has revealed some important variations. At
the time of writing my paper (1933), I was not able to secure Nicholls' paper
(1915-16) which deals with the classificatory importance of the vertebral
column in Anura. I have noted that in the case of Rhacophorus (Poly-
pedates) dubius and R. microtympanum, the 8th vertebra is procoelous, while
in R. maculattts and R. eqites it is diplasiocoelous, a term invented to
represent the double concave nature of the centrum by Boulenger. I have
Some Aspects of the Anatomy of Amir a (Amphibia) A Review 69
confirmed Nicholls' observation (1915-16) of the diplasiocoelous nature of
the centrum in Rhacophorus 'inaculatus. Mookerjee's observation (1932)
on Rhacophorus maximus was already articipated by Nicholls. And with
regard, to the genus Rhacophorus (Polypedates], I have stated (author, 1933)
that so far as the characters of the 8th and 9th vertebrae are concerned,
it could be split into two. Under the proccelous Rhacophoridse (Poly-
pedatidoc), we have to treat probably the procoelous species of Philautus
examined (Philautus chalazodes, P. nasutus and P. oxyrhynchus) and if
more species are discovered with pro- or diplasiocoelous type of centrum of
the 8th vertebra, then, this would be another feature, besides the prevomerine
teeth and sphenethmoid, to support Noble's view that Philautus is not a
single distinct genus but a polyphyletic assemblage. In passing, it may be
noted, that from the view-point of cranial anatomy, there is a vast body of
evidence disproving Noble's dictum that Philautus does not represent a
natural genus. In the family Ranidse, a large number of species exhibits the
diplasioccelous 8th vertebra. With regard to the genus Micrixalus, I have
stated (author, 1933) that in the possession of diplasiocoelous (incorrectly
stated as precocious, p. 1, column 2, line 27) 8th vertebra, " Micrixahis is
Ranid in every respect 1 '. The three species of Nyctibatrachus, the majority
of the species of Rana (except Rand curtipes Jerd., and R. tenuilingita
Rao) examined by me are diplasiocoelous. In R. curtipes, however,
the 8th and 9th vertebrae are fused and there is a single centrum and the
zygapophyses are a pair in number (anterior). The transverse processes are
Ranid in character, but in one specimen of R. curtipes, the right transverse
process of the 9th vertebra is absent and the ilium gains attachment
with the transverse process of the 8th vertebra. The fusion of the 8th
and 9th vertebrie is noticed in all the specimens of R. curtipes examined by
me, so much so that it is a distinguishing character of this species and
the position of this with reference to the other genera where .fusion of the
vertebral elements occurs, is discussed in my paper (author, 1933).
When Nicholls discovered the diversity in the vertebral column of the
genera Rana and. Rhacophorus of the family Ranidae [the Rhacophoridae
(Polypedatidae) being included under it], he noticed the difficulty of including
the genus Rhacophorus under the family Ranidse. He says, "The genus is one
in which Boulenger has merged the genus Polypedates, and it is represented
in the British Museum collection of skeletons by nine specimens. Of
these, four belonging to the species JR. maculatus, R. cruciger, R. macwtis
and R. robustus were diplasiocoelus. The remaining specimens R. maximus,
R. madagascariensis, R. schlegelii and R. reinwardtii were uniformly
procoelous". Noble (1931, p. 514) has treated the suborder diplasiocoela
'0 L. S* Ramaswami
(the true frogs, Raiiids ; old world tree frogs, polypedatids and narrow-
mouthed toads, brevicipitids) as comprising three families, viz., Ranidse,
Rhacophoridae (Polypedatidse) and Brevicipitidse. The Rhacophoridae
(Polypedatidse) are differentiated from the Ranids by the presence of an
intercalary cartilage between the distal and penultimate phalanges. Under
the subfamily Ratlines come the species of R&na, Nyct^batrach^ls and Nanno-
balrachus and under the subfamily Cornuferinse, is treated Micrixalus.
With regard to the two genera Philautus (Ixalus) and Micrixalus, Noble
(1931) states that " Philautus has arisen from Polypedates in many parts
of its range by the ofts repeated process, a loss of vomerine teeth " and
treats this under the family Rhacophoridse (Polypedatidse). I have shown
elsewhere (author, 1934) that cranial anatomy does not support the merging
of Philmttits with Rhacophorus except when the dentition of the prevomers,
the spheuethnioid and the nature of the vertebral centra of 8th and 9th
are taken into consideration (see p. 69). Micrixalus is described by
Noble (1931) as a group of small species of Hyla-mna lacking vomerine teeth.
Thus both these genera Micri-xalus and Philautus while possessing digital
expansions lack vomerine teeth ; but Philautus is a Rhacophorid (Poly-
pedatid) with intercalary phalanges, and Micrixalus a Ranid without them.
Therefore, the view " that the separation of Micrixalus from Ixalus is based
on arbitrary grounds and possibly when a large number of species is examined
the diagnostic characters of the two genera may be found to be too
slender for erecting two genera for their reception " (author, 1933) is not
tenable.
IX. The Vertebral Column of Megophrys major (author, 1935 a)
In the preparation that I have of the vertebral column of M. major,
the 1st and 2nd vertebrae are fused, and therefore, I have stated that it
differs from the ancestral form, the lyiopelmidae. The ancestral forms
possess ten vertebras and the first forms of Pelobatidae to be derived from
these arc the Megalophrynse (Megophrynse) and one of the species of this
group M. major examined by me, therefore, differs from the ancestral
stock in the possession of only 8 vertebrae. I have now examined another
specimen of the same species and have discovered that the 1st and 2nd
vertebra are not fused, and therefore, the normal form differs from the
ancestral stock in possessing 9 vertebrae. The one with fused 1st and
2nd vertebrae shows therefore, a variation. Further, in my 1st specimen,
the coccyx is partially united with the sacrum though Boulenger finds it
immovably united in his specimen of M. major. Parker, after an examina-
tion of a large number of species of Megalophrys including M. major,
Some Aspects of the Anatomy of Anura (Amphibia) A Revieiv 71
informs me (in litt.) that in normal cases of M. major, a small subcircular
cartilaginous omosternum is present. He also points out that in his specimen
of M. major, the coccyx is incompletely fused with the sacrum as in mine
and unlike that of Boulenger's. Probably Boulenger failed to see the non-
fusion.
In discussing the ancestry of Pelobatidse, I have followed the observa-
tions of Noble (1924, 1931). According to him, the I/iopelmidse are the
most primitive Anurans. This group has given rise on the one hand to the
Discoglossidse and on the other to the Pelobatidae, and cranial morphology
supports this origin of the Pelobatidse from the Liopelmid stock in 4 features.
They are the absence of a recessus saccif ormis, of a Bursa angularis oris and
the presence of a septomaxillary and the fusion of the mentomandibular
with the dentary. It is true that any one of these features may be found
in any other Anuran family but when we take all these four features
together, Noble's hypothesis is amply borne out by cranial anatomical
studies.
X. Summary and Conclusions
It has been known from a long time that Anuran families cannot be
distinguished systematically from one another by a single character but by
a set of features. With this in mind, we shall now proceed to examine the
various features that the cranial morphological investigations have revealed :
1. The cartilago prenasalis superior and inferior are uniformly met
with in the Anura. Exceptions, however, are very few where either
the superior is reduced (Breviceps fuscus) or the inferior cartilage is absent
(Hemisus). Thus, this character is of no diagnostic importance.
2. The cartilago obliqua is a lateral extension in the anterior narial
region from the tectum and from this a connective tissue projection the
plica obliqua depends into the cavum. Gaupp who studied Rana fusca
(S. temporaria] noticed that it depended from the cartilaginous roof and not
from the cartilago obliqua ''and the same feature as reported for Rana
fusca is also seen in Breviceps fuscus. Now, in the Rhacophorid
(Polypedatid) forms examined by me, it is noticed that in three species of
Philautus, the plica depends from the tectum, while in the species of
Rhacophorus (Polypedates), it depends from the cartilago obliqua. In the
Ranids, Buf onids and Pelobatids examined by me and in the forms studied
by the South African anatomists (with the exception mentioned on pp. 46
and 47), the plica depends from the cartilago obliqua. While the nature of
the dependence of the plica cannot be made use of for classificatory purposes
in other families, at any rate in the Rhacophoridae (Polypedatidse), it
72 L. S- Ramaswarni
appears to be useful in the present state of our knowledge, in distinguishing
Philautus from Rhacophorus (Polypedates).
3. The prechoanal sac which may occur as two sacs in the anterior
region into which the choanse open or as a single sac (Rhacophoms macu-
latus and Megophrys major) or as a vestigial structure (Uperodon systotna)
into which the choanae do not open, is also noticed in other forms like
Phrynomenis (double), Probreviceps, Breviceps, Spel&ophryne, Rhombo-
phryne and Rana grayi. It is absent from Glyphoglossus molossus, examined
species of Rhacophoms (except R. maculatus) and Philautus t Bufo, and from
Scaphiopus holbrookii and Nectophryne miser a. Since the occurrence of
this is noticed in genera, of different families, it is probably of independent
origin and does not show, therefore, any relationship. Obviously, this
cannot be used in systematic study.
4. The septum nasi which is either cartilaginous or is posteriorly
ossified gives rise to the tectum or the roof and the solum or the floor. The
solum is noticed to give rise to an elevated eminentia in a large number
of forms comprising Microhylidse, Ranidse, Bufonidse and Pelobatidae. It
was theorised by the South African anatomists that the occurrence of an
elevated eminentia was closely correlated with a terrestrial mode of life.
This is true in the case of forms like Uperodon systoma, Microhyla ornata,
Kaloula pulchra (K. p. taprobanica) , Glyphoglossus molossus, Breviceps,
Probreviceps, Spel&ophryne and other forms like Bufo and Hemisus, The
appearance of this structure in different forms can only be explained as due
to independent development. If only the terrestrial forms developed this
structure, of course the theory advanced would not have been found fault
with. In two typically aquatic forms of South India, Rana hexadactyla
and Rana cyanophlyctis, an elevated eminentia is met with, and it is there-
fore thought, that the elevation is purely in response to the sensory needs
of the animal. Whatever this aspect of the question may be, it is interest-
ing to note that the Microhylid genera so far studied, viz., Microhyla, Uperodon,
Kaloula, Ramanella, Breviceps, Probreviceps, Spelceophryne and Rhombo-
phryne (?), the elevated eminentia uniformly occurs and this character
therefore, can be utilised along with other features in distinguishing the
Microhylids.
5. The sphenethmoid bone : It has been noticed that in some examples
this bone is divided by means of a trabecular cartilage into a right and left
portions when the bone is described as paired. A complete sphenethmoid
is seen in Rhacophorus maculatus, Philautus petersi, P. o%yrhynchus y Rana
cyanophlyctis, R. hexadactyla, Megophrys major and Scaphiopus holbrookii
Some Aspects of the Anatomy of Anura {Amphibia) A Reviezv 73
while in U. systoma, K. pulchra (K. p. taprobanica], M. ornata, G. molossus,
Rhacophorus microtympamim, Philautits chalazodes, Phrynoments and
Rhombophryne it is paired ; in some, it may be absent like Spelceophryne,
Ascaphus and Rana curtipes, etc. We find that in the case of all Microhylid
examples so far studied, it is either paired or is absent (Spelaophryne,
Breviceps, Probreviceps, Kalophrynus, Callulina, etc). In the Ranidae, it
may be absent or it may be single or paired [though Parker (1934) says that
the ethmoid is single] and we meet with the same state of affairs in Rhaco-
phoridse (Polypedatidse). Hence, this character of the paired nature of the
sphenethmoid can be conveniently utilised with others in describing the
family characters of the Microhylidae, till we find other exceptions. With
regard to the Ranidse and Rhacophoridas (Polypedatidse), the sphenethmoid
can be described as either single or paired or absent (some Ranidse). Mord
examples should be studied before we can come to any conclusion with
regard to the sphenethmoid of the Bufonid and Pelobatid families.
6. The maxillae and premaxillse are not of great importance to us in
systematic study of the genera examined by me.
7. The prevomer, in the Ranid and Rhacophorid (Polypedatid)
families does not embrace the choana posteriorly and is not flat. This
observation is supported by Rhacophorus maculatus, R. microtympamim,
Philautus chalazodes, P. oxyrhynchus, P. peter si and Rana hexadactyla,
R. ciirtipes and R. cyanophlyctis. In the Microhylid forms studied by me,
U. systoma exhibits a well-developed prevomer and a degenerate palatine
(?) ; in M. ornata, the choana is not embraced posteriorly by the prevomer,
while in K. pulchra (K. p. taprobanica}, the posterior portion of the prevomer
overlies the palatine. In G. molossus, there is a postclioanal portion
(prevomer or prevonieropalatine) . Thus, in the Microhylid family, we meet
with conditions where the choana may be completely or incompletely
(Kaloula, Glyphogloss^ts } Rhombophryne, Spelceophryne, Breviceps, Pro-
breviceps) or may not be surrounded by the prevomer as in Microhyla.
Therefore, the nature of the prevomer is not only useful in distinguishing
the family but also in classifying the genera within it. Whether the state-
ment can be applied to the Bufonidce and PelobatidcC can only be settled
after examining some more genera.
8. The palatine bone is not so useful as the prevomer. This may be
degenerate (U. systoma 1 for the bone may also be described as a postchoanal
prevomer), or absent (M. ornata} or a pre-vomeropalatine arrangement
may be seen. Generally in the Rani das, Rhacophoridse (Polypedatidse),
Bufonidae and Pelobatidae, the bone is present ; however exceptions are not
74 L* S, Ramaswami
uncommon. Nectophryne misera and Scaphiopus holbrookii can be men-
tioned as Instances. Among the several species of Microhyla, some possess
it, while others do not. Thus, the nature or the disposition of this bone
cannot be utilised in taxonomy.
9. The septomaxilla is noticed to occur uniformly in all the forms
examined by me. This is not, therefore, of great systematic importance
except in one case. In Bufonidse, a limb of the septomaxilla appears in
the plica, a feature in which this family stands apart from the others. I am
aware of an exception to this ; in Bombinator, B rimer describes a limb of
the septomaxilla in the plica. Barring this exception, it will be found
useful to introduce this feature of the septomaxilla in describing the family
characters of the Buf onidae.
10. The middle ear and associated structures : It is very well known
that in widely different forms, the disappearance of the middle ear, eusta-
chian passage, tympanum and plectral apparatus may occur, as in Liopelma,
Ascaphus, Hemisus, Pelobates, etc. This may not establish any genetic
affinity and therefore, the disappearance of these structures fully or other-
wise should be considered secondary. Overlooking these genera, there is a
common plan on which the middle ear region with its associated structures
is built. The attachment of the columella (pars media plectri) to the dorsal
rim of the extraplectral (pars externa plectri plus extraplectral) cartilage
in the Microhylid examples examined by me, associated with a sickle-shaped
annulus tympanicus, is certainly different from the median attachment of
the columella with the pars externa noticed in Ranidse, Rhacophoridse
(Polypedatidse) and Pelobatidse. Since this feature is not uniformly
noticed in the other species of the Microhylid family, it cannot be used as a
safe criterion. The pars ascendents plectri deserves to be mentioned.
Gaupp described this commissural cartilage (laterohyal) in "Rana, but this has
not been confirmed by me in all the species of Rana examined or in the
Rhacophorid (Polypedatid) species studied. In Bufonid species (except
B-ufo vulgaris), a laterohyal is noticed, and this internal character can there-
fore be utilised in describing the family characters. It may be said that
Bufonids generally posses a laterohyal.
1L The pseudobasal joint: In the Anura, the pseudobasal joint
between the subocular shelf and the pseudobasal process (except in
Ascaphus} of the pterygoquadrate is common. In the Ranid, Rhaco-
phorid (Polypedatid) and Pelobatid families, the pseudobasal articulation
is found while in Bufonid species studied, there is a definite pseudobasal
connection. This is a distinguishing feature of the Buf onidse. Whether it is
Some Aspects of the Anatomy of Anura (Amphibia) A Review 75
an articulation or a connection it always lies posterior to the palatine
branch of the facial nerve and is ventral to the head vein. In describing
the family characters of Bufonidae, the possession of a complete
autosystyly should also be included.
12. The Bursa angularis oris or Muiidwinkeldruse is noticed in Micro-
liylidse (except Breviceps fuscus), Ranidse and Rhacophoridse (Poly-
pedatidae) and is generally absent from Bufonidse and Pelobatidse. In
describing, therefore, the family characters of the first three, the presence
of the Bursa may also be included.
13. The occurrence of a bony or cartilaginous beak between the
posteromedial processess of the hyoid (called basi-branchial by Parker, 1881)
is uniformly noticed in the South Indian Microhylid examples, but in the
foreign forms Breviceps, Probreviceps, Spelceophryne and Rhombophryne,
this is not present; all the same, Trewavas considers this as a distinguishing
character of the Gastrophryne group of the Brevicipitidse. This feature
is of no great systematic value in general but is of sufficient importance to
demarcate the examined Indian genera. To this may be added, the absence
of an omohyoid and the presence of only two petrohyoideus muscles
(Trewavas) and these three characters are very useful in distinguishing the
Microhylidae. The larynx of the Microhylidse exhibits two important
diagnostic features, viz., the absence of an independent apical cartilage and
of an cesophageal process from the cricoid annulus of both sexes. With
regard to the Pelobatidse, the presence of a lateral foramen in the Pelobatinae
would distinguish the Megophrynae (except M. fea t where also a lateral
foramen is present). Following Trewavas, the diagnostic features of the
Pelobatid family would be as follows :
(a) The occurrence of an arytenoid without an apical cartilage,
(6) cricoid ring incomplete dorsally [except in Scaphiopus hammondii
(male) where it is complete] (author, 1935 a), and
(c) hyalia more or less reduced.
H. The occurrence of a Meckelian epiphysis is so characteristic of
Microhylidse that it ought to be included as a taxonomic feature. How-
ever, we note the occurrence of a Meckelian epiphysis in Phryonomerus,
which has been treated as belonging to the subfamily Phrynomerinae under
Brevicipitidse by Noble (1931). Besides, Phrynomerus also shows other
Microhylid affinities, as dilated sacral diaphophysis, prechoanal prevomer
(Parker, 1934) as in Microhyla (but de Villiers (1930 a) describes the formation
of a prevomeropalatine in (Phyrnomerus) , divided ethmoid, etc. In spite
of all these features common with the Microhylidae, Noble says that, "The
76 ' L* S. Ramaswami
African Phrynomenis is not closely related to any other brevicipitid ".
Whatever may be the number of similarities between this genus and the
other genera of BrevicipiticUe of Noble, there is one important distinguishing
feature and that is, the presence of intercalary phalanges. No Microhylid
possesses these, and therefore, Parker (1934) has erected a new family
Phrynomeridae for accommodating this genus.
15. The nature of the centrum of the vertebrae has been utilised
largely in taxonomy. Within the family Ranidne and Rliacophoridju
(Polypedatidse), I have shown that both the procoolous and diplasioctrlous
type of centrum for the 8th vertebra occur. Therefore, in distinguishing
families this may not be a safe criterion, and it must be said that the pro-
priety of using the nature of the vertebra has been questioned from the time
of Gadow since the variations noticed have been largely fortuitous. At any
rate, I have no hesitation in following Parker (1934) for using the nature of
the centrum in the diagnosis of groups of genera within the families.
Conclusions
The study of cranial morphology and of the hyolaryngeal apparatus
and the vertebral column reveals the following features which can safely
be utilised along with others (Parker, 1934) in the classification of Anuran
families and subfamilies :
RanidcB :
(a) Vertebral column diplasioccelous or procoelous.
(b) Ethmoid entire or paired or absent.
(c) Eminentia generally flat except in Rana hexadactyla and Rana
cyanophlyctis.
(d) A Bursa angularis oris is present.
(e) A pseudobasal articulation is noticed.
Rhacophorida (Polypedatidce) :
(a) Vertebral column diplasiocoelous or procoelous.
(b) A Bursa angularis oris is present.
(c) A pseudobasal articulation is noticed.
Microhylida :
(a) Eminentia is invariably elevated.
(b) A Bursa angularis oris is present.
(c) A Meckelian epiphysis is present on either side in the lower jaw.
(d) An independent cartilago apicalis is absent from the arytenoid
cartilage.
(e) An cesophageal process is absent from the cricoid aimulus.
Some Aspects of the Anatomy of Anura, (Amphibia) A Review 77
Bufonidcv :
(a) Invariably a limb of the septomaxilla is seen in the plica obliqua.
(b) A Bursa angularis oris is absent.
(c) A pseudobasal connection is present.
(d) A laterohyal is invariably present.
Pelobatida :
(a) A Bursa angularis oris is absent.
(b) A pseudobasal articulation is present.
(c) The cricoid annulus is complete or incomplete.
(d) The liyoid of the subfamily Pelobatinae possesses a lateral foramen
while in Megophryme, it is wanting except in M. fece.
XL A cknowledgetnent
I must acknowledge my indebtedness to Professor C. R. Narayan
Rao, under whose guidance, the work embodied in the several papers, which
forms the basis of the present review was carried out from time to time.
I must also express my sense of gratitude to Professor E. S. Goodrich, M.A.,
F.R.S., of Oxford, to Dr. G. R. de Beer, D.SC., of L,ondon and to Mr. H. W.
Parker of the Birtisli Museum, for their constant guidance and help with
literature. My thanks are also due to Profesor A. Subba Rau, D.SC., for
helpful criticisms.
BIBLIOGRAPHY
Abel, O. . . " Stamrnengeschichte und Palaobiologie", 1929, Fischer
& Co., Jena.
Baini Prashad . . Rcc. hid. Mm., 1918, 25, Part III, 97.
Beddard, F. E. . . Proc. Zool. Soc. Land., 1911, p. 393.
de Beer, G. B. . . Quart. J. Micr. ScL* 1926, 70, 263 ;
" The development of the Vertebrate Skull," 1937,
Clarendon Press, Oxford.
Blumc, W, Ger/en. Morph. Jahrb., 1931, 65, 0(35.
Ibid., 1932, 71, 171.
Ibid., 1933, 72, 103.
Born, G. Gey en. Morph. Jahrb., 187(3, 2.
Boulenger, G. A. . Cat. .Bat. Sal., 2nd edition, London, 1882.
Fauna of Brit. Ind. (Reptiles and Batrachia), 1890.
*" The Tailless Batrachians of Europe," Ray Society,
1897.
Rcc. Ind. Mu8. 9 1920 20, 1.
Brock, G. T. Anat. Am., 1935, 80, Nr. 13/10.
Bruner, H. Z. . - Geyen. Morph. Jahrb., 1902, 29.
Devanesan, D. W. . . Proc. Zool. Soc. Land., 1922, 527.
Ecke, H. Z- Morph. Okol. Tiere., 1935, 29, 79.
Frazier, M. . . J. Morph. & Phy., 1924, 39, 285.
"8
L, S. Ramaswami
Gadow, II.
Gaupp, E.
Giiather, A.
Goodrich, E. S.
Goto, S.
Howes, G. B.
van Kampen, P. N.
Kingsbury, B. F., and Reed,
H. D.
Lapage, E. O.
Mahendra, B. C.
Mivart, St. G.
Mookerjee, H. K.
MiiUer, F.
Nicholls, G. E.
Noble, G. K.
-, and Parker, H. W.
Parker, H. W.
Parker, W. K.
Rao, C. R. Narayan
Ramaswami, L. S.
Cam. Nat. Hist.," Reptiles and Amphibia," Maemillan
& Co., London, 1901.
*Morph. Arb., 1893, 2.
Ibid., 1893, 3.
*Cat. Bat. Sal. Brit. Mus., London, 1858.
Proc. Zool. Soc. Lond., 1858, 339.
" Structure and Development of Vertebrates," Mac-
millan & Co., London, 1930.
Annotationes Zoolof/iccv Jap., 1906, 5, Part V, 267.
Proc. Zool. Soc. Land., 1891, 148.
" The Amphibia of the Indo- Australian Archipelago,"
Leiden, 1923.
J. Morph., 1909, 20, Nr. 4, 549.
J. Morph., 1928, 46, Nr. 2. 399.
Curr. Sci., 1936, 4, 744.
Ibid., 1937, 6, 13.
Proc. Zool. Soc. Lond., 1869, 280.
Curr. Sci., 1932, 1, 165.
Getjen. Morph. Jdhrb., 1932, 70, 131.
Proc. Linn. Soc. Lond., Session 128, 1915-16, 80.
Bui. Amer. Mus. Nat. Hist., 1922, 46, 1.
Amer. Mus. Novit., 1925, 165.
Ann. N.Y. Acad. Sci., 1927, 30, 31.
Bull. Amer. Mus. Nat. Hist., 1929, 58, 291.
" The Biology of Amphibia, " McGraw-Hill & Co., New
York, 1931.
Amer. Mus. Novit., 1926, 232.
Ann. May. Nat. Hist., 1928, Series 10, 2, 473.
*Arc7i. Zool. Torino., 1931, 16, 1239.
Ann. Mag. Nat. Hist., 1931 o, Series 10, 8.
" Frogs of the Family Microhylidse,"j Brit. Mus. Pub.,
Lond., 1934.
Phil. Trans. Roy. Soc. Lond., 1881, 172, Part III, 1.
J. Bombay Nat. Hist. Soc., 1920, 119.
Ibid., 1922, 439.
Proc. ZooZ. Soc. Lond., 1925, 587.
Proc. Ind. Acad. Sci., Bangalore, 1937, 6 B, 387.
Half-Yearly J. Mys. Univ., 1932, 6 ; 45.
Ibid., 1932 a, 6, 176.
Ibid., 1932 b, 6, 32.
Curr. Sci., 1933, 1, Nr. 10, 306.
Ibid., 1933 a, 2, Nr. 1, 7.
Proc. Ind. Acad. Sci., Bangalore, 1934, 1, 80.
Ibid., 1935, 2B, Nr. 1, 1.
Anat. Anz., 1935 a, 81, Nr. 4/6, 65.
Curr. Sci., 1935 b, 3, Nr. 7, 306.
Proc. Zool. Soc. Lond., 1937, 1137.
Some Aspects of the Anatomy of Anura (Amphibia] A Review 79
Ifcidewood, W. G.
Smith, M. A.
Hchoonees, D. A.
ctu Toit, O. A.
<lu Toit, G. P.
, and. de
Villiers, C. G. S.
Trewavas, B.
Versluys, J.
cle Villiers, 0. G. 8.
. . Proc. ZooL Soc. Loud., 1897, 577.
. . Bui. Raffles Mus., Singapore, 1930, Nr. 3.
. . 8. Afr. J. Sci., 1930, 27, 456.
. . 8. Afr. J. Sci., 1930, 27, 426.
Ibid., 1930 a, 27, 439.
Ibid., 1931, 28, 408.
Proc. Zool. Soc. Loud., 1933, 715.
Ibid., 1934, 119.
" A revision of the genus Heleophryne". Ann. Univ.
of Stellenbosch. S. Afr., 1934.
. . 8. Afr. J. Sci., 1933, 30, 394.
S. Afr. J. Sci., 1932, 29, 449.
. . Phil. Trans. Roy. Soc. Land., 1933, 222 B, 401.
. . Leerboek der vergl. ontleed. vertebrata, 2 deelen. Utrecht.,
1924.
. . 8. Afr. J. Sci., 1926, 23, 61.
Ibid., 1930, 27, 481.
Quart. J. Micr. Sci., 1930 a, 73, 667.
Ibid., 1931, 74, 275.
8. Afr. J. Sci., 1931 a, 28, 378.
Anat. Anz., 1931 6, 71, 305.
Ibid., 1931 c, 71, 331.
I&wL, 1931 <Z, 72, 164.
Ib-id., 1932, 74, 33.
Ibid., 1932 a. 75, 257.
BuL Mus. Comp. ZooL, Harvard, 1934, 67, 3.
Anat. Anz., 1934 a, 78, 295.
S. Afr. J. ScL, 1934 b, 31, 406.
.4wrf. 4ws. s 1936, 81,225.
tie Vos, 0. M. . . Anat. Anz., 1935, 80, Nr. 13/16, 241.
"Wagner, D. S. . . Anat. Anz., 1934, 79, 65.
Two more papers on Anuran Craniology have appeared since.
du Toit, C. A. Anat. Anz., 1938, 86, 388.
Hembach, W. . . Jenai. Zeitschr. Naturwiss., 1939, 72, 211.
* Papers not accessible to the author.
80 L. S. Ramaswami
ERRATA LIST
Half-yearly J. Mys. Univ., 1932, 6, Nr. 1.
Reprint Journal lAne
page page
3 34 I read "Microhyla oHnaveiisin and Cctcopus"
for "Microhyla and Cacopus"
3 47 last but one read ''like in all" for "like all".
11 55 23 read "Miss Lapage (13)" for h6 Lapago (11)".
93 67 15 read "crista parotica, which is to be seen" for
"crista parotica is to be seen".
23 67 35 read "Anura (Gaupp) is not accepted" for "Anura
(Gaupp).
Half-yearly J. Mys. Univ. 1932, 6 3 Nr. 2.
1 176 21,24 ]
4 179 22
5 ISO 10 j- read " 'de Villiers' ' fo r k " Villiers" .
6 181 28
7 182 12 J
Proc. Ind. Acad. JScL 9 1935, 2.
2 12 read "cartilago alaris" for "cartilago obliqua".
20 28 read "172" /or "162".
Proc. Ind. Acad. Sci., 1934, 1.
87 8 read "(See fig. 9)" for " (See fig. 8)".
93 34 read 61 1932" /or "1933".
94 25 read "CLXXII" for "CLXII".
Pate X. fig. 5. lettering . . . . read "U.B." for "V.B."
Proc. Zool. Soc. Lond., 1937, Part 4.
1165 7 read "B . melanostictus and B. parictalis" for
"B. melanostictus and B. pantherinus ".
1168 52 read "OLXXII" /or i CLXII".
Current Science, 1933, 1, Nr. 10.
Page 1 Column 1 Line 32 read "R. intermedlus (n. sp. Rao)" for "R. inter-
27 rc.cZ "amphicoelous (diplasiocoelous)" for
"procoelous".
10 read "a pair" /or "two pairs".
OXIDATION OF THIOLS AND ASCORBIC ACID IN
THE LATEX OF PAPAYA
BY C. V. GANAPAT'HY AND B. N. SASTRI
(From the Department, of Biochemi^tr)^ Indian Institute of Science, Bangalore}
Received July 10, 1939
IT was pointed out in an earlier paper 1 that the fresh latex of papaya (Carica
papaya, lyinti.) contains a remarkably large concentration of snlphydryl
compounds (amounting to nearly 2 per cent, calculated as glutathione)
and that practically the whole of it is in the reduced form. The actual
glutathione content of the latex is about 0-2 per cent. It has now been
found that vitamin C co-exists with glutathione and that the whole of it
is also in the reduced condition.
Considerable attention has been devoted to the elucidation of the
mechanism, present in the tissues, responsible for maintaining glutathione in
the SH-f orni. Hopkins and Elliot 2 showed that liver tissue contains thermo-
labile catalysts responsible for the reduction of glutathione. Glucose
dehydrogenase, 3 a system present in the intact mammalian erythrocytes 4
and the Warburg-Christian enzyme 5 are all known to be capable of reducing
GSSG to GSH. It is of interest to enquire whether the papaya fruit also
contains thermolabile catalysts, which function in an analogous manner
and incidentally ascertain the nature of protection afforded to vitamin C
against oxidation. A preliminary communication on the subject appeared
in Current Science.**
Experimental
I. THE SYSTEM GSSG ^t GSH
(1) The influence of aeration on the SH concentration of the papaya latex.
5 gm. samples of the fresh latex, drawn from plants grown in the Institute
nursery, were weighed into a series of wash bottles (300 c.c. capacity) dispersed
in 40 c.c. phosphate buffer (pH 7-4) and aerated by sucking in air with the
help of a filter-pump. At intervals of 2 hours the contents of the bottles
were transferred to a glass mortar, rubbed with trichloracetic acid (20 per
cent.) and sand, and filtered on a Buchner. The residue was repeatedly
extracted with trichloracetic acid until the filtrate gave a negative test
with nitro-prusside. The combined filtrate was made up to 100 c.c.,
sufficient water being added to bring the final concentration of the trichlor-
acetic acid to 10 per cent. 5 c.c. aliquots were used for the estimation of
SH. Two methods were employed for the purpose : (1) lodometric
titration according to the method of Kuhnau 7 and (2) the colorimetric
81
82
C. V. Ganapathy and Bo N. Sastri
method employing the photoelectric colorimeter. This method developed
in our laboratory has been described elsewhere. 8
The results obtained with different samples of latex are given in
Table I.* The figures represent c.c. of N/100 iodine required for titrating
the SH groups present in one gram of the latex. Temperature 25 C.
Aeration Time (Hrs.)
2
4
'6
8
10
12
Sample 1
8-2
8-4
8-0
4-4
2-4
2 ..
10-0
10-1
8-8
7-6
6-8
,,3 ..
7-5
7-5
6-7
5-1
,,4 ..
7-8
7-7
6-6
5-4
4-5
3-24
2-4
,,5 ..
6-8
6-7
5-1
3-2
,,6 ..
7-3
7-4
5-7
4-2
2-1
,,7 ...
6-7
6-6
4-9
3-3
,,8 ..
8-4
8-4
6-6
4-8
1-9
,,9 ..
7-1
7-1
6-3
5-1
2-5
,,10 ..
8-8
8-8
7-2
6-0
5-2 8
2-4
10f ..
7-5
7-4
5-9
4-8
3-8
(2) The influence of preliminary heating of the latex on the oxidation of
SH groups during aeration. In the second series of experiments the latex
was dispersed in phosphate buffer (pH 7-4) as before, raised to a temperature
of 50 C., at which temperature, it was maintained for 60 minutes, cooled
* The titre values tabulated in this section have to be corrected for the presence
of ascorbic acid. The main conclusions obtained from these experiments, however,
will not be affected, as the titre value for ascorbic acid remains constant until almost
all the thiols are oxidised, as will be shown in the subsequent part of the paper. The
values for the SH concentration determined by the photoelectric colorimeter are not
subject to this error.
t The figures refer to the values calculated in terms of ^ iodine obtained with
the photoelectric colorimeter ; only results for sample 10 are given here. The
difference between the values by the two methods is due to the ascorbic acid.
Oxidation of Thiols and Ascorbic Acid in the Latex of Papaya S3
and aerated. Subsequent treatment was the same as before. The results
obtained are shown in Table II.
II
Aeration Time (Hrs.)
2
4
6
8
Sample 1 . .
7-5
5-3
4-2
2-3
1-2
2
6-7
4.3
3-0
1-9
0-8
3 ..
7-0
5-1
4-0
2-4
4 ..
5-8
3-7
2-5
1-3
0-2
The results in the first series of experiments show that there is no fall
in the SH concentration during the first two hours of aeration, after which
there is almost a linear fall. In a few instances, there is a tendency towards
a slight increase in the SH concentration. When the latex is heated to 50,
prior to aeration, however, there is a steady fall in the SH content from the
very commencement of aeration. The SH content remains at a constant
level during aeration only if a reducing mechanism capable of reconverting
the SS to the SH form is present in the latex. If such a mechanism is
present, addition of fresh SS compounds should lead to an increase in the
SH concentration of the reaction mixture on incubation.
(3) Influence of the added SS compounds on the SH concentration of the
latex. The effect of addition of SS compounds prepared from the latex
itself, was examined in the third series of experiments. The thiols were
extracted by boiling the fresh latex with water and oxidised with iodine or
hydrogen peroxide. In the case of iodine the quantity required for oxida-
tion was determined by direct titration ; with H 2 O 2 , the aqueous extract
of the latex was raised to pH 9 by adding NaOH, then H 2 O 2 was added and
after oxidation was completed (as shown by negative test with nitro-
prusside) the excess of H 2 2 was removed by boiling. The resulting solu-
tion was finally brought down to pH 7 by hydrochloric acid and used.
A solution of the SS compounds (1 per cent, concentration) was mixed with
fresh latex and after 4 hours incubation, the SH content was again deter-
mined by iodometric titration. The results given in Table III clearly show
that there is an increase in the SH content over the initial value, thereby
showing that the SS compounds have been reduced. In a parallel series
of experiments, where the SS compounds were added to samples of latex
dispersed in buffer and previously heated to 50 C. for one hour, the increase
in the SH content was markedly less.
84
C. V* Ganapathy and B, N. Sastri
III
Fresli latex
Latex heated prior to incubation
Sample
Initial titre
Increase in
titre
Initial titre
Increase in
titre
1
8-8
0-8
8-9
0-3
2
7-2
1-0
7-3
04
3
7-1
1-1
7-2
0-4
(4) Experiments with the press-juice from papaya fruit pulp. 100 c.c.
of the fresh juice from the pulp of papaya fruit were incubated with a
solution containing SS compounds (2 5 per cent, concentration) for 4 hours
and the SH content estimated. In the control series, the juice was boiled
prior to incubation. The results tabulated in Table IV show that there is
a marked rise in the Iodine titre indicating the presence of reducing systems
in the juice. The thermolabile nature of the reducing systems is shown by
the fact that this increase in titre becomes markedly less when the juice is
heated prior to incubation.
TABLE IV
Sample
Time of
Hrs.
Increase in titre c.c. N/100 iodine
Fresh juice
Heated juice
1
2
2-0
0-8
2
4
5-1
1-3
3
4
5-2
1-4
4
2
2-4
0-9
5
2
3*7
1-1
6
4
7-2
1-8
7
2
4-1
1-2
8
4
6-0
1-8
shcorbic
in the Latex of Papaya 85
II. OXIDATION OK Aseoumc Acrn PUKSKNT IN Tins
Vitamin C exists aUmt'; wiih the thiols in the latex,. That glutatliione
affords protertioti ai;;iinst the oxidation of vitamin C by hexoxidase or
eoppei \\as sho\\ n by Hopkins and Morgan." The observations reported
in this stvtion of I ho paper support the main conclusions of Hopkins and
co workers,
(I) ///////r'jfev M/' tn'nititni on I he cvncentnilion of thiols and vitamin C
(/'///< /N7/uiV4i ///*' \. The lay-ont of the (experiments was exactly similar
to that tleseiibeti in the csiilier part of the paper. 5 gin. latex were weighed
unt into earh *f < wash bottles (.'WO c.e. cajmcily) and air bubbled through,
At intervals, the vitamin C (dye titralion) 10 and SH concentrations
were Si-paiately determined in aliqnots of the ticliloracetic acid extracts.
The results are tabulated in Table V. The values for vitamin C and thiols
are given in r,c. of X loo iodine t*ori\ i S]>onding to I gin. of latex,
TAIJI,K V
i
1
2
4
6
-0
0-0
-0
0-5
5 <5
5-7
r>-
3 -0
0-3
-3
-3
-3
-3
0-2
5-1
B -2
r> -3
5-1
3 -3
2-0
0-8
<>
O-K
0-8
()
0-0
-3
9-5
9-0
i) -()
7-0
5-1
Sample I Vitamin C'
Sli . .
"J Vitumin ('
sn .,
;i Viiumin ('
SI I , .
( % J) /t7/i'(7 <i/' /*/r//w/'>w>'y heating of the latex on the coarse of oxidation
of vitamin C ami //uW.s. The i)rcvious experiment was repeated with
samples of latex dispersed in phosphate buffer and heated at 50 C. for
one hour prior to aeration,
TAW,K VI
Time (Hrn.)
I
2
4
Sample 1 Vitamin r . ,
S1J
-0
5-5
('>
4 -0
(>
4 -1
0-5
2*1
0-3
0-6
,, 2 Vitumin C- . ,
0-8
0-8
0-8
0-7
0-6
811
0-1
5-0
3 -5
2-3
1-0
86 C, V, Ganapathy and B. N. Sastri
(3) Influence of thiol compounds of papaya latex on the oxidation of
ascorbic acid by copper,- To 50 c.c. of a boiled aqueous extract of the latex
was added pure crystalline ascorbic acid in solution (70 mg.) and aerated
after adding a trace of copper. The vitamin C content as well as the
concentration of the thiols were determined in aliquots at intervals of 10
minutes. Table VII gives the results obtained. The values for vitamin
C and SH are given in terms of N/100 iodine. 'There is complete protection
of ascorbic acid from oxidation by copper as long as there is even a small
amount of the thiols present in the solution.
TABLE VII
Aeration
time in
mins.
Vitamin C
SH
2-9
3-0
10
2-8
1-8
20
2 -8
1-0
40
2-2
0-0
60
0-6
0-0
Conclusions and Summary
The presence in the latex, and more especially in the pulp of the papaya
fruit, of a system responsible for maintaining the thiol compounds in the
reduced condition is clearly established. Its thermolabile character is
indicated by the observation that a preliminary heating of the latex dispersed
in buffer or the press-juice from the pulp to a temperature of 50 C. for a
period of 60 minutes, renders it inactive. The system present in the papaya,
in these respects, is analogous to that in the liver described by Hopkins.
Further work is called for in order to determine the exact nature of the
mechanism.
The significance of this finding lies in the fact that the reducing system
provides a regulating mechanism for the proteolytic processes in the plant,
the concentration of the SH compounds which function as the natural
activators of papain being conditioned by the activity of this system. It
also helps, indirectly, to maintain the ascorbic acid present in papaya in the
Oxidation of T/iiols and Ascorbic Acid in the Latex of Papaya 87
reduced condition, as the thiol compounds afiord complete protection to
tlie vitamin against 'oxidation by hexoxidase or copper as previously shown
by Hopkins, and confirmed by us.
1.
2.
3.
4.
5.
(>.
7.
8.
9.
10.
LITERATURE CITED
Ganapathy and Sastri
Hopkins and Elliots
M ann.
Mold rum
Ganapathy
JCuhnau
Ganapathy and Sastri
Hopkins and Morgan .
Proc. Ind. Acad. Sci. 9 1938, 8, 399.
Proc. Roy. Soc., 1931, 109B, 58.
tiiochcm. ,/., 1932, 26, 7S5.
Ibid., 1932, 26, 817.
Ibid., 1935, 29, 108.
Curr. Sci., 1938, 6, 451.
niochem. Z., 1931, 230, 353.
Proc. Ind. Acad. Sci., 1938, 8, 399.
Biochem. J. 1936, 30, 1446.
Birch, Harris and Ray Ibid.. 1933, 27, 590.
ON OF
IN
BY K. S. SRINIVASAN, B.Sc. (HONS.), M.Sc.
Eeceivecl June 11, 1939
[Communicated by Dr. M. O. P. lyengar, ar.A., Ph.D. (Lond.), F.L.S.]
Marchantia palmata is a very common liverwort growing in plenty at
Ootacamund on the Nilgiris. Most of the plants are female and bear plenty
of archegoniophores. The male plants (PL II, Fig. 5 c ; Text-fig. 2) are
extremely few, being about only 1 per cent, or less. In most cases, the
female receptacles show one or more proliferations from their under surface
(Pi. II, Fig. 5 a ; Text-fig. 3). A section of these receptacles shows the usual
archegonia on the underside as in all species of Mwchantia, but a section
of the proliferations shows, peculiarly enough, a large number of antheridia
on them (PI. II, Fig. 3). The receptacles in this Marchantia are thus bi-
sexual and androgynous.
A detailed account of the androgynous receptacles of this Indian liver-
wort has not been published so far. The author took advantage of the
large amount of material available at Ootacamund to make a detailed
study of these receptacles.
It is not easy to get a correct idea of the nature of these proliferations
with the aid of microtome sections alone, since their growth is generally
somewhat curved and twisted. It was only after examining numerous
hand sections along with the microtome sections of the proliferations in
all stages of growth and also after careful examination with the aid of a
Greenough Binocular dissecting microscope of both dissected and un-
dissected material that a correct idea of the nature of the proliferations
could be obtained.
The female receptacle is more or less disc-shaped with about 7 to 9
rays and measures 10 to 14 mm. across (Text-fig. 1). The upper surface is
* Front the University Botany Laboratory, Madras. This paper formed part
of a Thesis submitted for the Degree of Master of Science of the University of Madras.
It was read before the Annual Meeting of the Indian Academy of Sciences at Madras
on 20th December 1938.
88
Morphology of Androgynous Receptacles in M. palmata Nets 89
TEXT-FIGS. 1-11. Marchantia palmata Nees.
(1) A normal female receptacle. X 1-5. (2) A normal male receptacle, x 1-5.
(3) An androgynous receptacle showing the proliferations growing from its under
90 K. S. Srlnivasan
side. X 1-5. (4) A median vertical section of a normal female receptacle. x 7-5.
(5) Beginning of the proliferation. x 22. (0) Ventral view of the bisexual receptacle
with the velum and the perianth dissected out, showing the end of two female branches
beginning to grow out as proliferations P r P 2 . x 4. (") A section through a
proliferation showing an exerted antherkliimi by the side of an archegonium. X 96,
(8) & (0) Later stages in the growth of the proliferation. (8) X 13. (9) x 18.
(10) Same as Fig. 0, but with the proliferation (P) grown out a little more, x 7-5.
(11) A well-grown proliferation. X 11.
smooth and dome-shaped with or without a slight depression in the centre.
A number of archegonia is formed on each branch of the receptacle, and
a large number of the sporogonia develops to maturity. Each group of
archegonia is well protected by the pronounced development of the velum
which passes round the growing point also. Only when the velum is re-
moved by careful dissection can the growing point be seen clearly. At
first there is no indication of any proliferation from the female branch.
Each lobe of the receptacle bearing the archegonia, as in all species of
Marchantia, is bent downwards, the growing point being directed towards
the stalk of the receptacle. In the Nilgiri Marchantia, the lobe does not
grow adpressed to the ventral surface of the dome-shaped portion of the
archegoniophore, but grows a little free from it and is somewhat pendant
(PI. IT, Fig. 4; Text-fig. 4). The apical portion of the branch bearing the
archegonia, instead of ceasing its growth after forming a certain number of
archegonia as in the other species of Marchantia, becomes active again and
grows into a small cushion-like structure (Text-fig. 5). This structure
continues to grow forward and downward and forms a short, more or less
pendant, tongue-like outgrowth (Text-figs. 8, 9, 10). At first, the proli-
feration is made up of a solid mass of uniform cells (Text-fig. 5), but later
on air-chambers with assimilating filaments are developed in it (Text-fig. 9)
and the proliferation continues to grow gradually outwards and upwards
and at the same time turns on its axis as it w r ere about 180 or so and
finally brings the morphologically dorsal side to the upper side (Text-figs.
8, 11). It then continues to grow into a fiat green narrow lobe. This
outgrowth is somewhat narrow below but becomes gradually broader to-
wards its apical portion. As it grows, it goes past the velum, which then
appears as if attached to its flanks. Very frequently, it may branch di-
chotomously into two spreading lobes which may also occasionally divide
a second time dichotomously (PI. II, Fig. 5 a). Fully grown proliferations
show a certain amount of resemblance to the lobes of a normal antheridio-
phore. Their ventral surface bears amphigastria in definite rows together
with tufts of rhizoids. The amphigastria are produced rather early in the
development of the proliferation (Text-figs. 5, 8, 9, 11).
Morphology of Androgynous Receptacles in M. palmata Nees 91
Sections of these proliferations show a number of antheridia
stink in cavities on the upper side. But very often the first antheridium
that is formed is not sunk in any cavity, but is situated just on the surface
like an archegonium and is not surrounded by any tissue (PI. II, Figs. 3, 6 ;
Text-fig. 7). These exerted antheridia show a certain amount of resem-
blance to archegonia in that their walls are elongated radially (Text-fig. 19)
as in archegonia or are made up of two layers near the base as in the venter
region of the archegonium (Text-fig. 18). The later formed antheridia
grow in an acropetal manner with the youngest near the apical and the
18
TEXT-PIGS. 12-19. Marr.hantia palmata Nees
(12) A normal archegonium. X 513. (13) An abnormal archegonium with three
eggs, x 346. (14 & 15) Abnormal antheridia. (14) X 87. (15) X 97. (10) A normal
antheridium. X 55. (1.7) An antheridium with two chambers. X 63. (IS) Abnormal
antheridium. showing division of the cells of the wall-layer towards the basal
region. X 257. (10) Abnormal antberidium showing the cells of the wall-layer
elongated at right angles to the axis of the structure, x 170. Contents not shown
in Fig*. 17-19.
92 K. S. Srinivasan
oldest near the basal region. Most of the antheridia that are formed later
are normal and quite similar to those formed on normal male receptacles.
But the few antheridia that are formed on the proliferation near its base
are very peculiar in showing characters which are midway between those
of antheridia and of archegonia. Most of these resemble arcliegonia in
external shape, but contain inside a large number of sperniatogenous cells
(PI. II, Figs. 2, 6 ; Text-fig. 15). These sperniatogenous cells are invariably
found in a degenerate and aborted condition. A few are very similar to
antheridia in general structure but their upper portion is not broad and
rounded as in normal antheridia but are very narrow as In archegonia
(Text-fig. 14).
An interesting case of an abnormal archegoniiim was found in one of
the sections. This archegoniiim contained three eggs in the ventral region
and, out of these three, two were large and one was very small (Text-fig. 13),
This case suggests a tendency on the part of the cells inside the archegoniiim
to multiply in number.
One of the abnormal antheridia had a long and slender stalk, but the
antheridial portion was divided into two compartments by the development
of a partition-wall right across the antheridium (Text-fig. 17). The
chambers were empty and presumably contained sperniatogenous cells.
The significance of this was not clear. It is just possible that the two
chambers represent the development of the venter and the neck region
of an archegoniiim and were brought about by the division of the cells in
these two respective portions.
The proliferations during their growth would appear to start as a
definitely female structure, but to gradually change to a male structure.
And, during the stages of this transition, it forms sexual structures which
are intermediate between female and male.
Discussion
Androgynous receptacles have been recorded in a few genera of the
Marchantiacese (Preissid, Dumortiera and Marchantia] by various authors.*
As early as 1834, Taylor 14 recorded androgynous gametophores in
Dumortiera irrigua. In a later communication 15 as regards the gameto-
phores of this plant he states that " The fructification is commonly dioecious,
sometimes monoecious, and not rarely androgynous as observed in Marchantia
androgyna ".
* A short historical account of androgynous receptacles is given by Cutting 3 and
A. W. Haupt. 7
Morphology of Androgynous Receptacles in M. palmata Nees 93
In 1880, Goebel 6 described androgynous receptacles in Preissia
wnmutatd. He observed that in these receptacles the two anterior lobes
ore antheridia on the upper surface, while the two posterior ones had
rchegonia on the lower surface. He thinks that this does not represent
reversion to the primitive monoecious arrangement of the sexual struc-
ires, but is merely a case of replacement. In 1881, Leitgeb 12 also
^corded androgynous receptacles in Preissia commutata. These receptacles,
nlike those recorded by Goebel, bore two archegonial groups on the lower
irface of the anterior portion, while the posterior portion bore antheridia
n the upper surface. He expresses the view that the androgynous condi-
.011 is brought about by a delayed sexual differentiation of the branches
:iat bear the sexual structures, instead of such a change being initiated
arlier in the vegetative portion of the thallus itself. In 1899, Townsend 16
escribed androgynous receptacles in the same liverwort. These receptacles
ore antheridia on the upper side and archegonia on the under surface.
.. W. Haupt 7 states that the receptacles of Preissia quadrata are very fre-
uently bi-sexual. Some of them were like those described by I/eitgeb
nd others were like the ones observed by Goebel. He states that in most
ases " either both anterior groups of sex organs were of one sex and both
osterior groups of the other sex, or less frequently three were of one sex
nd one of the other. In a few cases both of the groups on one side of the
sceptacle were of the same sex, and both groups on the opposite side of the
ther sex". O'Hanlon 13 in his paper on Preissia quadrata, gives a
hotomicrograph of a longitudinal section of an androgynous receptacle,
i which the arrangement of the antheridia and archegonia were quite
.milar to that of Haupt's specimens. He states that " Preissia is not only
monoecious plant, but there also occur mixed heads".
Ernst 5 recorded androgynous receptacles in Dumortiem tnchocephala
nd D. velutina. He found that these receptacles were fairly common in
le former, but were found only very occasionally in the latter. In these
^ceptacles, one portion was completely male and bore antheridia on the
pper side and the other portion was completely female and formed arche-
Dnia on the under side. Campbell 1 also found bisexual receptacles in
^umortiera trichocephala and states that they were quite similar to those
mud by Ernst. Kashyap 11 states that androgynous receptacles occur
equently in Dumortiera hirsuta, especially in forms collected from very
Loist places, but does not give details regarding their structure.
Heberlein 9 found in an unnamed species of Marchantia from Peru-
ian Andes, two androgynous receptacles which had much the shape of
le female receptacles and had the archegonia on the under side in their
94 K. S. Srinivasan
usual position, but had antheridia on Its upper side sunk in cavities.
G. Haupt 8 found In Marchantia grisea that the receptacles were either
male or hermaphrodite. In the bisexual ones one portion of the receptacle
was male and the other female. Cutting 3 found in a species of Marchantia
which he got from Chelsea Physic Gardens, London, plenty of androgynous
receptacles. These receptacles resembled archegoniophores. But a number
of proliferations were growing from their lower surface. And on these
proliferations w r ere formed a number of antheridia sunk in cavities. He
states that these antheridia-bearing proliferations are " formed as out-
growths from a portion of the under surface of a female branch". He again
states that the " male outgrowth was formed secondarily as a kind of
proliferation, and is not a mere replacement of the normally female branch".
He suggests that " it would not be unlikely for the entire archegonia-
bearing portion of a branch to grow out into a protuberance", but states,
however, that " no such case has been seen" by him. Chopra 2
refers to some receptacles of Marchantia indica as androgynous and says
that they are quite similar to those described by Cutting.
Now, coming to the Nilgiri Marchantia, we find that its receptacle is
primarily an archegoniophore and its general form and course of develop-
ment are quite like that of any normal female receptacle. But the arche-
gonia-forming lobes of the receptacle, after producing the usual output of
female structures, instead of stopping further growth, become active once
again and grow out into antheridia-bearing proliferations. These prolifera-
tions produce at first, however, sexual structures which are intermediate
in character between an archegonium and an antheridium and only later on
do they produce typical antheridia. So we see that Cutting's suggestion
that " it would not be unlikely for the entire archegonia-bearing portion
of a branch to grow out into a protuberance " is now actually found very
commonly in the Nilgiri Marchantia, though Cutting did not find any such
case in the Chelsea material.
In all the previous cases mentioned above, the receptacles are andro-
gynous from the very commencement. Some portions of them form the
male structures and the other portions the female structures. The arche-
gonia and the antheridia are situated on the main receptacle itself, though
their positions varied in the different forms. But the present case is quite
different from any of the previously recorded types. Here the whole recept-
acle is completely female to start with and produces only archegonia up to
a certain stage. After that, some of the archegonia-forming lobes become
transformed into antheridia-forming ones. In other words, the lobes change
their sex from female to male during the course of their development . The
Morphology of Androgynous Receptacles in M. palmata Nets 95
difference between the previously recorded cases and the present one is this.
In all the previous cases the receptacle is bisexual from the very commence-
ment, some portions being male and the others female, whereas in the
present case the receptacle is unisexual (female) to start with and then in
its ultimate portions changes its sex from female to male.
What exactly may be the cause which leads to this change of sex is
not quite clear. It may be mentioned in this connection that a large
number of plants bearing these androgynous receptacles are infected with
a fungus. The vegetative portion of the thallus is very heavily infected
in almost all cases. In a few cases the mycelium was found to invade the
receptacle and to attack the sexual structures also. This fungus produces
on the dorsal surface of the receptacle and on the proliferations a number
of small pyciiidia (Pi. II, Fig. 1). I am unable to say at present whether
this fungus could in any way be the cause of the change of sex in the present
Marchantia.
G. Haupt, 8 while investigating the androgynous receptacles of Mar-
chantia grisea, found that the male plant had only 9 chromosomes whereas
the female and the hermaphrodite ones had 10 chromosomes, of which one
was very small and was called by him the " z " chromosome. But the
antheridia formed by these hermaphrodite plants had only 9 chromosomes.
He thinks that, during the formation of these antheridia, the " z "
chromosome is somehow lost. This " z " chromosome is considered by
him as being responsible for the female sex. In the Nilgiri Marchantia
the author found plenty of aberrations in the chromosome complement
of the normal female receptacle as well as of the proliferations. It could
not be decided how far these aberrations may be responsible for the change
of sex in the liverwort. A detailed account of these aberrations will form
the subject of a separate communication.
A word may be said here regarding the change of sex in the Nilgiri
Marchantia. The change of sex of the branch forming the archegonia is,
as seen already, not sudden. Before it begins to form normal antheridia, it
forms structures intermediate between antheridia and archegonia. The
change in the sex is gradual. A study of these abnormal antheridia brings
prominently to view the homologous nature of the two structures, viz., the
antheridia and the archegonia. Both these structures represent game-
tangia enclosing sexual cells or gametes within a common wall. In the
male structures are found quite a large number of sexual cells all of which
are functional, while in the female are found only a small number of sexual
cells of which only one (egg) is functional. Davis 4 suggests that antheridia
96 K. S* Srinivasan
and archegonia are derived from a common sporangial structure. Holferty 10
found In Mnium cuspidatum plenty of abnormal archegonia which showed
structures Intermediate between those of an antheridium and of an arche-
gonium. From a consideration of these, he conies to the conclusion that
antheridia and archegonia had a common origin, probably from a multi-
locular sporangial structure, as suggested originally by Davis. The various
intermediate sexual structures found in the Nilgiri Marchantia would also
appear to lend much support to this view.
Summary
1. Marchantia palmata which grows commonly at Ootacamund on
the Nilgiris forms plenty of androgynous receptacles.
2. These receptacles start as purely female receptacles, but soon one
or more proliferations bearing antheridia grow out from its under surface.
3. These proliferations are merely the archegonia-bearing lobes of
the female receptacle which become active again and continue to
grow further into narrow elongated outgrowths.
4. The proliferations produce at first sexual structures which are
intermediate between an archegonium and an antheridium and later on
typical antheridia. A consideration of these intermediate structures
suggests that antheridia and archegonia are homologous structures.
5. A fungus forming pycnidia is generally present in the liverwort.
It could not be dicided whether the change of sex in the receptacle is due to
this fungal attack.
I have great pleasure in acknowledging my indebtedness to Prof.
M. O. P. lyengar, M.A., Ph.D. (I,ond.), F.L.S., for his constant guidance and
help in the preparation of this paper. My thanks are also due to the autho-
rities of the University of Madras for the award of a research scholarship,
during the tenure of which the present investigation was carried out.
LITERATURE CITED
1. Campbell, D. EL . . t4 Studies on some East Indian Hepaticse," Ann. Bot.>
1918, 32, 319-38,
2. Chopra. R. S. . . ^ Notes on Indian Hopaticse, I. S. India," Proc. Ind.
Acad. ,S'ei., 103S, 7, No. 5, 243.
3. Cutting, E. M. .. "On Androgynous Receptacles in Marchantia^ Ann.
Bot. 9 1910, 24, 349-57.
4. Davis, B. M. .. "The origin of the archegonium," ibid., 1903, 17
477-92.
W. ///</. A cad. .S>/., />', :-/. .V, /'/. //
4 \
'^'llt,
Morphology of Androgynous Receptacles in M. palmata Nees 97
5. .Ernst, A.
Goebel, K.
8.
9.
10.
11.
12.
13.
U.
15.
1(3.
1.
o
3.
4.
,*
0.
7.
llaupt, A. W.
llaupt, GL
Jleberlein, FJ. A.
Ilolforty, G. M.
K ash yap, S. R.
Loitgeb, H.
O'llanlon, M. E.
Taylor, Th.
To \vnso nd, A,.
" IJntersuchungen uber Entwicklung Bau und Ver-
teilung der Inlloreszcnzen von Dumortiera," Ann.
Du. Jard. Bot. fa BuUenzorrj, ii, S., 1908, 22, 153-223.
" Uber die Verzweigung dorsiventraler Sprosse," Arb.
des. Bot. Instil, in Wurzburcj, 18SO, Bd. 2, Hoffc 3,
353 (from Cutting).
" Morphology of Preissia quadrc.ta" Bot. Gaz., 1926,
82, 30-51.
" Bcitrage y.ur Zytologie der Gattung Marchantia (L.),"
Kelts, fur. Ind. Aba. und Vereb., 1933, Bd. 63,
390-119.
' ' Morphological studies on a new species of Marchantia,"
Bot. Gas., 1929, 88, -127.
" The archegomum of Mnium cuspidatum," ibid., 1904,
37, 106-2(5.
Liverworts of the Western Himalayas and the Payijab
Plains, Pt. I, Lahore, 1929.
Untersuchunyen uber die Leber moose Graz., 1874-82.
ri. Die M arvhantiac>een, 1881.
" A study of Preissia quadrata," Sot. Gaz., 1927, 84,
208-18.
11 DC Marchantieis," Trans. Linn. Soc., 1834, 17, 375
(from Cutting).
Mackay's Flora Hiherniae, 1836, 2, 51 (from Cutting).
k< An hermaphrodite gametophore in Preissia commit.-
toto," Bot. Gaz., 1899, 28, 360-62.
EXPLANATION OF PLATE II
Marchantia palmala "Nces
Section of a proliferation showing a portion (darkly stained portion) attacked by
a. fungus. Note the pycnidia (py) of the fungus, x 25.
Section of a proliferation showing an abnormal anthericlium by the side of a
normal one. X 38.
Tangential section of a proliferation showing a superficial antheridium at base,
and a number of normal antheridia higher up. x 35.
Vertical section of a normal female receptacle. X 9.
Photographs of a bisexual (a), female (1) and male (c) receptacles, x 3.
Proliferation showing two abnormal antheridia one superficial and the other
inside a cavity. X 24.
Longitudinal section of a proliferation showing antheridia inside chambers. Note
the archegonia in their original positions in Figs. 6 and 7. X 36,
BY G. W. CHIPI^ONKKR, M.Sc.
(Department of Geology, Benares Hindu University}
Received June 20, 1939
(Communicated by 'Prof. L. Rama Eao, M.A., p.o.s.)
Introduction
IN the preliminary identification of the fossils from the Bagh Beds given
by Duncan, 1 the Bryozoa were represented by Eschara sp., and Escharina
sp., in the Deola-Chirakhan Marl. To this P. N. Bose added Ceriopora
dispar Stoliczka, a species described from the Ariyalur stage of the Creta-
ceous series of South India. 2 But like all the other groups of fossils from
these strata Bryozoa also were till now not studied in any detail. Subse-
quent to the study of the Echinoids and Rhynchonellids from this formation, 3
the Bryozoa were taken up for a detailed study with a view to investigating
into the alleged faunal affinities between the Bagh Beds and the Cretaceous
series of South India, 4 and to see what light they throw on the problem of
the age of the Bagh Beds.
The material studied here was collected by the late Prof. K. K. Mathur
and by the present writer from the various exposures in the neighbourhood
of Agarwara (lat. 22 16', long. 75 59'), Chirakhan (lat. 22 21' 30", long.
75 7' 30"), the type locality, Ajantar (lat. 22 19' 30", long. 74 55') and
Bagh (lat. 22 21' 30", long. 74 47' 30"). Though the Bryozoa des-
cribed below were obtained from all the fossiliferous divisions of the Bagh
Beds, 5 their main bulk comes from the Coralline limestones which are very
largely made up of their fragments. Due to the unusually hard nature of
the Coralline limestone and its saccaroidal appearance on the freshly
broken surface, it is only on the naturally weathered surfaces that
these fossils can be observed ; the fragments weathered out from the
Coralline I^imsetone always get mixed up with those from the Deola-
Chirakhan Marl ; the staining red due to iron oxide is more conspicuous in
the case of the specimens from the Marl ; but here the nature of the matrix
1 Q.J.G.S., 21, 354 ; Man. Geol. Ind., 250.
2 Mem. G.S.I., 21, 37, 40, 47 ; Pal. Ind., (), 2, 26, pi. 3, fig. 1-3 ; Man. Geol. Ind.,
250.
3 Proc. Ind. Acad. Sci., (I-}), 6, 60-71 ; ibid., 7, 300-10 ; ibid., 9, 236-46.
4 Mem. G.S.I., 21, 38-44.
5 Ibid., 21, 35-44 ; Curr. Sci., 4, 322.
98
Bvyozoa from the Bag/i Beds
99
is not always proved to be a dependable indication for the source of the
ssil ; and thus one is oftentimes very much "uncertain as to the exact
currence and the relative abundance of the different species in the Marl
.d in the Coralline limestone. Owing to this unfavourably hard nature
the rock, the method of washing and screening which is oftentimes
plicable for securing isolated Bryozoan specimens, is found to be absolu-
hy useless in the present case. This, besides the inherent difficulty in
i dying fossil Bryozoa has, very seriously hampered the present work.
is only in the three species of Ceriopora that isolated specimens were
tained from among the debris of the weathered Coralline Limestones ;
d only their internal structure could be studied under the microscope in
in sections taken along definitely known directions. In all the other species
Large amount of material could be examined only on the weathered sur-
ges of rock specimens, their longitudinal and transverse sections being
tained accidentally in rock sections.
The author very much regrets that due to weathering and the hard
;r actable material encrusting the fossils, the photomicrographs could not
better than what are reproduced here.
All the Type specimens are preserved in the Department of Geology,
mares Hindu University.
Description of Species
Order . . Cyclostomata, Busk.
Family . . Idmoniidse, Busk.
Genus . . Idmonea I/amouroux, 192L
Idmonea biserialis sp. nov.
(Plate III, Figs. 2, 9)
Dimensions. Outer diameter of the Zoacium
Inner
mm.
0-15
0-10
Description. The zoarium is branching dichotomously ; the apertures
i arranged in oblique alternating series on the right and the left ; aper-
:es are moderately projecting outwards and sub-circular in outline.
Comparison. This species closely resembles Idmonea francorum
rgens, 6 from the Senonian of Central France, in which, however, the
icia are a little contracting towards the aperture and thus have smaller
^rtural diameter.
6 Bull. Soc. Belg. Geo., 3, 343 ; pi. 13, fig. 1.
100 G, W, Chlplonker
In Idmonea controtilis Lonsdale 7 from the Senonian of New Jersey,
Charente, France, and Pooland, South Africa, the zoacia are without
any definite arrangements ; the upper part of the zoacia is free and much
projecting outwards.
Family . . Cerioporidse, Busk.
Genus . . Ceriopora Goldfuss, 1827.
Cenopora dimorphopora sp. nov.
(Plate III, Fig. 5 ; Plate IV, Figs. 2-3)
1 mm.
Dimensions. linger diameter of larger aperture . . 0-16
Shorter ..0-10
Longer diameter of smaller aperture . . 0-9
vShorter ,, ..0-6
Description. The zoarium is large, free, sub-cylindrical with lateral
branches ; the surface has sometimes feebly developed tuberosities. The
zoacial tubes are polygonal with rather thick walls. The apertures are
rather small, elliptical and in two sizes ; the larger apertures are often
surrounded by the smaller ones ; the larger apertures are more or less regu-
larly arranged in quincunx. In general the surface shows a heteroporid
aspect. The zonal lines are not much separated.
Comparison. By the nature of the zoarium this species resembles
Ceriopom lobifera Canu and Bassler, 8 from the Valangian of Sainte-Croix,
Switzerland, from which it can be easily distinguished by its elliptical aper"
tures in two sizes and much thicker zoacial walls.
From Cenopora dimorphocella Canu and Bassler, 9 from the Aptian of
Farringdon, England, this species can be distinguished by its elliptical
orifices and the zonal lines.
Cenopora conoformis sp. nov.
(Plate III, Fig. 3 ; Plate IV, Figs. 1, 5)
mm.
Dimensions. Longer diameter of the larger aperture 0-17
Shorter .. 0-12
Longer diameter of the smaller aperture 0- 10
Shorter ,, ,, .. 0-6
Description. The zoarium is conical with no lateral branches ; the
general surface gives a heteroporid appearance. The zoacial tubes are
7 Q.J.O.S., 1, OS ; Pal. Fr. t 5, 729 ; Prod. Pal. Strat., 2, 265 ; .-1m?. S. Afric. Mus.,
4, 286.
3 Proc. U.S. Nat. Mus., 67, 27, pi. 23, figs. 11-17.
9 Ibid., 29, pi. 24, figs. 1-6, pi. 31, figs. 7-8.
Bryozoa from the Bagh Beds 101
polygonal with the walls rather thick. The apertures are small, elliptical
and generally in two sizes. The larger apertures are surrounded by the smaller
ones and are generally arranged in a roughly quincunxial order. Not infre-
quently are found apertures of a third size roughly intermediate between
the two sizes, of which the micrometric measurements are given above. In
the case of such apertures the zoacial walls are found to be thicker than in
the case of others ; so that this third size of apertures is in reality a modifica-
tion of the larger ones.
Comparison. This species is distinguished from the associated
C. dimorphopora sp. nov., described above, by its conical zoarium without
lateral branches, the zonal lines more frequent and the occasional develop-
ment of the third size of apertures by thickening of the zoacial walls.
Ceriopora ellipsopora sp. nov.
(Plate III, Fig. 6 ; Plate IV, Fig. 4)
mm. mm.
Dimensions. Longer diameter of the orifice . . . . 16 -19
Shorter .. ..0.90-11
Description. The available material is in the form of cylindrical
pieces over 15 mm. in length ; so that it is uncertain whether it has a tendency
to branching laterally. The cross-section of zoarium is broadly elliptical.
The zoacial tubes are polygonal with rather thick walls. The orifices are
elliptical and arranged roughly in hexagonal pattern in such a way that
every orifice is at the centre of a hexagon.
Comparison. This species differs from the associated C. dimorphopora
sp. nov., described above, by the nature of its zoarium, only one type of
zoacia, and zonal lines rather crowded in the marginal region.
Ceriopora dispar Stoliczka 10 from the Ariyalur Stage of the South
Indian Cretaceous Series which Bose 11 had reported to be present in the
Bagh. Beds also, can be easily distinguished from the present species by its
more or less circular and smaller orifices.
Ceriopora lobifera Canu and Bassler 12 from the Valangin of Switzerland
differs from this species in having branching zoarium with oval to sub-
circular apertures and thinner zoacial walls.
Family . . Ceriocavidse Canu and Bassler.
Genus .. Ceriocava d'Orbigny, 1858.
10 Proc. U.S. Nat. Mus. 67, 29, pi. 24, figs- 1-6 ; pi. 31, f gs. 7-8.
" Pal. 2nd., (8), 4, pt. 2, 20, pi. 3, figs. 1-3 ; Ann. Pal, 11, fasc. Ill-TV, 27, pi. 11,
fig. 3 ; Cat. Cm. #??/., 1, 251.
12 Mem. G.S.I., 21, 37, 41, 43.
102 G. W. Cbiplonker
Ceriocava micropora sp. nov.
(Plate III, Fig. 1)
mm. mm.
Dimensions. Diameter of the orifice . . . . 10 14
Diameter of a large branch . . . . 2-3
Description. The zoarium is free and branching with the branches
cylindrical. The orifices are rather small, polygonal, arranged in quincunx,
but occasionally roughly in horizontal lines also. The ovicells are very
minute.
Comparison. This form resembles the Valangian species Ceriocava
grandipora Canu and Bassler 13 from Switzerland. The orifices in the present
species are, however, very much smaller (i.e., about one- third of those of the
Swiss species).
Order . . Cheilostomata, Busk.
Family . . Membraniporidse, Busk.
Genus . . Membranipora Blainville, 1834.
Membranipora, mathuri* sp. nov.
(Plate III, Fig. 10)
mm.
Dimensions. linger diameter of zoacium . . 0-27
Shorter ..0-19
Longer diameter of aperture .. 0-22
Shorter ..0-13
Description. The available specimens are preserved as small patches
each including only a few zoacia. The zoacia are arranged in more or less,
regular linear series, those of the adjacent series being nearly in horizontal
lines. The series have a common suture. The zoacia are oval to sub-
elliptical ; the apertures are oval. The zoacia are bordered by a more or
less distinct rim.
Comparison. M. vendinnensis d'Orb. 14 from the Cenomanian of Sarthe,
France, is closely related to this species ; but the French form has larger
zoacia and the disposition is less regular.
M. sub-ovalis Canu 15 from the Senonian of Tunis has the zoacia of the
same type and disposition as the present species, but the African form has
its zoacia larger and also has interzoacial pores.
13 Proc. U.S. Nat. Mus., 67, 07, pi. 0, figs. 14-17.
* This species is named after the late Prof. K. T\. Ma-.hur.
14 Prod. Pal. Strat., 2, 174 ; Pal. Fr., 5, 545, pi. GOO, figs. 9-10 ; null. Soc. Geo.
Fr., (4), 12, 349, pi. 13, fig. 1 ; ibid., (3), 28, 354 ; ibid., 25, 741.
15 Ibid., (1), 3, 660, pi. 21, fig. 2.
Bryozoa from the Bag/i Beds 103
Membmnipora pseudo-normaniana sp. nov.
(Hate in, Fig. 8)
mm. mm.
Dimensions. Conger diameter of zoacium . . 0-23
Shorter ..0-20
linger diameter of aperture . . 15 17
Shorter _ . 10 Q-12
Description. The zoaria are preserved as small patches ; the zoacia
are disposed irregularly and polygonal in outline ; the spaces between larger
zoacia are occupied by smaller ones which are probably due to overthicken-
ing of the zoacial wall of the larger ones. The aperture is oval to sub-
elliptical in outline and is nearly half of the zoacium in diameter.
Comparison. This form resembles very closely M. normaniana
d'Orb. 16 from the Senonian of France and North- West Germany; but the
Buropean species has wider apertures and the zoacial walls thinner.
M. cervicornis Brydone 17 from the B. muwonata zone (Senonian) of
Portsdown, England, has its orifices wider and more sub-circular and the
walls relatively thinner.
Family . . Onychocellidse, Jullien.
Genus . . Eschara I^amarck, 1801.
Eschara chimkhanensis sp. nov.
(Plate III, Fig. 7)
mm.
Dimensions. Length of the zoacium . . 0-36
Width ,, ..0-22
Description. The zoarium is encrusting on rhynchonellid shells, with
the zoacia arranged in regular linear series, the zoacia of the adjacent series
alternating. The aperture is oval with a low simple border which is clearly
seen on the anterior and on the two sides ; aperture is nearly half of the
zoacium in length.
Comparison. This species can be distinguished from E. cenomana
d'Orb. 18 from the Cenomanian of France, by its encrusting habit, larger
zoacia and the front wall and the aperture relatively a little narrower.
16 Prod. Pal. Strat. 9 2, 202 ; Pal. Fr., 5, 550, pi. 607, figs. 5-6 ; and 9-10, 551,
pi. 007, figs. 11-12 ; Icopoldina, -427, pi. 5, fig. 2.
17 Ceol. Mag., (5), 10, 198, pi. 7, figs. 3-4.
18 Prod. Pal. Strat., 2, 176 ; Pal. Fr., 5, 105, pi. 602, figs. 1-3, 246; pi. 687, figs.
14-16 ; Bull. Soc. Geo. Fr., (3), 25, 737 ; ibid., 28, 361 ; ibid., (4), 12, 353, pi. 14, figs.
3-4.
104 G. W. Chiplonker
The present species differs from E, sdntonensis d'Orb. 19 from the
Senonian of France, in having a larger and more oval aperture.
Eschara regularis sp. nov.
(Plate III, Fig. 4)
mm.
Dimensions. Longer diameter of zoacium . . . . 0-37
Shorter . . ..0-25
Width of aperture . . . . . . 12
Height of aperture . . . . . . 10
Description. The zoarium is encrusting ; the zoacia are hexagonal,
longer than broad and arranged in regular series with zoacia of the adjacent
series alternating ; they are provided with a raised prominent rim. The
front wall is feebly convex towards the middle and is much below the level
of the zoacial rim. The aperture is nearly semi-circualr with a low rim
slightly broader than high, anteriorly rounded, posteriorly straight and
rather large ; it is situated just below the zoacial rim, which is closely
followed by a very feeble, shallow depression after which the convexity of
the front wall begins.
Comparison. As compared to E. royana d'Orb. 20 from the Senonian
of France and Pondoland, South Africa, the present species is an encrusting
form with slightly broader zoacia.
E. parisiensis d'Ortb. 21 from the Senonian of the Paris basin, resembles
the present species in having a depressed groove along the inner side of the
zoacial rim and the regularity of disposition of the zoacia. The difference,
however, lies in the present species having shorter zoacia and the aperture
posteriorly straight and situated quite anterior to the centre of the
zoacium.
By the irregularity of disposition and shape of the zoacia and the oval
aperture with a tooth near its anterior margin E. nerei d'Orb., 22 a Senonian
species from Central France, can be easily distinguished from the present
species.
19 Pal. Fr., 5, 109, pi. 603, figs. 1-3 ; pi. 673, fig. 4 ; Prod. Pal. Slrat., 2, 261;
Bull. Soc. Geo. Fr. y (3), 28, 398.
20 Prod. Pal. Mrat., 2, 264 ; Pal. Fr. 9 5, 108, pi. 602, figs. 12-1.3 ; pi. 673, figs. 2-3 ;
Rev. ZooL, 112 ; Ann. S. Afrtc. Mu*\, 4, 280 ; Bull. Soc.. Geo. Fr. t (3), 28, 300.
21 Rev. ZooL, 112 ; Prod. Pal. StraL, 2, 264 ; Pal. Fr. 9 5, 110, pi. 603, figs. 4-6 .
pi. 673, figs. 5-6 ; Bull. Soc. Gin. Fr. t (4), 2, 13.
22 Rei\ ZooL, 112 ; Prod. Pal. StraL, 2, 264 ; Pal. Fr., 5, 111, pi. 603, figs. 10-13 ;
pi. 604, figs. 1-4 ; pi. 673, fig. 7 ; Bull. Soc. Geo. Fr., (3), 28, 391.
Bryozoa from the Bagh Beds 105
Eschara holkari* sp, nov.
(Plate III, Fig. 11)
mm.
Dimensions. Longer diameter of zoacium .. .. 0-3
Shorter .. .. 0-2
Description. The zoarium is encrusting; the zoacia are elliptical,
arranged in regular linear series, with the zoacia of the adjoining series alter-
nating with each other. The aperture is rather large, sub-circular to oval,
a little more than half the size of the zoacium and has an inconspicuous
rim ; the front wall is depressed towards the aperture. The space in the
interzoacial corners is trigonal and depressed.
Comparison. E. dejanira d'Orb. 23 from the Senonian of the Paris-
Pyrineese basin differs from the present species in having the zoacia oval
rather than elliptical, the aperture smaller and posteriorly slightly truncated
and the marginal rim incomplete.
Geological Age and Faunal Affinities
The accompanying table summarises the f aunal affinities of the Bryozoa
from the Bagh Beds with those from other parts of the world. Since there
are no species common to these beds and Cretaceous series of strata else-
where, of which the geological horizon is fixed on the basis of other fossils,
it is rather difficult to assign any definite horizon to these Bryozoa with
certainty. However, to discuss their relations with species from different
parts of the world, we find that these Bryozoa show, like the Echinoids
from these beds, 24 a mixture of affinities towards certain European species
ranging from the Valangian to the Senonian. Of the ten species described
above, to consider first the more abundant ones, we have Membranipora
mathuri, M. pseudo-normaniana, Eschara Chirakhanensis and E. regularis.
Of these Membranipora mathuri is related to M. vendinnensis d'Orb. from
the Cenomanian of Sarthe ; Eschara Chirakhanensis is allied to E. santonensis
d'Orb. from the Senonian of France on the one hand and to E. cenomana
d'Orb. from the Cenomanian on the other ; while the remaining two species
have their allies in the Senonian of France. Among the less abundant
forms, we have the three species of Ceriopora related to lower Cretaceous
* This species is named after His Highness The Maharaja Holkar in whose terri-
tory the Bagh Beds are nicely exposed in the type sections at Chirakhan.
23 Pal. Fr. 9 5, 161, pi. 675, figs. 17-19 ; Bull. Soc. Geo. Fr. 9 (3), 28, 401, pi. 6, figs.
15-16.
24 Proc. Ind. Acad. Sci., 6, 60-71 ; ibid., 9.
106 G* W. Chiplonker
Table showing Vertical Distribution and Affinity-Relations of
the Bryozoafrom the Bagh Beds
Xo.
Species from the
Bagh Beds
Allied species with
stratigraphical
position
No'lular
Limestone
Flower
Coralline
Limestone
Deol.a-
Chiraldian
Mar!
Upper
Coralline
Limestone
1
Idnioma hismaiis sp
nov.
Idmonia f ran com m Per-
jjons , SuiioniLi) of France
X
>
X
2
Cerlopora >!iinorplio-
pnra. sp. nor.
Cvriopora di morphocdta
Canu and Bassier ; Ap-
tiaii of Farrington
X
V
3
C. Gono/0r//i/ssp. nov
Do.
X
X
4
0. dli'psopora sp. nov
6'. lobifera Canu and Bas-
sler ; Valangian of Swit-
zerland
X
5
Ceriocava micropora
sp. nov.
Cerlocara yrandipora Canu
and Bassler; \'aiangian
of Switzerland
X
6
Memltranipora
tfiathari sp. nov.
Mtmbranijiora vendimien-
sis d'Orb. ; Cenoruanian
of Sarthe
X
v
v
7
M. psendo-nonnan-
iana sp. nov.
M. normaniana d'Orb. ;
Senonian of France
X
X
X
X
8
Eschara chirakhancn-
sis sp. nov.
Esc It ara sanlonen sis d* r b . ;
Senonian of France ; E.
Cenomana d'Orb.; Ceno-
X
X
X
X -
n jam an of France
E. regulari*^. nov.
K. parix i ensis d'Orb.;
Senoniaii of Paris basin
X
X
X
10
E. holhari sp. nov.
E. dejarina d'Orb.. Seno-
nian of Paris-Pyrineese
basin
X
<
species from Switzerland and Farringdon. Ceriocava micropora is allied to
C. grandipora Canu and Bassler from the Valangian of Switzerland. The
remaining two species, Idmonea biserialis and Eschara holkari, have their
near allies in the Senonian of France.
Thus to consider this Bryozoan fauna in general, it does not seem
unjustified to assign to them a horizon at about the Cenomonian, a conclu-
sion which is not contradictory to the results arrived at from the study of
the Echinoids and Rhynchonellids from this formation. 25
As is already mentioned in the Introduction, the Bryozoa described
here, are obtained in the main, both in number and kind, from the Upper
Coralline limestone, which is the uppermost member of the Bagh Beds.
Ibid., 7, 66-67 ibid., 7, 312-13 ; ibid., 9,
Bryozoa from the Bagh Beds 107
It is thus significant to note that the geological age assigned here to the
present Bryozoan fauna is based on the study of the species mainly from the
youngest member of the series.
As to the alleged direct faunal affinities between the Bagh Beds of
the Narbadha. Valley and the Cretaceous Series of South India, 26 the present
study of the Bryozoan fossils shows that no species are identical with, nor
even allied to any of the forms described from South India. 27 On the
contrary, these fossils are related to the Mediterranean biological province.
The present study thus supplies clear evidence, in addition to that furnished
by the Echinoids and the Rhynchonellids from these strata, 28 to show that
the Narbada Valley and the Trichinopoly District belonged to two different
basins of sedimentation with no direct connection between them and
supported their own faunas.
A cknowledgements
I take this opportunity of expressing my great indebtedness to the late
Prof. K. K. Mathur for kindly placing at my disposal his collection of fossils
which forms a part of the material for the present work. Very sincere
thanks are due to Dr. Raj Nath for his constant and keen interest in the
present work and for his valuable criticism. I am very much thankful to
the Director, the Geological Survey of India, for permission to work in their
Survey Museum and library. I take this opportunity of expressing my
great indebtedness to the Dhar, Indore and Gwalior Durbars for the kind
permission to collect fossils from their territories and for the facilities given
during the field-work.
BIBLIOGRAPHY
1 Bose, P. N. " Geology of the Lower Narbada Valley between
Nimawar and Ka\\anth." Mem. G'.S.Z., 18S4, 21,
pt. 1.
> Brydonc R. M. . . '" iNotes on new or imperfectly known Chalk Poly&oa,"
Gco. May., Deo. 5, 1013, 10, 190-99.
3 Cann F " Bryozoaires du Cenomaien de Sainte- Calais (Sarthe),"
fitdl. Soc. Geo. France, 1897, (3), 25, 737-54.
^ . . " Revision des Bryozoaires Cretaces figures par Ale.
d'Orbigny. II Teil : Cheilostomata," ibid., 1900,
(3), 28.
5 . . *< Bryozoaires fossiles," ibid., 1902, (4), 2, 10-14.
P . . " Contribution a 1'Btudes des Bryozoaires (Membrani-
pora)," ibid., 1905^, (4), 3, 059-61.
26 Mem. G.8.I., 21, 38-4-1.
27 Pal Ind., (8), 4-, pt. 2.
2* Proc. Ind. Acad. 8ci., (B), 6, 81-68 ; ibid., 7, 300-13 ; ibid., 9, 236-46.
108
G. W. Chiplonker
7. Canu, P.
g, -- 9 an( i Sassier, B. S.
9. Chiplonker, G. W.
10. -- : -
12. Duncan, M. P.
13. Gregory, J. W.
14. Lang, W. D.
15. Lecointre, G.
1(3. Lonsdale, W.
17. Oldharn, B. D.
18. Orbigny, Alcid cV
19.
20.
21. Persons, Ed.
22. Rode, K. P., and
Ohiplonker, G. W.
23. Stoliczka, P.
24. Voigt, E.
in Cottreaii- "Paleontologio de MarUigasca. X. Fossils
Cre"taee de la cote Orient-ale," Ann. Pal., ll)*J2
11, fasc. III. fit IV.
" Studies on the Cyclostomatous Bryozoa," Prac. II. ti.
Nat.Mus., 1926, 67 s Ait. 21; 1-12-1.
c< Echinoicls from the Bagh Bads, 1 ' Proc. I nil. Awl.
Sci. 9 1937, (B), 6, 00-71.
" Rhynchonellids from the Ba?h Beds?," -ibid., IMS,
(B), 7 5 300-10.
" Echinoids from the Bagh BedsPart II," ibid., UKW,
(B), 9, 236-46.
44 Description of the Echinodermata from tho strata
on the South-Eastern Coast of Arabia and at Ba^h
on the Narbada in the Collection of the Geological
Society/ 5 Q.J.C.S., 1805, 21.
Catalogue of the Fossil Bryozoa in the Department of
Geology of the British Musi-it ni< The Cretaceous
Bryozoa, 1809, 1.
in Wood, 11., " Cretaceous fauna of Pomloland," Ann.
South Afric. Muff., 1900, 4, 275-350.
" Sur quekjues Bryozoaires nouveaux on pen connus
du Oenomanien du Mans," 'Bull. Soc. (feo. Prance^
1912, (4), 12.
" Account of the six species of Polyparia obtained from
Timber Creek," Q.J.G.S., 1845, 1, 65-75.
Manual of Geology of Jndia, 1.893, 2nd ed.
Revue de tfooZ,, 1849.
Prod, de Pal. Sir at., 1850, 2.
Paleontoloyie Francaifse* terr. cret., Bryozoairos, 1850, 5.
' k Ke vision des Bryozoairos dxi CnStace' figures par
d'Orbip'ny Oyclostomata," Hull. Soc. lielg. GeoL,
18G9, S, 305-100.
u A Contribution to the Stratigraphy of the Bagh Beds,"
Curr.-Sci., 1935, 4, 322.
" The Cretaceous Fauna of Southern India," Pal. ln<l.,
1872, (8), 4, pt. 2.
i Morphologische und Strati graphisdic untcirsuoliungrn
uber die Bryozoenfauna tier oberer Kreide," T<M! I.
* Die Cheilostornen Bryozoen des jmi^ercn Ober-
kreide in Nordwest Deutschland iin Bo.lticum und
in Holland,' LeopoWinn Ber. K. J.eo. Dent. Acad.
Naiurfor. Halle, 1930, 6, 379-579.
G. W. Chifrlonker '
Proc. Ind, A cad. ScL, B, \ vol. X, PI, III
11
G. H'\ Chi plonker
Proc. Ind. A cad. &/., B, vol. X, PL IV
Bryozoa from the Bagh Beds 109
EXPLANATION OF THE PLATES
PLATE III
FTG. 1 .< Ccriocara micropora sp. nov. ; showing part of the zoarium. X 18 (Holotype ;
B.H.TJ. No. B/5).
FIG. 2. Jdmonea Inserialis sp. nov.; showing part of the zoarium. x!8 (Holotype;
B.H.U. No. B/l).
FIG. 3. Ceriopora co-noformis sp. nov. ; showing part of the zoarium. X 55 (Holotype ;
B.H.U. No. B/3).
FTO. 4. EsaJwra rc.gularis sp. nov. ; showing part of the zoarium. x IS (Holotype ;
B.H.IT. No. B/9).
FIG. 5. Ceriopora dimorpkopora sp. nov. ; showing part of the zoarium. x 18
(Holotype ; B.K.U. No. B/2).
FIG. 0. Ceriopora eUipsopora sp. nov. ; showing part of the zoarium. X 18 (Holotype ;
B.H.IJ. No. B/4).
FIG. 7. Eschara chirakhanensis sp. nov. ; showing part of the zoarium. x 18 (Holo-
type ; B.H.U. No. B/S).
FIG. 8. M enibranipora pseudo-normaniana sp. nov. ; showing part of the zoarium..
x 18 (Holotype; B.H.TJ. No. B/7).
FIG. 9.- Idmonea biserialift sp. nov. ; showing part of the zoarium. X IS (Paratype .
B.H.U. No. B/l-1).
FIG. 10. Membranipora -malhuri sp. nov. ; showing part of the zoarium. x .18
(Holotype ; B.H.U. No. B/C>).
FIG. 11. Eschara holkari sp. nov.; showing part of the zoarium. X IS (Holotype;
B.H.U. No. B/10).
PLATE IV
FIG. 1. Ceriopora conoformis sp. nov. ; showing cross-section of the zoarium. X 12
(Holotype ; B.H.U. No. B/3).
FIG. 2. Ceriopora dinwrphopora sp. nov. ; showing cross-section of a branch of the
Koarinm. X 12 (Paratype; B.H.U. No. B/2-1).
FIG. 3. Ceriopora dinwrphopora sp. nov.; showing part of the zoarium; natural
size (Holotype; B.H.U. No. B/2).
. 4. Ceriopora cllipuopora sp. nov.; showing part of the zoarium; natural size
(Holotype ; B.H.U. No. B/4).
!> 5. Ceriopora conoformis sp. nov. ; showing pait of the zoarium ; natural size
(Holotype ; B.H.U. No. B/3).
1116 39-Printed at The Bangalore Press, Mysore Road, Bangalore City, by G. Srinivasa Rao. Superintendent,
and Published by The Indian Academy of Sciences, Bangalore
STUDIES IN HELMINTHOLOGY*
Trematode Parasites of Birds
BY MAKUND BEHARI LAL, D.Sc.
(Lecturer in Zoology, The University, Lucknow, India]
Received June 2, 1939
(Communicated by Dr. G. S.Thapar, M.SC., ph.r>.)
CONTENTS
PAGE
INTRODUCTION AND ACKNOWLEDGMENTS . . . . . . . . 113
HISTORICAL REVIEW .. .. .. ..113
COLLECTION OF MATERIAL ... . . .. .. ..116
TECHNIQUE .. .. .. .. .. .. ..118
CLASSIFICATION.. .. .. .. .. .. .. 118
(a) Characters of Systematic Importance . . . . . . 118
(b) Systematic Description of the Families . . . . . . 118
FAMILY NOTOCOTYLID^ . . . . . . . . . . 118
Genus Notocotylus . . . . . . . . . . . . 120
Genus Hindia . . . . . . . . . . . . 120
Genus Naviformia . . . . . . . . . . 122
Genus Paramonostomum . . . . . . . . . . 122
Genus Neoparamonostomum . . . . . . . . . . 123
Genus Catatropis . . . . . . . . . . . 123
FAMILY CYCLOCCELID^ . . . . . . . . . . . . 125
Genus Hcematotrephus . . . . . . ... . . 127
Genus Cydoc&lwn . . . . . . . . . . . . 127
Genus Typhlophilus . . . . . . . . . . 3.31
Genus Contracalum . . . . . . . . . . 133
Genus Ophthalmophagus . . . . . . . . . . 134
FAMILY DICROCCELIID^ . . . . . . . . . . 134
Genus Lyperosomum . . . . . . . . . . 135
Genus Platynosomum .. . . * .. .. ..137
Genus Multivitelldria . . .. .. .. ..138
* Part of the thesis submitted and approved for the Degree of Doctor of Science
in the University of Lucknow, 1937.
111
II 2 Makund Behari Lai
PAGE
139
FAMILY OPISTHORCHIID^
Genus Opisthorchis
14-5
FAMILY LECITHODENDRIID^ . . - - " , ,
14:0
Genus Parabascus
FAMILY HETEROPHYID.E . " U9
Genus Ascocotyle
150
FAMILY MICROPHALLIDJ& " - n
Genus Levinseniella . * ' J
1 53
FAMILY CEPHALOGONIMID^ . . -
Genus Prosthogonimus . . . >
FAMILY ECHINOSTOMID^ .
Genus Echinostoma . . - * * . . loo
1 P\Q
Genus Echinochasmus
Genus Stephanoprora . . 164
Genus Parorchis . . * *
Genus Paryphostomum > ..168
Genus Petasiger
1 7A
FAMILY PSILOSTOMID^; . - 1 ' u
Genus Psilorchis . . - *
171
FAMILY STRIGEID^ . . -
Genus Cy&thocotyle . . - - 173
Genus Neodiplostomwn - ..173
Genus Proalaria .. ..178
1 70
Genus Neoalaria - - - - -
1 81
Genus Pharyngostomum - -
FAMILY SCHISTOSOMID^ . . 184
Genus Ornitholilharzia . . . . 185
Genus Chinhuta .. - 186
1 &7
Genus Gigantobilharzia * i0 '
HOST-PARASITE RELATIONSHIP . . . . 188
COPULATION IN TREMATODES . . . . . . . 188
GEOGRAPHICAL DISTRIBUTION .. . . - - ..190
191
REFERENCES
LIST OF ABBREVIATIONS - - 200
Studies in Helminthology 113
INTRODUCTION AND ACKNOWLEDGMENTS
BIRDS form one of the most important economic units of animals, some of
them even constituting the delicacies of our table, but very little attention
has so far been paid to the study of their Trematode parasites in India.
In view of their economic importance and also in order to remove some
existing confusion in the description and classification of these parasites,
I determined, at the suggestion of Dr. G. S. Thapar, to work out the morpho-
logy and the systematic position of Avian Trematodes from India. In
spite of the fact that birds are migrants, results have not been disappoint-
ing and I have been able to collect materials and data that add to our
knowledge of this group.
Part of the work has already been published and is referred to at its
proper place in the thesis.
The work was carried out in the Zoology Department of the I^ucknow
University during the years 1934-36. I wish to record here my deep
indebtedness to Dr. G. S. Thapar for his kindly advice and criticism through-
out the progress of my work. He further very kindly allowed me free
access to his valuable personal library and placed part of his own collection
of Avian Trematodes at nay disposal for which I express my sincere thanks.
Several other colleagues rendered vahiable help in my work Mr. J. Dayal
in going through the typed manuscript ; Dr. (Miss) Dorothy Speer and
Dr. A. C. Chatterji in the translation of some German literature ; Mr. K. S.
Iyer in going through some literature in French and I take this opportunity
to put on record my sense of gratitude for their help. I am also thankful
to Mr. G. N. Natu, our artist, for technical assistance in the preparation of
illustrations. My thanks are also due to the authorities of the Zoological
Survey of India for providing facilities to consult their Library. To Prof.
Birbal Sahni, F.R.S., I am indebted for the loan of a few journals from his
private library, and last but not the least I am thankful to the authorities
of the lyUcknow University for facilities given to carry out these investiga-
tions.
HISTORICAL REVIEW
The work on Avian Trematodes from India is hardly more than a decade
old. The earliest record, to which I have been able to refer, is that of
Bhalerao (1926) where he describes five species of Trematodes, viz.,
Echinochasmus corvus, Stephanoprora reynoldi, Phaneropsolus insolens,
Lyperosomum ka-ked and Platynosomum acuminatum from the Burmese
crow, Corvus insolens. Of these, Platynosomum acuminatum has previously
been reported from the liver of a Kestrel from Scotland by Nicoll (1915).
Makund Beharl Lai
Phadke and Gulati (1930) recorded a new genus Miiltivitellana from
the gall bladder of the common house crow, Corvus splendens and created
a new subfamily Multivitellarinse for its reception.
Moghe (1932) described two new species Echinostomum govindum and
Paramonostomum microstomim from the rectal cseca and the small intestine
respectively of the Indian Ruff, PhilomacJms pugnax.
Harshe (1932) described a new species Catatropis orientalis from the
rectal cseca of the pintail duck, Dafila acuta acuta.
Gogate (1934) recorded four trematodes Echinostoma revohtlwn, Pary-
phostomum testitrifolium, Petasiger minutissimus and Ornithobilharzia sp.
from wild ducks in Rangoon.
Srivastava (1935) described a new species, Ascocotyle intennedius, from
the intestine of the Indian Fishing Eagle, Hali&etus leucoryphus, and further
emended the diagnosis of the genus Ascocotyle. The same year (1935)
he described another species, Catatropis indicus from the rectal caeca of an
Indian fowl, Callus bankiva mitrghi.
Bhalerao (1935) described two species, Notocotylus babai from the
cgecum of the common kite, Milvs migrans govinda, and Cycloccelum sharadi
from the thoracic cavity of the western yellow-billed magpie, Urocissa
favirostris cucullata.
Khan (1935) described eight new species of the genus Cycloccelum,
viz., Cycloccelum lobatum, Cycloccelum nebularium, Cycloccelum straightum t
Cycloccelum indicum, Cycloccelum capellum, Cycloccelum mehrii, Cycloccelum
allahabadi and Cycloccelum erythropis from three different genera of the
Snipes.
Verma (1935) described Echinochasmus bagulai from the small intestine
of the Pond heron, Ardeola grayi and the Night heron, Nycticorax nycticorax,
and also added Echinochasmus ruficapensis from the intestine of the Indian
little grebe or diver, Podiceps ruficollis. In the same paper mention is
made of a small variety of Prosthogonimus and two other representatives
of the genus Prosthogonimus, but unfortunately no account of these forms
is so far available. A chambered excretory bladder was described by
Verma in the genus Echinochasmus and the diagnosis of the genus was
emended, with a suggestion to split it up into two sub-genera.
Patwardhan (1935) recorded Lyperosomum colorosum from the gall
bladder of a black -headed mynah, Temenuchus pagodarum ; Proalaria alceden-
sis from the intestine of the king- fisher, Alcedo atthis ; and Neodiplostomum
tytense from the intestine of a Barn owl, Tyto alba stertens.
Studies in Helminthology 115
Thapar and Lai (1935) added a new genus Psilorchis under the family
Psilostomidse from the intestine of the king-fisher and considered the
evolution of several structures in the family Psilostoinidae.
Later, Lai (1935) described a new species Notocotyhts indicus from the
intestinal cseca of the wigeon, Mareca penelope.
Again, Lai (1935) reviewed the genus Notocotyhis and discussed the
importance of the position of the genital opening. On the basis of this
character he created two new genera, Hindia and Naviformia.
L&1 (1936) described two new species of the genus Paramonostomum, viz.,
Paramonzstomtm querqweduhm from the intestinal caeca of the garganey,
Querquedula circia ; and Parawionostomtiwi casarcmn from the intestinal
casca of the Brahminy duck, Casarca rutila. The value of the position of
the genital pore as of generic importance in the classification of the members
of the subfamily Notocotylinse was further confirmed, and a new genus
Neoparamonostomum was created.
Verma (1936) recorded thirty-three new forms of trematodes from
Indian birds but gave no diagrams and added very meagre and incom-
plete descriptions. It is, therefore, not easy to diagnose and include
these forms with their present confused account at this stage, particularly
because the author himself, as appears from his paper, is in doubt regard-
ing the description of many of his forms. The same year (1936) he also
described Cyathocotyle calv'usi from the intestine of the King Vulture, Torgos
calvus.
Lai (1936) described a new species Par orchis snipis from the cloaca of
the common summer snipe, Totanus hypoleucos. The discovery of a species
of pamrchis from India in the snipe is interesting in so far as this genus
has only been recorded from gulls and restricted only to St. Andrews, North-
umberland coast, Millport and America. In the course of a discussion on
the systematic position of the genus a new subfamily Parorchinse was
created for its reception, and a suggestion was put forward regarding the
polyphyletic origin of the family Echinostomidae.
Lai (1936) while describing a new genus Typhlophilus shovellus from the
intestine of a shoveller duck, Spatula clypeata, demonstrated the presence
of a well-developed muscular ventral sucker in the family Cycloccelidse
and also discussed the advisability of retaining the subfamilies Cycloccelinse '
and Typhloccelinse. The significance of a ventral sucker as a basis for generic
divisions was also pointed out.
116
Makund Behari Lai
L,al (1.936) recorded a new species Levinseniella indica and suggested
the probable identity of several existing species of the genus with the genus
Maritrema owing to the presence of a cirrus sac.
It would, thus, appear that so far| only scattered contributions have
been made in India on the Avian trematodes and no comprehensive and col-
lective work has been attempted. I have, therefore, endeavoured to present
here the results of my investigations, together with summaries of descrip-
tions of other forms given by earlier workers. The latter is indicated by
small type to differentiate it from new and original work.
COU,ECTION OF MATERIAL
A large number of birds were subjected to post-mortem examinations
in normal salt solution during the years 1934-36, and the Trematodes
recovered from them are enumerated in the following table :
Locality
of the
host
Name of host
No.
examined
No.
infected
Location oi
parasite
Name of the trenmtode
Malhaur
Casarca rutila
(Brahminy duck)
G
2
Intestinal caeca
Paramanostomum casar-
cum Lai, 1936 '
Lucknow
Tot amis hypoleucos
(Summer snipe)
4
1
Cloaca
ParorcJiis snipes Lai,
1936
Gallinago yallinula
(Jack snipe)
3
1
Burs a Fabricii
Levinseniella indica Lai,
1936
Spatula clypeata
(Shoveller duck)
4
1
Intestine
Typhlophilus shovellus
Lai, 1936
"
Mareca penelope
(The Wigeon)
7
4
Intestinal caeca
Notocotylus indicus Lai,
1935 and Hindia luck-
nowensis Lai, 1935
Totanus fttscus
(Spotted red shank)
9
3
Intestine
StephanoproTa fusca
n. sp.
Dicrurus macrzcercus
(King crow)
4
Lucknow and
Sitapur
Upupa erops
(The Hoopce)
9
Lucknow
Ardeola grayi
(Paddy bird)
8
6
Intestine
Echinochasmus megavitel-
lus n. sp.
f Since submission of the thesis Mehra (1937) described Lepoderma bulbulii,
Lepoderma casarcii, Lepoderma ferruginum. Yidyarthi (1937) described Aphary ngostrigea
ardeolina, A. Indiana, Strijea orientali and Strigea nephronis. Chatterji (1938) mentions
Prosthogonimus sp. and Catatropis verrucosa. Pande (1938) describes Crassiphiala
stunkardii. The characters of these species are not summarized here.
Studies in Helminthology
117
Locality
of the
host
Name ol host
No.
examined
No.
infected
Location of
parasite
Name of the trematodc
Lucknow
Ardea cinerea rectiros-
6
1
Intestine
Pharyn gostomu m bagu lu m
tris
n. sp.
(Common grey heron)
?
Athene JBrama
4
2
Small intestine
Ncodiplostomum dilacce-
(Spotted owlet)
cu?n n. sp.
Passer domesticus
11
(House sparrow)
Barabanki and
Kittacincla melabarica
9
Sitapur
(Shama)
Lucknow
Milvus migrans
1
3
Liver
Opisthorchis cheelis n. sp.
(Common kite)
>f
Sarcogyps calcus
9
4
Opisthcrchis giddhis n.sp.
(King vulture)
Intestine
Neoalaria tha.paria n.g.,
n.sp.
Acridotheres tristis
2
1
Bursa f abricii
Prosthogonimus cuneatus
(The mynah)
Amausi
Querquedula circia
9
5
Intestinal coeca
Paramonostomum qu er-
(The garganey)
quedulum Lai, 1936
Small intestine
Echinostoma c7iasma
n.sp.
Lucknow
Halcyon smyrnensis
6
1
Intestine
Psilordiis indicus Thapar
(King-fisher)
and Lai, 1935
j
Psittacula krameri
10
(Green parakeet)
99
Coracias benghalensis
6
1
Intestine
N eodiplostomum sp.
(The blue jay)
3 }
Corvus splendens
10
2
,,
EcJiinocyasmus reniovarus
(Common house crow)
n.sp.
Lucknow and
Cinnyris zeylonicus
8
t
Sitapur
(Sun bird)
Ajgain and
Nettion crecca
5
2
Intestine
Psilorchis Ajgainis Lai,
Chinhut
(Common Teal)
1938
1
Main blood vessels :
Chinhuta indica Lai,
Kidney, Lung,
1937
Liver, etc.
Kukrail,
Sterba aurantia
8
Lucknow
(Common river tern)
Lucknow
Bulbucus ibis cow-
5
1
Renal vein
Qigantobilharzia egreta
man dus
Lai, 1937
(Cattle egret)
118 Makund Behari Lai
TECHNIQUE
Host of the Trematode material collected was fixed tinder slight pressure
of the coverglass In 90% alcohol. Some forms were fixed in Bouin's Fluid
and Chrom-osmic for section-cutting. In certain cases a special method of
narcotising and killing the worms in 5% alcohol and gradual transfer to
30% alcohol was found very useful. The worms, thus treated, were fixed
in a well-stretched condition without pressure and displacement of organs
or injury to parts. This was specially helpful in examining the ventral
glands of the family Notocotylidae. The material was preserved in all cases
in 70% alcohol or sometimes in 70% alcohol-glycerine. The specimens for
whole mounts were stained with Ehrlich's acid hsematoxylin in a dilution
of 1 : 20 of distilled water and differentiated in tap-water overnight. This
procedure gave a uniformly brilliant stain. Paracarmine stain was also used.
Although it stained the structures rather deep, it gave good results in differ-
entiating some of the deep-seated structures. Clearing was done in Clove
oil or Xylol. In certain cases sections, both transverse and longitudinal,
were cut, and for this double embedding Celloi din-Paraffin method was used.
The sections were stained with h^ematoxylin (Delafield's) and Eosin.
A special glycerine-alcohol method of clearing was found useful in
certain cases, specially in the Echinostomidae. The unstained material was
kept in 70% alcohol-glycerine in proportion of 3 : 1 and left for about half an
hour. Later it was transferred to 90% alcohol-glycerine in proportion of
4 : 1 and left overnight. The alcohol gradually evaporates leaving speci-
mens in pure glycerine which also thoroughly penetrates into the specimens.
The spines, reproductive organs, genital pore, etc., were easily seen by
this method.
CLASSIFICATION
(a) Characters of Systematic Importance
The author has already published a separate paper on this aspect of
the subject and has fully discussed the value of different characters in the
classification of Avian Trematodes (vide these Proceedings, 1937, 5, No. 2
Sec. B, 33-44). ,,,.-,
The systematic account of the various families given in the following
pages is on the lines of the views expressed in the above published paper.
(b) Systematic Description of the Families
Family Notocotylida I^iihe, 1909
The family Notocotylidse was formed by I,uhe (1909) to contain origin-
ally only three genera Notocotylus, Catatropis and Paramonostomum.
Studies in Helminthology 119
Later on several workers added to this list with the result that the family
was split up into three subfamilies, Notocotylinse, Nudacotylinse and Ogmo-
gasterinse.
The writer in the course of his investigations on the family observed
certain peculiar features, described later, which necessitates the modifica-
tion of the existing diagnosis of the family. The emended diagnosis is as
follows :
Diagnosis of the family Notocotylida I/iihe, 1909, emended.
Trematodes with flat body, without a head-collar. Skin smooth or
thickly covered with fine spines in the anterior part of the body and whole
of the ventral surface. Ventral papillce very rarely present. The ventral
surface provided in most cases with regular rows of unicellular glands.
Oral sucker simple. Ventral sucker absent. Pharynx not present. Oeso-
phagus short. Intestinal cseca simple and end blindly. Excretory pore
slightly dorsal. Excretory bladder with a short unpaired median stem and
a pair of long limbs which branch and anastomose dorsal to the oesophagus.
Testes symmetrical at the posterior end, lateral to the intestinal cseca.
Cirrus sac very long. Vesicula semindlis lying external or partly internal and
partly external. Ovary lies between the testes. Shell-gland in front of
ovary. Receptaculum serninis absent. lyaurer's canal present. Uterus
runs in fairly regular transverse loops between the intestinal cseca and in
front of the ovary. Vitellaria well- developed and lie in front of the testes,
mostly extra-csecal. Genital pore pre- equatorial and median, behind the
intestinal bifurcation or in front of it, sometimes far forward at the side of the
oral sucker or post-equatorial and lateral. Eggs with long thread-like filaments
at both ends. Parasitic in aquatic Birds and Mammals.
Of the three subfamilies, Notocotylinse, Nudacotylinse and Ogmo-
gasterinse, only Notocotylinse is recorded from avian hosts in India.
Diagonsis of the subfamily Notocotylinc& } Kossack, 1911, emended.
Notocotylidae, generally with 3-5 rows of ventral glands on the ventral
surface, sometimes absent. Vesicula seminalis lying external or partly
internal and partly external. Genital pore almost median, pre-equatorial,
behind or in front of the intestinal fork. Vitellaria well-developed in the
posterior half of the body and lateral to intestinal caeca. Testes at the
extreme end of the body, symmetrical. Ovary intertesticular, separated
from testes by intestinal crura. Uterine coils intercsecal.
Type genus Notocotylus Sens. Str.
B2 F
120 Makund Behari Lai
The author has already madet elaborate studies on the various genera
of the subfamily Notocotylinse and as a result of his investigations the
following genera have been recognized under the subfamily. J
Genus Notocotylus Sens. Str. (Lai, 1935).
Hindia Lai, 1935.
Naviformia Lai, 1935.
Paramonostomum Sens. Str. (Lai, 1936).
,, Neoparamonostomum Lai, 1936.
Catatropis Odhner, 1905.
Genus Notocotylus Sens. Str. (Lai, 1935)
Monostomes with two to five rows of protrusible ventral glands ; pharynx
absent ; testes extra-csecal and posterior ; ovary in between testes ; shell-
glands pre-ovarial ; uterine loops confined to the inter-caeca! region behind
the cirrus sac ; receptaculum seminis absent ; part of the vesicula seminalis
enclosed within cirrus sac ; cirrus sac never extending beyond half of body
length ; vagina about J- to as long as cirrus sac ; genital pore situated behind the
intestinal bifurcation ; vitelline glands extend upto the middle of the body ; eggs
018--022 mm. long with filaments at either end.
Type species Notocotylus attenuates.
Notocotylus indicus Lai, 1935
Monostome measuring 2-1.8 mm. x -63mm. Three rows of protrusible
ventral glands, 1.7, 16, 17. Oral sucker subterminal -12mm. x -13mm.
Oesophagus 075 mm. long. Intestinal caeca terminate at a distance of
2 mm. on the right and -1.9 mm. on the left side from the posterior end.
Right testis -36mm. long; left testis -38 mm. long. Cirrus sac -58mm.
long. Ovary 18 mm. x 16 mm. Eggs small and numerous with thin shell
and a filament at each end, measuring -018mm. X -Oil mm. without
filaments.
Host. Mareca penelope (Intestinal caeca).
Locality. Lucknow.
Genus Hindia Lai, 1935
Monostomes with three rows of protrusible ventral glands ; pharynx
absent ; testes extra-cgecal and posterior ; ovary in between testes ; shell-
glands pre-ovarial ; uterine loops lie in the inter-caecal region behind the
t Vide Proc. Ind. Acad. Sci. 9 1935, 2, No. 5, 419-23 and 457-66. Also 1936, 3,
No. 1, 25-34.
Studies in Helmint/iology 121
cirrus sac ; receptaculum seminis absent ; part of the vesicula seminalis
enclosed within the cirrus sac ; cirrus sac extending about |- to -|- body length ;
vagina never more than f of the cirrus sac ; genital pore at the intestinal fork ;
vitelline glands extend | to -J- of the body length from the posterior end ; eggs
014- -0209 mm. long., with filaments at either end.
Type species Hindia gibbus.
Hindia lucknowensis I^al, 1935
Monostome measuring 2 63 mm. x -74mm. Ventral glands 16, 15,16.
Oral sucker subterminal -15 mm. in diameter. Oesophagus -13 mm. long.
Intestinal cseca terminate at a distance of -2 mm. on the right and -15 mm.
on the left side from the posterior end. Right testis -35 mm. x -175 mm.
Left testis -375mm. x -2 mm. Cirrus sac -825mm. long. Cirrus about
- 3 mm. long., armed with several rows of spine. Ovary 175 mm. x 15 mm.
Eggs with thin shell and a filament at each end ; measuring 02 mm. x
-Olmm. excluding filaments.
Host. Mareca penelope (Intestinal c^eca).
Locality.- Lucknow.
Hindia babai (Bhalerao, 1935) I,al, 1935
(Syn. Notocotylus babai Bhalerao, 1935)
Body small, elongate, ellipticle, measuring 3- 83 mm. in length and -03 in maximum
breadth a little behind anterior end. Cuticle covered with minute spines. Three rows
of uniform, and eversible ventral glands present ; lateral ones with 17 glands each, and
middle one with 15 glands. Oral sucker is sub-terminal and measures -2 mm. X -16 ram.
Oesophagus mcasuies *12 mm. in length. The intestinal cticca end at a distance of
205 mm. from posterior end. Testes, elongate, deeply lobed, symmetrical ; extracaecal
and measure -535 -545 mm. x -24 -2mm. Cirrus sac, elongated, club-shaped
and measures 1-09 mm. Genital pore median, ventral to intestinal bifurcation.
Vesicula seminalis present inside the cirrus sac. Pars prostatica wide and elongated,
measures 435 mm. narrow muscular, eversible cirrus measures -575 mm. Ovary
deeply lobed, median, in between two testes. It measures -3 x -31 mm. Shell-gland,
prc-ovaiial and measures -235 mm. x -165 mm. Oviduct enlarges into uterus which
passes anteriorly in 18 transverse loops which are mostly intercaecal. Vagina measures
50 mm. Laurer's canal present. Receptaculum seminis absent. Vitellaria follicular,
extra-csecal, a portion intercsecal. They extend between anterior border of testes to
anterior 3/5 of the body. Yolk reservoir dorsal to shell-gland. Eggs measure
014 -017 mm. x 008 -Oil mm. with long filaments at either pole. A short
excretory bladder. Excretory pore dorsal and median near the posterior end.
Host. Milvus migrans govinda (Caecum).
Locality.. Rangoon.
Remarks. The form is transferred to the genus Hindia because of the
position of its genital pore and other characters.
122 Makund Behari Lai
Genus Naviformia I/al } 1935
Monostomes with three rows of protntsibU ventral gln-mls ; pharynx
absent- testes extra-caecal and posterior; ovary in between the tcst.es;
shell-glands pre-ovarial ; uterine coils lie in the intcr-aixnil region Muml
the cirrus sac ; receptaculum seminis absent ; part of the vesieuki semmahs
is enclosed within the cirrus sac ; cirrus sac extending about I of ihc body
length vagina f of the cirrus sac ; genital pore in front of the inlcstintd Jork ;
vitelUne glands behind mid-body, eggs .0178mm. X -0208 mm. long, with
filaments at either end.
Type species Namformia navifonnes.
No species recorded from birds from India.
Genus Paramonostomum Sens. Str. (I/al, 1936)
Monostomes without ventral glands and a pharynx ; intestinal cteca run
almost upto the posterior end of the animal ; testes may or may not be
lobed, extra-csecal in the posterior part of the body ; part of the vesicula
seminalis enclosed within the cirrus sac ; ovary in between the testes ; shell-
glands pre-ovarial ; uterine loops mostly confined to the iiiter-eflecal region
behind the cirrus sac ; receptaculum seminis absent ; genital pore always
opens in front of the intestinal fork and behind the oral sucker ; vitcllinc glands
never end behind the mid-body from the posterior end ; eggs with liUuneiits at
either end.
Paramonostomum querquedithtm I v al, 1936.
Monostome measures 3 -398 mm. x 1-11 mm. Oral sucker 17 mm. x
12mm. Oesophagus -165 mm. long. Right testis -(HJmin. x -124 mm.
Left testis -74 mm. x -235 mm. Cirrus sac -93 mm. long. Ovary -205 mm.
X -22mm. Eggs with thin shell and a filament at each end; measuring
.02mm. x -009mm. without filaments.
Host. Querquedula oircia (Intestinal caeca).
Locality. Amausi.
Pammonostomum casarcum I v al, 1936
Monostome measures 3 -8 mm. x -95 nun. Oral sucker H .mm. X
1mm. Oesophagus -14 mm. long. Right testis -8mm. x -15 mm. Left
testis -76mm. x -175mm. Cirrus sac -91mm. long. Ovary -38 mm. X
-22mm. Eggs thin-shelled and provided with a filament at each pole;
measuring -015mm. x -Olmm. without filaments.
Host. Casarca rutila (Intestinal caeca).
Locality. Malhaur, I/ucknow.
Studies in Helminthology 123
Genus Neoparamonostomum I/al, 1936
Monostomes without ventral glands and pharynx ; intestinal cseca run
almost upto the posterior end of the animal ; testes extra-csecal and posterior
and generally lobed ; part of the vesicula seminalis enclosed within the
cirrus sac; ovary in between the testes; shell glands pre-ovarial ; uterine
loops mostly confined behind the cirrus sac ; receptaculum seminis absent
receptaciilum seminis uterinum may be present ; genital pore always opens
behind intestinal fork ; vitelline glands end behind mid-body from the posterior
end ; eggs with filaments at either end.
Type species Neoparamonostomum ionorne.
(Syn. Paramonostomum ionorne Travassos)
Neoparamonostomum microstomum (Moghe 1932) Lai, 1935
(Syn. Paramonostomum microstomum Moghe, 1932)
Body elongated, 1 -348 mm.. -1 -624- mm. long. Maximum width *375 mm -409 mrn.
anterior to testes. Oral sucker, small, terminal, -003 mm. in diameter. Ventral
sucker absent. Very short oesophagus ; pharynx, and prepharynx absent. Intestinal
caeca extend up to -05 mm. from posterior end. Testes, extra-csecal at the posterior
end of animal, elongated, and lobed, -13 15mm. long. Cirrus sac in anterior
half of body, betAveen the intestinal caeca, -413 mm. long. It contains vesicula seminalis
straight and narrow, and opens at genital pore, immediately below the intestinal bifur-
cation. Ovary lies in the median line between the testes, intercsecal, shell-gland posterior
to ovary. Uterus with 13-15 loops on either side. Vagina as long as cirrus sac.
Uterus extends along more than J of the body. Vitellaria extend from anterior end of
testes to about middle of the uterus region. Eggs provided with long polar filaments
and measure -051 mm. long.
Host. Philomachus pugnax (Small intestine).
Locality. Nagpur.
Remarks. It is rather peculiar that the shell-gland has been described
by Moghe posterior to the ovary, a condition which is unique in the family
Notocotylidse. This apparently appears to be a case of misrepresentation
of structures as has been pointed out by some other authors also. But in
case the post-ovarial position of shell-gland is confirmed in other specimens^
it would not only affect our concept of the family Notocotylidse but would
also place it still nearer the family Pronocephalidae.
Genus Catatropis Odhner, 1905
Monostomes with three rows of weakly developed non-protrusible ventral
glands ; the middle row of ventral glands may be placed on a median keel or a
ridge ; pharynx absent ; testes extra-csecal and posterior ; ovary inter-
testicular ; shell-gland pre-ovarial ; vesicula seminalis is divisible into an
124 Makund Beharl Lai
external and an Internal portion ; receptaculum seminis absent; vagina is very
much developed, about as long as cirrus sac ; genital pore situated behind the
intestinal fork ; eggs with filaments at either pole,
Type species Catatropis verrucosa.
Catatropis orientalis Harshe, 1932
Remarks. Harshe while describing this new species does not mention
anything about the nature of the ventral glands present, whether they are
protrusible or non-protrusible. The question of protrusibility or non-
protrusibility is a vital one, as it is the most important difference by which
Catatropis is distinguished from Notocotylus, Hindia and Naviformia. This
species, therefore, should in the opinion of the writer, be kept as a species
inqitirendum till more light is thrown by a reinvestigation of the form and
hence the characters of Catatropis orientalis, sp. inq. are not summarised
here.
Catatropis indiciis Srivastava, 1935
Worms light brown in colour, -i-6 mm. long and 1-2 mm. broad. Dorsal surface
convex, ventral surface concave with 3 longitudinal rows of non-protrusible unicellular
glands glands in median row contiguous, those in lateral rows distinct and 1012 in
number in each row. Excretroy system similar to that seen in Catatropis. Oral sucker
14- -2 mm. in diameter; oesophagus -2- -26 mm. long; pharynx absent. Testes
deeply lobed, -75- -99 mm. x -2- -3 mm., extracaecal ; vesicula seininalis enormous,
outside the cirrus sac. Cirrus sac flask-shaped, -87-1 -2 mm. long and -17- -2 mm.
broad containing cone-shaped pars prostatica, -35 mm. X -09 mm. surrounded by
prostate cells. Genital pore close behind oral sucker. Ovary lobed, -26- -35 mm.
size, intercsecal, in level with testes and " posterior to shell-gland. Laurer's canal
present. Receptaculum seminis absent. Vitellaria, irregular follicles. Receptaculum
seminis uterinum present. Uterus in transverse coils. Metraterm muscular as long
as cirrus sac. Eggs thin-shelled, with long polar filaments. They measure -017- -02
mm. x -008- -01 mm. without filaments.
Host. Callus bankiva murghi (Rectal cseca).
Remarks. This form is remarkable in having the genital pore far
forwards at the posterior margin of the oral sucker. This position of the
gential pore appears to be unique for the genus. The vesicula seminalis
also is wholly external in Catatropis indicus. The importance of this
character was emphasized by Chatter ji (1933) who created a new genus
Canada in the subfamily Lepodermatinse on the presence of an external
vesicula seminalis, besides the usual internal vesicula seminalis. Owing
to the unique position of the genital pore, the absence of a median ridge or
keel for bearing ventral glands and lastly in the absence of an internal
vesicula seminalis the writer feels inclined to remove this species from the
Studies in H elm intho logy .125
genus Catatropis. Probably a re-study of more material of this form will
necessitate the creation of a new genus for this species.
Remarks on the family Notocotylidte. In the preceding pages and
also earlier in the pages of these Proceedings* the writer has given an
account of the subfamily Notocotylinse. The sub-families Nudacotylinse
and Ogmogasterinae possess such distinctive characters when compared
with Notocotylinae that there could be no doubt for their retention as
distinct stibfamilies. Yamaguti (1933) described a new genus Cymbiforma
from the intestine of a mammal, Sika nippon and created the subfamily
Cymbiforminse for its reception. The genus shows affinities with the
members of the subfamily Nudacotylinse but differs in the relative position
of the cirrus sac and uterine coils. Yamaguti also mentions, that Barker
(1915), in describing the subfamily Nudacotylinae, covered variations in
these characters as well. It would, thus, appear that the difference in the
relative position of uterine coils and cirrus sac should not form a sufficient
basis for the erection of a new subfamily Cymbiforminse. The question of a
mammalian host lodging this form may rise a little doubt but as already
discussed earlier in the Host-parasite relationship (vide these Proceedings,
1937, 5, No. 2, Sec. B, 33-44) this point may be negligible. The sub-
family Cymbiforminae may, therefore, be dropped and be regarded as a
synonym of the subfamily Nudacotylinse, and the genus Cymbiforma thus
comes under the subfamily Nudacotylinae.
Family Cydoccelidcz Kossack, 19H
Stossich (1902) created a subfamily Cyclocoelinae for the reception of
four genera, Cyclocoelum, H&matotrephus, Qphthalmophagus and Typhloccelum.
Kossack (1911) raised it to the status of a family, Cyclocoelidse and added to
it two genera, Hyptiasmus and Spaniometra. He also included the genus
Bothriogaster under this f amity. Harrah. (1922), while revising the mono-
stomes of North America, divided the family into three subfamilies :
(1) CycloccBlinse with Cyclocalum, H&matotrephus and Hyptiasmus.
(2) Typhloccelinse with Typhloccelum and Tracheophilus.
(3) Ophthalmophaginse with Ophthalmophagus, Bothriogdster and
Spaniometra.
Witenberg (1926) fused the subfamilies Ophthalmophaginse and Cyclo-
coelinse, leaving only two subfamilies, Cyclocoelinse and Typhloccelinae. The
* Vide 1935, 2, No. 5 ; and 1936, 3, No. 1.
126 Makund Behari Lai
main point on which Witenberg maintained only two subfamilies is tlu*
absence or presence of caaeal diverticula.
Family diagnosis. ^Txe.nmtodvs, large to medium size with strong
muscular flat body; small ventral sucker sometimes present. Month
terminal or subterminal, oral sucker rudimentary ; mostly absent,
Pharynx non-muscular. Intestinal caeca join each other at the posterior
end. They are either simple or provided with lateral diverticula on their
inner margin. Excretory pore dorsal and short, situated at the posterior
end. Genital pore median, not very much behind the mouth opening.
Copulatory organ present but feebly developed. Seminal vesicle lies inside
the cirrus sac. Vitellaria, lateral, between body-wall and intestine, some-
times run continuously along the intestine and meet at the posterior end.
Gonads intercsecal. Testes two, simple or pressed, and lie near each other.
Ovary, always entire and spherical. I/aurer's canal and receptaculum,
seminis exceptionally present. Uterus very strongly developed with regular
peculiarly pointed loops and extends from the posterior end up to the
intestinal bifurcation. Eggs numerous, without polar filaments and produc-
ing miracidia with two eyespots while still inside the uterus. Generally
parasitic in the body cavity, air-sacs and nasal opening of aquatic birds.
Exceptionally present in the intestinal canal.
Type genus Cyclocalum.
The family is divided into two subfamilies :
(1) Cyclocoelinse without lateral intestinal diverticula.
(2) Typhlocoelinse with lateral intestinal diverticula.
The Avian Trematodes reported from India under the subfamily Cyclo-
ccelinse belong to the following genera :
7. Hcematotre-phus.
2. Cycloccelum.
Key for the identification of the Indian genera of the subfamily Cyclo-
coelinse :
Ovary in front of both the testes which are not separated by uterine
coils Hcematotrephus.
Ovary in between two testes which are always separated by uterine
coils Cycloccelum.
Studies in Helminthology 127
Genus Hcematotrephus Stossicli, 1902
Only one species has been reported from avian host in India.
Hcematotrephus nebularium (Khan, 1.935)
(Syn. Cycloccelum nebularium Khan, 1935)
Trematodes with medium size, 10-13 mm. long and 2-3-5 mm. broad. Pharynx
well-developed. Oesophagus S-shaped or straight. Intestinal caeca devoid of lateral
diverticula and meet posteriorly in an arc -285 mm. in front of the posterior end.
Excretory bladder between the intestinal arc and posterior end of the body, with
dorsally placed excretory pore. The two round tostes lying obliquely behind the ovary
measure 0-7-1 mm. in diameter and are not separated by uterine, coils. Cirrus sac
tubular, dilated posteriorly, extending from the genital pore to the anterior end of the
intestinal bifurcation. Ovary rounded, smaller than testas, and measures -31 41 mm.
in diameter. Laurer's canal absent. Uterus in transvere loops with large thick-shelled
eggs. Vitellaria, confined to the extreme edges of the body. A small yolk reservoir
is present. Eggs, without operculum. showing miracidia with characteristic double
eyespot, and measuring -12 mm. X 087 mm. in size.
Host. Glottis nebularia (Abdominal air-sacs).
Locality. Allahabad.
Remarks. Khan who included this species under the genus Cycloccelum
has described a small structure lying inside the ovary which he calls a
receptaculum seminis. This position of receptaculum seminis is rather
peculiar. It is further interesting to note that Khan does not find any
sperms in it and he mentions that Morishita (1924) had termed a similar
structure as ootype. The present writer considers that the structure is
possibly not of the nature of receptaculum seminis which evidently is absent
in this form. Owing to the pretesticular position of the ovary, the species
is removed to the genus H&matotrephus.
Genus Cycloccelum Brandes, 1892
Nine species of this genus have so far been recorded from Avian hosts
in India. Out of these one Cycloccelum nebularium has just been allocated
to the genus Htzmatotrephus. Of the remaining eight, seven have been
described to possess a receptaculum seminis which is absent in the genus
Cycloccelum.
Cycloccelum erythropis Khan, 1935
Length 7-5-17 mm. Maximum breadth 1-7-2-3 mm. Oral sucker feeble
15 mm. x -1 mm. Pharynx -15 25 mm. in diameter. Oesophagus more or less
straight. Intestinal arc 15 mm. in front of the posterior end. Excretory bladder
12 -15 mm. by -IQ-^S^ mm. in size. Excretroy pore median, dorsally situated
at hinder end. Gonads in posterior fifth of the body. Testes spherical ; anterior testis
separated by four or five uterine coils from posterior testis. Vesicula seminalis straight
128 Makund Behari Lai
and inside the cirrus sac. Genital pore, vjentral, just behind the pharynx. Cirrus sac
club-shaped, reaching middle of intestinal "bifurcation. Ovary very small 19- -29 mm.
in diameter ; shell-gland mass spherical ; uterus runs forward in closely situated coils ;
nietraterm short. Vitellaria laterally pressed against body-wall. Thin-shelled ova.
Host. Tringa erythropus.
Locality.- Allahabad.
Remarks. This species definitely conforms to the characters of the
o-enus Cyclocahim. It does not possess a receptaculum seminis and differs
in this important feature from the other species which have been described
to possess a receptaculum seminis.
Cycloccehtm sharadi Bhalerao, 1.935
Trematodes with body tapering towards both ends ; medium size, measuring
10-5-11 mm. in length and 41- -5 mm. in maximum breadth. Oral sucker, muscular
measuring '44 mm. X -32 mm. Prepharynx short. Pharynx well developed,
muscular, and measures -425 mm. X -38 mm. Oesophagus short. Excretory bladder,
simple, flat sac, excretory pore opens dorsally by a short duct. Nerve ganglia on either
side of the pharynx, a dorsal nerve-band joins them. Genital glands in posterior fourth
of the body. Testes separated by uterine coils. Posterior testis in contact with intestinal
arc and measures 2 15 mm. X 1 -32 mm. Anterior testis measures 2 -07 mm, X 1 -7 mm.
and is in contact with left intestinal caecum. Cirrus sac small with cirrus in the form of
horizontal S. Vesicula seminalis large, fills up the cirrus sac. Pars prostatica non-
cellular ; prostate glands present. Male opening in the genital atrium. Genital atrium
large and situated posterior to the pharynx, genital pore slightly behind pharynx.
Ovary, between two testes, oval, measures -5 mm. x -43 mm. Shell-gland, larger
than ovary on the left of the latter. Receptaculum seminis, oval, on the left of ovary
between it and testis. It measures -36 mm. x -275 mm. Vitellaria between ca3ca
and margin of body, extending upto hinder border of pharynx. Laurer's canal absent.
Uterine coils in posterior rd, intercsecal, metraterm strongly muscular, surrounded with
gland-cells. Long and elliptical eggs -123- -14 mm. x- 06--OSlmm.
Host. Urocissa favirostris cucullata (Thoracic cavity).
Locality. Muktesar.
Cycloccelum allahabadi Khan, 1.935
Size 17 mm. X 2-5-3 mm. Oral sucker flat. Pharynx -28 mm. in diameter.
Oesophagus S-shaped. Testes separated from each other by uterine coils ; Genital pore
at posterior end of pharynx. Cirrus sac club-shaped, extending upto middle of
intestinal bifurcation. Ovary, -3-35 mm. x -2- -25 mm. in size. Recepta-
culum seminis inside the ovary and well developed. Receptaculum seminis uterinum
absent. Shell-gland posterior and left to ovary. Uterus fills up the body between
posterior testis and intestinal bifurcation. Vitellaria, extending over c^eca at places
and reaching uterine coils. Ova thin-shelled and operculate, -119 mm. x -08 mm.
Host. Tringa erythropus (Thoracic air-sac).
Locality. Allahabad.
Studies in Helminthology 129
Cycloccelum capellum Khan, 1.935
Body 17-25 mm. X 3'5-lG mm. Oral sucker rudimentary. Pharynx *275 mm.
*n diameter. Oesophagus S-sliaped or straight. Excretory pore dorsal and terminal.
Tostes separated from each other by uterine coils ; anterior testis to the right side and
8mm. X (5 Sinm. in size ; posterior, median, much-lobed, 1-1 mm. X -7- -88 mm.
in size. Genital pore at posterior margin of pharynx. Cirrus sac flask-shaped and
hardly reaching intestinal bifurcation. Ovary to the left side. -37- -55 mm. x
37- -5 mm. Shell-gland behind ovary. Receptaculum seminis pear-shaped on the
inner side of the ovary. Reeeptaeuhun seminis uterinum filled with sperms. Uterus
thrown into double loops, never overlapping craca. Vitellaria, irregular in extension
on two sides ; yolk reservoir present, 15 mm. long. Ova with fully developed miracidia,
13 mm. X ()(.> 8 mm.
Host. Capella gallinago (Cervical air-sacs).
Locality. Allahabad.
Cycloccelum straightum Khan, 1935
Body long, 25 mm. X 4 -3 mm. in size ; oral sucker feebly developed, ventral sucker
absent. Pharynx -344 mm. X -425 mm. Oesophagus S-shaped or straight. Excre-
tory pore dorsal and terminal. Testes separated from each other by uterine coils ;
anterior to the right side and 1 -05 mm. x '68 mm. ; posterior mesial, 1 -25 mm. x
99mm. Cirrus sac nearly ilask-shaped reaching to 15 mm. in front of intestinal
bifurcation. Pars prostatica, prostate glands and cirrus not observed. Ovary, median,
45 mm. X -41. mm. in size. Shell-gland to left of ovary ; receptaculurn seminis imme-
diately behind ovary, Laurer's canal absent. Receptaculum seminis uterinum large,
filled with sperms. Uterus filling up the space upto intestinal bifurcation. Vitellaria
from middle of intestinal bifurcation to excretory bladder. A small yolk reservoir
present. Ova, thick-shelled, non-operculate, with fully developed miracidia, 130 mm.
X -008 mm.
Host. Glottis nebularia (Abdominal air-sac).
Locality. Phulpur, Allahabad.
Cycloccelum indicum Khan, 1935
Length 20-27 rain. Breadth 4-4 -5 mrn. Rudimentary oral sucker. Pharynx
28mm. in diameter. Oesophagus straight. Excretory pore dorsal at hinder end.
Testes separated from each other by uterine coils ; anterior, to the left and -85 mm. in
diameter; posterior, in the intestinal arc -85- -93 mm. in diameter. Cirrus sac
club-shaped. Genital pore behind pharynx, median. Ovary -5- -6 mm. in diameter;
shell-gland near right wall of ovary ; receptaculurn seminis elongated inside the ovary.
Receptaculum seminis uterinum present. Uterus with widely separated loops, extending
over cseca and reaching body wall. Vitellaria irregular in extension on the two sides.
Ova, thin -shelled with fully developed miracidia, measure 0-12 mm. x -068mm.
Host. Glottis nebularia (Body cavity).
Locality. Allahabad.
130 Makund Behari Lai
Cycloccelum mehrii Khan, 1935
Length 18-28 mm., breadth 3 4-5 mm. Oral sucker very rudimentary. Pharynx
muscular 27 mm. in diameter. Oesophagus S-shaped. Excretory pore posterior
and dorso-median. Testes separated from each other by uterine coils ; anterior, lateral,
and -85 mm. x 0-7-1 mm. ; posterior in intestinal arc, -9-1 3 mm. in. diameter.
Cirrus sac retort-shaped, extending behind anterior wall of intestinal bifurcation.
Genital pore, ventral to middle of pharynx. Ovary 4-6 mm. in diameter ; shell-
gland close behind ovary ; racsptaculum seminis, small pear-shaped inside the ovary.
Receptaculum seminis uterinum large. Uterus, with well-separated loops, overlapping
ca&2a at various places. Vitellaria, dense, extending from middle of intestinal bifurca-
tion to excretory bladder. Yolk reservoir small. Ova, thin-shelled, with fully
developed miracidia, -12 mm. X -068mm.
Host. Capella gallinago gallinago.
Locality. ?
Cycloccelum lobatum Khan, 1935
Length 13 mm., breadth 2 -4 mm. Oral sucker not visible in whole mount.
Pharynx muscular, -27 mm. in diameter. Oesophagus straight. Excretory pore
median and dorso-terminal. Testes separated from each other by uterine coils ; anterior
testis, lateral and equal in size to the posterior; posterior median, 6- -75 mm. X
55- 7 mm. Genital pore at middle of pharynx. Cirrus sac more or less flask-
shaped with tubular anterior end and sac-like posterior part reaching just behind anterior
end of intestinal bifurcation. Ovary, -4- -5 mm. x -35- -55 mm. Shell-gland
present. Receptaculum seminis inside the ovary. Receptaculum seminis uterinum
absent. Uterus between posterior end of shell-gland and intestinal bifurcation.
Vitellaria from posterior end of cirrus sac to excretory bladder. Ova, thin-shelled and
non-o perculate, with fully developed miracidia, 119mm. x -068 mm.
Host. Glottis nebularia (Thoracic cavity).
Locality. Allahabad.
Remarks on the genus Cycloccelum. The species Cycloccelum sharadi,
Cycloccelum allhabadi, Cycloccelum stmightum, Cycloccelum capellum, Cyclo-
ccelum indicum, Cycloccelum mehrii and Cycloccelum lobatum have been
described to possess a receptaculum seminis. This structure is not present
in the type species of the genus Cycloccelum. In view of the opinion already
expressed (I^al, 1937) on the systematic value of this character it may be
considered desirable to exclude all these species which possess a recepta-
culum seminis from the genus Cycloccelum. It is suggested that a new
genus Receptoccelum, characterised by the presence of receptaculum seminis,
may be created for* their reception.
Subfamily Typhloc&lina Harrah, 1922
Under the subfamily Typhlocoelinae, the author discovered a genus
Typhlophilus I/al, 1936, full details of which have already been published
in a previous number of these Proceedings (vide 1936, 4, No. 1, Sec. B).
Studies in Helminthology 131
Genus Typhlophilus I^al, 1936
Distome, with a flat ribbon-like body, and witli an extremely feeble
funnel-shaped oral sucker and a small muscular ventral sucker. Curved
prepharynx, globular pharynx and extremely small oesophagus. Intestinal
caeca, provided with about 10 diverticula on the inner margin, meet in the
middle posteriorly to form the Intestinal Bow. Excretory bladder crescent-
shaped, excretory pore dorsal and sub-terminal. Testes, two, in grape-like
bunches separated from each other by the first coil of the uterus. The
anterior testis lies on the left and near intestinal csecum, the posterior
fills the arch near the Intestinal Bow. Ovary oval, on the right side at the
level of the left testis. Ootype, shell-gland and a small receptaculum
seminis present between the ovary and the posterior testis. Genital pore
situated ventral to the intestinal bifurcation, immediately behind the
pharynx. Eggs small, thin-shelled, without filaments.
Type species Typhlophilus shovellus.
Typhlophilus shovellus I^al, 1936
Ribbon-like and grey coloured trematode ; 3-6 mm. long and 1-1.5 mm.
broad. Oral sucker extremely feeble, -19mm. x -195mm. Ventral
sucker circular and muscular, about the middle of the body, 125 mm. in
diameter. Prepharynx -2mm. long, Pharynx thick- walled, -19mm. x
-21mm. Oesophagus extremely small. Intestinal caeca provided with
10 diverticula on their inner margin and join posteriorly. Excretory
bladder -35mm. long. Anterior testis 125mm. x -06mm. Posterior
testis -1.5 mm. X -07mm. Cirrus sac -25mm. long. Ovary at the level
of the left testis, -055mm. x -07mm. Uterus full of small eggs, eggs
measure -02mm. X -01 mm. with thin shell and not provided with any
polar filament.
Host. Spatula clypeata (Small intestine).
Locality. Lucknow.
Remarks on the family Cycloccelidce. The retention of the two sub-
families Cycloccelinse and Typhlocoelinse by Witenberg (1926) on the basis of
caecal diverticula was questioned by Joyeux and Baer (1927) who regard
the c^ecal diverticula of only generic importance in the same way as the
relative position of gonads. The writer (I/al, 1936) has previously shown
that the various genera of Cyclocoelidse can be arranged under two distinct
groups on the basis of csecal diverticula. Both groups have forms which
show variations in the relative position of the gonads, the nature of uterine
132 Makund Beliarl Lai
coils and the position of genital pore, and thus the presence or absence of
c^cal diverticula seem to form the natural subdivisions of the iaimiy.
Cameron (1934) and other earlier workers have considered this character m
the division of the family Pasciolidse into two subfamilies, Fasciohme and
Pasciolopsinse. It is, therefore, desirable to retain here Witenberg's sub-
divisions into Cycloccelinse and Typhloccelmse.
The subfamily Typhlocoelinse has three distinct genera but, as men-
tioned by Witenberg (1926), the subfamily Cycloccelinse has got a vast
assemblage of forms. Joyeux and Baer (1927) have been rather conser-
vative and do not consider the various genera of Witenberg as valid. They
suggest the fusion of all the genera of the subfamily Cycloccelinse into two
forms, vis., Cydoccehm and Spaniometra which they distinguish by the
relative position of the ovary and testes. The genus Cycloc&lum, according
to Joyeux and Baer, contains all those forms which have their ovary either
pre-testicular or intertesticular. Although Witenberg's classification of
the family into genera is not free from defects the writer does not agree
with Joyeux and Baer in their contention. As already stated the relative
position of the gonads has been greatly emphasised in several cases and
should here be regarded as a character of at least generic importance. The
position of the uterine coils is another character of sufficient importance
and combined with the relative position of the ovary merits consideration
in establishing the genera of the subfamily Cycloccelinse. Therefore, the
members of the subfamily Cyclocoelinse can be grouped into four distinct
divisions, as follows :
1. Ovary in front of the two testes, not separated by uterine coils,
e.g., Hcematotrephus.
Other forms are Wardianwn, Htematoprimum, Corpopyrum and
Uvitellina.
2. Ovary in between the two testes separated by uterine coils, e.g.,
Cycloccelum.
Other forms are Cycloprimum, Harmhium, Prohyptiasmus, Hyplias-
mus, Allopyge and Transccelum.
3. Ovary behind both the testes, not separated by uterine coils, e.g.,
Contraccelum.
4. Ovary behind both the testes which are separated by uterine coils,
e.g., Ophthalmophagus. The genus Spaniometra also comes under
this.
Studies in Helminthology 133
Each division is represented by a well-established genus and the other
previously known genera put under each group may be regarded as mexe
synonyms. Now, there remain only two genera, viz., Bothriogaster* and
Morishitium which have not been accounted for here. Both of these possess
ventral sucker and it would be better to keep them as distinct genera till
more information is available about them.
Diagnosis of the genus Hcematotrephns Stossicli, 1902, emended.
(Syn. Wardianum Witenberg, 1926 ; H&matoprimwn Witenberg, 192G ;
Corpopyrum Witenberg, 1926 and Umtettina Witenberg, 1926.)
Cyclocoelinae with flat body ; anterior end narrower than the posterior ;
oral sucker not well developed, subterminal ; ventral sucker absent ;
intestinal cseca smooth, without any lateral projections, extend up to the
posterior end of the animal, meeting in the middle line ; testes two, almost
spherical, lying in the posterior broad end of the animal and not separated
by uterine coils ; ovary spherical, smaller than testes, always in front of both
testes, vitelline glands extend along the intestinal cseca.
Diagnosis of the genus Cycloccelum Brandes, 1892, emended.
(Syn. Cycloprimum Witenberg, 1926 ; Harrahium Witenberg, 1926 ;
Prohyptiasmus Witenberg, 1926 ; Hyptiasmus Kossack, 191.1 ; Allopyge
Johnston, 1913 and Transccclum Witenberg, 1926.)
Cyclocoelinae with Hat body ; anterior end narrower or may be as broad
as posterior ; oral sucker more or less funnel-shaped, weakly developed ;
ventral sucker absent ; intestinal caeca, smooth without projections, run
almost up to the posterior end of animal where they meet together in the
middle ; testes two, almost spherical, the posterior one lying in the " In-
testinal Bow," the anterior further forwards, separated from each other
by uterine coils ; ovary spherical, smaller than testes, lying always in
between the two testes ; vitelline glands extend along the intestinal oeca.
Diagnosis of the genus Contracoelum Witenberg, 1926, emended.
Cycloccelinse with flat spindle-shaped body ; anterior and posterior ends
both narrower than the middle ; oral sucker quite terminal ; ventral sucker
absent ; intestinal cseca, smooth without any lateral projection, run almost
* The name Boihriogasler is not available for the generic name since it is preoccupied
by Bothriog aster Sselivanow, 1879 (Myriapoda). In view of this, the name will have to
be changed.
134 Makund Behari Lai
up to the posterior end of the animal where they meet together in the middle ;
testes two, almost spherical, lying side by side, more or less conti-
guous and not separated by uterine coils, in the middle of the body ; ovary
oval, smaller than testes, tying behind both testes in the " Intestinal Bow "
vitelline glands extend along the intestinal caeca.
Diagnosis of the genus Ophthalmophagus Stossich, 1902, emended.
(Syn. Spaniometra Kossack, 1911.)
Cyclocoelinse with flat body ; anterior end narrower or as broad as the
posterior end ; oral sucker weakly developed ; ventral sucker absent ;
intestinal cseca smooth, without lateral projections, run almost up to the
posterior end of animal where they meet in the middle ; testes two, almost
spherical situated in the intercaecal region and separated by intervention of
uterine coils ; ovary spherical, smaller than the testes, lying always behind
both the testes ; vitelline glands extend along the intestinal cseca.
Family Dicrocoeliidcz Looss, 1907
The family Dicrocoeliidse contains numerous genera and has been a
subject of investigation by a large number of workers. The latest compre-
hensive work is that of Poche (1925) where he has recognised not less than
20 different genera. The main features on which the generic division is
based in this family is the relative position of the testes. The distinction
and arrangement of the various genera under the family is rather confused
and a revision of the family will have to be done. The family Dicrocoeli-
idse has the following diagnosis :
Trematodes small to medium in size with a more or less elongated,
flattened, translucent and non-muscular body. Pharynx and oesophagus
present. The intestinal cseca are simple and do not quite reach the posterior
end of the body. The excretory bladder is tubular or sac-like. The genital
pore is situated in the middle line between the oral and ventral suckers.
The testes are situated in level with or behind the ventral sucker and in
front of the ovary, opposite to each other or one behind the other. The
cirrus sac is small and does not extend behind the anterior margin of the
ventral sucker. The ovary is situated behind the testes. The vitelline
glands are well developed and are laterally situated, partly overlapping the
intestinal caeca. The uterus occupies most of the space behind the genital
glands. Eggs relatively small.
Type genus Dicroccelum.
Studies in Helntinthology 135
Among birds in India, the family is represented by three genera :
1. Genus Lyperosomum Looss, 1899.
2. Genus Platynosomum L,ooss, 1907.
3. Genus Multimtellaria phadke and Gulati; 1930.
Genus Lyperosomum I v ooss, 1899
Looss created the genus Lyperosomum for the reception of L. porrectum,
L. longicauda, L. strongylosum and L. plesiostomum. Braun (1901) added
three species to the genus, viz., L. corrigia, L. salebrasum and L. rudectum.
Von I v instow (1906) described L. squamatum but this was later removed
from the genus by Skrjabin (1913). I,ooss again (1907) included under the
genus, L. lobatum, L. alssoni and L. dathratwn, the last one, however, was
assigned later to the genus Platynosomum. Skrjabin (1913) added L. fili-
forme. Nicoll (1914) described L. sdtulum and L. direptum of which the
latter was transferred later to the genus Oswaldoia by Travassos (1919).
Johnston (1917) described three species L. parvium, L. megastomum and
L. harrisoni. In the same year, Travassos (1917) added L. obliqum, L. trans-
versum, L. rarurn, L. lari and L. sinuosum. Isaitschicoff (1919) described
L. donicum and L. attenuatum. Layman (1922) added three species
L. rnagnitestium, L. vanellicola and L. transverso-genitalis. The same author
(1923) described L. fringillce and again (1926) L. lanicola, L. asowi,
L. loossi andL. alaudcz. Baylis (1927) transferred Distoma vitta as L. vitta.
The latest comprehensive work is that of Skrj,abin and Udinzew (1930) who
also described a new species L. papabejani and besides have given a useful
key for the identification of the various species. Price (1935) described
a peculiar species L. monenteron which shows only a single intestinal caecum.
He thinks that probably this condition of the intestine may be present in
some other, species. The writer considers this feature as very important
and thinks that such forms should not be included in the genus. In fact,
it appears necessary to create a separate genus for the form described by
Price.
Diagnosis of the genus Lyperosomum LOOSS, 1899.
Dicrocceliida. Body elongated ribbon-shaped; spherical in section.
Testes tandem behind the ventral sucker. Vitellaria behind the testes.
Other characters as of the family.
From India two species have so far been described under the genus, one
by Bhalerao (1926) and the other by Patwardhan (1935).
B3
136 Mafeund Behari Lai
Lyperosomum kakea Bhalerao, 1926
Body elongated, tapering at both ends ; cuticle without spines. Length 3 -36 mm.
and breadth (maximum at the level of ventral sucker) -33 mm. Oral sucker terminal,
13 mm. X -15 mm. Ventral sucker, situated at l/5th of the distance from the anterior
end, measures '2mm. in diameter. Prepharynx absent. Pharynx, globular, measur-
ing -06 mm.* in diameter. Intestinal caeca up to the posterior end of body. Tastes,
oval, one behind the other some distance behind ventral, sucker placed centrally with
their axes oblique and measure -IS-- 18 mm. x -12- -125 mm. Genital pore some
distance "behind pharynx. Cirrus sac pear-shaped, lies centrally, about -28 mm.
anterior to ventral sucker and contains a coiled vesicula seminalis. Pars prostatica,
ductus ejaculatorius and cirrus small. Ovary ovoid, behind testes, in central line and
measures -165mm. X 12mm. Receptaculum seminis immediately behind it.
Laurer's canal present. Shell-gland, post-ovarial. Uterine coils fill up posterior part
of body behind the ovary. Vitellaria small, extending a little behind ovary. Eggs,
oval, operculate measuring -028- *03 mm. X -016- -018 mm.
Host. Corvus insolens (Liver).
Locality. Rangoon .
Lyperosomum colorosum Patwardhan, 1935
Body elongated, tapering at both ends and measures 1*7-3 -0mm. in length.
Maximum breadth -192 --34 mm. at the level of acetabulum. Integument smooth;
parenchyma with scattered pigment particles. Oral sucker, subterminal, elliptical,
076 -134mm. X 09216 mm. Pharynx muscular and spherical, -043--075 mm.
in diameter. Oesophagus -06- -1mm. long. Intestinal caeca extend up to 4. /5th
of the worm, posteriorly. Acetabulum * 138- -24 mm. in diameter, and is 356-- 63 mm.
behind the anterior end of the body. Testes, two, post-acetabular, one behind
the other. Anterior testis spherical '127- -22 mm. in diameter. Posterior testis,
oval, separated from anterior by a uterine loop and measures 108- 208 mm., -123-
. 217 mm. Genital pora midway between acetabulum and anterior end of body.
Cirrus pouch elongated, pyriform -12- -21 mm. X -04- -07 mm. It contains large
vesicula seminalis. Ovary spherical, smaller than testes, '04 mm. behind pos-
terior testis, in the median line. It measures -084- -148 mm. in diameter.
Receptaculum. seminis dorsal and lateral to ovary and measures '023 04 mm.
in diameter. Shell-gland is similar in position but slightly ventral. Vitellaria, post-
ovarial, in two lateral rows of 6-7 follicles on each side. Uterus extends backwards
as far as posterior end of body. Eggs large, elliptical, thick-shelled and measure
0125 -022mm. X -025 -04mm.
Host. Temenuchus pagodarum.
Locality. Nagpur.
Key for the identification of the species of Lyperosomum :
Ovary larger than both testes L. kakea.
Ovary smaller than both testes L. colorosum.
* In the original paper it is mentioned as 6 mm. which is probably a misprint.
Studies in Helminthology 137
Genus Platynosomum Looss, 1907
The genus Platynosomum was created by I,ooss (1907) for a trematode,
obtained from Circatus gallicus, which he named P. Semifuscum. Later
on several authors added to this genus. Kossack (1910) described
P. fastesum, Nicoll (1915) recorded P. acuminatum horn Kestrel and also
transferred two species of Dicwccelium as P. defiectens and P. petiolatum.
Travassos (1916) added P. microchis and later again (1918) P. arietis.
Tubangui (1928) reported P. phillippinerum. The latest work on the genus
is that of Cameron (1928) where he described P. planicipites from a tiger
cat.
Diagnosis of the genus Platynosomum Looss, 1907.
Dicrocceliida. Body lancet-shaped, greatest width at the level of the
testes. Anterior end more pointed than posterior. Suckers almost of
equal size. Testes symmetrical, at equal height behind ventral sucker.
Cirrus sac plump, sac-shaped. Ovary lies behind the testes. Other
characters as of the family.
The only form recorded from India is P. acuminatum by Bhalerao
(1926) which was previously obtained by Nicoll (1915) from Kestrel in
Scotland.
Platynosomum acuminatum Nicoll, 1915
Host. Corvus insolens (L,iver).
Locality. Rangoon.
Remarks. Bhalerao gives no description or figure but mentions that
the only variations from Nicoll's description, that he finds in his specimens,
are in the dimensions of the body and the posterior extension of vitellaria.
Since the writer has no specimens, he cannot add any observations on the
subject. A summary of Nicoll's description is, however, given below :
Length 6 -3 mm., maximum breadth just behind ventral sucker is 1-5 mm. Oral
sucker twisted to right, -45 mm. X -40 mm. Ventral sucker is -6 mm. X -75 mm.,
situated 1-97 mm. from, the anterior end. Pharnyx contiguous with oral sucker and
measures 17mm. X -15mm. Oesophagus short. Intestinal diverticula long and
narrow. G-enital pore ov r er the pharynx, twisted to right. Cirrus pouch -7 mm. X
16mm. contains vesicula seminalis, pars prostatica and ductus ejaculatorius. Testes
symmetrical, immediately behind ventral sucker and measure -25 mm. X -36 mm.
Ovary behind left testis, oval. Vitellaria lateral, between leva! of testes, and 2-3 mm.
from posterior end ; follicles small. Uterus fills up greater portion of post-acetabular
space ; convolutions entirely confined behind ventral sucker. Vagina weakly
developed. Eggs numerous, measure -033- -039 mm. X -018- -02 mm.
138 Makund Behari Lai
Genus Multivitellaria Pfaadke and Gulati, 1930
Body thick and opaque, ventrally convex. Pharynx present. Ovary
post-testicular and median above the fork of the excretory bladder. Testes
lateral and lying under the caeca. Twenty-eight pairs of vitellaria lying
exterior to the caeca. Follicles of the vitelline glands tubular. Genital
pore sucker-like close to and in front of the acetabulum and lying behind
the gut-fork. Bxcretory bladder Y-shaped.
Type-species Multivitellaria hewletti.
Multivitellaria hewletti Phadke and Gulati, 1930
Body thick and opaque and ventrally convex. Length 6 -0-16 -6 mm., breadth
4 -0-8 '7mm. Oral sucker smaller and less prominent than acstabulum. Pharynx
present. Oesophagus *5 mm. long ; intestinal caesa, straight or slightly curved
in a zigzag manner. Testes, two, lateral, pre-ovarial, lying under the caeca. Longi-
tudinal sections of the fluke showed the presence of seminal vesicle and pars
prostatica in close proximity of the cirrus sac. Genital pore, sucker-like, close to and
in front of acetabulum, and lying b3hind the intssbinal fork. Ovary, situated medially
adjacent to shell-gland, recsptaculum seminis, and Laurer's canal. Vitellaria, 28 pairs,
lying exterior to cfleca. Excretory bladder Y-shaped. Uterus with loops ; eggs
78-88^ by 34-53^.
Host. Corvus splendens (Gall bladder).
Locality. Bombay.
Remarks. This form shows the usual characters of the family Dicro-
cceliidae but differs from all the genera in possessing a thick and fleshy body,
much greater extension of the vitellaria, and a sucker-like genital pore. The
writer, therefore, fully agrees with the authors in regarding this form as a
distinct genus. Phadke and Gulati have also suggested a new subfamily
Multivitellarinse for the reception of this form.
Key for the identification of Avian genera of the family Dicrocoeliidae
reported from India :
1. Testes tandem . . Lyperosomum.
Testes connubial . . 2
2. Vitellaria extend throughout the
body length genital sucker
present . . . . Multivitellaria.
Vitellaria confined to the middle
of the body genital sucker
absent . . . . . . Plcttynosomum.
Studies in Helminthology 139
Family Opisthorchiidce Braun, 1901
The family Opisthorchiidae contains about a dozen genera and attempts
were made to divide the family into subfamilies but Morgan (1927) has
pointed out the undesirability of this course of action owing to absence of
any constant differences. Further, Morgan (1927) has suggested the syno-
nymy of Notauhis to Opisthorchis and the writer agrees with him. In fact
several other genera, viz., Amphimerus and Cyclorchis which are recognised
by Morgan as distinct from Opisthorchis also do not show those distinctive
features which should be taken as characters of generic importance. Morgan
(1927) further suggests the fusion of the genera Clonorchis and Opisthorchis
on the basis of the branching of the testes but the writer thinks that the
difference in the excretory bladder in the two cases is rather much pro-
nounced. It would, therefore, be desirable to keep the two genera as
distinct. Recently Yamaguti (1933) described a genus Oesophagicola
from the oesophagus of a marine snake and created a new subfamily Oeso-
phagicolinse for its reception, under the family Opisthorchiidae. The genus,
in question, no doubt shows important differences from the members of the
family Opisthorchiidae with which it resembles in the absence of a cirrus
sac and the relative position of the gonads. It is, however, not possible,
for the writer at present to make any comments on the systematic position
of this genus.
Diagnosis of the family Opisthorchiidce Braun, 1901 ; emended Morgan,
1927.
Body flat and transparent, sometimes much thickened, narrowing
anteriorly. Suckers near each other (except in Microtrema), often only
moderately developed or atrophied. Pharynx present and usually followed
by a short oesophagus. Intestinal caeca long and unbranched, not always
reaching the posterior end. Excretory vesicle Y-shaped with proportion-
ately long stem and short branches, opening at the posterior end or on the
ventral surface in the region of the testes. In the former case the stem winds
S-sliaped between the testes or may lie dorsal to the testes. The genital
opening is immediately in front of the ventral sucker. A cirrus sac is
absent. The testes lie near the hind end of the body, obliquely or directly
behind one another ; sometimes they lie laterally. They are simple, lobed
or dendritic. The ovary lies immediately in front of the testes and may be
simple, lobed or multilobed. lyaurer's canal present ; receptaculum seminis
usually strongly developed. Vitellaria moderately well developed, divided
into acini or continuous and lie between the caeca and the margins of the
body. Uterine folds in front of the testes and mostly in front of the ovary,
140 Makund Beharl Lai
extending to or in front of the ventral sucker. Eggs numerous, small and
light-brown in colour. In the gallbladder and bile ducts of mammals,
birds, reptiles and fish.
The family is represented among birds in India by a single genus
Opisthorchis.
Genus Opisthorchis Blanchard, 1.898
A very large number of species have been described under the genus
and their diagnostic characters are based on the nature of the cuticle, ratio
of oral to ventral sucker, disposition of the gonads, nature of the excretory
bladder, the position of the ventral sucker, etc. In fact, there are some
species which do not show any clear differentiation and will probably be
proved as synonymous. But in the present state of our knowledge and
also in the absence of data on the life-history, it is not possible yet to suggest
any change in their nomenclature, although Price (1932), on morphological
grounds alone, has emphasised the identity of several species.
Morgan (1927) described 0. dendriticus from the liver of a Saras crane,
Antigone imported into the I/ondon Zoological Gardens from India.
The writer has been able to collect two species of the genus from the aviaii
hosts in India ; and although he is fully aware of the undesirability of
multiplication of species he finds it necessary to do so as the specimens
collected present certain important differences from the existing species
of the genus. They are, therefore, being described here as new species.
The diagnosis of the genus Opisthorchis given by Morgan (1927) is also
slightly emended.
Diagnosis of the genus Opisthorchis emended.
Opisthorchiidse ; body distinctly, often very greatly elongated, some-
times pear-shaped, anterior end attenuated ; posterior end broader. Skin
generally smooth. Excretory vesicle usually Y-shaped with long sigmoid
stem winding between the testes or passing dorsal to them. Copulatory
organs absent. Testes in the posterior portion of the body, and placed
either obliquely or directly behind one another. . Ovary simple or lobed.
Laurer's canal present. Receptaculum seminis prominent. Uterine coils
extend from the ovary to the ventral sucker, sometimes slightly overlap the
intestinal cseca. Vitellaria moderately developed, lateral of the intestinal
caeca and not extending anterior of the ventral sucker, ending posteriorly
at the level of the ovary. Vitellaria do not always form one region.
Type species Opisthorchis /elineus.
Studies in Helminthology
141
Opisthorchis gliddhis n. sp. (Figs. 1 and 2)
About a dozen specimens of this trernatode were obtained from the
liver of Sarcogyps calvus shot by the writer in Badshahbagh, I^ucknow.
The parasites appeared pink in fresh condition and their anterior end was
very contractile and movable.
The body is more or less pear-shaped, and measures 3 - 9 mm. x 1 5 mm.
The oral sucker is ventro-terminal and measures -15 mm. x -2mm. The
ventral sucker lies at a distance of 1-275 mm. from the anterior end and
measures -175mm. x -2 mm. The mouth leads into a short prepharynx
05mm. in length. The pharynx is elliptical and measures -15mm. x
llmm. The oesophagus is thick-walled and measures -3mm. in length.
The intestinal cseca extend almost up to the posterior end of the body.
ph.
ut.
OS mm.
FIG. 2
e.p,
FIG.
FIG.
FIG. 1
1. Opisthorchis giddhis n. sp., entire worm. dorsal view.
2 . Opisthorchis giddhis eggs .
The excretory bladder is Y-shaped and opens by the excretory pore at
the postero-dorsal end of the body. The median stein of the ' Y ' passes
in between the testes and bifurcates into the lateral horns between the
anterior testis and the receptaculum seminis,
142 Makund Beharl Lai
The testes are ovoid bodies with notched margin and lie obliquely one
behind the other at the posterior end of the body. The anterior testis
which lies on the left side measures -4 mm. x -15 mm. The posterior testis
measures -475mm. x -16 mm. The vasa deferentia unite to form a coiled
vesicula seminalis in the region of the metraterm. The vesicula seminalis
runs along the right margin of the ventral sucker and opens immediately
in front of it at the genital pore.
The ovary is an elongated, flattened, transversely placed body measuring
42mm. x -1mm. A well-developed receptaculum seminis lies behind
the ovary and measures -2mm. x -15mm. The duct from the recepta-
culum seminis forms a loop near the left end of the ovary before entering
the ootype. A small oviduct leads from the ovary to open into the ootype
which lies in front of the ovary and is surrounded by small unicellular shell-
glands.
The vitellaria consist of small follicles which stretch extra-caecally in
the region of the uterus. The vitelline zone on the left side is broken up
into groups while on the right side it is more or less continuous except for
a slight anterior part. The transverse vitelline ducts run in front of the
ovary and meet to form a very small yolk reservoir from which a small
duct opens into the ootype.
The uterus arises from the left of the ootype and runs forward in peculiar
transverse loops. It ends in an elongated metraterm and opens in front of
the ventral sucker. The eggs are numerous but small and measure
0225 mm. x -01 mm.
Remarks. This form differs from all the known species of the genus
Opisthorchis in having a peculiarly thick- walled oesophagus. The nearest
ally of this form is 0. obsequens from which it differs in possessing distinct
prepharynx, very feebly lobed testes, nature and extent of vitellaria and
much smaller eggs. It is, therefore, regarded as a new species.
Opisthorchis cheelis n. sp. (Figs. 3 & 4)
A large number of these trematodes were collected by Dr. G. S. Thapar
from the liver of the common kite, Milvus migrans. The body of the animal
is elongated and leaf -like and measures 6 8 mm. x 1 . 5 mm. The cuticle
is thin and sparingly spinose, specially in the anterior half of the body.
The oral sucker is terminal and measures 2 mm. in diameter. The
ventral sucker which lies at a distance of 1-65 mm. from the anterior end is
-2 mm. x -15 mm. in size.
S Indies in Helminthology
143
pfe
o.s..
oes.
5 /77/T7.
FIG. 4
e.p.
FIG. 3
FIG. S.OpisthorcMs checlis n. sp., entire worm, ventral view. Body spines
not shown.
FIG. ^.Opisthorch'is checlis eggs.
The niouth leads into an extremely small prepharynx which, is followed
by a globular pharynx -095mm. X -085mm. in size. The oesophagus is
thin- walled and measures 3 mm. The two intestinal caeca extend upto the
posterior end of the body.
The excretory bladder is Y-shaped. The median stem is not S-shaped
but slightly sinuous and runs in between the two testes. It divides into
the two horns just behind the receptaculum seminis. The excretory pore is
dorsal and terminal.
144 Makund Behari Lai
The two testes are obliquely placed behind each other. The anterior
testis which, is situated slightly towards the left is four-lobed and measures
48 mm. X -42 mm. The posterior testis is ovoid and measures -4 mm. X
3 mm. The vasa deferentia unite to form the vesicula seminalis which is
very slightly coiled and runs along the right border of the ventral sucker
to open at the genital pore which lies immediately in front of it.
The ovary is somewhat bilobed and median in position. It measures
4mm. x -275 mm. The receptaculum seminis is an elongated body,
slightly constricted in the middle and lies immediately behind the ovary.
It measures 3 mm. x 1 mm. A small duct leads from the ovary into the
ootype which is situated a little to the left of the ovary and is surrounded
by unicellular shell-glands.
The vitellaria are more or less continuous follicles, extra-csecal, and lie
in the middle region of the body. The vitelline zone on the right side is
slightly smaller than the left. The vitelline ducts from the two sides run
backwards in the form of a ' V ' and meet to form a very small yolk reservoir
from which a narrow duct leads into the ootype.
The uterus arises from the right side of the ootype and runs forward in
inter-csecal transverse loops and opens at the genital pore immediately
in front of the ventral sucker. It contains a large number of eggs which
measure 025mm. x -Olmm.
Remarks. This species differs from all the existing species of the genus
except 0. viverrini, 0. piscicola, 0. caninus, 0. noverca and 0. pedicellata in
having its cuticle armed with minute and sparingly arranged spines.
From 0. viverrini it differs, however, in having the oral sucker slightly
larger than the ventral, much longer oesophagus, slightly lobed ovary,
disproportionately developed vitellaria and smaller eggs.
From 0. piscicola it differs in having the ventral sucker smaller than
the oral, oblique position of testes, disproportionate vitellaria confined to
almost middle third of the body, and in having larger eggs.
From 0. caninus it differs in having a non-pedicled ventral sucker
situated at double the greater distance from anterior end, a very small
prepharynx, much longer oesophagus, extension of intestinal caeca up to the
posterior end, more or less straight (not coiled) vesicula seminalis, very
slight lobation of ovary, elongated and post-ovarial receptaculum seminis
and the non-pedicled genital pore.
From 0. noverca it differs in having a very long oesophagus, extension
of intestinal caeca to the posterior end, anterior testis lobed and posterior
Studies in Helminthology 145
with almost entire margin, ovary ahead of the horns of the excretory bladder,
much smaller range of vitellaria and much smaller eggs.
From 0. pedicellata it differs in having smaller ventral sucker, non-
pedicled genital pore, absence of looped terminal ends of the male and
female ducts, extra-csecal vitellaria, elongated receptaculum seminis and
smaller eggs.
Key for the identification of the avian species of the genus Opisthorchis
reported from Indian hosts :
1. Vitellaria in middle of body,
testes slightly lobed . . 2
Vitellaria confined to posterior
half of the body, testes dendritic 0. dendriticiis.
2. Oesophagus thickwalled, cuticle
non-spiny . . 0. giddhis.
Oesophagus thinwalled, cuticle
spiny . . 0. cheelis.
Family Lecithodendriidce Odhner, 1910
Owing to the close affinities between I^ecithodendriinse and Pleuro-
genetinse, Odhner (1910) created the family lyecithodendriidae for their
reception. This has not been questioned, but some confusion has been
created about the grouping of genera into subfamilies. It may be pointed
out here that this is not based on any definite grounds and apparently
different characters have, been considered in this connection ; e.g., the length
of the intestinal caeca, the extent of the vitellaria and the position of the
testes as well as the position of the genital pore and the host-relationship.
Mehra (1935) attached undue importance to trivial characters and thus
added confusion by further dividing the family Ivecithodendriidae into six
subfamilies, viz., Anchitreminse N. Subfam. Lecithodendriinae, Pleuro-
genetinae, Phaneropsoliiise N. Subfam. ; Exotidendriinae N. Subfam., and
Eumegacetinae N. Subfam. On a review of his work it appears that too
much stress has been laid on the posterior extension of intestinal caeca and
position of cirrus sac or on the median or lateral position of the genital pore.
The present writer considers that all these features show variations in
the family and should not form the basis of subfamily divisions. The only
character which appears to be constant is the cirrus sac. It is either present
or absent. The presence or absence of cirrus sac should, as has already
been discussed (I^al, 1937), form a character of subfamily importance.
46 Makund Behari Lai
It is, therefore, considered desirable to keep only two subfamilies, viz.,
ecithodendriinse with forms in which the cirrus sac is absent and Pleuro-
enetinae containing forms which possess a cirrus sac. The various sub-
imilies of Mehra can easity be accommodated under these two heads,
[uch emphasis has been laid on keeping the genus Ganeo in the subfamily
leurogeiietinse although it lacks a cirrus sac. The chief argument advanced
. the position of the genital pore which is sinistral in Ganeo as in other
leurogenetinse while it is median in I,ecithodeiidrimse. The writer considers
lat the shifting of the genital pore from median to marginal position in
bout the same level is not a character of subfamily importance, at any
ite, certainly not of a greater significance than the presence or absence of a
irrus sac. It would, therefore, be desirable to transfer the genus Ganeo
) the subfamily Lecithodeiidriinse as suggested earlier by Travassos (1930).
Diagnosis of the family Lecithodendriidce Odhner, 1910.
Digenea, with variable shape of body. Body spines either present or
Dsent. Ventral sucker at about the middle of the body. Pharynx, oeso-
tiagus and intestinal cseca of variable length. Excretory bladder V- or
-shaped. Testes generally symmetrical, sometimes, close together, at
ariable level in the body. Cirrus sac present or absent. Ovary, dorsal,
Lostly dextral, at varying levels of the body. Receptaculum seminis small,
aurer's canal present. Vitellaria situated on either side, branched follicles,
endritic mostly in front of the middle body. Uterus well-developed, with
igular loops, mostly pressed to the hinder end of the body. Genital pore
L the anterior part of the body, median or left-sided and sometimes dorsal.
;ggs numerous, small. Parasitic in intestine of Mammals, Birds, Reptiles
nd Amphibia.
The family is divided into two subfamilies :
1. Lecithodendriinae without a cirrus sac.
2. Pleurogenetinse with a cirrus sac.
The only avian species under this family reported so far from India
omes under the subfamily Pleurogenetinse as it possesses a cirrus sac.
Genus P#ra&#scws LOOSS, 1907
(Syn. Pleuropsolus Mehra, 1935)
Diagnosis. Body small, about 1 mm. long with blunt ends. Skin
trongly and thickly spinose. Oral sucker subtermiiial, globular. Aceta-
ulum almost pre-equatorial, about twice as large as oral sucker. Pharynx
mall, oesophagus long. Caeca extend post-testicular. Excretory pore
Studies in Helrninthology 147
caudo-termiiial. Copulatory organ muscular. Cirrus sac large, thick, club-
shaped, extends to left around acetabulum. Ductus ejaculatorius, pars
prostatica and prostatic cells present. Genital pore at the side of acetabulum.
Testes two, large, oval, post-acetabular. Ovary elongate pre-testicular, right of
median line. Shellgland post-acetabular. Vitellaria chiefly pre-acetabular.
Uterine coils broad, almost entirely post-acetabular. Eggs numerous,
light -brown and operculate.
Parabascus insolent (Bhalerao, 1926)
(Syn, Phaneropsolus insolens Bhalerao, 1926 ;
Pleuropsohis insolens Mehra, 1935)
Body pear-sliapsd, cuticle covered with spines. Length -585- '73 mm. Breadth
29 -3(55 mm., maximum being at the level of t3sbes. Oral sucker -115mm. x 1 mm.
Ventral sucker smaller than oral, ! mm. in diameter. Pharynx, globular, muscular
03 -035 mm. in diameter. Intestinal casca short and somewhat anterior to the
tastes. Testes, oval with entire margins, lie symmetrically, and measure -097-
0-12 mm.* X -07- -093 mm. Cirrus sac, zigzag or often horse-shoe-shaped. It is
dorsal to ventral sucker, partly anterior to it, lying between it and the intestinal fork.
On its posterior side it contains a large vesicula seminalis and pars prostatica. Ductus
ejaculated us small followed by fairly long muscular psnis capable of protrusion through
genital pore, situated immediately anterior to the ventral sucker, on the left side of
middle line. Ovary oval, measures 6-7- mm. X -04- -05 mm., right of the ventral
sucker and is often overlapped by the right test-is. Shell-gland behind ventral sucker
in the central line. Receptaculuin seminis varies in size. It lies on the inner side of
the right tcstis. Laurer's canal present. Uterus long, coils mostly post-testicular.
Eggs, brown, operculated, -018- -022 mm. X -097- -0105 mm. The vitellaria are
situated at the sides of or overlapping intestinal caeca; follicles, small, 8-12 on each side.
Excretory pore at the hind end of body leads into V-shaped bladder whose arms
diverge and end slightly behind tcstes.
Host. Corvus insolens (Intestine).
Locality. Rangoon.
Remarks. Bhalerao (1926) described this species under the genus
Phaner op solus. Mehra (1935), on small size of cirrus sac and acetabular
position of genital pore, suggested the creation of a new genus Pleur op solus
for it. On looking through the figure given by Bhalerao it appears that the
size of the cirrus sac is not small but owing to its bent character, it does not
extend beyond the intestinal bifurcation. The position of the genital pore
in the present form is pre-acetabular and this is certainly different from the
position of genital pore in Phaneropsolus where it is far forwards just behind
the pharynx. The present form also resembles the genus Parabascus in
the shape of cirrus sac, the pre-acetabular position of genital pore, in the
* In the original account it is " -97-0 12 mm. " which evidently is a misprint.
148 Makund Behari Lai
extent of vitellaria and in the position of the excretory pore. It differs,
however, from Parabascus in having shorter Intestinal caeca, smaller ventral
sucker and an avian host-features which are not of very great importance.
It is true that the species should be removed from the genus Phaner op-
solus, but it should not be taken as a basis for the creation of a new genus
Pleuropsohis as suggested by Mehra (1935). It is suggeted to place it under
the existing genus Parabascus with which it resembles in many important
points discussed before.
Family Heterophyidce Odhner, 1914
Odhner (1914) suggested the name Heterophyidse to replace the older
names Cotylogonimidse and Ccenogonimidse. Ciurea divided the family into
five subfamilies, viz., Heterophyinse, Metagoniminae, Centrocestinse,
Apophallinse and Cryptocotylinse. This division has subsequently been
modified by Nicoll (1923), Faust and Nishigori (1924), Poche (1925) and
Witenberg (1926). The family Heterophyidse as it stands to-day has the
following diagnosis :
Minute forms, usually not exceeding 2 mm. in length. Anterior por-
tion of body thinner and more movable than the posterior portion. Skin
covered with small scale-like spines that are reduced posteriorly and may
even disappear towards the posterior end of the body. Intestinal caeca
simple, usually extending to the posterior end of the body. Genital pore
in the immediate neighbourhood of the acetabulum ; genital ducts usually
open into a genital sinus, which may be variously modified and contains a
cirrus-like body or gonotyl (genital sucker). Acetabulum usually median
but may be displaced to the right of the median line ; sometimes it is
partially or completely atrophied and enclosed in the genital sinus. Cirrus
sac absent ; seminal vesicle well developed, U- or S-shaped ; the vas def erens
surrounded proximally by a mass of prostatic cells. Testes two or one,
oval, globular, or slightly lobed, near the posterior end of the body, side
by side, or obliquely one in front of the other. Ovary oval, globular or
slightly lobed, generally pre-testicular sometimes post-testicular usually to
the right of the median line. Seminal receptacle and I^aurer's canal present
near the ovary, usually near its posterior border. Vitellaria, mainly in the
lateral fields, may extend anteriorly to or beyond the genital aperture.
Uterus usually restricted to the intercaecal field between the ovary and
genital pore, but may extend to posterior end of body. Adults parasitic
in the intestine of birds and mammals.
Type genus Heterophyes Cobbold, 1866.
Studies in Helminttwlogy 149
Only one genus Ascocotyle of the family Heterophyidse has been report-
ed from birds in India.
Ascocotyle lyooss, 1899, emended Srivastava, 1935
Diagnosis. Minute distomes, body thickly spinose ; oral sucker armed
with a single or double crown of straight cylindrical spines. Oral sucker
continued posteriorly into a distinct appendage ; prepharynx long, pharynx
well developed and muscular, oesophagus present or absent, intestinal caeca
long or short. Acetabulum median, situated in association with the genital
sinus in a depression of the ventral body surface. Testes situated one on
each side at the hinder end of body ; vesicula seminalis and ejaculatory duct
well developed. Citrus sac is absent. Ovary median or slightly to one
side, pre-testicular ; receptaculum seminis large, situated in level with ovary
or behind it. Vitellaria lateral, usually post-acetabular sometimes extend-
ing as far forward as the pharynx and meeting mesially near the intestinal
bifurcation. Uterus usually post-acetabular, rarely extending as far for-
ward as the pharynx ; eggs large, operculate, measuring 0-01. 5-0 -035 mm.
X 0-008-0- 01. 7 mm. in size. Parasitic in birds and mammals.
The only avian species recorded from India is Ascocotyle inter medius
Srivastava, 1935.
Body pyriform, -ft 9mm. X -2 -38 mm. in size with backwardly directed
spines. Oral sucker terminal, () 4 -05 mm. in diameter, surrounded by double crown
of cylindrical spines about 28-30 in number. Ventral sucker feeble, -066 -077 mm.
in diameter, in middle of body, lying together with the genital pore in a shallow depres-
sion the vcntrogenital sinus. Genital opening guarded by the gonotyls. Excretory
pore terminal. Oral sucker has an oral caecum, which lies dorsal to prepharynx. Pre-
pharynx long. Pharynx -03- -04 mm. X -02 - -03 mm. Oesophagus short.
Intestinal ca&ca terminate in level with the anterior margin of ovary. Testes, posterior,
symmetrical, measure -12 - -17 mm. X -07 --12 mm. Vesicula seminalis, enor-
mous, retort-shaped. Cirrus sac absent. Ejaculatory duct joins with uterus forming a
genital sinus. Ovary between right testis and ventral sucker irregular and measures
^08 -1 mm. X -11 -14 mm. Receptaculum seminis, yolk reservoir and Laurer's
canal present. Vitellaria extend up to pharynx. Uterus never extending beyond
genital sinus. Eggs, operculate, -03 -035mm. x -015 --017mm.
Host. HalicBetus leucoryphus (Intestine).
Locality. Allahabad.
Remarks. -The genus Ascocotyle was created by lyooss in 1899. Faust
(1920) described the genus Phagicola but subsequently (1926) considered it as
a species of Ascocotyle. Meanwhile Stuiikard and Haviland (1924) split
up the genus Ascocotyle into two sub-genera Ascocotyle and Parascocotyle.
150 Makund Behari Lai
Witenberg (1929) recognized ' Parascocotyle 'of Stunkard and Havlland
and suggested that a redescription of new material of Phagicola pitheco-
phagicola could settle the question whether Parascocotyle is synonymous
with Phagicola or they both are valid genera. Price (1935) proved the
identity of Phagicola and Parascocotyle and considered that Phagicola and
Ascocotyle should be regarded as distinct genera, differing from each other
in the arrangement of spines on the collar in one or two rows, presence or
absence of oesophagus, length of intestinal caeca, position of vitellaria and
the extent of the uterus. Srivastava (1935) in describing a species Asco-
cotyle (Phagicola) intermedius has pointed out that this species connects the
two genera of Price as regards the extent of vitellaria. He has, therefore,
regarded Phagicola and Ascocotyle as sub-genera of Ascocotyle and based the
separation of the species Ascocotyle intermedius under the sub-genus Phagi-
cola on the grounds that it possesses a long oesophagus, long intestinal caeca,
and has its uterus confined behind genital sinus. This does not appear to
be sound. In fact in his description Srivastava says (page 271), " The
wide intestinal c^eca are moderately long/' In his diagram of the species
a few eggs are shown a little ahead of the region of the genital sinus. There
appears, therefore, to be some confusion in the matter. Owing to the
arrangement of oral spines the extent of vitellaria and spiiiation of the body,
Price (1936) still holds that the sub-genera Ascocotyle and Phagicola should
be given a generic status. There is, in fact, a great deal of variability in
the extent of vitellaria and in the size and extent of oesophagus and intesti-
nal caeca throughout the genus, and in the absence of constant characters,
the writer considers that this differentiation into genera and sub-genera
be abandoned.
Family Micvophallidce Travassos, 1920
Ward (1901) created a subfamily Microphallinse for the reception of the
two genera Microphalhis and Levinseniella and kept it close to the sub-
families Brachycoelinae and Pleurogenetinse. A few other isolated genera,
viz., MonoccBCum Stafford, Spelotrema Jagerskiold, Spelophallus Jagerskiold
and Maritrema Nicoll were added to this subfamily. Odhner (1.91.1) trans-
ferred the subfamily Microphallinse to the family Heterophyidas and was
supported by Nicoll (1924), Poche (1926), Fuhrmann (1928), Stunkard (1929),
Faust (1.929) and Sprehn (1932). Other workers, viz., Ransom (1920),
Ciurea (1924), Viana (1924), Witenberg (1929) and Mueller and Van Cleave
(1932) emphasized its exclusion from Heterophyidse. Travassos (1920)
while agreeing with these latter workers raised it to the status of a family
Microphallidse, to which view the present writer fully subscribes.
Studies in Helminthology 151
The family Microphallidee as it stands to-day has the following diag:
no-
Trematodes, with small oval or elliptical or pear-shaped or conical
body with blunt apex. Oral sucker, terminal or subterminal. Ventral
sucker situated in the mid-body or hind-body. Pharynx present, prepharynx
present or absent. Oesophagus long ; intestinal caeca, with one csecum
sometimes longer than the other, very short and never reaching behind
ventral sucker. Excretory bladder V-shaped. Genital opening either ad-
jacent or away from the ventral sucker, often provided with a genital atrium
which may have sometimes thimble-shaped pockets on its lateral walls.
Cirrus sac present or absent. Pars prostatica lying free or covered when
cirrus sac present* Vitelline glands, in separate follicles either in groups
or irregular or in the form of a horse-shoe-shaped band. Testes behind
ovary ; the two testes are almost on the same level. Ovary near the aceta-
bulum. Receptaculum seminis absent. I^aurer's canal present. Uterus
fills up the posterior broad portion of the body. Eggs, small and numer-
ous.
Type genus Microphallus.
Only one genus of the family, viz., Levinseniella has been reported from
India.
Genus Levinseniella Stiles and Hassall, 1901
Diagnosis. Microphallidas ; excretory bladder small and not reaching
up to testes. Vitellaria in groups of 7-8 follicles on each side. Genital
pore close to the ventral sucker. Genital atrium shows pocket-like thicken-
ings. There are always four thimble-shaped papillae associated with the
male apparatus. The " Female papillae " are thin-walled, between the
ventral sucker and the male atrium.
Type species Levinseniella brachysoma.
Levinseniella indica Lai, 1936.*
Body roughly triangular, covered over with minute spines upto the
level of the ventral sucker ; measures - 93 mm. x 5 mm. Oral sucker
1 mm. X -12 m,m. Ventral sucker -08 mm. x -078 mm. Prepharynx
02 mm. long. Pharynx -056 mm. x -055 mm. Oesophagus -22 mm.
long: Right testis -105 mm. x -058 mm. I^eft testis -12 mm. x -06mm.
Vesicula seminalis -1 mm. x -065 mm. Pars prostatica -135 mm.
* Vide Proc. Ind. Acad. Sci., 1936, 4, No. 2, Sec. B., 92-96.
B4
152 Makund Behari Lai
long. Male genital opening lies at about -09 mm. from the ventral sucker.
Ovary -105 mm. x -06 mm. Eggs numerous with thick shell and without
filaments, measure -017 mm. X -006 mm. Vitelline glands in groups of
5 or 6 just behind testes.
Host Gallinago gallinula (Bursa fabricii).
Locality Lucknow.
Remarks on the family Microphallida. The subfamily Microphallinse
Ward (1901) was raised to the status of a family Microphallidse by Travassos
(1920). This family shows distinctive differences from the family Hetero-
phyidae in the copulatory apparatus, the absence of a receptaculum seminis
and the nature of the genital pore. It has, therefore, been rightly excluded
from the latter family and kept as a separate family.
The type genus of the subfamily Microphallinae is Microphallus which
is chracterised by the absence of a cirrus sac. The genus Mantrema has
also been included under the same subfamily although it shows a well-
developed cirrus sac. The presence of a cirrus sac alone, as indicated earlier
(Lai, 1937), is sufficient for the exclusion of the genus Maritrema from this
subfamily. The writer, therefore, considers it desirable to retain the sub-
family Microphallinse for only those forms in which a cirrus sac is absent,
and to create a new subfamily Maritreminse for the reception of other forms
which possess a cirrus sac.
The family Microphallidae is divided into two sub-families thus :
1. Subfamily Microphallinge.
2. Subfamily Maritreminse. N. Subfam.
Diagnosis of the subfamily Microphallince.
Microphallidse without a cirrus sac. Vesicula seminalis and pars
prostatica lying free in the parenchyma. Vitellaria in groups of 5-7 follicles
on each side behind the testes.
Type genus Microphallus.
Diagnosis of the subfamily Maritremince. N. Subfam.
Microphallidae with a cirrus sac. Vesicula seminalis and pars pros-
tatica inside the cirrus sac. Vitellaria in the form of a band.
Type genus Maritrema.
Studies in Helminthology 153
Family Cephalogonimidce Nicoll, 1914
Nicoll (1914) erected the family Cephalogonimidse for the reception of
subfamily Cephalogoniminse lyooss, 1899. Pande (1932) and Bhalerao
36) think that the family should be brought back to the subfamily status
1 should be included in Plagiorchidse. In support of this view, Bhalerao
36) has pointed out that Nicoll (1935) has himself included it in Plagi-
hidee. He has also tried to show that the position of the genital pore in
ae genera of Plagiorchidse, viz., Renifer, Ochetosoma and Lechriorchis is
eral to pharynx and forms a near approach to the condition met with
Cephalogonimidse and further that there is a similarity in the nature
the excretory bladder in the two cases. The writer, however, does not
ee with Bhalerao in his contention. The mere fact, that the
ginal author does not mention Cephalogonimidse as a distinct family
a later stage, should not be enough to prove the non-existence of the
aily. The condition of the genital pore and cirrus sac as also some other
iracters are very distinctive in Cephalogonimidae and it would be more
propriate to regard it as a distinct family. Some of the genera of Plagi-
:hidse mentioned by Bhalerao should be simply regarded as connecting
ks showing a possible transition from one family to the other. This does
t, however, invalidate the existence of two distinct families, Cephalogoni-
de and Plagiorchidse.
ignosis of the family Cephalogonimidte Nicoll, 191.4.
Skin spiny. Prepharynx and pharynx present ; oesophagus variable,
ira |- to more than f th length of the body. Excretory vesicle Y-shaped,
netimes with a caudal vesicle. The median stem of the ' Y ' is shorter
in the branches. Genital pore marginal, dorsal or anterior to the oral
:ker. The cirrus sac long and sinuous or saccular, extending to or
yond the intestinal bifurcation. Testes behind acetabulum and ovary,
rary close to the acetabulum and on the right side of the median line of the
dy. Receptaculutn, semiius and I^aurer's canal present. Uterine coils
ascending and descending rami, chiefly in posterior region behind the
stes. Eggs thick-walled and numerous.
Type genus Cephalogonimus.
The family is represented by the genus Prosthogonimus among birds in
dia.
Genus Prosthogonimus I/uhe, 1899
Diagnosis. Body small or medium in size, flattened, with the maxi-
um breadth behind the middle. Skin spiny. Prepharynx short. Oesophagus
154 Makund Behari Lai
of medium size. Excretory bladder Y-shaped. Testes symmetrical,
behind ovary and acetabulum. Ovary close to acetabulum. Cirrus sac
long, slender and sinuous with convoluted vesicula seminalis. Genital pore
marginal at the level of the oral sucker. Receptaculum seminis and I v aurer's
canal present. Vitellaria extra-caecal. Uterus in coils in posterior part of
the body, chiefly behind the testes. Eggs numerous.
Remarks on the genus Prosthogonimus. The genus Prosthogonimus was
transferred to the f amity Plagiorchidse by Poche (1.925). Fuhnnann (1928)
and Sprehn (1932) also keep it under Plagiorchidse. Sinha (1932) has
discussed at length the differences between the genus Prosthogonimus and
the family Plagiorchidae and removed the genus to the family Cephalo-
gonimidse to which view the writer fully subscribes.
Prosthogonimus cuneatus (Rudolphi, 1809) Braun, 1901.
A single specimen of this parasite was obtained by Dr. G. S. Thapar
from the bursa fabricii of the common mynah, Acridotheres tristis.
The specimen measures 4-95 mm. x 2-375 mm. The oral sucker is
terminal and measures -2 mm. x -25 mm. The pharynx measures
.1.75 mm. X -1 mm. This is followed by a long oesophagus -3 mm. in
size which bifurcates at its posterior end into the two intestinal cseca. The
ventral sucker is well developed and lies at a distance of 1 8 mm. from the
anterior end. It has a circular shape and measures 6 mm. in diameter.
The testes lie at the same level behind the ovary and the ventral sucker.
They are 6 mm. apart from each other and oval in outline. They measure
75 mm. X -4 mm. The cirrus sac is a slender and sinuous structure,
containing coiled vesicula seminalis. It opens at the lateral margin of the
body by the side of the oral sucker.
The ovary is greatly lobed and lies in between the right testis and the
ventral sucker. It measures -6 mm. x -4 mm. The receptaculum seminis
is a more or less spherical body lying just behind the ovary 011 the inner
border of the testes. The ootype occupies a small area just behind the
ventral sucker.
The vitellaria are confined almost to the middle third of the body.
They are extra-caecal, and consist of a large number of distinct small and
rounded follicles.
The uterus fills up the entire space behind the testes. It has both
ascending and descending convolutions which pass in between the two
Studies in Helminthology 155
testes. In front of the ventral sucker trie uterus runs more or less straight
to fofrn the metraterm which runs along the cirrus sac and opens at the
genital pore. The eggs are numerous but small and measure -02 mm. x
01 mm.
Family Echinostomidoe LOOSS, 1902
The family Echinostomidse was erected by I,ooss (1.902) for the recep-
tion of his subfamily Echinostominse and some other genera. Dietz (1909)
described a large number of forms from avian hosts. Odhner (1910) at-
tempted to classify the family into three subfamilies, Echinostominse,
Himasthlinse and Echinochasminse. A large number of genera have been
described under the family which are not included in any of the existing
subfamilies. Recently the writer (I^al, 1.936) erected a new subfamily
Parorchiiise for the reception of the genus Par orchis. The family Echino-
stomidse has the following diagnosis :
Body more or less elongated, small or very large, usually much flattened
anteriorly, less so, or even cylindrical posteriorly. Oral sucker small and
weak, surrounded dorsally and laterally, but not ventrally, by a collar-like
[old, bearing one or two rows of spines, which are continued laterally to
ventral corners, the corner spines sometimes large or specialized ; acetabu-
Lum large and powerful, usually pre-equatorial and sometimes near oral
sucker. Cuticle usually spinose, specially anteriorly. Excretory vesicle
Y-shaped, with lateral twig-like branches. Pharynx and oesophagus present;
intestinal cseca extend to posterior end of body. Genital aperture pre-
acetabular ; genital sinus present or absent. Cirrus pouch usually present.
Testes post-equatorial, tandem or connubial in p.osition. Ovary pre-testi-
cular, usually to right of median line ; L,aurer's canal present. Recepta-
culum seminis present or absent. Vitellaria lateral, rarely extending
anterior to acetabulum ; sometimes confined to middle and consist of dis-
tinct and big follicles. Uterus in transverse coils, rarely extending beyond
intercsecal field. Parasites of intestines or bile ducts of vertebrates.
The family is divided into four subfamilies :
1.. Echinostomiiise I v ooss, 1.899 Type subfamily.
2. Himasthlinse Odhner, 1.91.0.
3. Echinochasminse Odhner, 1910.
4. Parorchinse I>1, 1936.
Subfamily Echinostomince I v ooss, 1899
Diagnosis. Echinostomidae, cirrus sac generally extends up to the
centre of the ventral sucker and is never pushed behind. Cirrus long,
156 Makund Behari Lai
generally not covered with spines and when contracted it appears to be
coiled. Seminal vesicle coiled and undivided. Collar spines in one or two
rows, continuous on the dorsal side.
Type genus Echinostoma.
Echinostoma revolutum LOOSS, 1899
Host A nas pcecilorhyncha (Intestine).
Locality. Rangoon.
Remarks. Gogate (1934) obtained this species from the duck, Anas
pcecilorhyncha but gives no description or diagram. He mentions that the
only feature in which his form differs from the description of lyUhe (1909)
is the size. The biggest of them is only a little longer than the smallest
mentioned in the description. The writer has no specimens of this species
in his possession and cannot add to the description. A summary of I/tihe's
description is, however, given below :
" Body 10-21 mm. long. Mouth sucker -25 -50 mm. Pharynx -21 -35 mm.
Ventral sucker -67-1 -69 mm. Cuticle spiny. Collar spines 37. There are 27
1 Randstacheln ' and 5 * Eckstachem ' on either side. Sometimes spines 35 in number.
Testes of very variable shape, in post-equatorial body. Vitellaria extend up to the
level of the acetabulum from the posterior end. They do not meet in the middle in the
region of uterus or testes. Eggs -097 -126 mm. x -059 -071 mm. "
Echinostomum govindum Moghe, 1932
Body 4 '6 -4 -94 mm. x -92 rum. Cuticle with spines numerous anteriorly but
absent in the post-test icular region. Oral sucker -149mm. X '133mm., surrounded
by reniform collar of 32 spines. There are 4 spines on each side placed close together.
Short prepharynx. Muscular pharynx -22mm. X -36 mm. followed by long- oeso-
phagus. Intestinal caeca run up to '14mm. from posterior end. Ventral sucker sub-
globular lies about '96mm. from anterior end and measures -6mm. in diameter.
Testes, ovoid, post-ovarial, -4 mm. X -24 mm. in size, lie one behind the other. Cirrus
sac ovoid, lies between intestinal bifurcation and anterior border of ventral sucker. It
contains transversely elongated vesicula seminalis surrounded by prostatic glands.
Gfenital pore behind the intestinal bifurcation. Ovary pre-testicular, -12 mm. in dia-
meter. Shell-gland lies between ovary and anterior testis. Uterus runs up to ventral
sucker. Vagina runs dorsal to ventral sucker and cirrus sac. Vitellaria, rounded
follicles, extend from posterior end of intestinal caeca to anterior level of ventral sucker.
Eggs -047mm. X -023mm.
Host. Philomachus pugnax (Rectal caeca).
Locality. Nagpur.
Echinostoma chasma n. sp.
Two immature individuals of this species were obtained from the smal 1
intestine of the garganey, Querquedula circia at Amausi. The most re-
markable feature of this form is the excretory bladder which is greatly
Studies in Helmintkology
157
sinuous and showed peculiar periodical expansions and contractions of its
stem.
The body which has spiny cuticle is elongated with a broad posterior
end and measures 2 58 mm. x 64 mm. The head with collar of spines is
marked off clearly from the rest of the body. The collar spines are 34 in
number and are arranged in one regular row without break on the dorsal
side. The dorsal spines measure -075 mm. and the lateral and marginals
05 mm. in length.
-co/.
2 mm.
FIG. 6
FIG. 5
FIG. 5. Echinostoma eft-asm//, n.sp., enitre worm ventral view. Body spines not shown.
FIG. 6. Echinostoma chasma, head collar showing spinas. (Sketched from glycerine
mount).
The oral sucker is -15 mm. in diameter. There is a small prepharynx,
and a muscular pharynx measuring -175 mm. X -125 mm. The oesophagus
is -25 mm. long and bifurcates at its posterior end into two intestinal cseca
158 Makund Behari Lai
which, are wide and have greatly crenated margins and run close by the
side of the acetabulum up to the posterior end of the animal.
The ventral sucker is -45 mm. X 41 mm. and lies at a distance of
. 65 mm. from the anterior end.
The testes are ovoid bodies, the anterior measures -13 mm. x -15 mm.
and the posterior -11 mm. X -15 mm. The cirrus sac, which partially
overlaps the ventral sucker on the left side, contains an incompletely bi-
lobed vesicula seminalis. The cirrus is fairly long and opens at the genital
pore just in front of the acetabulum and behind the intestinal bifurcation.
The ovary is oval and pretesticular and measures -85 mm. x -55 mm.
The shell-gland, ootype, etc., could not be observed. The uterus runs in more
or less sinuous but straight course dorsal to ventral sucker in the median
* line and opens at the genital pore. No eggs are present in the uterus.
The vitellaria are not fully developed and consist of small follicles of
diffuse nature, extending from the posterior end up to the level of the
middle of the ventral sucker.
Remarks. The form described above comes under the genus Echino-
stoma owing to its having a collar of spines uninterrupted on the dorsal
side, cirrus sac anterior to the ventral sucker and a non-spiny cirrus. But
the form is peculiar in possessing an incompletely bilobed seminal vesicle,
a feature in which it differs from all the existing species of the genus and
resembles the subfamily Echinochasminse. Although greater details of
anatomy are not known owing to immature state of the specimens, the
writer feels inclined to regard it as a new species of the genus Echinostoma
showing a transitional stage in the lobation of the vesicula seminalis.
Subfamily Himasthlinte Odhner, 1910
No representative of this subfamily has been reported so far from,
avian hosts in India.
S^ibfam^ly Echinochasmincz Odhner, 1910
Echinostomidcz. Cirrus sac is pear-shaped and does not reach even
up to the centre of the ventral sucker. Seminal vesicle is not coiled and is
sharply divided into two parts. Pars prostatica present. Ducttis eja-
culatorius is extremely short. Collar spines generally in one row but some
show arrangement in a double row. They are discontinuous on the dorsal
side and 20-26 in number.
Type genus Echinochasmus.
Sludies in Helminthology
Echinochasmus megavitellus n. sp.
159
A large number of these parasites were obtained from the intestine of
the paddy bird, Ardeola grayi. The worms exhibited slow movements of
their body when liberated in normal salt solution,
vit.gl
Fia. 7
FIG. 7. Ecliinochasmus megavitellus n. sp.
spines not shown.
enitire worm ventral view. Body
The body is elongated oval and measures 1-05 mm. X -45 mm. The
cuticle bears spines. The anterior end has a prominent head which bears
a collar of spines. The collar spines numbering 24, are arranged in one single
row, leaving a little space on the dorsal side. They are arranged in two
groups of 12 spines each and are of uniform size measuring -02 5 mm,
long.
160
Makund Beharl Lai
/ mm.-
FIG. 8
FIG. 8. Echinochasmus megavitellus n. sp. Head collar showing spines. (Sketched
from glycerine mount.)
The oral sucker is sub-ventral and measures -06 mm x -05 mm. The
mouth leads into a thin, prepharynx -075 mm. in size. This is followed by a
muscular pharynx measuring -09 mm. X -1 mm. The oesophagus is short
and bifurcates at its posterior end into the two intestinal caeca which run up
to the posterior end of the testes. Their terminal ends are curved inwards
and come near each other but do not meet.
The ventral sucker is strong and powerful and measures -18 mm. x
17 mm. It lies at a distance of -38 mm. from the anterior end and
09 mm. behind the intestinal bifurcation.
The testes lie very close to each other in the posterior third of the body
and are interc^ecal in position. The anterior testis measures -2 mm. x
11 mm. and the posterior -17 mm. X -112 mm. The vesicula seminalis
lies in front of the ventral sucker and is a bilobed body of 1 mrn. length.
It ends in a small ductus ejaculatorius which is surrounded by prostate
gland cells. The vesicula seminalis, ductus ejaculatorius, a short cirrus
and prostate gland cells are all enclosed in a thin-walled cirrus sac which
lies in a transverse position just in front of the ventral sucker.
The ovary which lies on the right side of the median line in front of
the two testes is an ovoid structure and measures -08 mm. x -07 mm.
Studies in Helminthology 161
The ootype lies in the median plane a little to the left of the ovary in
front of the anterior testis and is surrounded by small unicellular shell-
glands. There is no receptaculum seminis in these forms. The uterus
arises from the left side of the ootype. It is a short tube, thick in the
middle, and contains very few eggs. It is confined between the
ootype and the ventral sucker and ends in a thick- walled metraterm which,
runs along the left border of the ventral sucker and opens at the genital
pore which lies at a distance of -02 mm. in front of the ventral sucker.
The eggs are fairly large and thick-shelled and measure 07 mm. x 05 mm.
The vitellaria consist of very large and compact follicles lying in the
posterior half of the body behind the ventral sucker. They are extra-
caecal and meet together behind the testes filling up all the post-testicular
space. The two transverse vitelline ducts run just in front of the anterior
testis and open together into the ootype.
Remarks. The present form differs from all the described species of
the genus Echinochasmus except E. mordax, besides several other features,
in the position of its cirrus sac which is transversely placed and is slightly
ahead of the ventral sucker. Prom E. mordax it differs in possessing two
extra collar spines, ratio of oral to ventral sucker, in having bigger testes
and smaller ovary and in having big massive extra-caecal vitellaria. The
present form, therefore, appears to be new to science and because of its
large vitellaria is named Echinochasmus megavitellus, n. sp.
EcAwocA^sm-ws reniovarus n. sp. (Figs. 9 & 10)
Several specimens of this Trematode were recovered from the intestine
of the common house crow, Corvus Splendens. The worms are more or less
oval in shape and measure I -17 mm. x -425 mm. The cuticle bears
minute spines which extend up to the anterior testis. The head is very
prominent and possesses a well-developed collar beset with spines. The
collar spines are 24 in number and are arranged in two groups of 1.2 spines
each leaving a short space on the dorsal side in the region of the oral sucker.
They show a slight variation in size increasing in length towards the dorsal
surface and form a single row except for 4 pairs of spines towards the
pharynx which show an alternating arrangement.
The oral sucker is I mm. x 095 mm. The prepharynx is extremely
small. The pharynx measures -09 mm. x -06mm. and leads into an
oesophagus of moderate size which bifurcates into the two intestinal caeca
at its posterior end.
162
Makund Behari Lai
OY.
e.p,
co/.
ph.
V.S.
*2 mm.-
FIG. 10.
FIG. 9
FIG. 9. EcMnocJiasmus reniovarus n. sp., entire worm ventral view. (Body spines
not shown.)
FIG. 10. Schinochasmus reniovarus n. sp. Head collar showing spines. (Sketched from
glycerine mount.)
The ventral sucker, larger than the oral, is circular in outline and
measures -16 mm. in diameter. It lies at a distance of -46 mm. from the
anterior end and -13 mm. behind the intestinal bifurcation.
The excretory bladder is peculiar inasmuch as it shows five tubular
chambers opening one after the other into the bladder. These are just the
lateral branches of the stern of the bladder which have become dilated at
their proximal ends. This gives a chambered appearance to the excretory
bladder. The excretory pore is dorsal and sub-terminal.
Studies in Helmmthology lf)3
The testes, two in number, much broader than long, are contiguous and
occupy the posterior third of the body. The anterior testis measures
25 mm. x -1.4mm. and the posterior -26 mm. x -145 mm.
The vesicula seminalis is distinctly bilobed and lies on the right side
of the median line immediately in front of the ventral sucker. The cirrus
is short and is enclosed in a cirrus sac which also encloses the vesicula
seminalis and prostate gland cells. The genital pore lies at a distance of
016 mm. in front of the acetabulum in the middle line.
The ovary is pre-testicular, kidney-shaped and dextral in position.
It measures -11 mm. X -05 mm. The ootype complex lies on the left
of the ovary just in front of the anterior testis. The uterus is short and
contains a few eggs which are thick-shelled, large and measure 075 mm. x
045mm.
The vitellaria which consist of small rounded follicles extend from the
posterior end of the body right up to the level of the pharynx. They
meet both anteriorly and posteriorly and form a wreath of follicles across
the body.
Remarks. The present form differs from all the species of the genus
except E. bagulai in possessing a chambered arrangement of the excretory
bladder. But it differs from . bagulai in the greater anterior extension
of the vitellaria and shape of the ovary and testes. In its general external
appearance it stands nearest to E. corvus from which it differs in the position
of the cirrus sac and possessing a distinctly chambered excretory bladder.
It is, therefore, regarded as a new species which is designated E. renio-
varus because of its possessing a kidney-shaped ovary.
Echinochasmus corvus Bhalerao, 1926
Body 1 -06-1 '08 mm. X -405 -49 mm. Collar kidnoy-shaped with 24 spines,
disposed in one row, broken dorsally by a short space. II -\- 3 ' Eokstacheln ' and 9 -|- 9
4 Randstacheln '. Oral sucker 070- 080 mm. in diameter. Ventral sucker measures
13 '14: mm. in diameter. Prepharynx present, short -027 mm. in length. Pharynx
058 mm. Intestinal caeca simple, extend to the posterior end of the body. Testes,
one behind the other, separated by a broad band of: vitellaria from the posterior end.
Anterior testis, transversely oval, entire margin, and measures 20 mm. x -15mm.
Posterior testis somewhat round but flattened anteriorly, and measures -23 mm. x
-175mm. Cirrus sac lies dorsal to ventral sucker, in the median line, and extends
posteriorly almost to the hind margin of ventral sucker. Genital pore, pre-acetabular
and behind intestinal bifurcation. Vesicula seminalis well developed. Pars prostatica
and ductus ejaculatorius very small. Ovary immediately anterior to testes, dextral,
oval, measures I-* 13 mm. X -08- -105 mm. Rcceptaculum seminis round,
present on its innerside. Laurer's canal present, shell-gland in front of anterior testis.
Uterus small, containing very few eggs, . between anterior testis and ventral sucker.
164 Makund Behari Lai
Eggs oval, operculated, with brown shell, and measure 07- '085 mm. X -045- 06 mm.
Vitellaiia consist of large rounded follicles, situated dorsally along the side of body.
Vitellaria meet centrally In front of ventral sucker and posteriorly behind the testis.
They do not extend up to pharynx.
Host. Corvus insolens (Intestine).
Locality. Rangoon.
Echinochasmus -ntficapensis Verma, 1935
Body length 2-5 3 -5 mm. ; maximum breadth, in front of acetabulum, !-
-0 mm. Oral sucker spherical -16 mm. or oval -15 mm. X '1 mm. Collar large,
transversely elongated, reniform or triangular, -3-0 -45 mm. X 5-0 -(55 mm. ;
collar spines 24, dorsally interrupted in one regular row on each side, ventral-most of
each side smaller, others nearly equal. Prepharynx 0-1-0 -2 mm. ; pharynx oval,
0-15 mm. x 0-17 mm. ; oesophagus long, 0-25-0* 5 mm. Ventral sucker sub-
globular, -35 mm. in diameter, at one-third the body length from anterior end. Ovary
small, oval, along mid-transverse line or slightly ahead of it ; testes median, close behind
one another, in posterior half of body, margin smooth ; anterior subquadrate or ovo-
spherical, broader than long 0-18-0 -22 mm. X -19-0 -35 mm. ; posterior ovo-
spherical or sub-triangular 0-18-0 -25 mm. x -18-0 -2 mm. Ova -07-0 -08 mm. X
-039-0 -052 mm.
Host. Podiceps ruficollis.
Locality. Allahabad.
Echinochasmus bagulai Verma, 1935.
Length 1 -0 mm. to 1 75 mm. ; maximum width, about middle of body, -25 mm.
to 0-5 mm. Oral sucker 0-07 mm. in diameter. Collar reniform with 2-1 spines,
interrupted dorsally, in two rows, alternating with one another ; spines of outer row
-025-0 -035 mm. long, of inner row 0-021 - 0*027 mm. long. Prepharynx about
-05 mm. long ; pharynx -042-0 -59 mm. in diameter ; oesophagus 2J- to 3 times as long
as pharynx. Ventral sucker -2 mm. in diameter. Ovary and testes in posterior half
of body ; ovary rounded, -05 mm. in diameter, to right of median line ; testes median
close behind one another ; transversely elongated, variable in form and outline. ; anterior
testis usually somewhat smaller in dimensions than the posterior. Eggs large, 7 to 15
or more and measure -067-0 -0756 mm. x -050-0 -0588 mm.
Hosts. Ardeola grayi & Nycticorax nycticomx (Small intestine).
Locality. Allahabad & Nagpur.
Stephanopr or a fusca n. sp.
Two specimens of this trematode were obtained from the small intestine
of the spotted red-shank, Totanus fuscus. The worms are elongated and
cylindrical in shape and measure 4-775 mm. x -45 mm., the greatest width
being a little behind the posterior testis. The head is marked off from the
rest of the body and bears a collar of 22 spines. The spines are arranged in
two groups of 11 each, leaving a short space on the dorsal side in the region
Studies in Helminthology
165
of the oral sucker. The spines are of uniform size, measuring 05 mm. x
02 mm. and are arranged in one row except for four pairs of spines towards
the pharynx which show an arrangement in double row.
ov.
ooi.
tes.
tes.
yjt.gl.
FIG. 11
FIG. 11. Stephanoprora fusca n. sp., entire worm ventral view.
FIG. 12. Stephanoprora /usca head collar showing spines. (Sketched from glycerine
mount.)
166 Makund Behari Lai
The oral sucker is sub-ventral and measures 1 mm. in diameter. There
is a small prepharynx measuring -15mm. and this leads into a muscular
pharynx which measures -125mm. X -1mm. This is followed by an
oesophagus -275 mm. long which bifurcates at Its posterior end into the
two intestinal caeca which run up to the posterior end of the body.
The ventral sucker is strong and muscular and measures -25 mm. in
diameter. It lies at a distance of 9 mm. from the anterior end and 3 mm.
behind the intestinal bifurcation.
The testes are oval with entire margin and lie at about the middle of
the body. The anterior testis measures -375nim. x -25 mm. and the
posterior -4mm. x -22 mm. The vesicula seminalis is bilobed and lies
on the right side of the median line. It is enclosed in a thin-walled cirrus
sac which also contains a small cirrus surrounded by prostate gland cells.
The cirrus sac measures 3 mm. in length and bears the genital aperture at
its anterior tip.
The ovary is situated at a distance of -75 mm. behind the ventral
sucker in front of the two testes in the median line. It is oval in shape and
measures -15 mm. X -11 mm. The ootype complex lies between the ovary
and the anterior testis. There are a large number of small unicellular
shell-glands present around the ootype. The uterus arises from the left side
of the ootype, curves behind and after forming a loop runs dorsal to
ovary towards the anterior end. There are a large number of eggs present
in the uterus which measure -05 mm. x -025 mm.
The vitellaria consist of elliptical follicles and extend from in
between the testes to the posterior end. The follicles of the two sides
approach each other but do not meet in the middle. The two vitelline ducts
run parallel to the body in a longitudinal direction to open into a very
small yolk reservoir. A small and narrow duct from the yolk reservoir leads
into the ootype.
Remarks. The form described above differs from all the existing
species of the genus Stephanoprora. It differs from 5. polycestus and
S. reynoldi in the absence of a receptaculum seminis. From 5. pseudo-
echinatus, S. ornata and S. spinosa it differs in the position of the cirrus sac.
It stands nearest, however, to S. merulcB, S. denticulatus and S. conciliatus.
But it differs from S. merula in having smaller eggs, well convoluted uterus
and in the absence of a receptaculum seminis uterinum. From S.* denti-
culatus it differs in possessing smaller eggs and in not having its cirrus sac
contiguous with the ventral sucker. It differs from S. conciliate in having
Studies 171 Helminthology 167
a dextral position of the cirrus sac, smaller eggs, longer prepharynx and non-
contiguous testes. It is therefore, a new species for which the name
Stephanoprora fusca n. sp. is proposed.
Stephanoprora reynoldi Bhalerao, 1926
Body elongated, cuticle covered with spines up to the posterior margin of anterior
testis. Collar kidney-shaped with 22 spines in one row and broken dorsally. 2 ' Ecksta-
cheln ' and 9 ' Randstacheln ' on each side. Oral sucker -21 mm. X -68mm. Pre-
pharynx short. Testes with entire margin one behind the other, in the middle of body.
Anterior testis -39 mm. X -33 mm., posterior -50mm. x -33 mm. Cirrus sac situated
sinistral to ventral sucker and extends from the intestinal fork to the posterior end of
ventral sucker, pear-shaped and contains two-lobed vesicula seminalis. Pars prostatica
and ductus ejaculatorius small. Ovary, anterior to testes, oval, slightly dextral and
measures -15mm. X 115mm. lleceptaculum se minis post-ovarial, and shell-gland
behind it. Laurer's canal present. Uterus, with few ova, between anterior testis
and ventral sucker. Eggs measure -05o 0895 mm. X 031- -049 mm. Vitellaria con-
sist of large follicles. Yolk reservoir present. Vitellaria extend from the centre of the
anterior testis to the posterior end and run lateral to the body, approaching each other
closely at the posterior end. Excretory pore terminal.
Host. Corvus insolens (Intestine) .
Locality. Rangoon.
Subfamily Parorchince- I/al, 1 936*
Genus Parorchis Nicoll, 1907
This genus is of rare occurrence and is recorded for the first time in
India and the species Parorchis snipis I/al, 1936, forms the third valid
species of the genus.
Par orchis snipis I y al, 1936*
Body beset with spines for the anterior 2/3rd part ; 2-58 mm. X 1- 11 mm. Head
inconspicuously marked, ridge-like collar with single row of minute spines. Four
pairs of spines at the lateral margin of collar larger, -025mm. long. Oral sucker
24mm. X -27 mm. Ventral sucker - 58 mm. X -55 mm. Prepharynx -07mm.
Pharynx 12 mm. X 1.8 mm. Oesophagus 19 mm. long. Excretory bladder -26 mm. X
3mm. Bight testis -35 mm. X 27mm. Loft testis -36 mm. X -225mm. Vesicula
seminis behind the ventral sucker -2mm. X 13 mm. Cirrus sac -09mm. long.
Ovary -185mm. X -14mm. Reccptaculum seminis behind ovary -035mm. X
032mm. Shell-gland post-ovarial. Male and female openings lie together in a
genital atrium behind intestinal bifurcation. Eggs in a single row, show segmentation,
and measure -05-- 06 mm. x 025mm. Vitelline glands, about a dozen follicles on
each side, partly extra-caecal and partly intercaecal.
Host. Totanus hypoleucos (Cloaca).
L o cality . I/uckno w .
B5 F
168 Makund Behari Lai
The author also discussed the systematic position of the genus Par orchis
and came to the conclusion that this genus should be placed in a separate
subfamily Parorchinse Lai, 1936, in the family Echinostomidse. In the same
communication* evolution of Echinostomes was discussed and a poly-
phyletic origin of Echinostomidae was suggested.
Isolated Genera of the Family Echinostomidce
The genera Paryphostomum and Petasiger are not included in any of
the existing subfamilies of Echinostomidse. The writer is not in possession
of material of these forms and cannot definitely express his opinion. But
on looking up the available literature, he finds that both these genera should
be included in the subfamily Echinostominae, because of the arrangement
of collar spines in an uninterrupted row and non-extension of its cirrus
sac behind the ventral sucker.
Paryphostomum testitrifolium Gogate, 1934
Body elongated 3-5-5 mm., armed with spines in. front of ventral sucker. Maxi-
mum width in the region of testes 555 '952mm. Head collar strongly developed
single row, 27 spines, unbroken dorsally. Ventrally collar spines gathered into two
' end groups ' of 4 spines each. Oral sucker sub-terminal, measures -115 mm. x
128 mm. Prepharynx small, -027 mm. Pharynx globular -135 mm. X '093 mm.
Oesophagus -464 mm. long. Intestinal ca3ca run to posterior end of body. Ventral
sucker -547 mm. x *539 mm. enclosing a spacious cup-shaped cavity. Genital pore
in between ventral sucker and intestinal bifurcation, median. Testes, branched, tri-
foliate, two antero -lateral and one postero -median branches lying one behind the other ;
posterior larger than anterior. Cirrus pouch 197 mm. x -0945 mm., dorsal and
oblique to ventral sucker. Ovary globular, -17 mm. X -165 mm., pre-testicular.
Shell-gland in between ovary and anterior testis. Receptaculum seminis, posterior
to ovary. Vitellaria extend between ventral sucker and posterior end of body, filling
the post-testicular portion with loosely packed follicles. Uterus, short not much
coiled, pre-testicular, intercsecal. Vagina is dorsal to ventral sucker. Ova, numerous,
small -0771 mm. X -0409 mm.
Host. Dendrocygna javanica (Intestine).
Locality. Rangoon.
Petasiger minutissimus Gogate, 1934
Body small, elongated 948-1 -338 mm. Well-developed head collar with 23 spines,
of which 17 are arranged in a single dorsally uninterrupted row, measuring '0368
-0613 mm. x 01- -0171 mm. Remaining 6 spines, in two ventral ' end groups ' each
with 3 spines, measuring 0552- -073 mm. X -015 -0225 mm. Maximum width
* Vide Proc. Ind. Acad. Sci., 1936, 4, No. 1, Sec. B, 27-35.
Studies in Helminthology 169
behind ventral sucker -291- -349 mni. Body anterior to]ventral sucker covered with
spines. Oral sucker -045- -053 mm. Prepharynx -05- -06 mm. Pharynx globular
0495- -05 mm. Oesophagus thin -3- -33 mm. Intestinal caacal ends hidden in the
vitellaria. Ventral sucker 177 195 mm. X 198-*226 mm. Genital pore, ventral
to intestinal bifurcation. Testes oval, overlapping, with long axes at right angles.
Cirrus pouch -113 mm. x -121 mm. dorsal to ventral sucker and half portion extending
between it and the intestinal bifurcation. Ovary ovoid, -0644- -064 mm. X 066mm.
close to ventral sucker, lateral. Becsptaculum seminis and shell-gland obscured by
vitelline follicles. Vitelline follicles behind posterior margin of ventral sucker to
posterior end of body and filling loosely the po^t-tesbicular region. Uterus, pre-
testicular, short. Ova, few large -0592- -093 mm. X -0368- 048 mm.
Host. Dendrocygna javanica (Intestine).
Locality. Rangoon.
Key for the identification of the Avian Genera and Species of the family
Echinostomidse reported from India :
1. Testes connubial.. .. .. Par orchis snipis.
Testes tandem . . . . . . 2
2. Spines on collar broken on the dorsal
side . . . . . . . . 3
Spines on collar uninterrupted on
dorsal side . . . . . . 4
3. Collar spines 24 .. . . . . Echinochasmus (see 6).
Collar spines 22 .. . . . . Stephanoprora (see 10).
4. Seminal vesicle undivided . . 5
Seminal vesicle incompletely bilobed. Echinostoma chasma.
5. Collar spines 27 .. . . . . Paryphostomurn testitrifolium.
Collar spines 23 .. . . . . Petasiger minutissimus.
Collar spines 32 .. . . . . Echinostomum govindum.
Collar spines 37 . . . . . . Echinsotoma revolutum.
6. Excretory bladder with chambered
diverticula . . . . . . 7
Excretory bladder without chambered
diverticula . . . . 8
7. Vitellaria extend preacetabular . . Echinochasmus reniovarus.
Vitellaria behind acetabulum . . Echinochasmus bagulai.
8. Vitellaria meet in the pre-acetabular
region. . . . . . . Echinochasmus corvus.
Vitellaria behind acetabulum . . 9
170 Makund Beliari Lai
9. Cirrus sac, transverse, ahead of
ventral sucker . . . . . . Echinochasmus megamtellus.
Cirrus sac, vertical, touching ventral
sucker . . . . . . Echinochasmus ruficapensis.
10. Cirrus sac dextral . .. Stephanoprora fusca.
Cirrus sac sinistral . . . . Stephanoprora reynoldi.
Family Psilostomidce Odhiier, 191.3
Odhner (1.913) raised the subfamily Psilostominse lyiihe (1909) to the
status of the family Psilostomidse and described several new genera under
it. Travassos (1921) added the genus Lyperorchis and Bhalerao (1931)
described another genus Testifrondosa^ from the intestine of pig. Thapar
and I/al (1935) described from King- fisher the genus Psilorchis, the only
Avian genus of Trematode recorded from India under the family. I^al (1938)
described the second species, Psilorchis ajgainis, of the genus from Nettion
crecca.
Family Diagnosis. Worms with flattened leaf -like body of variable
length. Head indistinct without any collar. The internal structure is
comparable to that of the family Echinostomidse. Skin smooth or some-
times covered with scale-like or simple spines. Pharynx strongly developed.
Intestinal bifurcation in front of the ventral sucker. Intestinal caeca reach
almost up to the posterior end of the animal. Excretory bladder Y-shaped.
The two limbs of the bladder join each other in the region of the ventral
sucker to form a big unpaired sinus. Ovary is anterior to both the testes.
The disposition of the genital apparatus is on the same plan as that in
Echinostomidse. Vitellaria consist of distinct follicles and present on the
lateral sides of the body ventral to intestinal caeca. They do not always
meet together in the middle line posteriorly. Eggs similar to those of the
Echinostomes, about 088- 125 mm. long. Parasitic in the intestine of
birds and mammals.
Genus Psilorchis Thapar and 1>1, 1935
Psilostomidtz with leaf-like body. Ventral sucker much larger than
the oral. Short Y-shaped excretory bladder. Genital pore, dextral, in
front of the ventral sucker. Testes more or less bean-shaped and tandem ;
each testis provided with a well-developed funiculus which leads forward
t Chatterji (1938) mentions that Testifrondosa cristata Bhalerao (1931) is synonym-
ous with Paryphostomum sufrartyfex Lane (1915) which comes under E chinos tomidse.
Studies in Helminthology - 171
into a vas deferens. Receptaculum seminis absent. A yolk reservoir
present. Uterine coils lie in front of the testes. Vesicula seminalis retort-
shaped, situated in front of the ventral sucker ; cirrus short. Vitellaria
behind the ventral sucker and do not meet those of the other side
posteriorly.
Type species Psilorchis indic^ts.
Key for the identification of the species of the genus Psilorchis
reported from India :
Cirrus sac adhering to the ventral
sucker ; ovary spherical . . P. ajgainis.
Cirrus sac separate from the ventral
sucker ; ovary oval . . P. indicus.
Psilorchis indicus Thapar and I^al, 1935
Body 8 -57 mm. X 1-17 mm. ; smooth cuticle without spines. Oral sucker
17mm. X -1 mm. Ventral sucker situated in anterior 2/9th part of body, measures
75mm. X -65mm. Prepharynx 18 mm. long ; Pharynx -18mm. X -16mm. Oeso-
phagus 09 mm. long. Anterior testis - 75 mm. X 42 mm. Posterior testis 75 mm. X
36 mm. Ovary 41 mm. x 25 mm. Eggs large, oval, 125-130 mm. X 08-- 1mm.
Host. King-fisher (Intestine).
Locality. Lucknow. .
Psilorchis ajgainis I^al, 1938
Body 6 -65 mm. X -75mm.; cuticle without spines. Oral sucker -2mm. in
diameter. Ventral sucker 65 mm. X 55 mm. Pharynx 1 mm. in diameter. Oeso-
phagus 125mm. long. Anterior testis -65 mm. X -375mm. Posterior testis
7 mm. X -425mm. Ovary -25mm. in diameter. Eggs large, oval, -I--. 125 mm. X
04- -05 mm.
Host. Nettion crecca (Intestine).
Locality. Ajgain.
Family Strigeidcz Railliet, 191.9
The family Strigeidse was erected by Railliet (1919) for the forms
previously included under Holostomidse Blanchard, 1847 and Hemisto-
midse Brandes, 1888. He further created a new superfamily Strigeoidea
for the whole group having only one family, Strigeidse. More recently a
comprehensive study of the group in the form of a working classification
172 Makund Behaii Lai
has been given by I/aRue (1.926). Under the family Strlgeidse, he has
listed five subfamilies, Strigeinse, Braunininae, Cyathocotylinse, Polycotylinse
and Alariinae.
Poche (1925) excluded the subfamily Cyathocotylinae from the family
Strigeidse on the basis of presence of a cirrus sac and absence of the division
of the body into two regions and erected the family Cyathocotylidse. Szidat
(1936) has further elaborated this view and has divided the family into
several subfamilies. On the other hand, Bosnia (1931) recognized the Alari-
idse of Tubangui (1922) as a distinct family and Dubois (1933) divided it
into three subfamilies.
But in spite of these modifications, the writer is still inclined to main-
tain the classification of I^aRue (1.926) as he considers that the differences
between the various subfamilies of Strigeidse are not of such a great import-
ance as to warrant their separation into different families.
Diagnosis of the family Strigeid& Railliet, 1919.
The body is usually divided into two regions by a constriction, an
anterior part bearing chiefly the special organs of attachment and a posterior
part containing the major portion of the genitalia. The forebody is flatten-
ed or incurved ventrally or cup-shaped ; the hind body is more or less
cylindrical, ovoid or conical. The oral sucker is terminal or sub-terminal ;
the acetabulum is generally present but it is weak. The holdfast organ
is situated posterior to the ventral sucker and is sometimes covering the
latter. The intestinal caeca are without diverticula, usually extending to
the posterior end of the body. Cirrus and cirrus sac present or absent.
The vitelline glands are strongly developed. The uterus consists of few
coils containing but few eggs.
The writer is giving below a key to the subfamilies of Strigeidse Railliet,
1919:
1. Cirrus sac present . . . . 2
Cirrus sac absent . . . 3
2. Body not divided into anterior and
posterior part . . . . . . Cyathocotylinse.
Body more or less divided into anter-
ior and posterior part . , . . Braunininae.
3. Anterior part of the body cup-shaped Strigeinse.
Anterior part of the body flattened. . 4
Studies in Helminthology 173
4. Uterus confined to hindbody and does
not enter the holdfast organ . . Polycotylinse.
Uterus usually enters the holdfast
organ . . . . . . . . Alariinae.
Siibfdmily Cyathocotylince Muehling, 1.898
Diagnosis. The body is short and broad, without division into distinct
anterior and posterior regions. Cirrus and cirrus sac are present. The
genital pore is situated at the posterior end of the body.
Type genus Cyathocotyle.
Only one species of the genus Cyathocotyle has been reported from
India.
Cyathocotyle calvusi Verma, 1936
Body 2*06 mm. X 1-02 mm. Oral sucker 0-15 mm. x 0-19 ram., pharynx
0*09 mm. x 0*097 mm., oesophagus arid prepharynx abservt. Holdfast organ 0-32
4 mm. in diameter, wall covered with very peculiar characteristic, hair-like, fibrous
outgrowths. Testes very conspicuous, much elongated ; anterior 0- 56 mm. X -18 mm.,
alongside holdfast organ ; posterior -65 rnm. x -18 mm. in narrower part of body,
separated from the hind end by about half its length. Ovary roundish, -28 mm.
in diameter, masked by vitelline follicles, anterolateral to hind testis. Vitellaria
grouped in large follicles of irregular shape along intestinal caa'ca meeting in front.
Cirrus sac 0-60 mm. x 0*14 mm., contains coiled seminal vesicle, prostatic cells and a
long tubular, eversible cirrus. G-enital atrium short, with male and female ducts open-
ing side by side into it ; genital pore postero-terminal. Eggs about twice as long as
thick, not many, -081-0 -092 mm. x -041-0 -047 mm.
Host. Torgos calvus (Intestine).
Locality. Allahabad.
Subfamily Polycotylince Monticelli, 1892
Diagnosis. Body divided into two regions, forebody flattened, hind-
body cylindrical. Cirrus sac and cirrus absent. Holdfast organ round,
elliptical or bulbous. Lateral wing-like suckers present or absent. Genital
pore posterior. Uterine coils confined to hind body and never enter the
holdfast organ.
Type genus Polycotyle.
*Neodiplostomum dilaccecum n. sp.
A large number of these parasites were obtained from the small
intestine of the spotted owlet, Athene Brama. The body is flat and broad
* A number of specimens of this form were also obtained from Dr. Thapar's
collection.
174
Makund Behari Lai
and shows a clear division into a long anterior and a short posterior part.
The posterior margin of the anterior portion of the body is produced into
a thin flap which is placed transversely across the body and divides it into
two parts.
OiS.
ph.
v.s.
vii.gl.
tes.
i.e.
FIG. 13
Neodiplostomum dilaccecum n. sp., enitre worm ventral view.
The length of the specimen is 2*35 mm., and its breadth is -9 mm.
The anterior portion of the body measures 1*35 mm. and the posterior por-
tion 1-0 mm. The antero-ventral oral sucker measures -1 mm. X -05 mm.
The mouth leads into a narrow prepharynx which passes into a thick- walled
Studies in Helminthology
175
and muscular pharynx -075 mm. X -05 mm. in size. The oesophagus is short
and bifurcates into the two intestinal caeca which run almost up to the
posterior end of the animal terminating in greatly dilated, club-shaped ends.
The ventral sucker is muscular and measures -I mm. in diameter.
The holdfast organ is a heart-shaped structure with its apex pointing
anteriorly. It encloses a wide cavity in its posterior part. There is an
inconspicuous adhesive-gland mass at the posterior end of the holdfast
organ.
The gonads occupy the posterior half of the body. The ovary is ellipti-
cal and measures -25 mm. X -15 mm. The ootype with shell-gland lies
in between the testes. The uterus, starting from the ootype, takes an
ascending course up to the level of the ovary. It then turns back in a
descending loop and runs dorsal to the testes to the posterior end where it
opens into the genital atrium. The eggs measure -1 mm. x -05 mm.
ut.
2 mm.
FIG. 14 -
Neodiplostomum dilaccecum, Reproductive organs enlarged dorsal view.
176 Makund Beharl Lai
The testes, two in number, are broad and transversely elongated bodies.
The anterior testis measures -525 mm. x -275 mm. and the posterior mea-
sures -5 mm. X -3 mm. The vesicula seminalis is a thick convoluted
structure lying behind the posterior testis and opening into the genital
atrium along with the female duct.
The vitellaria are distributed throughout the anterior and the posterior
part. Anteriorly they extend a little beyond the ventral sucker and fill
up the entire space of the holdfast organ. Posteriorly they stretch up to
the dilated csecal ends.
Remarks. The form described above differs from all the species of
the genus except N. spathulaforme, N. gavialis, N. orchilongum, N. cochleare,
and N. Thomasi in having a larger broad and flat anterior part of the body.
But it, differs from all these species in several points. From N. spathulce-
forme and N. gavialis it differs in the absence of papillae around the hold-
fast organ. It differs from N. tytense, in greater posterior extension of the
vitellaria and the shape of the anterior testis. From N. orchilongum it
differs in the position of the ootype which in N. orchilongum is rather
peculiarly situated in front of both the testes. From N. cochleare it is
distinguished in having a broad and flat posterior part of the body and in
the absence of a cup-shaped union of the margin of the anterior part of
the body. From N. Thomasi it differs in the shape of the posterior part
of the body, in the shape of the testes, and in the absence of receptaculum
setninis.
It is, thus, evident that this form stands apart from all the known
species of the genus. It is further distinguished by the possession of a
heart-shaped holdfast organ and greatly dilated and club-shaped csecal ends.
It is, therefore, designated as a new species.
Neodiplostomum sp.
A single immature specimen of this trematode was obtained from the
intestine of the blue jay, Coracias benghalensis. The animal is 1-675 mm.
long and has the maximum breadth of 575 mm. in the level of the holdfast
organ. The body is distinctly divided into two parts. An extension of
the posterior margin of the anterior portion overhangs the anterior end of
the posterior portion of the body. The anterior portion of the body is
905 mm. long including the marginal expansion of 05 mm. The posterior
part measures 82 mm. long.
The oral sucker is oval in outline and measures 05 mm. X -07 mm.
The mouth leads into a very short prepharynx which passes into a globular
Studies in Helmintkology
177
muscular pharynx -031 mm. in diameter. The Oesophagus is rather small
and immediately divides into the two intestinal caeca.
FIG. 15
Neodiplostomum sp. ventral view.
The ventral sucker, which lies at a distance of -425 mm. from the
anterior end, measures -07 mm. X -071 mm. The holdfast organ
consists of two hemispherical pieces enclosing a cavity between them and
measures -217 mm. x -185 mm.
The ovary lies at a distance of -08 mm. from the anterior margin of
the posterior portion of the body. It is an ovoid body, measuring - 145 mm.
X -11 mm.
The anterior testis which lies close to the posterior is -085 mm. from
the ovary and measures 17 mm. X -085 mm. The posterior testis measures
-185 mm. X -11 mm.
The vitellaria extend almost throughout the body of the animal and
consist of small follicles. No other details could be made out in the
specimen.
Remarks. From the description given above, it is clear that the trema-
tode belongs to the genus Neodiplostomum because of the division of the
178 Makund Behaii Lai
body into two distinct parts, overhanging margin of the anterior part and
absence of any lateral suckers at the anterior end. But in the absence of
greater details of structure, it is not possible to assign any definite specific
position to this trematode.
Neodiplostomiim tytense Patwardhan, 1935
Body 2 -82 mm. long, divided into two unequal regions. Forebody flattened and
measures 1 -62 mm. X 1 -23 mm. Its lateral margins unite posteriorly to form a spoon-
shaped depression. Hind body cylindrical and measures 1 -2 mm. X -03 mm. Cuticle
smooth. Oral sucker small, and measures -056 mm. in diameter. Pharynx, small
globular -062 mm. in diameter. Oesophagus short. Acetabulum -087 mm. in diameter.
Holdfast organ, sub-circular, small, -294 mm. in diameter and lies -22 mm. behind
ventral sucker. Ovary in front of testes at the junction of fore and hind body, ovate
and measures -25 mm. x -19 mm. Uterus and eggs not developed. Vitellaria consits
O f a large number of small follicles, scattered around the holdfast organ, and extend
into the anterior third of hind body. Shell-gland situated near the left side at a level
between the two testes. Testes, one behind the other, in the middle-third of hind
body. Anterior testis pear-shaped, slightly to left and measures -375 mm. X -225 mm.
Posterior testis deeply bilobed, and measures -45 mm. X -225 mm. Vesicula seminalis
large sac behind posterior tsstis. Genital pore at posterior end of body.
Host. Tyto alba stertens (Intestine).
Locality. Nagpur.
Proalaria alcedensis Patwardhan, 1935
Body 2 -24 mm. long, divided into two parts. Forebody flattened measures
72 mm. x 66 mm. Hind body, cylindrical, and measures 1-52 mm. X -5 mm.
Oral sucker, terminal and measures -035 mm. in diameter. On its either side are situa-
ted sucker-like prominences. No prepharynx. Pharynx spherical, muscular and
-037 mm. in diameter. Oesophagus short. Acetabulum, transversely ovate,
056 mm. X -048 mm. in size. Holdfast organ slightly oval and invaginated posterior-
ly. It measures *175 mm. x -15 mm. A pair of adhesive glands are situated behind
the holdfast organ. Ovary transversely oval situated considerably behind the junction
of fore and hind body. It measures -125 mm. x -1 mm. Vitellaria, of closely packed
follicles, confined to anterior half of the hind body. Uterus extends a short distance
in front of the ovary and contains a few but large eggs measuring -075 mm. X -092 mm.
Genital pore at posterior end of body. Testes lie in middle-third of hind body and
occupy entire width. Anterior testis measures -5 mm. x -325 mm. and posterior
testis -4 mm. X -41 mm. Vesicula seminalis is voluminous and sac-like and lies
behind the posterior testis. Details of cirrus, vesicula seminalis, pars prostatica not
given.
Host. Alcedo atthis (Intestine).
Locality. Nagpur.
Studies in Helminthology
Neoalaria thaparia n. g., n. sp.
179
About half a dozen specimens of this tretnatode were obtained from
the intestine of the king vulture, Sarcogyps calvus at lyucknow. The
animals showed active movements of the body which is flat and marked
o.s.
tes.
Sot,
g.p.
bur.
FIG. 17
FIG. 10
FIG. 16. Neoalaria thaparia n. g., n. sp., ventral view.
FIG. 17. Neoalaria thaparia reproductive organs enlarged ventral view.
off into two regions by a very inconspicuous constriction. The posterior
end of the animal has a bursa which is shaped like an inverted cup. The
writer has not been able to observe any sucker or papillse inside the bursa.
The length of the specimen is 2-45 mm. Its maximum width in the
region of acetabulum is 9 mm. The anterior part of the body is roughly
square in outline and is much smaller than the barrel-shaped posterior part.
The oral sucker which projects a little beyond the anterior margin of
the body measures -15 mm. x -125 mm. The mouth leads into an ex-
tremely small prepharynx which passes into the muscular pharynx
180 Makiind Behari Lai
125 mm. X -1 mm. The oesophagus is short and thin and bifurcates Into
the intestinal cseca at its posterior end.
There are two wing-like lateral suckers situated by the side of the oral
sucker. They measure -21 mm. in length. The ventral sucker, almost
circular in shape, is well developed and muscular and measures -25 mm. in
diameter. The holdfast organ is trough-shaped with thick wall and lies at
the level of the demarcation of the body. There is no adhesive gland in
this trematode.
The gonads are confined to the barrel-shaped posterior part of the body
and lie in transverse plane parallel to each other.
The testes are peculiarly dumb-bell shaped and lie at a distance of
3 mm. apart from each other. The anterior testis measures 5 mm. X
25 mm. and the posterior -5 mm. X -275 nun. The vesicula seminalis is
thick and convoluted behind the posterior testis and opens into the bursa.
The ovary is elongated and is placed mesially, -15 mm. ahead
of the anterior testis. It measures -3mm. x -125mm. The ootype
complex lies in between the two testes. The uterus runs in an ascending
and a descending loop, a little ahead of the ovary but is confined to the
posterior body and does not enter holdfast organ. It opens into the bursa
along with the male duct. The eggs are thick-shelled and measure 07-
075mm. X -045- -05 mm.
The vitellaria also show peculiar distribution. They extend up to
the level of oesophagus anteriorly and a little behind the testes posteriorly.
They are closely arranged around the ventral sucker and the holdfast organ
but towards the periphery they extend in chains, cutting clear parenchy-
matous spaces in between the chains. In the hinder part they are confined
to the margins and do not extend into the middle of the body.
To sum up, the genus Neoalaria is characterised thus :
1. Trematodes with body, not distinctly divided into two regions.
There is no union of the posterior lateral margins of the fore-
body. The anterior part of the body is roughly square ; posterior
barrel-shaped.
2. Ventral sucker larger than oral ; lateral wing-like suckers present.
3. Holdfast organ trough-shaped ; adhesive glands absent.
4. Gonads lie parallel to each other in transverse plane.
5. Testes peculiarly dumb-bell shaped.
6. Bursa without any sucker or papillae.
Studies in Helminthology 181
7. Uterus, with ascending and descending loops, not entering the
holdfast organ.
8. Ootype complex in between the testes.
9. Vitellaria marginal in posterior body ; in anterior body, dense
around ventral sucker and holdfast organ and extending in
transverse chains towards the periphery and cutting clear
parenchymatous tissue in between the chains.
10. Eggs, thick-shelled.
From the above, it is evident that the present form stands apart from
all the genera of Strigeidse except Alaria and Proalaria in possessing lateral
wing-like processes at the anterior end. It differs from Alaria in having
its uterus behind the holdfast organ and in possessing a small trough-shaped
holdfast organ and the absence of any union of the posterior lateral margins
of the forebody. It differs from Proalaria in the absence of a clear division
of the body into two parts and in the absence of adhesive gland. From
both these genera it differs in having peculiar disposition of the vitellaria,
dumb-bell shaped testes, bursa without sucker or papilla and peculiar shape
of the body.
It is, therefore, regarded as a new genus for which the name Neoalaria
is proposed with Neoalaria thaparia as the type species.
As the new genus resembles Proalaria in the disposition of the uterus,
it is being included under the subfamily Polycotylinae.
Subfamily Alariince, Hall and Wigdor, 1918
Body indistinctly or distinctly divided into two portions, anterior part
flattened, posterior part broad or cylindrical. Cirrus and cirrus sac absent.
Holdfast organ cordate or elongated oval. 1/ateral suckers, tentacles or flaps
usually present near the oral sucker. Uterus usually enters the holdfast
organ.
Type genus Alaria.
Pharyngostomum bagulum N. Sp.
Two specimens of this Trematode were obtained from the intestine of
the eastern grey heron, Ardea cinerea rectirostris. The body shows division
into two parts although the constriction is only superficial. The anterior
part of the body is flat and longer than the posterior part which is short
and more or less ovoid in shape. The anterior part measures 63 mm. and
the posterior -35mm. in length. The lateral margins of the anterior part
of the body do not show any union.
182
Makund Behari Lai
tes.-
ves.senl,
g.p.
PIG. 18
PJiaryngostomum bayulum n. sp. ventral view.
The mouth, is surrounded by an oral sucker measuring 05 mm. by
03 mrn. and leads into an extremely small prepharynx which is followed by
a globular pharynx -05mm. x -04 mm. in size. The oesophagus is thin
and divides at its posterior end into the two intestinal cseca which run up
to the holdfast organ and are not visible behind it.
The holdfast organ is a peculiar thick-walled C-shaped structure with
the mouth of the ' C ' directed forwards. The writer has been unable to
locate any adhesive gland which is apparently absent.
The ovary is median in position and lies at the constricted region of
the body. It measures -085 mm. X -075mm. Close behind it are the
two testes, the anterior is elongated and bilobed and measures -26 mm. X
15 mm., and the posterior is smaller and measures -175 mm. X -12 mm.
Studies in Helminthology 183
The vitellaria extend anteriorly up to the level of oesophagus ; posteriorly
they are scattered and less numerous behind the testes.
The uterus and vesicula seminalis open together into a genital atrium
which lies a little distance in front of the posterior end. A mass of
indistinct tissue behind the ovary is probably of the nature of ootype,
shell-gland, etc., but it has not been properly identified. There are no
eggs in the animal which appears to be an immature individual.
Remarks. The form described above comes under the family Strigeidse
but does not possess either a cirrus sac or cup-shaped anterior body and
hence cannot be included under the subfamilies Cyathocotylinse, Brauniniae
and Strigeinse. The absence of both, clear demarcation into anterior and
posterior part of the body and the union of the lateral margin of the fore-
body as also the presence of a thick and short posterior body, exclude it
from the subfamily Polycotylinse.
The form, however, resembles the genus Pharyngostomum Ciurea, 1922,
of the subfamily Alariinse in having a short and broad posterior body,
the absence of united margin of the forebody, in the extension of the vitel-
laria up to oesophagus, in the nature of the intestinal caeca, position of the
ovary, and a reduced acetabulum. Although the course of uterus could
not be traced in the specimen, it seems probable that the uterus may enter
the holdfast organ as it generally extends beyond the ovary which in the
present case touches the holdfast organ.
But the present form differs from the described species of Pharyngo-
stomum in having a posterior extension of the vitellaria, in the shape of the
holdfast organ and in the almost entire margin of the testes. It is, there-
fore, being described as a new species.
Key for the identification of the Strigeidae included here :
1. Cirrus sac absent . . 2
Cirrus sac present . . . . Cyathocotyle calvusi.
2. Uterus does not enter the holdfast
organ . . . . . . 3
Uterus enters the holdfast organ . . Pharyngostomum bagulum.
3. lateral suckers present . . . . 4
lateral suckers absent . . . . Neodiplostomum (see No. 5).
4. Body not clearly divided into two
parts ; adhesive gland absent ;
testes dumb-bell shaped . . . . Neoalaria, dumbellata.
B5a IT
184 Makund Beharl Lai
Body clearly divided Into two parts ;
adhesive gland present . . . . Proalaria alcedensis.
5. Vitellaria extend up to the posterior
end of the hind body ; anterior
testes broad and elongated . . Neodiplostomum dilacacum.
Vitellaria extend only up to the
anterior third of the hind body ;
anterior testis pear-shaped . . Neodiplostomum tytense.
Family Schistosomidce I/>oss, 1.899
(Syn. Schistosomatidse Poche, 1907 ; Bilharziidae Odhner, 1912)
The author has already elaborated on this family in a recent paper
in the pages of these Proceedings* and has given a detailed history of the
family.
Diagnosis of the family Schistosomidce.
Trematoda. Sexes separate, pharynx absent ; oesophagus short, termi-
nating posteriorly in a bifurcation to form intestinal branches or caeca which
join caudally at the caecal union to form a single, slender intestinal caecum
terminating near the posterior end of the body. Suckers present or absent ;
acetabulum, when present, cephalad of the genital pore. Body of male
may be widened caudad of the acetabulum and have the sides incurved
ventrally, forming a gynaecophoric canal in which the female lies. Testes
consist of four or more follicles. Cirrus pouch present or absent. Female
more slender than male. Ovary elongate, sometimes spirally curved, and
lying cephalad of the csecal union. lyaurer's canal present or absent.
Vitellaria extensive, extending from the distal pole of the ovary to the
posterior end of the body. Parasitic in the blood-vessels of birds and
mammals.
The family Schistosomidae is divided into two subfamilies :
1. Schistosominae Stiles and Hassall, 1898.
2. Bilharziellinae Price, 1929.
Key to the subfamilies of the family Schistosomidae :
Caecal union caudad of equator of the
body .. .. .. .. Schistosomincz.
Caecal union cephalad of equator of
the body .. .. .. Bilharziellince.
* Vide Proc. Ind. Acad. Sci., 1937, 6, No. 5, 274-83.
S /tidies in Helminthology 185
Subfamily Schistosominat Stiles and Hassall, 1898
l . ---Males flattened and with the lateral edges
of the body infolded ventrally to form a gynsecophoric canal. Suckers
present. Intestinal cimi long, usually uniting caudad of the equator of
the body ; common caxnnn relatively short. Testes situated in the anterior
or posterior half of the body, always cephalad of the cascal union. Females
slender, thread-like, either longer or shorter than the males. Uterus usually
contains many eggs.
Type guimsSchistosoma Weinlancl, 1858.
Genus Ornithobilharzia Odliner, 1912
Diagnosis Schhtosomintc. ........ Female shorter than male. Male with well-
developed gynuxophoric canal, formed by an infolding of the lateral edges
of the body. Suckers present. Cuticle covered with spines. Digestive tract
similar to that of Schistosoma ; intestinal cseca long, showing a tendency to
form several anastomoses before finally uniting to form, the common
aeeum. Testes numerous, (JO or more, commencing a short distance
caudad of acetabulum, and extending into posterior half of the body. Cirrus
pouch rudimentary or absent. Seminal vesicle free in the parenchyma ;
prostate absent. Genital pore situated immediately behind the acetabulum.
Female elongate, slender and flattened. Ovary elongated, loosely or tightly
coiled, and situated in anterior third of the body. Vitellaria extensive,
occupying about two-thirds of body length. J y aurer's canal present (at
least in some species). Uterus short, containing only one egg at a time.
Type species Ornithobilharzia intermedia.
Ornithobilharzia^. Gogate, 1934
Length 9 '5 mm., breadth "4 88 mm. Spines and tubercles present on the body.
Oral wicker -1.00 mm. in diameter. Ventral sucker pedunculated, -472 mm. X
117 mm. Intestinal ctnca itmtf* joining to form a common c*Bcurn -52 mm. from the
posterior end. Tester about 70, circular, i'eobly developed. Cirrus pouch rudimentary,
posterior to ventral sucker. Uyrueeophorio canal well developed.
Host.Dendrocygnajavanica (Clotted blood).
Locality. -Rangoon.
Remarks. The author of this species found only two immature forms
and gives no diagram of the specimen^. In the absence of an adequate
account, the writer is unable to comment on this form.
B6 F
186 Makund Behari Lai
Subfamily Bilharziettince Price, 1929 emended
Schisiosomidce. Suckers present or absent. Gymecophoric canal
absent or imperfect!} 7 formed or sometimes well developed. Paired intestinal
ceca short uniting cephalad of the middle of the body ; common caecum
long, with or without lateral dendritic branches. Testes numerous and
situated behind the aecal union along the course of the common caecum.
Uterus short, containing a single egg.
Type genus Bilharziella I v ooss, 1899.
Key for the identification of the Avian Bilharziellinse reported from
India :
Gynsecophoric canal well developed ;
ventral sucker present . . . . Chinhula indica.
Gynaecophoric canal absent ; ventral
sucker absent . . . . Gigantobilharzia egreta.
Genus Chinhuta I,al, 1937
Male with a well-developed gynsecophoric canal, extending from a
little behind the oral sucker up to the posterior end ; female with flattened
body and smaller than the male. Suckers present. Oesohagus provided
with unicellular oesophageal glands. The two intestinal caeca unite a little
in front of the middle of body ; common caecum long, without lateral
branches but provided with angular thickenings of its wall. Testes vary
between 70-80 in number and extend from the caecal union to the posterior
end of the animal. Cirrus pouch well developed, enclosing a part of vesi-
cula seminalis, prostate gland-cells, pars prostatica and the cirrus. The
terminal end of the cirrus sac is slightly bent to the left side. The male
genital pore lies slightly to the left of the median line near the middle of
the body. The ovary is elongated and sinuous, situated at the level of the
caecal union. Uterus short and straight. The female genital opening is a
little behind the acetabulum, Vitellaria situated on the sides of the com-
mon caecum in distinct follicles. A vitelline reservoir is present.
Type species Chinhuta indica.
Chinhuta indica I^al, 1937
Male. 2-95 mm. long, -375 mm. broad; lateral edges of body rolled
inwards to form a deep groove. Oral sucker weak -i mm. in diameter.
Ventral sucker muscular -15 mm. in diameter. Oesophagus -375 rn.ni.
long, with unicellular oesophageal glands. Intestinal cseca join together
Studies in Helminthology 187
the pre-equatorial region at a distance of -625 mm. behind the
aitral sucker and 1-275 mm. from the anterior end. Testes 70-80, oval
dies of variable size; largest -125 mm. x -04 mm.; smallest -05 mm. x j
'25 mm. Vesicula seminalis divided into two lobes ; larger -117 mm. X.
'875 mm. and smaller -087 mm. x -07 mm. ;
Female. 1-8 mm. long, -19 mm. broad. Oral sucker -04 mm. in dia-
eter. Ventral sucker -075 mm,, in diameter. Oesophagus -225 mm.
tig with unicellular oesophageal glands. Intestinal cseca join together at
distance of -675 mm. from the anterior end. Ovary lies -27 mm.
hind ventral sucker and measures -155 mm. x -05 mm. No eggs were
uncl in the uterus. Vitellaria distinct and large follicles posterior to ;
p ary. ',
Host. Nettion crecca (Main blood- vesels and internal organs).
Locality. Chinhut, I^ucknow.
I
Genus Gigantobilharzia Odhner, 1910 4
: ,agnosis of the gemts Gigantobilharzia Odhner, 1910. ,
BilharziellincB. Female cylindrical and shorter than the somewhat ;
ttened male. Posterior extremity of both sexes provided with lateral \
De-like projections. Cuticle without spines or tubercles. Oral sucker
esent or absent. Ventral sucker always absent. Gynsecophoric canal
sent or reduced to a short groove, situated in anterior part of the body. jj
imentary canal similar to that of Bilharziella. Testes originate behind \
testinal union and extend almost up to the posterior end of the body. |
rrus pouch present. Genital pore situated between the intestinal bifurca- ;
>n and reunion. Ovary moderately long and spiral. Vitelline follicles
cupy about nine-tenths of the body length. Uterus short, containing a
' v eggs. ;
Type species Gigantobilharzia acoiylea Odhner, 1910. (
Gigantobilharzia egrela Lai, 1937
Male. 38-85 mm. long, -275 mm. broad. No oral or ventral sucker,
esophagus 1-2 mm. long. Csecal union takes place -4 mm. behind caecal
Furcation. Testes more than 600, filling 35-6 mm. of body ; size varia- j
le, largest -15mm. x -1 mm. and smallest -075 mm. x -05mm. Vesicula j
cninalis -2 mm. in length. I
Female. Not obtained. )
Host. Bulbulcus ibis coromandus (Renal vein) . I
Locality, Lucknow. j
188 Makund Behari Lai
Remarks. Only two more species of this genus are recorded G. cicotylea
by Odliner (1910) from Sweden and G. inonacotylea by Szidat (1930) from
East Prussia.
HOST-PARASITE RELATIONSHIP
From a review of the habitat of trematodes in hosts, it is clear that the
same species of trematode may sometimes infect more than one host. It is
now a considered fact that the host-specificity theory of older hclmintlio-
logists is no more an axiom. Cases have been reported of non-specificity
of hosts in the trematodes affecting various groups of vertebrates. The
writer (I v al, 1937) has already emphasised the non-specificity of hosts in
trematodes and also referred to a w r ork of similar nature on cestodes by
Meggitt (1934). Even in the case of Avian Trematodes there appears to be
no specificity of hosts in several cases. During his own investigations on
the subject, the writer has come across examples where one species of trema-
tode recorded from India infects more than one host. Thus, Platynosomum
acuminatum has been described from Kestrel by Nicoll (1915) and from Crow
by Bhalerao (1926). Echinochasmus bagulai has been obtained by Verma
(1935) from two different genera of the Herons. Prosthogonimus cuneatus
which is now reported from Acridotheres tristis has already been described
from a large number of birds from Asia, Africa and Europe. Echinostoma
revolutum has also been reported from an exceedingly large number of
different hosts. It is, thus, evident that the host-specificity hypothesis
does not strictly hold good for at least the Avian Trematodes.
COPULATION IN TREMATODES
The writer had no opportunity of making observations on the develop-
ment of the Avian Trematodes described in the present communication,
but he has been able to record a case of copulation in the genus Levinseniella.
The process of copulation has been observed in a large number of trema-
todes, both ecto-parasitic and endo-parasitic. Previous records reveal
that the time of copulation varies with the individuals, and climatic
conditions do not affect it specially in those cases where the host happens
to be a warm-blooded animal. Several different methods of copulation
have been observed from time to time in the case of Digenetic Trematodes.
The fertilization is affected either through cross-copulation or self-copula-
tion. The cross-copulation may be mutual or reciprocal, e.g., in Prosotocus
(lyooss, 1885), Diwoc&lium (I+instow, 1890) and Nematostrigea (Nitzsch,
1819), where the terminal portion of the uterus or the metraterm serves as
the vagina; or it may be one-sided copulation, e.g., in Paragonimus (Eerbert,
Studies in Helminthology
189
1881), Liolope (Cohn, 1902) and Collyriclum (Jegan, 1916) in which the
Laurer's canal is said to function as vagina. In the case of self-copulation
which also has been observed in some treruatodes, e.g., in Apophavynx
(Odhner, 1911) there is the introduction of the cirrus of the individual into
its own metraterm.
o.s,
or,
2 mm.
FIG. 20
FIG. 19. Levinseniellcb indica copulating worms.
FIG. 20. Levinseniella indica showing retracted cirri and grooves.
In the case of Levinseniella indica (Fig. 1.9) in which mutual copulation
takes place, a pair of individuals were obtained in sexual congress when the
bursa fabricii of the Jack snipe, Gallinago gallinula, was opened in normal
salt solution. These trematodes being of small size, the fact of -their
recovery in a copulating state could not be definitely ascertained till the
peculiarly shaped speck of tissue was subjected to examination under a
low power of the microscope. The copulating individuals lie side by side,
with their anterior ends pointing in opposite directions. One individual
lies on its dorsal while the other on its ventral surface. The union takes
place by the sides of the individuals which also partially overlap each other.
A complete contact between the couple in the affected region is accom-
plished through the help of deep undulations in the neighbourhood of the
cirrus. These undulations of the skin which take the form of grooves and
papillae are developed on the affected sides of the individuals in j the region
of contact of their genital organs ; and they appear to be temporary
structures formed only during copulation as they are entirely absent in other
specimens. Each papilla has one or two fxirrows on its side and the papilla
190 Makund Behari Lai
of one Individual fits into the groove of the other and thus affords a firm
hold for the trematodes. The cirrus of these individuals bears a large
number of minute spines and those also help in maintaining the contact
during copulation. In the present case the trematodes were so strongly
attached to each other that the pressure of the cover-glass did not seem to
have an}' effect.
The cirrus of each individual which is a thick cylindrical body appa-
rently lengthens out very much and is thrust into the metraterm of the other.
The male genital atrium lies at the base of the cirrus and during copulation
its cavity, which normally contains the cirrus, is very much reduced by the
contraction of its muscular thickenings. This exerts a pressure on the
atrium and the cirrus is thereby pushed out.
An attempt was made to fix the specimens in the copulating state but
the couple separated out and only their half retracted cirri (Fig. 20) could
be observed in the fixed specimens. These observations on the copulation
were made for about 20 minutes during which a camera lucida diagram was
made. Since the parasites were recovered already in a copulating state,
the author is unable to say anything definite regarding the duration of
copulation.
GEOGRAPHICAL DISTRIBUTION
With regard to the distribution of the various genera of Avain Trema-
todes of India in relation to those found in other parts of the world, it may
be mentioned that the distribution seems to be governed by the migration
factor. The birds, most of which are migrants, foreign as well as local, can
carry with them parasites which are peculiar to certain countries. These
cases of hosts as carriers are very common. Certain cases of peculiar distri-
bution are worth mentioning. The genus Parorchis was reported from
Northumberland coast, St. Andrews Millport, and America. It has now
been obtained in India. Similarly the genus Gigantobilharzia which was
reported only from Sweden and East Prussia is now being reported from
India. The occurrence of Parorchis, Gigantobilharzia and certain other
trematodes in India throws important light on the effect of the migratory
hosts on the spread of helminthic infections. Some of these trematodes
which were more or less endemic in particular countries have now been
discovered in India, and as an explanation of this peculiar distribution it
may be said that the parasites were brought to this country by the migra-
tory birds. Some of these parasites would in course of time produce the eggs
which may hatch out into miracidia and may infect the snails of a similar
or closely similar variety as the original intermediate host. It is not
Studies in Helminthology
191
surprising that some of these miracidia may become adapted to their new
environment and would thus produce cercaria which may infect new hosts.
The usual barriers in case of land and sea animals do not control the infection
in case of birds which can fly across continents and oceans and wander to
distant places.
Baer, J. G.
Barker, F. D.
Baylis, II. A.
Bhalerao, G. D.
Brandes, G.
Braun, M.
Brown, F. J.
Cameron, T. W. M.
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The cercaria of . Schistosomum spindalis," Ind. Jour.
Med. Res., 1921, 9.
Lehrbuch der Helminthologie," Berlin, 1932.
198
Makund Behari Lai
Srivastava, H. D.
Stiles, C. W., and Hassall, A.
Stossich, M.
Stunkard, II. W.
Stunkard, II . W., and
Haviland, C. B.
Stunkard, H. W., and
Cable, B, M.
Stunkard, H. W., and
Dunihue, F. W.
Szidat, L.
Szidat, L., and Ursula
Tanabe, B.
Thapar, G. S., and Lai, M. B.
Travassos, L.
' Studies on tlie family Heterophyidre Part I, "
Proc. U.P. Acad. Sci. 9 1935, 4.
; On a 113 w species of Catatropis from an Indian fowl,
Gallus bankiva muryhi" Ibid., 1935, 4.
! Index-catalogue of medical and veterinary Zoology."
Hyy. Lab. .Bull, 1908, 37.
' II Monostomum mutabile ele sur forme offiiie," Boll.
Soc. Adriat. Sci. Nat. Triest., 1908, 37.
: The parasitic worms collected by the American Museum
of Natural History Expedition to the Belgian Congo,"
Bull. Amer. Mus. Nat. Hist., 1929, 58.
' Further observations on the occurrence of anal oj:>en-
irigs in di genetic trematodes," Zeitsch. fur Para-
site-nit., 1931.
' The life-history of Cryplocotyle lingua," Jour. Morph.
and Phy&ioL, 1930, 50.
; Trematodes from the rat," Amcr. Mus. Novitats.,
1924, 126.
' The life-history of Par orchis avitus," Biol. Bull.,
1932, 62.
' Notes on trematodes from a Long Island duck with
description of a new species," Ibid., 1931, 60.
Giyantobilkarzia monacotylea, n.sp., ein iieuar Blut-
parasit aus Ost-Preussischen Wasservogeln, "
Zeitsch. fur Parasitaik., 1930, 2.
' Parasiten aus Seeschwalben. 1. Ueber neue Cyatho-
cotyliden aus der Darm von Sterna hirundo und
Sterna par adisea," Ibid., 1936, 8.
Beitrage zur Kenntniss der Trematoden der Mono-
stomiden Grattung Notocotylus," Zentrabl. Bakl.,
1933, 129.
; The life-history of a new schistosome, Schistosoma-
tiuni pathlocopticiim Tanabe, found in experi-
mentally infected mice," Journ. Paras-it., 1923,9.
' On the Morphology of a new ganus of Tremafcodo
parasite from the Kingfisher from Lucknow,"
Proc. Ind. Acad. Sci., 1935, 2.
' Contribuicao Para Conhecimento Da Fauna Helminto-
logica Brasileira, IX," Arch, de Agric. e Med. Vet.,
Nichtheroy, 1920, 4.
1 Reprint : Informacoes sobre a fauna helmintologica de
Matto Grosso," Folha Med. Rio de Janeiro, 1923, 3.
: Cyclocoelidse Brazileiros," Reimpresso do Brazil
Medico, 1925, lAnn. 25, 1.
: Trematodoes Novos," Ibid., 1935, Anno 25, 1.
Studies in Helminthology
199
Tubangui, M. A.
Vernia, S, C.
Vevers, G. M.
Vidyarthi, K-. D.
Ward, II. B.
Ward, H. B., and Hirsch
Witenberg, G.
Wright, S.
Yamaguti, S.
" Trematods Parasites of Philippine Vertebrates,"
Philipp. J. Sci. 9 1928, 36.
" Trematode Parasites of Philippine Vertebrates V.
Flukes from birds," Ibid., 1932, 47.
" Trematode Parasites of Philippine Vertebrates VI.
Descriptions of new species and Classification,"
Ibid., 1933, 52.
" Studies on the Indian Species of the Genus Echino-
chasmus, Parti, and on an Allied New Genus Epislho-
chasmus," Proc. Ind. Acad. Sci., 1935, 1.
" Notes on trematode parasties of Indian birds Part I,"
Alld. Uni. Studies, 1936, 12.
" A new strigeid parasito of the rare genus Cyatho-
cotyle," Nature, London, 1936, 138, 34-35.
" Observations in the genus Paragonimus Braun with
a re-description of P. compactus," Jour. Helminth.,
1923, 1.
" New Strigeids from Indian Birds," Proc. Nat. Acad.
Sci., India, 1937, 7, No. 3.
" Note on the Parasites of the Lake Fish 3," Trans.
Am. Micro. Soc., 1901, 31.
" On the structure and classification of the parasitic
worms," Jour. Paras-it., 1917, 4.
" Trematode," in Ward and Whipples's Fresh-water
Biology, New York, 1918.
" The species of Parayonimus and their differentia-
tion," Ann. Trap. Med. and Paraslt., 1915. 9.
" Die Trematoden der Fainilie Cyclocoelidae," Zool.
Jahrb. AU. Syst., 1926, 52.
" Notes on Cyclocoelidze," Ann. Mag. Nat. Hist.,
1928, 10.
" Studies on Trematode family Heterophyidae, " Ann.
Trop. Med. Parasit., 1929, 23.
" Corrections to my paper on Heterophyida3 (1929),"
Ann. Mag. Nat. Hist., 1930, 10.
" Notes on the anatomy of Microphallus opacus,"
Trans. Amer. Micr. Soc., 1912, 31.
" Studies on Helminths of Japan Part I," Jap. Jour.
Zool., 1933, 5.
" Studies on Helminth Fauna of Japan Part 3,"
Avian Trematodes," Ibid., 1934, 5.
" Studies on Helminth Fauna of Japan Part 5, " Ibid.,
1935, 6.
" Studies on Helminth Fauna of Japan Part 16.
Trematodes of fishes," Kyoto, 1937, 3.
200 Makund Behari Lai
LIST OP ABBREVIATIONS USED IN THE FIGURES
bur, . . Bursa. m.g.a. . Male genital atrium.
dr. . . Cirrus. oes. . . Oesophagus.
c.s. . . Cirrus sac. Sot. . . Ootype.
col. . . Collar. o.s. . . Oral sucker.
c.6Z. . . Excretory bladder. ov. . . Ovary.
e.ch. . . Excretory chamber. pk. . . Pharynx.
c.p. . . Excretory pore. r.s. . . Receptaculum saminis.
f.g.a. . . Female genital, atrium. tes. . . Testes.
y.p. . . Genital pore. ut. . . Uterus.
gr. . . Groove. v.s. . . Ventral sucker.
ft.o. Holdfast organ. ves. sem. . . Vesicula seminalis.
i.e. . . Intestinal cffica. vit. d. . . Vitelline duct.
l.s> - Lateral sucker. vit. gl. . . Vitslline glands.
met. . . Metraterm. vit. res. . . Vitelline reservoir.
LITTLE LEAF A TRANSMISSIBLE DISEASE OF
BRINJAL
BY K. M. THOMAS, B.A., M.Sc., D.I.C.
AND
C. S. KRISHNASWAMI, B.A., B.Sc.Ao.
(From the Agricultural Research Institute, Coimbatore)
Received May 30, 1939
(Communicated by Rao Bahadur G. N. Ran^aswami Ayyangar, I.A.S., F.N.I.)
Introduction
THE brinjal or egg plant (Solanum melongena) is an important vegetable crop
in South India, its importance perhaps, being second only to that of the
plantain (M^isa spp.). It is grown all over the Presidency as an irrigated
crop, either alone or mixed with other vegetables and often as a subsidiary
crop in betel gardens. As a pure crop it is grown in two seasons, one trans-
planted in July-August and the other transplanted in December-January*
There are many local varieties which vary considerably in size, shape,
colour and flavour. Being a vegetable fancied by all classes of people, it
commands a good market and a good crop will fetch in normal years a return
of not less than Rs. 300 per acre. The crop is subject to the attack of a
number of insect pests and fungus diseases, but so far, no virus disease has
been recorded in South India. In recent years a disease of obscure origin
resulting in a gradual diminution in the size of the leaves and sterility of
plants has been causing considerable damage to the crop in a number of
places in the presidency, and this paper describes the disease and gives an
account of the experiments carried out at Coimbatore in connection with its
investigation.
History
The disease was first observed in the Central Farm, Coimbatore
(Thomas) in 1.937 when the investigation was first started. Subsequently
reports of the disease have been received at the Mycological Section from
various parts of the presidency. In 1938 the disease was observed in a
virulent form at Nilampur, a village near Coimbatore, causing damage
to an extent of nearly 50% of the crop in the field. The disease would appear
to be of virus origin. Savulescu (1934) mentions a virus disease of brinjal
in Rumania, which causes mottling of foliage and dwarfing, but the disease
is apparently entirely different from the one described in this paper.
201
B7 F
202 K. M 9 Thomas and C. S, Krishnaswami
Smith (1937) does not mention the disease in the list of virus diseases
attacking solanaceous plants. Uppal (1929) mentions a mosaic disease of
chillies which causes reduction of leaves and a similar disease of chillies has
been noticed by Park (1934) in Ceylon. Uppal's disease however would
appear to be sap transmissible, while the Coimbatore disease is not.
Coleman's (1917) photographs of Datura stramonium indicate the possibility
of a similar disease having been present in South India at the time. But
it is believed that no record of the disease as such, or a description of the
disease has been made before.*
Etiology
Macroscopic and microscopic examination of a large number of affected
plants did not reveal the presence of any constantly associated insect,
fungus or bacterium that might be held to be the causative organism of the
disease. The absence of any living visible organism, was suggestive of a
virus being the cause of the disease.
Symptoms of the Disease
The disease has been noticed on brinjal and Datum fastuosa in nature.
Brinjal. In brinjal the most characteristic symptom is the reduction
in the size of the leaves. As the disease progresses the new leaves produced
become smaller and smaller with the result that in the course of a month the
leaves are very considerably reduced. The average size of a full grown leaf
of a two-month old healthy plant is 10cm. x 16 cm. and the maximum
size of fully developed leaves of infected plants is 2 cm, x 5 cm. (Plate V,
Fig. 6). Both the petiole and the lamina are involved in the reduction, the
leaves becoming almost sessile. There is also considerable modification in
the texture of the leaves. In healthy plants the leaf is thick, leathery,
hirsute, and dark green in colour. In diseased plants the leaf becomes thin,
soft, glabrous, and pale green in colour. In thorny varieties, the thorns
in diseased plants tend to get attenuated and in some cases entirely
disappear owing to the disease.
Another characteristic symptom is the stimulation of growth of axillary
buds including the latent buds, accompanied by the shortening of internodes
of the branches. In course of time, the suppressed branches with numerous
reduced leaves get crowded at the axils and the plant presents a character-
istic bushy appearance (Plate V, Fig. 5). At this stage the appearance of
the plant is so entirely changed that it is difficult to recognise the original
brinjal plant.
* Dr. K. M. Smith in a letter to the senior author believes this to be so and has
suggested that the virus be termed Datura Virus 2.
Little Leaf A Transmissible Disease of Brinjal 203
The third characteristic of the disease is the modification in the floral
parts (phyllody) (Plate V, Fig. 7). In many affected plants there is no
trace of any floral parts being present, but whenever found these undergo
teratological modifications, the corolla, androecium and gynoecium turning
completely green (virescent). In cases where the disease makes its appear-
ance after the flowers are normally formed, the flowers are shed and no fruits
are set. In very rare cases one or two fruits may develop normally. As
a rule however, affected plants are sterile and do not bear fruit at all. The
little-leaf affected plants are therefore absolutely worthless to the cultivator.
The disease affects the plants in all stages of growth, viz., from the seedling
stage till the period of maturity.
Datura fastuosa (Purple variety). The disease occurs on this host in
nature. The symptoms are similar to those described in brinjal, namely,
reduction in the size of leaves, shortening of internodes, crowding and
stimulation, of axillary branches (Plate VI, Figs. 5 and 9). Phyllody is some-
times noticed but flowers are often normal and viable seeds are produced.
Datura fastuosa (White variety). In the white variety also the disease
occurs in nature manifesting similar symptoms but phyllody of floral parts
is common, and viable seeds are seldom formed. The crowding of leaves
in the axils is very noticeable (Plate \ 7 I, Fig. 10).
Tomato. The disease has not so far been observed on this plant in
nature, but has been induced by grafting. The reduction in the size of
leaves, stimulation of axillary buds and phyllody are characteristic sym-
ptoms. Fruits when formed get mummified. No viable seeds are formed. The
petioles and young shoots assume a purplish tinge (Plate VI, Figs. 5 and 7).
Tobacco. The disease has not been observed in nature on tobacco but
has been successfully induced from brinjal to tobacco by grafting (Plate VI,
Fig. 11). The changes brought about by the disease in this host are very
marked. The affected plants show all the characteristic modifications of
virescence, reduction in the size of leaves and stimulation and crowding
of axillary buds as in the case of brinjal, tomato and datura.
Varietal Susceptibility
So far as our present experience goes, all South Indian varieties of brinjal
appear to be susceptible to the disease. It was not possible to distinguish
varieties of brinjal by morphological characters as genetically pure types
were seldom found. In a preliminary experiment designed to test the
varietal susceptibility of brinjal to the disease, 28 samples of seed from one
source labelled as separate varieties and 8 samples from a firm of reputed
seedsmen, were sown in pots, and the plants so raised were exposed to natural
B7a P
204
M. Thomas and C. S* Krishnaswami
Infection. Plants which escaped infection were further artificially Infected.
The results (Table I) showed that all the varieties tested were susceptible
to a greater or less degree.
TABLE I
Susceptibility of varieties of Solatium melongena to infection
Variety
Number
of plants
tested
Number show-
ing little
leaf
Central Farm 8
7 each
2
9 ..
?3
3
10 ..
35
1
11 ..
3?
2
13 ..
3?
3
14 ..
33
1
10 ..
33
2
17 ..
33
3
18 ..
33
2
19 ..
2
20 ..
33
2
21 ..
33
2
22 ..
3?
2
23 ..
3?
5
24 ..
33
1
25 ..
33
3
27 ..
33
4
28 ..
"
2
31 ..
3?
3
32 ..
3?
2
Little Leaf A Transmissible Disease of Brinjal
TABI, I(Contd.}
205
Variety
Number
of plants
tested
1ST umber show-
ing little
leaf
Central Farm 33
77
3
34 ..
7?
2
35* ..
77
T.C.K.
77
3
S. 235
20 each
3
S. 237 L.W. . .
7?
3
S. 238
77
2
S. 239
77
2
S. 240
77
3
S. 241
77
3
S. 242
7?
4
S. 245
77
3
* Proved to be susceptible by grafting.
Transmission Studies
Material and Methods.
Original source of virus. Two affected plants collected from the
Central Farm, Coimbatore, formed the original source of the virus.
Plants. In a set of preliminary experiments the plants used were
seedlings raised from seed obtained from a previous crop, which was not
diseased. The plants were kept exposed and no attempt was made to
keep them under insect-proof conditions. But in later trials, the seed was
treated with ceresan, and sown in autoclaved soil in seed pans which were
enclosed in muslin nets to prevent the entry of insects (Plate VI, Fig. 4)
and throughout the period of growth all possible precautions were taken
to grow the seedlings under insect-proof conditions.
Inoculum. Virus extract was obtained by crushing the leaves and
tender shoots of affected plants in a sterilised agate mortar and expressing
206 K. 1VL Thomas and C. S. Krishnaswami
the juice through folds of sterilised muslin. Holmes' (1929) method of
swabbing the surface of the leaf with the virus extract mixed with
carborundum powder and Sein's (1930) pin-prick method were adopted for
inoculation.
Grafting. This was done by the usual inarching method and the stock
and scion were tied with raffia. After the lapse of a few days the stem of
the scion was severed just below the graft.
Budding. This was done by the usual method of making a T-shaped
cut in the bark of the stock and inserting a scion bud with a piece of wedge-
shaped bark attached and tying up with raffia. Budding was successful only
in a few cases and was given up for later experiments.
Insect transmission. The seedlings raised were transplanted after
28 days in earthenware pots filled with treated soil* and immediately
enclosed in glass lamp chimneys covered with muslin tops. The insects
were either transferred directly from a diseased plant, or were enclosed
in a glass tube and fed on a diseased plant for a period of not less
than 24 hours prior to transfer. Aphids and tingids were transferred by
means of a sterile camel hair-brush, taking care to see that the insects
were not injured. No precaution was taken not to touch the leaf with the
brush, as it was proved by sap inoculation experiments that the disease was
not sap transmissible.
In the case of jassids, the insects previously fed on diseased plants
were transferred to muslin-topped lamp chimneys which were subsequently
placed over the healthy plants. It was found that sooner or later the insects
settled down on the plant to feed. Six days after the transfer, the insects
were killed by fumigating the plants with nicotine fumes or spraying them
with a solution of nicotine sulphate. Care was taken to see that only one
kind of insect was introduced at a time and in any series where an unknown
insect was found inadvertently introduced, the series was rejected. As soon
as the first symptoms of the disease were noticed the chimneys were removed
and the plants were allowed to grow, under normal conditions. Consider-
able difficulty was experienced for want of an insect-proof glass-house but
this was partially overcome by transferring the experimental plants, to
the verandah of the laboratory in the main building which is 30 feet high
from ground level and where owing to the absence of other vegetation
nearby, infestation of insects other than those introduced was negligible.
The controls remained healthy.
* The soil used was treated with boiling water poured twice over to kill all insects.
Little Leaf A Transmissible Disease of Brinjal 207
The number of plants used in each series of experiments throughout
was not less than five of each kind unless otherwise mentioned. And each
series was repeated thrice, so that the total number of plants used was not
less than fifteen in any series.
Experiments and Results
Sap inoculation. Inoculation done as per the method described supra
on the following host plants gave negative results :
Solanum melongena, Datura fastuosa (purple and white), tomato,
tobacco and Nicotiana glutinosa.
The experiment was repeated four times during different seasons of the
year, and no case of transmission was noticed in any of the series.
Grafting. It was found that the disease was easily transmitted from
brinjal to brinjal by means of grafting healthy stocks with affected scions.
In the preliminary experiment in 1937 out of 23 plants grown in the pot-
culture house four were grafted with diseased and 4 with healthy scions
and 15 plants were left undisturbed. The four plants grafted with diseased
scions took infection and developed symptoms of disease within 28 days,
while the others remained free. Since then over 90 grafting transfers have
been effected in the pot culture house at Coimbatore.
Insect transmission. Attempts were made to see if the following
sucking insects found on brinjal in Coimbatore were vectors :
1. Aphis rumicis I,inn.
2. Aphis malvoides Das.
3. Aphis gossypii (Glov.)
4. Empoasca devastens Dis.
5. Eutettix phycitis Dis.
6. Thrips sp. (unidentified}.
7. Urentius echinus Disk.
8. Phenococcus insolitus Gr.
Transmission experiments with the aphids and tingids gave negative
results repeatedly.
208
K* M. Thomas and C. S Krishnaswami
TABLE II
Results of Insect Transmission Experiments
' Source
S?ries ' of
virus
Trans-
ferred
to
Name of the insect
Number
of plants
used
Number of
plants
infected
1
Brinjal
Brinjal
Aphis rumieis
6*
2
??
7?
Aphis malvoides
6*
3
??
7?
Aphis gossypii
6*
4
?>
7?
Urcntius echinus
6*
'
5
7?
77
Ewpoasca devast ens
6
1
6
??
77
Entettix phycitis
6
2
r-
i
7?
7?
do.
6
3
8
77
77
do.
6
3
9
7?
7?
do.
6
4
10
77
77
do.
3
2
11
JJ
77
Thrips sp.
5*
12
7?
Tomato
Umpoasca devastens
3
13
77
7?
Eutettix phycitis
8
14
77
Chillies
do.
4
15
JJ
Datura
do.
4
16
3?
8. ccantho-
do.
6
3
17
Tomato
carpum
Brinjal
Thrips sp.
3
18
7?
Tomato
do.
3
19
7?
77
Phenococcus insolilus
3
* Repeated four times.
The results show that the two jassids Empoasca devastens and Eutettix
phycitis could transfer the disease from brinjal to brinjal. Of these the
Little Leaf A Transmissible Disease of Brinjal
209
number of successful transfers effected by Eutettix phycitis was greater
than those effected by Empoasca devastens. Both the jassids are of common
occurrence in the brinjal fields in Coimbatore, and possibly are chiefly
responsible for the spread of the disease in the field.
Host Range
The disease has been successfully transferred from brinjal to brinjal
(cultivated and wild) tomato, tobacco Datura fastuosa and Solanum
trilobatum by means of grafting. The results of these experiments are given
in Tables III nd IV.
TABI,E III
Results of Grafting Infected Scions on Different Hosts
Eesult
Source of virus
Host plant
Method
Number of
healthy
plants
used
Number of
plants
infected
Brinjal
Brinj a]
Grafting
90
90
7?
(wild)
7?
7
6
77
Datura (purple)
77
6
5
7?
Tomato
7?
15
13
Tomato
7?
7?
3
3
Datura
Brinjal
7?
5
3
(purple)
Tomato
77
5
o
(white)
77
77
5
nil
Brinjal
Tobacco
77
5
4
>?
Santalawi album
7?
2
nil
77
Solanum trilobatum
11
1
1
210
9 M. Thomas and C. S. Krishnaswami
Period required for development of symptoms
Method
Time talc en
Nature of infection
of trans-
mission
for symptoms
to appear
Brinjal to brinjal
Insect
24 to 45 days
jj jj
Grafting
21 to 30
datura
jj
9,
,, tomato
jj
15 to 22
Tomato to ,,
16 to 20
Brinjal to tobacco
40
Seed Transmission
An attempt was made to see whether the disease is transmissible through
seed. Two lots of seed collected from partially diseased brinjal and
Datura fastuosa plants were sown under insect-proof conditions. No disease
was noticed in the seedlings. It would appear therefore that the disease
is not seed transmissible.
Control
Destruction of all solanaceous weeds from, gardens and prompt removal
of affected plants would appear to be the only possible means of keeping the
disease in check until resistant varieties are found. In a field where the
weeds were completely eradicated and the diseased plants rogued out the
subsequent incidence was reported to be markedly less than in the other
fields in the same locality.
Summary
A transmissible disease of Solanum melongena has been found to cause
considerable damage to the crop in many parts of the Madras Presidency.
The general nature of the symptoms together with the absence of a visible
associated organism was suggestive of a virus being the cause of the disease.
Owing to the complete suppression of the productive phase in the affected
plants and the high percentage of incidence the loss is very great. The
disease is found to be transmissible to Datura fastuosa, tomato, tobacco
and wild brinjal, S. xanthocarpum and S. trilobatum.
Little Leaf A Transmissible Disease af Brinjal 211
The disease is not transmissible by sap inoculation but is easily trans-
mitted by means of grafting. Two species of jassids would appear to be
vectors of the disease.
A cknowledgements
The authors are greatly indebted to Dr. K. M. Smith, F.R.S., for
examining preserved material and photographs of the disease and to
Mr. M. C. Cherian, Entomologist to the Government of Madras, for identi-
fying the insects mentioned in this paper.
REFERENCES
Cbleman, L. C. . . Department of Agriculture, Mysore State, Mycological
Series Bulletin No. 3, 1917, Plate 4.
Holmes, F. O. . . Botanical Gazette, 1929, 78, 56-33.
Park, M. . . Administration Report, Director of Agriculture, Ceylon,
1034-35, D/124-D/131.
Savuloscu, T. . . Int. Cer. Agron. al. Rornaniei, 1934.
Sein, F. . . Journal, Department of Agriculture, Porto Rico, 1930,
14, 49-08.
Smith, K. M. . . A Text-Book of Plant Virus Diseases, 1937, 560-97.
Thomas. K. M. . . Administration Report of Government Mycologist, Madras.
for 1937-38, 157.
Uppal, B. N. . . International Bulletin of Plant Protection, 1929, 3, 99.
DESCRIPTION OF PLATES
PLATE V
p IO- x. View of a brinjal field in a village near Coimbatoro, showing large number of
affected plants.
FIG. 2. Healthy and naturally infected brinjal plants of the same age.
FIG. 3. A healthy brinj.al plant.
FIG. 4. Brinjal plant showing symptoms of disease induced by grafting.
p IG> 5. Same plant showing advanced stage of disease.
. 6. _ Terminal shoot of a diseased plant showing the reduction in the size of leaves.
Leaves from the healthy plant of the same age on the right.
FIGS. 7, 8 and 9. Modification of floral parts brought about by the disease in brinjal.
Healthy flower on the left in Fig. 7.
FIG. 10. Modification of floral parts in Datura fastuosa.
;p IGL 11. _ Mummified fruit of infected tomato plant showing characteristic splitting.
]? IG> 12. _ Modification of floral parts in tobacco and tomato :
Top : Tobacco flowers healthy (left) and infected (right).
Bottom : Tomato flowers infected (left) and healthy (right).
9 12 K* M. Thomas and C. S. Krishnaswami
PLATE VI
FlG j ^ brinjal se3cliing (left) showing symptoms of disease transferred by the
Insect Eutettlx pliycMs fed on diseased shoots. Control healthy plant on
the right.
p IG 9 Same plant two months later (Control not in, the picture).
p IG> 3^ (i) Brinjal seedling 28 days old (control).
/9\ 9J ,, ,, ,, infected by Eutettix pJiycltis.
/3) ^ 2 months old infected by Eutettix phycltls.
p IG _| View of insect-proof cages in which seedlings were raised for th:. k experiments.
FlG< 5. Showing diseased plants (left to right) of brinjal, tomato, Datura faxtuosa
(purple) and Datura fastuosa (white).
p IG> Q^ Solanum melongena (wild) showing- ths disease induced by grafting.
FlG 7^ Showing healthy (left) and diseased (right) tomato plants (var. Golden queen).
Disease transmitted from brinjal by grafting.
p IG> 3. Diseased tomato plant (var. Dwarf giant).
FlG< g p Datura fastuosa purple variety showing healthy plant (left) and infected
plant (right).
FIG. 10. Datura fastuosa (white) healthy (extreme left), showing symptoms in early
stage (middle) and showing symptoms in advanced stage (extreme right).
PIGS. 11 and 12. Tobacco plants (variety : Harrison special), showing disease induced
by grafting.
C. 6". Krishnaswami
10
K. M. Thomas ami
C. S. Kris/mtisMi/i
1'rof. lint. .-/W. .V/.. />', '"/. .V. /'/. /7
TESTICULAR OVA IN URAEOTYPHLUS
NARAYANI SESHACHAR
BY B. R. SBSHACHAR
(From the Department of Zoology, University of Mysore, Central College, Bangalore)
Received August 2, 1939
( Communicated by Prof. A. Subba Ran)
OVA in the testis of Amphibia have been reported by a number of workers
both normally as well as during implantation experiments. In the toads,
however, they have been more frequently found than in other Amphibia.
The development of the Bidder's organ in many species of Bufo is probably
the culmination of this faculty to develop ova by the male. The develop-
ment of the Bidder's organ and its probable function have formed the
subjects of numerous memoirs to which that of Witschi (1933) is probably
the latest contribution helping to elucidate many obscure points in regard
to this peculiar organ.
While the development of ovarian structures in species of Amphibia
other than Bufo is comparatively rare, Swingle (1917), Crew (1921) and Rau
and Gatenby (1923) have described in various species of Rana isolated ova
in the testis of both adults and larvse. The last-mentioned authors noticed
in a male specimen of Rana temp or aria a structure resembling the Bidder's
organ.
The presence of ova in relation to the testis is more common in experi-
mental work. Meyns's (1910) classic experiments on transplantation of
testis in frogs yielded results which pointed generally to the fact that in
transplanted and regenerating testis fragments, eggs are commonly found.
These observations have been substantiated by the work of Ponse (1924)
on Bufo vulgaris and later by Welti (1928) in the same species. But
contrary results have been reported by I^auche (1915) in Rana, Witschi
(1925) in Bufo and Moszkowska (1932) in Bombinator. No eggs were found
by these authors in the testicular implants.
Champy (1921) has produced precisely similar results by different
means. In a specimen of Triton alpestris, starvation produced a suppression
of spermatogenesis but when later the specimen was fed, the gonocytes of
the starved male gonad had metamorphosed into oocytes and the individual
in every way resembled a female.
213
B8 F
214
B. R. Seshachar
A review of the existing literature reveals nothing regarding this prob-
lem in Apoda. So far as I am aware, a Bidder's organ has not been
noticed in any example of this group nor have testicular eggs been reported,
either normally or as a result of experimental work.
So the interest attached to the discovery of testicular eggs in a member
of the Apoda is considerable. During my study of the spermatogenesis of
UrcBotyphhts narayani, I came across a set of sections of the testis showing
eggs. Three such ova were found in the different lobes of the testis of
the same animal. The following is a brief description of the ova.
FIG. 5
The ova are all intratubular. They are large and all the three are of
about the same size and in about the same stage of development. The cells
present all appearances of typically developing ova of the female. The
nucleus appears to be in the germinal vesicle condition. No distinct
chromosomes nor a nuclear network can be seen. The nucleus takes an
almost uniform dark stain. An interesting feature is the presence in the
nucleus of each ovum, of a large number of nucleoli. They vary from 15 to
27 in number. This is another feature in which the testicular ova resemble
normal ova of the female, where also multiple nucleoli are met with. At least
in one of the ova, the nucleoli are extruded into the cytoplasm and a number
of them occur scattered all over the cytoplasm (Fig. 6). An extrusion of
nucleoli into the cytoplasm is common in normal ova also. In one of the
Testicnlar Ova in Uneotyphlus narayani Seshachar 215
FIG. 6
oocytes a cap of granules is found in relation with the nucleus (Fig. 7). I
believe these are mitoch.ond.rial granules. It is obviously hazardous on my
FIG. 7
216
B. R Seshachar
part to attempt to discuss the cytology of these cells, seeing the paucity of
the material at my disposal and also the absence of any particular methods
for this type of cytological work (the material was fixed in Flemming's
fluid with acetic acid and later the sections were bleached in hydrogen
peroxide for the study of the chromosomes). However, a few tentative
conclusions can be arrived at as a result of the examination of the material
at my disposal. The Apoda obviously fall in a line with other Amphibia,
especially with the Anura, in the occurrence of oocytes normally in the
testis. Witschi (1934) has discussed at length the origin of such oocytes
in normal adult testis in Anura and assigns two important reasons for their
occurrence : (1) A passive conveyance of the eggs from the cortex to the
medulla by the sex cords in young specimens belonging to sex races of the
undifferentiated type ; and (2) A transformation of primitive gonia into
ovicells by their enlargement and by a change in their nuclear organization.
Ova which have been derived b}^ the latter method are usually intratubular.
This transformation has been called ' oviform degeneration '. I believe the
ovicells found in the testis of Uraotyphlus narayani belong to the second
type and have been formed by a transformation of primitive gonia. The
oocytes probably degenerate.
Cliampy, C.
Crew, F. A. E.
Lauche, A.
Meyns, R.
Moszkowska, A.
Ponse. K.
Ran, A. S., Gatenby, .T. B.
BIBLIOGRAPHY
Sur les correlations entre les caracteres sexucls
males et les divers elements du testicule chez les
amphibians (etude sur Triton alpestris) ," Comptes
Rendus de I'acad. Sci. 9 1921, 172, 482.
"A descripion of certain abnormalities of the Reproduc-
tive system found in frogs and a suggestion as to
their possible significance/' Proc. Roy. Phys. Soe.,
Edinburgh, 1921, 20, Pt. 5, 236.
" Experiment. TJnt2rsuch. an der Hoden Ei:'st:>cken
und Brunstorganen erwachssner rind jugendlicher
grasfrosche (Ranafu-tca), " Arch. Mikr. Anat., 1915,
2, 86, 51.
" Ueber froschhoden transplantationeii," Pfluyer's Arch.
f d. ges. Physiol., 1910, Bd. 132.
" Etules endD3:laologiqu3s (besticule et hypophyse)
chez le Bombinafcor," Biol. Bull. P ranee et Bely.,
1932, 66, 502.
L'organe Bidder et le determinisme des caractei'es
ssxuels sescniaires du Crapaul (Rufo vulgar is), "
Rev. Suisse de ZooL, 1924, T. 31.
" Notes on the distribution, morpliology and cytology
of the organ of Bidder, " Journ. Roy. Micros. Son.,
1923, No, 262, Pt. 1, 19.
B. R. Seshachar
Proc. Ind. Acad. Sd., B, vol. X\ PL VII
FIG. 1
FIG. 2
FIG. 3
FIG. 4
Testicular Ova in Uraotyphlus narayani Seshachar 217
Swingle, W. W.
Wclfci, E.
Wit-sclii, E.
FIG. 1.
FIG. 2.
FIG. 3.
FIG. 4.
FIG. 5,
FIG. 6.
FIG. 7.
. . " The accessory chromosome in a frog possessing
marked hermaphrodite tendencies, " Biol. Bull.
1917, 33, 70.
. . " Evolution des greffes de glandes genitales chez le
crapaud (Btifo vulgar is), auto-, homo-, heterogreffes,"
Rev. Suisse Zool., 1928, Bd. 35, H.I. 75.
.. " Studien ueber Geschlechtsumkehr end sekundaere
Geschlechtsmerkmale der Amphibien," Arch. Jul.
Klaus Stiftung, 1925, Bd. 1, 127.
... _~ _ "Studies in sex differentiation in Amphibians. VI.
The nature of Bidder's organ in the toad/' Amer.
Journ. Anat., 1933, 52, No. 3, 461.
t f Genes and inductors of sex differentiation in Amphi-
bians, " Biol. Rev., 1034, 9, No. 4, 460.
EXPLANATION OF FIGURES
Photomicrograph of the longitudinal section of a testis lobe of Urocotyphlus
narayani showing an ovicell in one of the locales of the testis. x 38.
The ovicell enlarged to show the multiple nucleoli. X 165.
Photomicrograph of another ovicell showing an aggregation of what are prob-
ably mitochondria in relation with the nucleus, x 165.
Photomicrograph of the third ovicell. X 165.
Drawing of the ovicell seen in Figs. 1 and 2. x 533.
Drawing of the ovicell seen in Fig. 4 showing the distribution of the extruded
nucleoli in the cytoplasm. X 533.
Drawing of the ovicell seen in Fig. 3 showing the distribution of the mito-
chondrial granules in relation with the nucleus, x 533.
TWO NEW TO THE LIST OF THE
ASPERGILLI
BY Coiv. I. FROII^ANO DK MBU,O
(From Medical College, Nova C6<t)
Received May 30, 1989.
To the list of the Indian Aspergilli given by Chaudhuri and Umar 1 I will
add two species, hitherto not recorded in Indian flora.
They have been found as saprophytes in the scrapings of the first case
of chromoblastomycosis found by me in India, and provenient from
Karachi, where the patient saw the beginning of his lesions since sonic ten
years ago.
The cultures were obtained primarily in petri-dishes of glucosed and
maltosed Sabouraud and have been sent to the mycological collection of
Central Bureau voor Schimmelcultures, Baarn Holland, where they are
kept.
I. Aspergilhts japonicus Saito (Bot. Magazine, Tokyo 1900, 20).
Cultures. Maltosed Salowaud. On the 2nd and 3rd day the culture
has a white dirty, somewhat brownish tone. On the 4th day the colour
resembles coffee powder. From the 5th day forwards the culture takes a
somewhat violet stain.
Glucosed Sabouraud idem.
Plain agar very weak development.
Potato idem ut Sabouraud.
Milk : on 4th day a greyish membrane at the surface, soon covered by
a coffee violet powder.
Measurements: ConidiopJiores max. 572-6 micros., inin. 264;
Columella (expanded extremity of the conidiophore which supports the
Sterigmata) generally ovoid, with the large diameter measuring 2(5 to 43
microns and the small one a little more than half of these dimensions ;
Sterigmata, single, 6-5 ; Conidia, round 2-6 ; Head (columella -] sterigma
+ conidial chain) max. 118, min. 46.
This species which resembles A. niger and A. atropurpiweits Zimm.
differs from them by its sterigmata which are simple and by its colour
218
Two New Additions to the List of the Indian Aspergilli 219
which, being, first, dirty white with a brownish tone, becomes at least of
coffee powder colour with violet reflexes.
The work of the cultures, drawings and measurements has been done
with the collaboration of my pupil Barboza Barreto, to whom my thanks
are due.
11. Aspergillus carbonarius Bain. (Bull. Soc. Bot. Fr. } 1880, 27).
Cultures. Maltosed Sabouraud whitish, cotton-like on the 2nd day,
with some black points scattered on the surface, since the 3rd day, and
becoming entirely black with erect conidiophores of 1 to 2 -5 mm. visible
at naked view, supporting blackish heads resembling chimney black.
Glucosed Salouraud idem. Plain agar nihil. Potato, simple and
glycerinaled idem. Carrot idem. GlycerinateA carrot nihil. Liquid media
(milk, I v angeron, Bouillon, Vegetable Bouillon) -surf ace soon covered by
black powdery and very thick layer of heads and spores.
Measurements. Conidiophores max. 2500 microns, min. 350 ; Columella
round and ovoid. Diameter of the round ones 90. Dimensions of the
ovoid ones 75/50 ; Conidia : membrane warty ; 4- to 5 microns diameter ;
Sterigmata : Primary 6-7 microns ; Secondary, very small and hardly
visible ; Heads max. 300, min. 200.
N.B. In both these species perithecia were not found.
Inoculated to rats and rabbits subcutaneously, intraveinously and
intraperitoneally, both these Aspergilli were found innocuous.
The cultures, drawings and measurements of this species have been
done with the collaboration of my pupil Estevam Afonso, to whom my best
thanks are due.
Resumd. To the list of Indian Aspergilli, recorded by Chaudhuri and
Umar in 1938, two more species are added : A. carbonarius Bain, 1880, and
A. japonicus Saito, 1906.
BIBLIOGRAPHY
1. Ghaudhuri, II., and Mohcl. l Molds of tho Punjab. I. The Aspergilli," Proc. Ind.
Umar Acad. Sci., (B), 1938, 8, No. 2.
1126-39-Prmted at The Bangalore Press! Mysore Road, Bangalore Chy. by O. Srinivww Rao, Stiiufrintrmlent.
and Published by The Indian Academy of Science*, Bangalore
COMPOUNDS OF PHOSPHORUS IN MILK I
BY B. K. ACHARYA, M.Sc.
AND
S. C. DEVADATTA, D.Sc. (EDIN.)
(From the Chemistry Laboratories, Wilson College, Bombay, 7)
Originally received October 14, 1938
Received in revised form August 5, 1939
ALTHOUGH phosphorus compounds of milk have been investigated by
many in the field, their classification and general nature are still incom-
plete and obscure.
Jordan Hart and Patten 1 have estimated the total phosphorus in the
whole milk and in its acid-soluble portion ; they have also estimated the
amount of organic phosphorus in the acid-soluble portion.
Lenstrup 2 states that there are four different forms of phosphorus
compounds in milk which can be estimated.
Total phosphorus
Acid-soluble P Acid-insoluble P
Inorganic P Organic P Protein P Lipoid P
He separated the acid-soluble phosphorus compounds from the acid-
insoluble by picric acid. Inorganic phosphorus is then precipitated from
the acid-soluble fraction by adding ammonia-magnesia mixture, leaving
organic phosphorus in the solution. He analysed the acid-soluble fraction
and the portion containing organic phosphorus.
Liiddecke 3 found that picric acid in cold slowly attacks lecithin on
standing. Therefore Graham and Kay 4 used trichloracetic acid as a preci-
pitant instead of picric acid. In their paper (loc. cit.) they state that
their values for inorganic phosphrous are vitiated by the possibility of the
1 Amer. J. Physiol, 1906, 16, 268.
2 J. Biol. Chem., 1926, 70, 193.
3 Inaugural Diss. Munich, 1905.
4 J. Dairy Res., 1933, 5, 54-62-63-74.
221
Bl
222 B. N Acharya and S. C. Devadatta
organic phosphorus from compounds of the type phosphagon, decomposing
In presence of strong acids.
W. Hochheimer 5 found hexose mono-phosphoric acid, pyrophosphate
and adenosine triphosphoric acid in cow's milk.
An attempt to find the nature of phosphorus compounds in the buffalo's
milk available in Bombay City, is made in this communication.
Experimental
Compounds of phosphorus in the acid-soluble portion of milk were
estimated by employing the methods developed by Eggleton and Kggleton 6
for muscle tissue, with some modifications.
To obtain the acid-soluble portion of milk, 0-5 c.c. of milk was taken
in a centrifuge tube, 2 -5 c.c. of water and 2 c.c. of 25 per cent, trichlor-
acetic acid were added, and centrifuged. The acid extract was neutralised
by the addition of finely ground baryta until neutral to phcnolphthalein
(i.e., pH =9). The mixture was centrifuged and decanted. The liquid
separated from the precipitate is called Fraction B. The precipitate (Frac-
tion A) was dissolved in a drop of concentrated hydrochloric acid and
diluted to 10 c.c. with water.
Fraction A. The orthophosphate was detected by Brigg's method
before any hydrolysis had set in. The pyro- was estimated by Tollman's
method. 7 The organic phosphorus was looked for after the solution was
oxidised by sulphuric acid and 100 volume hydrogen peroxide, and hydro-
lysed. In all estimations a quantity of solution expected to contain
0.15 to 0-2mg. of phosphorus was used.
Fraction B. This contains hydrolysable and non-hydrolysable organic
phosphorus compounds soluble in barium hydroxide at pH 9.
Hydrolysable phosphorus. To estimate the amount of phosphorus
associated with the hydrolysable organic compounds, the solution was
hydrolysed with 2 c.c. of 5 .5 N. Sulphuric acid for sixty minutes at ordinary
temperature before the application of Brigg's method.
For Non-hydrolysable phosphorus, the total amount of phosphorus
in this fraction was estimated by the method of oxidation and hydrolysis
referred to already. The difference between this amount and that due to
the hydrolysable variety gives the amount of phosphorus associated with
the non-hydrolysable organic phosphorus compounds.
5 Kinderhelik, 1932, 54, 49-64.
6 J. PhysioL, 1929, 68, No. 2, 193.
7 Biochem. Zeit., 1928, 203, 172.
Ccimfxnuids of Phosphorus In Milk / 223
Whole milk and the acid-insoluble portion of milk were separately oxi-
dised and after complete hydrolysis, Brigg's method was applied for the
estimation of phosphorus. We have been able to check up the values
obtained by direct estimations by comparing with those obtained indirectly,
i.e., by difference.
The acid-insoluble portion of milk was analysed for lipoid phosphorus
and case-in phosphorus.
For the estimation of lipoid phosphorus the method adopted was that
of Graham and Kay (loc. cit.).
Casein in milk was obtained by Hammerstein method, 8 and the phos-
phorus content in it was estimated by subjecting it to oxidation and hydro-
lysis as in the case of whole milk.
Results and Discussion
TABLE I
Composition of Milk
(In gm. per 100 c.c. of Milk)
Kxpt.
ftp. gr.
at300.
Total
Solicit*
Fat
S.N.F.
Casein
LactoFo
Ash
Chloride
I
L029
17-80
8-50
0-30
3 --12
4-02
0-721
0-0648
2
1-028
10-36
8-30
8-00
3-13
4-05
0-703
0-0623
3
1-030
20-44
0-24
11-20
3-02
4-00
0-810
0-0720
4
1-030
21-03
0-03
12-00
4-18
4-55
0-802
0-0766
6
1-030
20-3(1
0-45
10-01
3-06
4-05
0-803
0-0723
1-031
21-06
0-70
11-08
4- 20
4-20
864
0-0778
7
1-031
21-00
0-70
11-08
4-28
4-23
-0-861
0-0775
8
1-030
18-03
7-37
10-03
3-08
4-08
- 765
0-0689
9
1-030
10-20
0-48
11-88
3-83
4-08
0-703
0-0720
10
1-031
21-30
8-91
11-03
4-00
4-04
0-801
0-0723
Average
1-030
10-03
0-03
10-01
3-87
4-74
0-708
0-0717
Standard deviation
0-000
0-20
1-8
1-015
1-048
0-24
0-048
0-0025
S.N.F. = Solids not fat.
Z. Physiol Cham., 1883, 7, 227 ; 1885, 9, 273.
224
B. N. Acharya and S. C. Devadatta
TABLE II
Analysis of Ash of Milk
(In gms. per 100 c.c. of Milk)
Expt.
Ash
Calcium
Phosphorus
Ratio
CaO/P 2 O r>
1
0-721
0-1521
0-0800
1-101
2
0-703
0-1482
0-0710
1-276
3
0-810
0-1717
0-0820
1-091
4
0-862
0-1828
0-0873
1-279
5
0-803
0-1723
0-0803
1-309
6
0-864
0-1830
0-0875
1-278
7
0-861
0-1828
0-0874
1-279
8
0-765
0-1603
0-0775
1-272
9
0-793
0-1673
0-0790
1-290
10
0-801
0-1098
0-0802
1-294
Average
0-798
0-2325
(as CaO)
0-1895
(as P 2 S )
1-253
Standard deviation
0-048
0-20
0-036
0-071
TABLE III
Analysis of Compounds of Phosphorus
(Percentage of Total Phosphorus)
I
II
III
No.
Acid-soluble
Fraction B
Fraction A
P
Easily hyd.P
Non-hyd. P
Non-hyd. P.
Ortho-P
Pyro-P
1
78-27
9-22
4-88
9-50
42-39
13-81
2
75-50
7*88
4-68
8-91
39-87
12-59
3
75-90
9-49
4-77
8-64
39-82
H-13
4
78-50
9-90
4-65
8-72
41-09
12-88
5
77-26
9-66
4-90
9-57
40-10
13-06
Compounds of Phosphorus in Milk /
TABUS IV
Concentration .of Different Types of Phosphorus
(In mg. per 100 c.c. of Milk)
225
-- 1
2
3
4
!
5
Kxpt.
Total phos-
phorus
Total acid-
soluble P
Total acid-
insoluble P
FRACTION B
FRACTION A
Easilv
hyd. P
Non-hyd.
P
Non-hvd.
P ^
Ortho-
P
Pvro-
P
1
110- (33
86-10
24-20
10-20
5-40
10-52
46-90
15-28
2
128-56
97-10
31-26
10-14
6-02
11-50
51-26
16-18
3
120-10
91-20
27-32
11-40
5-73
10-38
47-83
16-97
4
106-83
83-90
21-98
10-50
7-10
9-32
43-89
13-76
5
124-90
98-50
27-63
12-07
6-02
11.95
50-09
16-32
6
98-33
12-30
6-14
12-20
51-26
15-20
7
78- 6G
9-83
4-91
9-76
40-56
12-15
8
76-44
9-56
4-77
9-48
39-84
11-81
9
83-32
..
10-42
5-19
10-32
42-89
12-86
10
74-46
9-31
4-65
9-24
38-45
11-15
11
-
78-90
9-86
5-01
9-79
41-29
12-20
12
79-96
9-99
5-99
9-999
41-99
12-36
13
86-40
..
10-87
5-39
10-72
45-10
13-36
14
96-32
12-04
6-01
11-95
50-12
14-89
15
Average
. 118-20
95-45
12-00
5-96
11-84
49-62
14.75
86-87
26-48
10-70
5-62
10-60
45-4
13-95
Standard
deviation
" 2-65
2-98
0-96
0-63
0-95
L34
1-61
226
B 9 N. Acharya and S. C Devadatta
V
Acid-insoluble Fraction of Milk
Concentration of Different Types of Phosphorus
(In mg. per 100 c.c. of Milk)
Expt.
Total P
Total acid-
insoluble P
Casein P
Lipoid P
I
126-2
25-36
22-98
3-59
2
113-4
22-03
20-20
4-47
3
128-6
30-08
27-26
4-20
4
123-4
25-23
22-96
3-94
Average
122-9
25-68
23-35
4-05
VI
Concentration of Creatine
(In mg. per 100 c.c. of Milk)
Expt.
Creatinine
Creatine
Creatine
phosphorus
acid
1
0-734
1-287
2-073
2
0-629
1-206
1-942
3
0-593
1-324
2-132
4
0-746
1-543
2-485
6
0-483
1.704
2-744
6
0-599
1-654
2-663
In order to see the average variation in the composition of milk
several samples were examined. Analyses of some of the samples are
given iu Tables I and II. These variations however do not affect our
results of the detailed analyses of milk, as the relative amounts of phos-
phorus compounds do not change (vide Table III).
Table IV gives the different types of compounds of phosphorus in mg.
per 100 c.c. of milk. It will be seen that 73-6 per cent, of the total
)hospliorus in the whole milk is soluble in trichloracetic acid, which is in
agreement with that obtained by Graham and Kay (loc. cit.).
Compounds of Phosphorus in Milk^I 227
In the acid-soluble portion of milk Graham and Kay have found
65.2 per cent, of the total phosphorus in the inorganic form and 9.8 per
cent, as organic, which they call esters. From our method, the results of
which are given in Tables III and IV, it is seen that only 50.24 per cent,
(ortho + pyro) of phosphorus constitutes inorganic variety and as much as
24-27 per cent, is in the organic form. The lower value for the inorganic
variety was expected as the easily hydrolysable organic phosphorus com-
pounds in Fraction B were separated from the inorganic by adjusting the
acid extract of milk to a pH 9 with baryta. A study of the nature of this
variety accounts for the incorrect values obtained by Graham and Kay.
Fraction B. The phosphorus in Fraction B exists in two forms :
(1) directly estimable by Brigg's method and (2) obtained after oxidation
and hydrolysis. The former constitutes 9.1 per cent, and the latter
4.8 per cent, of total phosphorus (vide Tables III and IV).
The easily hydrolysable phosphorus changes into ortho condition of
Fraction A, in presence of trichloracetic acid, on heating or on dialysing
the milk. This organic variety is, therefore, mistaken for the inorganic
type.
In order to find the nature of these organic phosphorus compounds
various substances were looked for. But, except for creatine and lactose,
the indentity of other substances could not be established. Creatine in
milk was estimated according to Masayoshi Sato and Kuchi Murata. 9
The amount of creatine found in milk is given in Table VI, column 2. It
is likely that creatine might exist as creatine phosphoric acid which is
hydrolysed by acid. This creatine phosphorus unlike in the case of muscle
(Eggleton, loc. cit.) accounts only for a part of the hydrolysable variety.
Therefore some other substance may be associated with this organic
phosphorus.
It is not clear in what form lactose exists in milk. It has been sug-
gested by Mai Monatsschr 10 that lactose is bound up with phosphorus in
loose combination. If so only a very small amount of lactose present in
milk will be used in this combination. This point requires further investi-
gation.
Fraction A. In Tables III and IV are given the amounts of the
compounds of phosphorus, insoluble at pH 9 in presence of barium
hydroxide. It will be seen that this fraction is divided into three types :
9 J. Agri. Chem., Soc. Japan, 1933, 9, 1-5.
10 Kinderhelik, 1932, 51, 391-92.
228
B. N a Acharya and S* C. Devadatta
ortho, pyro and non-hydrolysable phosphorus. From Table IV, column
5, it is seen that 3842 per cent, of the total phosphorus is in the ortho-
and 11-8 per cent, in the pyro- condition. The non-hydrolysable variety
forms 9-1 per cent, of the total phosphorus.
The amount of phosphorus in the acid-insoluble portion of milk is
given in Table IV, column 3. It may be noted that 22-4 per cent, of the
total phosphorus, which is present in the acid-insoluble portion, consists of
lipoid phosphorus 3-7 per cent., and casein phosphorus, 11) per cent,
(refer Table V).
It will be seen thus from the above discussion that we have been able
to establish the existence of five independent types of phosphorus com-
pounds : ortho, pyro, organic phosphorus compounds insoluble in barium
hydroxide at pH 9 and hydrolysable and non-hydrolysable organic phos-
phorus compounds soluble at pH 9 ; over and above these the acid-insolu-
ble phosphorus consists of casein phosphorus and lipoid phosphorus. The
amount of phosphorus present in these seven varieties are estimated
directly, as shown below in mg. per 100 c.c. milk:
Total phosphorus
11.8-20
Acid-soluble P
86-87
Baryta
Acid-insoluble P
20-48
Casein P. J,ipoicl P
23 -35 4 -05
Soluble
Insoluble
Non-hydro-
lysable P
5-62
Hydro-
lysable P
10-70
Ortho P
45-4
Pyro P
13-95
Non-hydro-
lysable P
10 -GO
PHOSPHORUS, CALCIUM AND MAGNESIUM
IN MILK II
BY B. N*. ACHARYA, M.Sc.
AND
vS. C. DEVADATTA, D.SC. (EDIN.)
(From th& Chemistry Laboratories, Wilson College, Bombay, 7}
Originally received October 14, 1938
lleceived in revised form August 5, 1939
CALCIUM, magnesium and phosphorus are capable of forming both soluble
and insoluble salts, but the mode of formation of the latter will be of great
interest. Some workers have drawn their conclusions regarding the salt
formation in milk from the analysis of the scurn formed after heating the
milk ; others generalised merely on the strength of their experiments of
dialysis or of ultra-centriftiging and so on.
Van Slyke and Bosworth 1 have shown that milk contains CaHPO 4 on
the results of their ultra-centrifuge experiments.
Soldner and others 2 have found that on heating milk a scum is deposited
which is mostly tricalcium phosphate Ca 3 (PO 4 ) 2 , concluding its presence
originally in milk.
Palmer 3 however thinks that CaHPO 4 , stabilised with gelatine, forms
a precipitate of colloidal calcium phosphate on heating.
Various workers 4 have studied the effect of dialysis of milk and tried
to draw inferences about the composition of salts in milk which do not seem
to be definite. Considerable divergence in their observations is to be
attributed to the fact that the experimental conditions were different in
each case.
Casein a phospho-protein body in milk was also subjected to various
experiments. Somer and Hart 5 state that casein forms some kind of loose
1 J. Biol. Chem., 1915, 20, 135 ; 191(5, 24, 199.
2 Soldner, Landw. Versuchis-Stat, 1S85, 35, 351 ; de Vries and Boekhout, Ibid.,
1901, 55, 201; Purvis, Brehaub and M'Hattie, J.Roy. San. Inst., 1912, 33, 154
Grosser, Biochem. Z., 1913, 48, 422 ; Diffloth, Bull. Sci. PharmacoL, 1904, 10, 278.
3 Proc. Soc. Exp. Biol. Med., 1921.
4 G-yorgy, Biochem. Z., 1923, 142, 1 ; Mattick and Hallett, J. Agric. Sci., 1929, 19,
452 ; Wardlaw, J. Roy. Soc., N.S.W., 1914, 48, 253, etc.
5 J. Biol. Ckem., 1919, 40, 137.
229
230
B. N. Acharya and S. C. Devadatta
compound with calcium. It is reported that casein on suspension in water,
or on heating for some time, loses some of its phosphorus. 6 Berggren 7 is of
opinion that the phosphorus of casein is more loosely bound than is usually
supposed.
To get an insight into the distribution of calcium, magnesium and phos-
phorus in milk the problem of the composition of milk is to be approached
in several ways. In the present investigation estimations of phosphorus,
calcium and magnesium were made under varying conditions. First of all
milk was divided into two portions soluble and insoluble in trichloracetic
acid (2 c.c. of 25'% acid for 0-5 c.c. of milk diluted to 3 c.c. with water).
Next the amounts of the constituents of whole milk which are dialysable
were differentiated from the undialysable. lastly fresh milk was heated
after removing the fat, till scum was formed. The scum was analysed-
The residual milk free from scum was divided into two portions soluble and
insoluble in trichloracetic acid.
For the estimation of phosphorus Brigg's method was used. For
calcium, the method of Masayoshi Sato and Kuchi Murata 8 with a few
modifications was adopted. Magnesium was estimated by the usual gravi-
metric method.
Results and Discussions
TABLE I
Amount of Total, Acid-Soluble and Acid-Insoluble Phosphorus, Calcium
and Magnesium
(In mg. per 100 c.c. of Milk)
1
2
3
No.
Total
Acid-Soluble
Acid-Insoluble
P
Ca
Mg
P
Ca
Mg
P
Ca
Mg
1
126-2
148-2
18-42
100-30
99-26
17-23
25-36
48-00
1-19
2
113-4
146-3
14-31
92-36
98-24
13-30
22-03
48-28
1-01
3
128-6
152-4
19-63
97-10
101-34
18-31
30-08
50-48
1-32
4
123-4
149-2
18-83
98-50
98-34
17-52
25-23
50-50
1-31
122-95
149-0
18-00
96-57
99-30
16-59
25-68
49-32
1-21
6 Loubavin, Ber., 1877, 10, 2237 ; 1879, 12, 1021.
7 J. B'ioL Chem., 1922, 95, 451.
8 J. Agr. Chem. Soc. Japan, 1933, 334-36.
Phosphorus^ Calcium and Magnesium in Milk //
TABLE II
Analysis of the Scum and the Milk after removal of Scum
(In nig. per 100 c.c. of Milk)
231
1
2
3
Total
Acid- Soluble
Acid-Insoluble
P
Ca
Mg
P
Ca
Mg '
P
Ca
Mg
Whole milk . .
126-4
148-20
18-42
100-30
99-26
17-23
25-36
48-CO
1-89
Milk heated and
scum re moved
99-64
86-20
10-10
78-64
71 (53
9-60
21-06
15- eo
0-50
Scum
27-40
64-00
8-00
-
Scum is derived from the acid-soluble and acid-insoluble fractions.
TABLE III
i
Amounts of Dialy sable and Undialy sable Phosphorus, Calcium and
Magnesium
(In mg. per 100 c.c. of Milk)
I
II
III
Time in
hours
Total
Dialysablc
Undialysable
P
Ca
Mg
P
Ca
Mg
P
Ca
Mg
120-4
148-i2
18-42
6
43-20
44-32
10-42
82-68
102-94
8-00
12
47-63
48-06
12-39
77-35
99-83
6-03
18
53-93
53-58
14-00
71-96
94-63
4-42
24
60-32
CO- 00
16-10
64-90
98-32
2-32
30
60-32.
60-20
16-32
63-19
86-28
2-10
The above results show that even bafore twenty-four hours all the (maximum) dialyaable con-
stituents have passed out in the dialysate.
232
B e N. A chary a and S 9 C. Devadatta
IV
Amounts of Dialy sable and Undialy sable Phosphorus, Calcium
and Magnesium
(In mg. per 100 c.c. of Milk)
(Results of 30 hours Dialysis of the same Four Samples of Milk used in
Table I, respectively]
I
2
3
4
DIA.LYSABLE
UNDIA.LY8A.IiLE
! UNLHALYSA'BLE ACID-
SOLUBLE
U N I) r A L YS A BL K At 1 JJ>-
iNSOMTHLK
o
K
P
Ca
Mg
P
Ca
Mg
P
Ca
Mg
P
Ca
%
1
02-23
G'3-20
10-32
03- 13
80 -2S
2-10
4-8- 8(>
51-48
1-02
14-20
33-40
Truces
2
52-90
50.30
12-23
59-17
89-10
2-08
40-38
47-90
1-07
12-20
40-34
3
02-40
03-40
10- 8(>
05-32
01-00
2-77
48-20
50 - 50
1-10
10-87
40-83
4
55*35
55-10
15-93
00 -OS
82-40
2-90
47-86
49-20
1-20
18-80
32-82
,.
58-32
58-00
15-30
03 05
87-35
2-40
47-83
49-82
Ml
15-52
30-80
j\ r ..B.- Sum of 1 and 3 is more than the acid-soluble of column 2 in Table 1. For explanation
see text.
Similarly 4, acid-insoluble contents on dialysis are less than that in column 3, Table 1.
E V
Analysis of Milk Dialy sed for 6 and 30 Hours. (Undialy sed Acid-
Soluble and Insoluble Fractions.} Concentration of Phosphorus,
Calcium and Magnesium
(In mg. per 100 c.c. of Milk)
1
o
3
4
Casein
in gm.
TOTAL
AC ID -SOLUBLE
A CUD
-I.V.SOLU
Ca
!LK
?
Ca
Mg
P
Ca
Mg
P
JVltf
Whole milk
3-2
120-4
148-2
18-42
100-30
99-20
17-23
25-36
48-00
1-19
Milk dialysed for
Ghrs.
2-9
78-40
103-04
61-00
63-00
10-86
40-14
Dialysate after
hrs .
44-46
44-02
10-42
..
Milk dialvsed for
30 hrs.'
63-13
86-28
2-10
48-86
51-48
1-20
14-13
33-46
Dialysate
62-23
60-20
16-32
N.B. In Tables II, LIT and V, the values obtained in the case of an experiment, i.e., one sample
of milk are given. The process of heating and dialysis being semi-quantitative, the
other three sets of readings obtained are not identical with these values as is to be
expected, but are similar to them and lead to the same conclusions, They are nQt
reproduced here.
Phosphorus, Calcium and Magnesium in Milk 1 1
TABLE VI
Amount of Phosphorus in the Undialy sable Acid-Soluble Fraction
(In mg. per 100 c.c. of Milk)
As Tricalciuni and Magnesium Phosphate
233
a
b
a b c
Kxpt,
Free phosphorus,
as undialysable
acid-soluble
phosphorus
Phosphorus as
Ca 3 (P0 4 ) 2
Phosphorus as
as Mg 3 (P0 4 ) 2
Phosphorus in
organic com-
pounds
1
^8G
26-61
1-030
21-22
2
46-38
24-77
0-922
21-56
3
48-20
26-12
0-999
21-08
4
47-86
25-45
1-030
21-38
1. Acid-insoluble Phosphorus, Calcium and Magnesium. The amounts
of phosphorus, calcium and magnesium in milk, acid-soluble, and acid-
insoluble fractions are recorded in Table I. It can be seen that the amounts
of acid-insoluble phosphorus, calcium and magnesium are 25-68 mg.,
49*32 mg. and 1-21 mg. respectively. It is clear that the concentration
of the acid, or more accurately pH of the resulting solution is such that all
the calcium and the magnesium salts such as carbonates, phosphates, etc.,
must be soluble. The only source of phosphorus, calcium and magnesium
must be, therefore, from those substances which are precipitated b} 7 the
acid, i.e., the proteins mainly casein (lactoalbumin and lactoglobulin) . It
has been shown that the acid-insoluble phosphorus consists of casein
phosphorus plus lipoid phosphorus. 9
The next question is to trace the source of calcium and magnesium.
It is suggested that calcium must be in the form of a loose compound like
calcium caseinate. 10 But if we look at the molecular proportions of
calcium and magnesium and that of casein, no direct relationship is appa-
rent. A more plausible explanation, however, appears to be the adsorption
of the calcium and the magnesium ions by casein complex molecules, to
form a colloid system and that the acid coagulates this colloid and the ions
of calcium and magnesium are adsorbed by the coagulum.
9 Acharya and Devadatta, Proc. Ind. Acad. Sci., same volume.
10 Van Slyke and Bosworth, J. Biol. Chem., 1915, 20, 135 ; 1916, 24, 199.
234 11 N. Acharya and S. C. Devadatta
2. Scum of the Milk. Fresh Milk on mere heating gives rise to a scum
which when dry has the following composition: Phosphorus 274mg.,
Calcium 64 -00 nig., and Magnesium 8-OOmg. per 100 c.c. of milk (refer
Table II). The explanation of this scum formation is (1) the coagulation
of some constituents which are dispersed colloidally and (2) the formation
of insoluble salts both as a result of heating. The former phenomenon
is evident and needs no comment. If this scum was derived exclusively
from, the colloidally dispersed constituents, we should expect that the acid-
insoluble portion of milk, free from scum on analysis, should show a decrease,
corresponding to the amounts of phosphorus, calcium and magnesium
found in the scum ; but it should not affect these in that portion of milk
which is acid-soluble. It is observed that there is a decrease in the acid-
soluble constitutents also. Some colloidal substances like calcium and
magnesium phosphates have been coagulated. There is also the precipita-
tion of acid phosphates, and carbonates of calcium and magnesium, which
before heating, in the form of acid salts, are more soluble in water than the
normal salts. On heating milk for a long time in presence of few drops
of methyl orange, a decrease of acidity was observed.
3. Water-soluble and Water-insoluble Constituents !t is seen from
Table I that the amounts of phosphorus, calcium and magnesium that are
soluble in acid are respectively 96-67 mg., 99-30 mg., and 16 59 mg. per
100 c.c. of milk. Milk may contain calcium and magnesium as carbonates,
phosphates, lactates, etc., both in the form of normal salts and of acid salts.
To estimate the amounts of acid salts soluble in water, milk was dialysed
for 30 hours against distilled water, and the dialysate was analysed. In
Table IV are given the amounts of dialysable and undialysable phosphorus,
calcium, magnesium, in mg. for 100 c.c. of milk, i.e., soluble salts P = 58-32,
Ca = 58*00 and Mg = 15-36 nig. ; and insoluble salts P = 63-65,
Ca = 87 -35 and Mg -= 2 -46 mg.
For a particular sample shown in Table V it can be seen that the
amounts of phosphorus, calcium and magnesium that pass out of the bag
during dialysis for six hours are, 44-46 mg., 44-62 mg. and 10-42 mg.
respectively. The constitutents which are not dialysable but soluble in
acid are : phosphorus 61-00 mg., calcium 63-00 mg. and magnesium only
traces. Similar values are given for 30 hours dialysis. On comparing the
total of dialysable and acid-soluble undialysed constituents, Table IV,
columns 1 and 3, with the total acid-soluble constituents, Table I, column 2,
there is an increase in the amounts of phosphorus, calcium, and magnesium
present in the acid-soluble (dialysable plus acid-soluble undialysable)
portion, on dialysis. It may be noted also that there is a corresponding
Phosphorus ) Calcium and Magnesium in Milk // 235
decrease in the acid-insoluble portion (undialysable) . The increase in the
amounts of acid-soluble constituents of phosphorus, calcium and magnesium
amounts to 9-73, 8-52, and 1-11 mg. respectively, on dialysing milk for
30 hours (Table V). This is due to the fact that casein, which had adsorbed
calcium and magnesium ions, on coming in contact with water becomes
partly soluble, otherwise quite insoluble in acid. Further, it can be seen
that casein decreases in undialysable milk from 3 2 to 2 -9 gm. per cent. It
is also possible to detect some casein in the dialysate. Therefore our as-
sumption that the acid-insoluble phosphorus, calcium and magnesium as
being associated with casein is corroborated (vide below). 9
4. Acid- soluble Phosphorus, Calcium and Magnesium. The insoluble
or undialysable fraction is further subdivided into acid-soluble and acid-
insoluble parts. From Table IV, column 3, it will be seen that the amounts
of phosphorus, calcium and magnesium which are soluble in acid but
insoluble in water are as follows : phosphorus 47 -83 mg., calcium 49-82 mg.
and magnesium 1-11 rng. per 100 c.c. of milk (cf. Table V). These amounts
refer to such salts as tricalcium and magnesium phosphates. If we look to
the amounts of phosphorus we find that it is much more than what should
correspond to tricalcium and magnesium phosphates as noted in Table VI.
That surplus phosphorus may be the portion of phosphorus combined to
the acid-soluble organic phosphorus discussed elsewhere. 9
Summary and Conclusions
In conclusion, it may be said that the portion of milk insoluble in acid is
mostly casein which had adsorbed some ions of calcium and magnesium.
The portion soluble in acid but insoluble in water consists of tricalcium and
magnesium phosphates and organic phosphorus. Scum which is collected
after heating the milk is derived both from the acid-insoluble and from the
acid-soluble portions. On dialysis some acid-insoluble portion becomes
water soluble. The water-soluble portion of milk consists of acid phos-
phates of calcium and magnesium. The amounts of phosphorus, calcium
and magnesium present in water, acid-soluble and insoluble fractions and
in scum of 100 c.c. of milk are recorded.
We are grateful to Mr. P. M. Barve for his helpful suggestions.
1457 39 Printed at The Bangalore Press. Mysore Road, Bangalore City, by G. Srinivasa Rao, Superintendent,
and Published by The Indian Academy of Sciences. Bangalore
STUDIES IN SORGHUM SUDANENSE, STAPF-THE
SUDAN GRASS
BY G. N. RANGASWAMI AYYANGAR, F.N.I., I.A.S.
Millets Specialist and Geneticist
AND
B. W. X. PONNAIYA, B.Sc., AG.
(Assistant, Millets Breeding Station, Coinibatore)
R3c?.iv3d Dssember 7, 1938
Introduction
SUDAN GRASS is a Grass Sorghum introduced into America from the Sudan.
Its importance is because of its high forage value in America and also in
other parts of the world. S. sudanense belongs to the series Spontanea,
sub-section Arundinacea, in the Eu-sorghum section of the genus Sorghum. 1
Snowden after an intensive examination of several sorghums, both wild and
cultivated, comes to the conclusion that the cultivated sorghums belonging
to the sub-series Bicoloria in the series Sativa (also of sub-section Arundinacea}
are the products of some combination of S. cethiopicum (Hack.) Rupr. ex.
Stapf and S. sudanense, Stapf. He thinks it probable that in view of the
close affinities, S. dochna (of the sub-series Bicoloria) owes its origin very
largely to S. sudanense.
The reasons for this probable contribution of S. sudanense to the origin
of some races of cultivated sorghum are (1) its ready crossing with culti-
vated sorghums and the large number of grain sorghum natural crosses that
occur in it, (2) its tough racemes, (3) the persistent pedicelled spikelets, 2
(4) the absence of deciduousness of sessile spikelets through callus formation, 3
(5) size and shape of sessile spikelets, (6) the arrangement of the spikelets
in the panicle, (7) time of anthesis, 4 and (8) absence of hard seeds.
The studies presented in this paper represent experiences gained in the
course of the examination of crops raised from fifty seed samples from
various sources, including forty from Russia, during the last five years.
Agro-Botanical Description
Duration 80 to 110 days ; Seedlings coleoptile deep purple, purple,
or green 5 ; Seedling-leaves bluish green or green 6 ; Leaf-sheath blackish-
purple 7 ; Nodal band purple, stray cases green 8 ; Axil of leaf-sheath above
237
238 G. N. Rangaswami Ayyangar and B. W. X. Ponnaiya
nodal band purple, stray cases green ; Auricular junction purple, stray
cases green; Midrib white 9 ; Awn long 9 to 1.1mm., stray cases nil;
Stigma very light yellow ; Anther (fresh), very light yellow, (dry) very
light brown 10 ; Grain brown in colour, enclosed in glumes ; Glume
bleached blackish purple, stray cases unbleached ; Panicle loose conical,
secondary branches adpressed (occassionally diverging), tertiary branches
absent in some cases ; Spikelets arranged in a cymose fashion ; Emer-
gence good, about 15 cm. ; Peduncle about 40 cm. long, hollow just
immediately below the panicle ; Tillers 1.5 to 20, the main and primary
tillers (almost contemporary to it) are both unimodal in internodal disposi-
tion, the late tillers arising from the primary tillers are however of the
uniform increasing type 11 ; Bloom very sparse, stray cases apparently
nil. 12
The studies on this Sudan grass could be grouped into (1) rare charac-
ters that are common to 5. sudanense and 5. dochna (more a fodder than a
grain sorghum), (2) characters experienced so far in Sudan grass only, and
(3) those that are of evolutionary interest.
Rare Characters that are Common to S. sudanense and S. dochna
E-ligulate and Non-auriculate Condition of the Leaves. In a previous
publication it has been recorded that the g-ligulate and non-auriculate
condition in Sudan grass (gene Ig) is a simple recessive to the ligulate and
auriculate condition 14 (lyg). In many of the extracted types of Sudan grass
from Russia an ^-ligulate condition prevails. When it occurs, the plant
assumes an erect habit and looks compact and rigid (Fig. 1) and the earheads
are rod-like. As in cultivated sorghums, in Sudan grass also, when the
ligule and auricle are absent, the pulvinus is absent (Fig. 3) and there is a
shortening in the spikelet-free area in the base of panicle branches and
branchlets. 15 This leads to a compact head with panicle branches over-
crowded with spikelets. The contrast between a loose head from a ligulate
and auriculate plant and a compact head from an ^-ligulate non-auriculate
plant, is well brought out in Fig. 2. The occurrence of the ^-ligulate
condition has been recorded in S. sudanense and in Broom corn (S. dochna
var. technicum). This concurrent manifestation of the same rare character
in both these sorghums is very significant in the support that it gives about
the contribution of S. sudanense to Ihe evolution of S. dochna.
Compact- spindle Panicle. Being a ' wild ' sorghum the panicle in
Sudan grass is usually loose and conical. A rare heteiozygous mutation
was met with in family No. S. 75 (Guntur collection) in which there occurred
a segregation for panicle shape only, giving 1.61 plants with loose conical
Studies in Sorghum sudanense, Stapf~The Sudan Grass 239
panicles and 48 with compact spindle-shaped ones. The unusual occurrence
of this compact spindle (not the compactness associated with the E-ligulate
condition) in this grass is significant. This type of panicle (PaJ has been
reported upon in S. dochna This lends a second evidence to its affinity
to 5. dochna.
Pcdicelled Spikelets with a Wash of Purple. The pedicelled spikelets
in sorghum are usually unpigmented. They are green when fresh and
straw-coloured when dry. In red-grained sorghums where the sap colour
invades even the pedicelled spikelets, a tap of the ripe head will dislodge a
number of red coloured pedicelled spikelets. This is one of the characters
in the sap colour series associated with red-coloured grains. 8 In the case
of the P and Q factor manifestations of purple pigments, i.e., reddish-purple
and blackish-purple of leaf -sheath and glume, 7 the pigment never shows
well on the glumes of pedicelled spikelets. There is another type of mani-
festation of purple pigment which is related to P and Q and which shows
in certain African varieties only, even on the emergence of the panicle from
the boot. 17
In most of the Sudan grasses a new type of manifestation of purple
pigment (not of the P type) is met with. It is the presence of this type of
pigment that gives the characteristic purple-washed appearance to a fresh
panicle beginning to flower. The purple-wash is seen on the first glume of
the pedicelled spikelet which is the one that is most exposed to the sun.
When so exposed, the second glume is also coloured. The colour is deeper
at the base of the glume and is best seen at flowering time, but it disappears
at about the dough stage of the grain. This special manifestation of purple
pigment is peculiar to Sudan grass. A gene designated PW is responsible
for the manifestation of the purple pigment mostly on the exposed first glume
of the pedicelled spikelet in Sudan grass. In the F 2 , of family No. S. 75
(Guntur collection) there was a 3 : 1 segregation with 1.54 purple-washed
spikelets and 58 green spikelets. It is interesting to note that this additional
purple dominant gene which has dropped out in most of the cultivated
sorghums 17 - 18 ' 19 has been met with in Africa, an additional evidence of its
being the home of sorghum. This character was also met with in certain
varieties of S. dochna from China. The occurrence of this phenomenon in
S. dochna is an additional and third evidence for the probability of S. suda-
nense being a progenetor of the 5. dochna group of cultivated sorghums.
Characters Experienced in Sudan Grass Only
Banded-seedlings. Seedliugs with chlorophyll deficient bands- have
occurred in Sudan grass (Guntur collection). The bands are prominent in
240 G* N. Rangaswami Ayyangar and B* W. X, Ponnaiya
the broad first seedling-leaf. 20 They are white and are devoid of chloro-
phyll. The band may be at the tip, top, middle or at the sides of the first
seedling leaf. It is usually in a single and occasionally in more than one
band. The bands are 1 to 2 mm. wide. They may show in the second
seedling leaf also. This character expresses in seedling-leaves only. In
stray cases the lower leaves in a tiller have repeated this experience . The
adult plants do not give a clue to this character. In crosses between banded
and normal varieties, the normal green condition proved a simple dominant
character. The F 2 segregation gave 317 non-banded seedlings and 100
banded seedlings. A gene designated cb is responsible for inducing
albinotic bands in the seedling leaves of Sudan grass.
Seedling Habit. A study of seedlings in seed-pans shows that sorghum
seedlings differ in habit. These differences are minute in grain sorghums.
In Sudan grass with its characteristic tillering, seedling habit differences
have however been perpetuated graphically. There are pure lines in which
the seedlings have a spread-out habit, the tillers making an angle of about
45 with ground level. There are other pure lines in which the tillers are
practically erect. This seedling habit is best seen when the seedlings are
about 4 to 5 weeks old. It gets obscure on and after flowering. In crosses
between the two types (Guntur collection) the open habit has proved a
simple dominant to the erect habit. In the F 2 generation (family No.
S. 86) 86 open and 30 erect seedlings were obtained. A gene designated
SO produces seedlings with an open habit.
Striping of Leaves. Regular striping in leaves is often pathological in
origin. 21 Irregular and erratic stripes are aspects of maternal inheritance. 22
True breeding types with faint and regular white longitudinal stripes on the
leaf-blades have been recorded by Russian workers. 23 They note that such
plants were weak. Unlike their experience, which was probably due to
poor populations good monogenic segregations have been obtained
between normal green and striped leaves, the figures from three segregating
selections from family No. S. 1.73 from Russia being 130 and 32, 200 and 65,
and 125 and 40 respectively. A gene designated cs produces leaf-blades
with thin white stripes. It is interesting to record the fact that when the
leaves are striped, a few glumes exhibit a similar striping.
Bloom. In a previous paper the occurrence and inheritance of waxy
bloom has been recorded. 12 Heavy bloom is a simple dominant to sparse
bloom. It was then stated that the wild sorghum group in which was
included S. sudanense shows a distribution of bloom that is more sparse
than in the grain sorghums. The examination of many lines of Sudan grass
Studies in Sorghum sudanense, StapfTlie Sudan Grass 241
shows that, whereas the bloom is very sparse in the majority of types, there
occur types in the Guntur collection with a still sparser manifestation of
bloom, bordering on absence. On the leaf-sheath (especially on the boot at
flowering time) and on the internode, the normal Sudan grass has a very
sparse coating of bloom. In the extremely sparse bloom condition, the
Internode is practically blooniless while the boot retains traces of bloom.
Segregations have been obtained for the very sparse and extremely sparse
condition of bloom, the total of five families in both F 2 and F 3 being 219
very sparse to 70 extremely sparse. Another dilution gene seems to
determine the lightness of manifestation of the bloomy condition. The
relationship of the four types is being worked out.
Panicle. In this grass the primary branches of the panicle have
pulvinii and make an angle with the central stalk. The secondary branches
lack the pulvinii and are therefore practically adpressed to the primary
branches. Whereas the majority of Sudan grasses have such adpressed
secondary branches, there were found two pure lines one from Russia and
another from Guntur in whichrthe secondary branches had well marked
pulvinii resulting in their being at about a right angle to the primary
branches giving the earhead a fulness in look in contrast to the sketchy
appearance of the usual type (Fig. 4). In crosses between the two types,
the angular condition of the secondary branches has proved a simple domi-
nant to the adpressed condition. In the F 2 generation a monogenic segre-
gation of 49 angular to 17 adpressed plants was obtained. This behaviour
was in a loose conical type (PaJ, in which the primary branches with their
marked pulvinii ramified from the central stalk and made it easy to pursue
this character. The sparse-headed ' wild ' sorghum has thus given a
helpful clue to one of the constituents in panicle structure. A gene
designated Pa 2 thus determines the angular disposition of the secondary
branches to the primary branch. In grain sorghums that are more highly
evolved, other factors like compactness of the panicle and heaviness of the
grain make it difficult to pursue easily the effect of this gene.
Another contribution that this ' wild ' Sudan grass has to make to the
understanding of panicle structure, is the light that it throws on the arrange-
ment of the spikelets on the panicle. The sessile and pedicelled spikelets
are arranged in the form of a cyme. All the sessile spikelets represent the
main flower of the cyme and the pedicelled spikelets the lateral ones. The
whole arrangement is dichasial but with an alternate development (Fig. 6).
This explains why every sessile spikelet has a pedicelled one and the
terminal sessile spikelet two pedicelled spikelets, one of which the ultimate,
242 Go N. Rangaswanil Ayyangar and B W X 8 Ponnaiya
has always a slightly longer pedicel than that of its mate. All the sorghums,
whether wild or cultivated examined so far conform to the above descrip-
tion. In the cultivated sorghums this is not quite clear owing to the over-
crowding of the spikelets. In a cyme the middle flower is the oldest and
flowers first. This fact explains the second wave of anthesis of the pedi-
celled spikelets in the cultivated sorghums which has been described in
great detail in a previous paper. 24
A proliferated earhead occurred in family S. 52, a Sudan grass pure line
from Guntur (Fig. 5). In this the lower panicle branches turned into
vegetative shoots and thus evidenced the evolution of the panicle from
foliar organs. It is noteworthy that some of the vegetative shoots however
give out panicles once again from some of their axils.
Characters of Evolutionary Interest
Tillering. The tillering habit is an important attribute of the
Graminese. A tussock is a product of heavy tillering. In the evolution of the
cereals with their definite sowing and harvest time, a fairly uniform matu-
rity of earheads is a requisite. Under crowded conditions, cereals can be
few headed, provided they are fairly uniform in duration. In sorghum a
practically single stalked condition has been bred up from a many tillered
condition by not giving full scope for the tiller buds to develop. Although
tillering is affected by spacing, the capacity of sorghum varieties to tiller
varies with varieties. Among the cultivated varieties, S. durra is normally
non-tillering and Feterita belonging to S. caudatum is usually a tillering
variety. Tillering connotes wild vigour and non-tillering condition is
brought about by cultural and selective operations, until the whole plant
puts forth its undeviated vigour into the production of one large earhead.
The following experiences met with in Sudan grass help in understanding
this valuable attribute of grasses, viz., tillering.
Among the many pure lines of Sudan grass that are being grown at
the Millets Breeding Station, S. 77 is one belonging to the collection from
Guntur. This was true to the tillering habit in 1935. In 1936 the single
plant selection carried forward from the 1935 crop, instead of coming pure
as expected, segregated throwing out a few strange plants, anaemic, tillerless
and absolutely single stalked (Fig. 7). There were 78 plants with tillers
like the parent and 20 of the strange weak group. It was obvious that
there had occurred a mutation heterozygous in nature. From this F 2 ,
an F 3 generation was raised and the behaviour of the selections is
given below :
Studies in Sorghum sudanense, S tap f The Sudan Grass 243
TABUS
Selection
Number
Character of the
F 2 Selection
F 3 Behaviour
Tillering
Single-
stalked
S. 103
Single-stalked
. .
Pure
S. 105
Tillering . .
Pure
. .
S. 103
??
Pure
S. 104
?)
49
14
S. 106
77
70
26
S. 107
77
96
22
S. 109
J?
TOTAL . .
134
48
349
120
Calculated 3:1 . .
351 -75
117 -25
X 2 = -086 P > -7
It will be noted that 4 of the 6 tillering plants were heterozygous and
segregated again. It is clear that the tillering habit is a monogenic domi-
nant to the absolutely single-stalked non-tillering condition which has been
met with for the first time in this sorghum. Out of the 1.20 single-stalked
plants only 8 survived till maturity, the rest succumbing to the attack of the
shoot borer. In the pure line S. 1.03, out of the 1.7 seedlings that germinated
only one reached the stage of maturity. Being weak, the main shoot dying,
and unable to produce tillers (basal buds absent), these anaemic plants are
of very poor survival value. Before discussing this phenomenon, the
contrasting characters in the two groups, viz., tillering and absolutely
single-stalked, are given below :
244 G. N. Rangaswami Ayyangar and B. W. X. Ponnaiya
Character
Tillering plants
Single-stalked plants
Tillers
12 to 15
Nil (basal buds absent)
Vigour
Healthy
Sickly
Seed-setting
Good
Poor (10 to 15%)
Panicle ;
Shape
Loose and conical
Compressed and rod-like (Fig. K)
Top
Normal
Ending in a a pur
Number of whorls
16 to 18
10 to 12
Pulvirms
Present
Absent
Primary branches
About 55 to the axis
Adpressccl to the axis
Spikelets and their distribution
Sessile spikelets
Many, evenly distributed
Few, crowded at tips of
branches (Fig. 9)
Number
1000 to 1200
Less than 200
Size
6-0 X 2-75mm.
7-25 X 225 mm.
Sessile spikelets
Nature
Coriaceous and short-nerved
Papery and long-nerved
Length of nerves on glumes
2 -5 mm.
5-5 mm.
Pedicelled spikelets
Number
1300 to 1500
Loss than 50
Size
5-0 X 1-25 mm.
Mere scales or absent
Length of pedicel
2-0 mm.
Do.
Flowering
Ordinary
Partly cleistogamous
All these connote a primitiveness of equipment in the single stalked
plant that has naturally not helped in the survival of this type in nature :
The ready susceptibility to attack from the seedling-borer and the poor
setting of seed make it difficult to perpetuate the single-stalked pure line.
With the help of this experience it has now become possible to give a
genie background to this common phenomenon of tillering. A gene
designated TX is at the back of the tillering habit in S. sudanense.
Gene tx gives rise to an absolutely single-stalked (tiller-less) plant. The
capacity to produce tillers is absent through the absence of buds at ground
level. This is different from the single stalk of the grain sorghum in
which the buds are present and could be activated. The higher axillary
Studies in Sorghum sudanense, StapfThe Sudan Grass 245
leaf-buds could however be stimulated into activity, but since the frame of
the plant is so frail this potentiality leads to no fruitful result. The spur
that plumes the panicle seems to be the long central axis prolonged. The
tillering habit, short coreacious glumes, short nerves on these and the
presence of pedicelled spikelets attributes of modern sorghums show the
great advance in their evolution.
Whereas the above experience chronicles a case of tiller versus no tiller,
Sudan grass has afforded a second interesting experience in the nature of
advance in the tillering habit. Among the pure lines of Sudan grass there
exist two distinct tillering types. The maj ority of pure lines have the first
shoot well ahead of the later straggling tillers. In a few types the tillers
flower practically along with the main stalk. At the flowering time the
two types are very readily distinguishable, the latter being weak. In
crosses between these two types the comparatively un-uiiiform flowering
type proved a simple dominant to the dead uniform flowering type. In
family No. S. 200, also from Guntur, there occurred a segregation giving
64 plants with tillers maturing much later than the main head and 22 plants
with tillers maturing almost with the main head. The latter are shorter,
their stalks are thinner, and the panicles are smaller, and there are no late
tillers. It looks therefore that the lack of vigour in this group has been
brought about by the over rapid activation of buds and the unspaced
development of tillers arising from such activation.
A gene TU seems to be responsible for a gradual activation and delayed
and spaced growth of tillers in Sudan grass. Gene tu gives rise to a uniform
activation, uniform growth and maturity and consequent weakness of the
tillers and the plant in general. These experiences in the tillering habit of
Sudan grass are valuable in giving a clue to the fact that behind the vigorous
tillering and the suppression of such tillers resulting in the single-stalked
condition of Sorghum the Great Millet there exist genetic factors
whose detection is difficult, under the obscuring effects of continued
cultivation.
Weak Midrib. Another phenomenon of great evolutionary interest
is the weak midrib. In a family raised from selection S. 92 received from
Trivandrum, which must have been a mutant (heterogygous), there occurred
a number of plants very abnormal in appearance. In them, instead of the
characteristic rigid leaves, the leaves were ribbon-like, weak, supple and
bent down. Counts taken showed that there were 36 normal plants with
a normal midrib in the leaves and 10 plants with a weak midrib. The weak-
midrib-plants had many disabilities with the result that they did not produce
246 G* N. Rangaswami Ayyangar and B. W. X Ponnaiya
any seed. The occurrence of this rare character necessitated a large
number of selections being carried forward to the third generation. Out
of the 36 normal plants available, except the two that were very poor,
all the others were carried forward and a third generation raised. Of
the 3-i selections, 1.1 were pure for normal plants and 23 segregated
repeating the F 2 experience. The figures are given below :
TABI,E
F z from S. 92 Family
Select! en
dumber
Character of
Selection
F 3 Behaviour
Midrib
formal
Weak
S. 118
Normal Midrib
27
9
119
j?
24
9
120
7? . .
70
22
121
77
46
13
122
77 '
119
34
123
77
50
17
124
77 "
59
23
125
7?
106
33
126
77
174
54
128
77 '
57
19
131
7? *
153
52
132
77 * -
67
22
133
77 -
172
55
134
77
17
4
135
77 *
85
28
Studies in Sorghum sudanense, Stapf The Sudan Grass 247
Selection
Kumber
Character of
Selection
F 3 Behaviour
Midrib
Normal
Weak
S. 136
Normal Midrib
141
50
137
?>
107
34
138
>> *
42
11
141
?> -
41
.13
142
?? * *
33
11
144
?
74
26
147
>?
69
29
149
j?
TOTAL . .
122
4:6
1855
614
Calculated 3:1 . .
1851-75
617 -25
X 2 = -023
P < -8
A gene designated MD is responsible for producing a strong and normal
midrib in the leaf-blade of sorghum. Gene md results in a weak midrib,
which is the cause of the ribbon-like leaf -blades that lack erectness (Fig. 10).
Among the families which segregated and gave the weak-midrib there
was also a segregation for the bluish-green 6 and green colour of (seedling)
leaves in 14 of them. The cross-collated tabulation given below shows
that the factor MD is independent of the factor CBL '
J J C3
Selection dumber
Strong Midrib
Weak Midrib
Eluisli-
green
Green
BIuLsIi-
Gi-cen
8. 118
19
8
8
1
119
17
n
I
6
3
120
53
17
17
5
122
97
22
27
7
125
86
20
27
6
126
135
39
39
15
128
47
10
13
6
132
55
12
18
4
133
130
42
36
19
134
12
5
2
2
135
64
21
20
8
144
57
17
19
7
147
49
20
23
6
149
TOTAL (actual) . .
Calculated (9:3:3:1) ..
97
25
33
13
918
265
288
102
884 -7
294 -9
294 -9
98-3
X 2 = 4-58 P> -20
The weak-midrib is weak up to 5 cm. from the top of the leaf -sheath
after which it loses its individuality and becomes like one of the nerves in
the blades (Fig. 1.1). Even this trace of the midrib is lost in the top leaves
and the flag is midribless. Detailed measurements taken in 1.0 plants in
each of the groups, normal and weak-midribs, show that in height of plants,
thickness of iuternodes, number of leaves, length and breadth of leaf -blades,
duration and in general panicle shape, both the groups are alike. Besides
the weakness in the midrib, the only difference is a distinct pull-down in the
tillering capacity which is inevitable owing to the disability imposed upon
the photosynthetic equipment. In the normal plants the thickness of the
midrib of the fourth leaf from the top is 2-0 mm. at the base, 1-2 mm. at
the middle and 0-027 mm. at the top which is the same as the thickness
of the blade. In the case of the same leaf with the weak-midrib it is - 7 mm.
at the base, 0-5 mm. at the middle, and 0.27mm. at the top. These
measurements explain the drooping down of the ribbon-like leaves.
The consequences attendant on the presence of the weak-midrib are
of still greater interest. Owing to the rarity of this occurrence, practically
every plant with a weak midrib was carefully examined in all its parts.
Along with the suppression of the midrib there is a suppression of the style
and stigma (Fig. 12). In most cases these organs are completely absent
and in stray cases vestiges of the style are present. Concurrent with the
suppression of the style and stigniatic tissue there is a suppression of the
tissue of the awn. Whether the spikelets are 'nil '-awned or long-awned
the normal lemma is distinctly bifid, with a strong awn-base which protrudes
only a short distance if it is ' nil '-awned (Fig. 13) or to a greater distance
if it is long-awned (Fig. 14). In the weak-midribbed plants when the style
and stigma are completely absent, the lemma is entire (bifid nature not seen)
(Fig. 15) and there is no specialised tissue connoting the awn-base. When
occasionally the vestiges of the styles are present, there is concurrently
a faint manifestation of the awn-base (Fig. 1.6).
The genetics of the awn has been worked out and the ' nil '-awned
condition (extreme reduction in length) is dominant to the long-awned
condition. In a dihybrid segregation from normal and weak-midrib, and
'nil'- and long-awned condition instead of the theoretical 9:3:3:1 ratio,
there -occurred a 9 : 3 : 4 ratio as will be seen from the following table.
TABI,
Normal-midrib
Weak-midrib
Selection Number
TSTil'-awn
Long-awn
(9-11 mm.)
'Nil '-awn
Long -awn
(9-11 nun.)
S. 118
18
9
9
. .
122
89
30
34
12-4
43
16
23
125
77
29
33
126
134
40
54
128
39
18
19
132
46
21
22
133
128
44
55
135
63
22
28
137
84
23
34
141
29
12
13
144
57
17
26
TOTAL (aeutal) . .
807
281
350
Calculated (9:3:4)
809-1
269/7
359-6
X 2 =0-74 P>0-50
The awn segregation in the weak-midrib plants is not felt, due to the
lack of awn tissue (homologous to the midrib). Gene md which affects
the midrib affects also the homologous awn tissue. The very intimate
connection between the midrib and the awn easily explains this phenomenon.
It has been shown, that when the leaf-blade is forked, the awns are forked
likewise. 25 When a severe attack of Sclerospora occurred and affected the
leaf-blade, there was a suppression of the awn. 26 The effect of the weak-
midribbed condition does not stop with this concurrent effect on the
liMt' blatlf and the awn, but also extends to the style and stigma. The
lsninultn;y between stigma and awn in sorghum has been given in great
<li;t"il in a immU. r of publications. 1 '. ~ 7 2H - Gene mcl affects the style and
stigma vitally and they are entirely absent or only the rudiments of the
si vie an* pivse-nt.
This accounts for the non-setting of seed in the flowers of the weak-
nndiiblu'd plants, though the earliead is normal and the anthers emerge
t|tiitf normally. Tlu- otlier effects of tliis wcak-midribbed condition are the
occasional presence of a fully developed dorsal lodicule 30 and in stray
casrs a srcond set of three anthers.
These experiences are of very great interest in the evolution of plant
organs. The homology between the leaf-blade and awn, and between style
and stigma and the column and subtile of the awn has been well established.
This experience in Sudan grass threads up all the three homologous organs,
riz. t midrib, awn and stigma. A serious disturbance in vegetative equip-
ment produced catastrophic effects on the homologous reproductive
equipment/* 1 v Such abnormal characters, resulting in sterility, could be
produced by X-ray treatment, but their occurrence in nature is possibly
to be explained through mutation induced in a new environment.
Coming to grain sorghums an instance is on record in which a cross
between S. durra of Coimbatore and S. nigricans of Tanganyika (with
pcclicelled spikelets fertile) 32 gave in the F 2 a few plants whose side-shoots
showed midribless leaves. Another interesting experience is worth
recording here. Bulbils in carheads are a rare occurrence. 33 One such
bulbil was nursed till it produced an carhead. In that plant of abnormal
origin two leaves had no midribs. The first seedling-leaves in all sorghums
do not; elaborate a midrib. 20
These interesting experiences are useful in throwing helpful light in the
evolution of the Gnuninece, their leaf-sheath, leaf-blade and ligular and
auricular equipment, glume and awn structures, lodicular, stylar and
stigmatic elaborations, and help us to appreciate the big advance they have
made in having their present equipment which has helped them to become
the premier group among crop plants.
Summary
An agro-botanical description of the Sudan grass (S. sudanense, Stapf)
is given and the many affinities to cultivated sorghums are mentioned.
Sudan grass is believed to have contributed to the origin of the cultivated
sorghum S. dochna. The rare and simple recessive characters, viz., 0-ligulate
leaves and compact-spindle panicles and the rare dominant character, purple-
washed pedicelled spikelets (gene PW) occur in both S. sudanense and in
S. dochna. Besides these, characters experienced only in Sudan grass so
far, are also described. Seedlings with banded chlorophyll deficient areas
have been met with. This character has proved a simple recessive to the
normal whole green. A gene cb is responsible for this character. There
are two types of seedling habits, viz., open and erect. The former is a
simple dominant to the latter and the gene SO is behind this character.
Rare types with faint longitudinal stripes on leaves have occurred and have
proved a simple recessive (gene cs) to the normal non-striped condition
(gene CS). There is a great pull down in the waxy bloom condition in this
grass. Even in this poverty of manifestation the very sparse condition
(on leaf-sheath and internode) is a simple dominant to the extremely sparse
condition (traces on the boot only). The gene controlling the angle which
the primary branches make with the central stalk of the panicle does not
determine a similar disposition of the secondary branches to the primary
one. A new gene Pa 2 makes the secondary branches pulvinate and diver-
gent to the primary branch. Gene pa 2 results in the absence of the pulvinus
in the secondary branches (and consequently the secondary branches are
adpressed to the primary branch). The Sudan grass has thrown light on
the evolution and arrangement of the spikelets in sorghum in general, as
the spikelets are sparse. The spikelets are arranged in a dichasial cyme
with an alternate development. The sessile spikelets represent the older
ones and explain the two waves of anthesis in sorghum. A proliferated
earhead in which the lower panicle branches turned foliate is recorded.
Cereals, which are also grasses, tiller. Tillering is an aspect of vigour. In
Sudan grass the gene responsible for tillering TX could be isolated and is
a simple monogeiiic dominant to the non-tillering condition tx. Gene tx
produces weak single-stalked plants in which the basal and underground
axillary buds are absent. Their panicles are weak, spurred and rod-like,
and the spikelets are few and cluster at the terminals. The glumes are long,
papery and develop long nerves. The pedicelled spikelets are practically
absent being reduced to mere scales. The plants do not survive the shoot-
borer attack and this is due to the inability to produce tillers in replacement
of the main shoot. Gene tx has however no effect on the higher axillary
buds. Next to the tiller-less single-stalked condition is the uniform tillering
habit. Gene tu is responsible for this. Though many tillered, their gush
of growth (want of a slow spaced development induced by gene TU) results
in exhaustion and consequent weakness. TU is a simple monogenic domi-
nant to tu. A strong midrib is an essential equipment in the long
auud,uciiae,
/ ne Sudan Grass
253
k>aW)lacies in grasses, especially in cereals. Gene MB produces the normal
strong midrib. Gene md results in a weak-midrib that makes the leaf
ribbon-like mid droop down. The midrib being homologous to the awn
and stigma, there is a concurrent suppression of the awn and of the stylar
and stigma! ie organs. Gene MI) has been found to be independent of gene
C m . which produces bluish-green seedling leaves.
Hnowdon, ?. I).
;as\vami Ayyangar,
and
Ponnaiya, B. W . X.
, ct al.
7.
, et al. . .
8.
- - - -
9.
-"--- --
10.
- --~ ,/rfoZ. ..
II.
-- - , ct al*
12.
- - , el al.
i:>,
and Rao,
V. P.
M.
, et al. . .
15.
, et al*
Ml.
- - and
Ayyar, M. A. S.
17.
- - - - - and
Ponnaiya, B. W. X.
18.
- - - "-- -, el al.
19.
..-. - , etal. ..
20.
--' ----- - , et al.
21.
Ghorian, M. G., and
Kailasam, M. B.
22.
Ilaixgaswarni Ayyangar,
G. N., and Ayyar,
M. A. S.
23.
Favorov, A. M., and
Il'avcnsclman, P. 8.
Bla
LITEIiATiTRB CITED
The Cultivated Races of Sorghum, 1936.
Curr. Sci., 1937, 5, 538-39.
Ibid., 1936, 5, 299-300.
Ibid., 1937, 6, 158.
Agric. J. Ind., 1930, 25, 2,62-63.
Curr. Sci., 1938, 6, 556-57.
Ind. J. Agric. Sci., 1933, 3, 589-94.
Madras Agriv. J., 1934, 22, 1-11.
Ibid., 1935,23,350-52.
Ind. J. Agric. Sci., 1933, 3, 605-08.
Proc. Ind. Acad. Sci., 1938, 7, 161-76.
Ibid., 1937,5,4-15.
Curr. Sci., 1935, 3, 540-42.
Ind. J. Agric. Sci., 1935, 5, 539-41.
Proc. Ind. Acad. Sci., 1938, 7, 286-88.
Ibid., 1938, 8, 100-07.
Curr. Sci., 1037, 5,590.
Ibid., 1938, 6, 612-13.
Proc. Ind. Acad. Sci., 1938, 8, 317-23.
Ibid. 9 1938, 8, 151-56.
Proc. Assoc. Econ. Biolgists, 1936, 1-7.
Madras Agric. /., 1934, 22, 407-08.
Herb. Jlev., 1934, 2, 143-47.
254 G* N* Rangaswami Ayyangar and B e W. X. Ponnaiya
24. Rangaswami Ayyangar, Indian J. Agric. Sci, 9 1931, 1, 445-54,
G. N., and Rao, V. P.
25. ,etal. . . Curr. Sci., 1935, 4 , 316-17.
213. and Hariharan, Ibid., 1935, 3 5 363-64,
P. V.
27. and Rao, V. P. /6id., 1935, 4, 176-77.
28. and Reddy, T. V. Ibid., 1936, 4, 817-20.
29. . . Madras Agric. J., 1938, 26, 1.23-26.
30. Long, B. . . Bot. Gas., 1930, 89, 154-68.
31. Anderson, E., and De Annals of Bot., 1935, 49 9 671-88.
Winton, D.
32. Rangaswami Ayyangar, Curr. Sci., 1935, 3, 433-34.
G. N., and Rao, V. P.
33. . . Ibid. 9 1935, 3, 362-63.
(,\ /V. .Kaugas-wawi Ayyaug-ar and Proc. Ind. Acad. ScL, B, vol. X, PL VII f
./*, ll\ .A*. Pounaiya
K-lfcuUte
1 aiuclcs
Central Axis of Panicles
K-Hgulatc Plant of Sudan Grass
G. N, Rangaswami Ayyangar and
B. W. X. Ponnaiya
Proc. 2nd. A cad. Sci., B, vol. X, PL IX
FIG. 4
Divergent
Adpressed
FIG. 5
Panicle lower branches turning foliate
FIG. 6
Arrangement of Sessile and Pedicelled Spikelets
Panicle of Sudan Grass
N. Rcingaswami Ayyangar and
B. JJ 7 . X. Ponnaiya
Proc. hid. A cad. -Sri., B, vol. X, PI,
FIG. 7
Whole Plants
Spur
FIG. 9
Panicle Branches
. A 7 ". Rangaswami Ayyaugar ami
B. IP. X. Ponnaiva
Proc. fntt. Acad. Sc/., //, vol. A, /'/. A./
FIG. 10. Whole Plants
Fio. 12. Ovaric-s
FIG. 13
' Nil '-awn
Top of lemmas lobed
FIG. 14
Long-awn
Fully lobed
FIG. 15
No awn traces
No lobes
FIG. 16
Faint awn base
Feebly lobed
LAMELLIBRANCHS FROM THE BAGH BEDS
BY 0. W. CiiipivONivER, M.Sc.
(/'Vow I he De})(trhnmt of Geolotjy, Benares Hindu University)
Received September 12, 1939
(Communicated by Prof. L. Kama Ran, M A.., F.G.s.)
I, IKK the rhyuchouellids 1 and bryozoa 2 from this formation, lamellibranchs
also have not uptil now, received proper attention. As a species useful
for purposes of correlation Duncan 3 had mentioned among the Bagh fossils
the presence of Ncithea quadricostata, Sowerby, a species marking the Upper
(reeti Sand horizon in England and many other countries. P. N. Bose 4
tfiivc a list of lamellibranch species, which, however, as mentioned by him,
were only tentatively diagnosed, and as such have helped us little in
assigning any definite age to the Bagh Beds. Irately P. N. Mukerjee 5 has
studied a small collection of molluscan fossils from the Jliabua and AH
Rujpur States. But the material available for him, being much ill-pre-
served has contributed little to our knowledge of the lamellibranch fauna
of this formation. Protocardium pondicherriense d'Orb., and Cardium
(Trachycardium) incomptum Forb., both from the Trichinopoly stage of the
vSouth Indian Cretaceous vSeries, are the only lamellibranch species which
lie could definitely identify from among his collection ; while the rest of
them are either diagnosed only genetically or names of the comparable
species are mentioned.
The material collected by the present writer being, however, quite
extensive and well preserved has yielded a large number of species with
more or less well-marked Cenomanian affinities, thus confirming the results
of the study of other groups of fossils from this formation.
Besides the species described below, there are in the author's collec-
tion some specimens of Ostrea, which are, however, too fragmentary to be
described.
1 (Jlii plonker, 1938, pp. 300-16.
2 Oliiplonkor, 1939 a, pp. 98-109.
3 Duncan, 1865, p. 354 ; 1887, p. 84.
4 Hose, 1883, pp. 37-43.
5 Mukerjee, 1938, p. 196.
255
B2 , F
2
256 G, W. Chiplonker
Description of Species
Family . . Pinnidse.
Genus . . Pinna I^inneus, 1758.
Pinna mathuri sp. nov.
(Plate XIII, Fig. 4)
Description. This species is represented by two incomplete specimens.
The surface of each valve is angulated in the middle thus giving a rliom-
boidal cross-section. The valves are not fissured medianly. The dorsal
part of each valve is almost flat and the ventral one feebl} 7 " convex. The
growth lines are very fine and are seen as low indistinct folds near the
ventral margin. The surface of the valves is ornamented with seven or
eight ribs on the dorsal part and six or seven ribs of the same strength on
the ventral part. Each of the shallow concave interspaces carries an
intercalary rib. All the ribs are slightly but conspicuously flexuous.
Comparison. This species can be distinguished from Pinna laticostata
Stoliczka, 6 from the Utatur and Ariyalur stages of the Cretaceous Series
of Trichinopoly District, by the presence of intercalary ribs in all the inter-
spaces. P. arata Forbes 7 can similarly be distinguished from the species
described here, by the former having the intercalaries only on the dorsal
part of its valves.
As compared to P. vanhcepeni Rennie, 8 an upper Cretaceous species
from Pondoland, the present species has its valves less angulated in the
middle, the ventral part of the valves carries more ribs and there is an
intercalary in each of the interspaces. But with all these differences these
species appear to be closely related.
Occurrence. Nodular limestone and Deola-Chirakhan Marl.
Family . . Pernidae.
Genus . . Inoceramus Sowerby, 1819.
Inoceramus pseudo-latus sp. nov.
(Plate XII, Fig. 3)
Description. The shell is rather tumid and oval, and obliquely elong-
ated. The anterior side is short and rounded. The shell is elongated
postero-dorsally. The surface of the shell is covered with thick fairly close-
set folds and concave depressions which are much wider than the folds.
6 Stoliczka, 1871, p. 385, pi. 25, figs. 2-3 ; pi. 26, fig. 4.
7 Forbes, 1846, p. 153, pi. 16, fig. 10 ; Stoliczka, 1871, p. 384, pi. 24, fig. 5 ; pi. 25,
fig. 1 ; pi. 26, fig. 5.
8 Bennie, 1930, p. 172, pi. 18, fig. 9.
Lamellibranchs from the Bagk Beds 257
The umboues are acute and slightly turned. The greatest inflation of the
shell is situated a little above the middle. The shell has in its posterior
third, a ridge-like elevation extending from the umbo to the postero- ventral
extremity.
Comparison. In comparison to Ino. multiplicatus Stoliczka 9 from the
Trichinopoly stage of the South Indian Cretaceous, the present species
has its shell more oblique, longer and more tumid ; its umbones are more
anteriorly situated ; the concentric folds on the surface are closer and the
shell has a ridge-like elevation.
Ino. kilns Mantell 10 from the Cenomanian of Central Europe and England
very closely resembles this species. The difference, however, lies in the
latter having the ridge-like elevation and a less apicate shell.
Occurrence. Deola-Chirakhan Marl.
Inoceramus lamarcki var. indicus var. nov.
(Plate XII, Fig. 1.)
Description. The shell is very high and obliquely prolonged postero-
ventrally. The umbones are produced and slightly incurved. The surface
of the shell is covered with distant and rather prominent folds, and indis-
tinct concentric striations.
Comparison. This species is comparable with Ino. simplex Stoliczka 11
from the Ariyalur stage of South India. The latter species has, however,
a shell less oblique and shorter, and the posterior margin is less convex.
As compared to Ino. lamarcki var. cuvieri Sowerby, 12 from the Upper
Chalk of England to which it is very closely related, .the present form has
its umbones less incurved and its shell less inflated
Occurrence. Deola-Chirakhan Marl.
Inoceramus sp. A.
(Plate XII, Fig. 5)
Description. The shell is oval to pear-shaped, nearly as high as long
and not much inflated. The anterior side is short and rounded ; the ventral
margin is strongly convex and has a weakly developed angulation. The
9 Stoliczka, 1871, p. 406, pi. 28, fig. L
10 Sowerby, 1829, pi. 582, figs. 1-2 ; Mantell, 1822, p. 216, pi. 27, fig. 10 ; d'Orbigny,
1844-48, p. 513, pi. 408, figs. 1-2 ; 1850, p. 197 ; Morris, 1854, p. 170 ; Zittel, 1866,
p. 100, pi. 13, fig. 7 ; Geinitz, 1872-73, p. 45, pi. 13, figs. 4-5 ; Woods, 1904-12, p. 279,
pi. 48, figs. 5-6, text-fig. 36.
11 Stoliczka, 1871, p. 40,8, pi. 28, figs. 3-4.
12 Woods, 1904-12, p. 320, pi. 53, fig. 7, text-figs. 73-84.
B2a F
258 G, W. Chiplonker
surface is covered with rather distant, prominent, concentric folds alter-
nating with broad depressions. The umbones are pointed and slightly
turned. The thickness of the shell is approximately one-third of the length
and is situated a little above the middle of the shell.
Comparison. As compared to Inc. crispianus Stoliczka 13 (non Mantell)
from the Ariyalur stage of the Cretaceous Series of Southern India, the
present species is much shorter and flatter.
Ino. cf. percostatus Miiller 14 from the Trigonia Sandstone of the North
Saghalin, in comparison to the present species, is longer and has its con-
centric folds a little further apart.
Occurrence. Nodular limestone and Deola-Cliirakhan Marl.
Inoceramus sp. B.
A few large-sized specimens in fragmentary condition are comparable
to Ino. striatus Mantell 15 from the Cenomanian and Senonian of Bohemia.
Occurrence. Deola-Chirakhan Marl and probably also the Nodular
Limestone.
Family . . Pectinidse.
Genus . . Neithea Drouet, 1824.
Neithea morrisi Pictet and Renevier
(Plate XII, Fig. 7)
1845 Pecten quinque-co status Forbes, Q.J.G.S., Vol. I, p. 249 (in
part).
1858 Janira morrisi Pictet and Renevier, Mater. Pal. Suisse.,
Ser. 1, p. 128, pi. 19, fig. 2.
1865 Janira morrisi Coquand, Aptien Espagne, p. 341.
1870 Janira morrisi Pictet and Campiche, Mater. Pal. Suisse.,
Ser. 5, p. 244.
1901-02 Vola morrisi Choffat, Faune Cret. Port., Vol. 1, Ser. 4,
p. 147, pi. 4, figs. 5-6.
1903 Pecten (Neithea) morrisi (Pictet and Renevier) Woods, Mon.
Cret. Lam. t Vol. 1, p. 201, pi. 39, figs. H-13.
1916 Neithea morrisi (Pictet and Renevier) Douville, Mem. Acad.
Sci. t Vol. 54, Ser. 2, p. 171, pi. 22, figs. 17-18.
1934 Pecten (Neithea) morrisi (Pictet and Renevier) Nagao, J. Fac.
Sci. Hok. Imp. Univ., (4), Vol. 2, p. 206, pi. 26, figs. 2-6.
13 Stoliczka, 1871, p. 405, pi. 25, figs. 1-3 ; pi. 28, fig. 2.
14 Yabe and Nagao, 1925, p. 115, pi. 28, figs. 7-8 ; pi. 29, fig. 10.
15 Geinitz, 1871-73, p. 210, pi. 46, figs. 9-13 ; 1872-73, p. 41, pi. 13, figs. 1, 2, 9, 10.
Lamcllibranchs from the Bagh Beds 259
Description. 'Phis species is a small race, the largest of the specimens
measuring 24 -4 mm. in height. The shell is fan-like and a little higher
than long. The ears are small and smooth except for the faint growth
lines. The convex valve carries six, convex, major radial ribs with two
secondary ribs much smaller than the major ones, in each of the five inter-
spaces. The major ribs are flanked by very fine thread-like costse on
either side. The interspaces are concave and narrower than the ribs in
the dorsal region, but become nearly equal to them near the ventral margin.
The lateral areas on the convex valve carry two or three, rarely four, very
fine radial ribs.
Comparison. This is the most abundant of the lamellibranch species
in the Chirakhan area. These specimens from the Bagh Beds agree essen-
tially well with the Aptian-Albian type of Pictet and Renevier, except
that the fine thread-like costse on either side of the primary ribs are absent
in some of the specimens, and the areas carry two or three radial rib-lets
such as are found in the Japanese specimens, 16 though Woods 17 says that
in N. monisi the areas are typically smooth.
Duncan 18 and P. N. Bose 19 had reported the presence of the widely
distributed species Neithea quadricostata Sowerby and N. quinquicostata
Sowerby, in the Bagh Beds. An examination of the material in the collec-
tion of the writer as well as in the collection of the Geological Survey of
India, has not revealed any specimens that could be assigned to either of
these almost world-wide species of Sowerby. All these specimens, however,
clearly belong to this widely occurring species of Pictet and Renevier.
Occurrence. Deola-Chirakhan Marl.
Family . . Spondylidae.
Genus , . Plicatula I,amarck, 1801.
Plicatula spini-costata sp. nov.
(Plate XIII, Fig. 3)
Description. It is a small-sized species having the shell of an oval
outline, the lower valve very feebly convex and the upper one nearly flat.
The surface of the shell is ornamented with numerous rather coarse spiny
ribs all of sensibly the same size ; as a common feature the ribs do not
increase by division at the successive growth stages.
16 Nagao, 1934, p. 207.
17 Woods, 1900-03, p. 202.
18 Duncan, 1865, p. 354 ; 1887, p. 84.
19 Bose, 1883, p. 40.
Comparison. This species might at first sight be confused with Plica-
tula nmlticostata Stoliczka 20 from the Trichinopoly stage of the South Indian
Cretaceous Series. The present species has, however, its upper valve flat-
tened and the lower one is comparatively much less convex ; its ribs do not
generally divide and are of sensibly the same size.
The species described here is very similar to PI. auressensis Coquand 21
from the Cenomanian of Tunis and Egypt. The North African species
has, however, in some individuals its upper valve feebly concave.
The present species shows the closest resemblance to PL aspera
Sowerby 22 from the Alpine Turonian and Carnpanian. But the latter species
has its lower valve more convex.
It would thus clearly appear that Plicatula spinicostata is related on
the one hand to PI. aspera Sow. from the Alpine Turonian and Carnpanian
and to PL auressensis Coquand from the Cenomanian of North Africa on
the other hand.
After Neithea morrisi Pictet and Renevier, this is the next most abund-
ant species in these strata, of the Narbada Valley.
Occurrence. This species is recorded from the Deola-Chirakhan Marl
and the Coralline Limestones.
Plicatula batnensis Coquand
(Plate XII, Fig. 5)
1889 Plicatula batnensis Coquand Thomas and Peron, Moll.
foss. Tunisie, p. 205, pi. 26, fig. 16.
1912 Plicatula batnensis Coquand Pervinquiere, Etude Pal.
Tunisienne, p. 1.62, pi. 9, fig. 21.
1917 Plicatula batnensis Coquand Fourtau, Catalogue invert.
foss. Egypt., p. 22.
Description. The shell is broadly oval with both the valves very
feebly convex. The surface is covered with numerous and rather coarse
ribs without spines. The ribs do not increase by division. The growth
stages are very scabrous.
20 Jorler, 1.846, p. 155, pi. 17, fig. 3 ; Stoliczka, 1871, p. 44(5, pi. 34, figs. 15-18 ;
pi. 46, figs. 5-6.
21 Lartet, 1880, p. 137, pi. 11, figs. 21-22 ; Thomas and Peron, 1890, p. 204 ;
Pervinquiere, 1912, p. 156, pi. 11, figs. 2-18; Fourtau, 1917, p. 20.
22 d'Orbigny, 1844-48, p. 686, pi. 463, figs. 11-12; 1850, p. 254; Zittel, 1865,
p. 120, pi. 19, fig. 1.
Lamellibranchs from the Bagh Beds 261
Comparison. These specimens from the Bagh Beds agree essentially
with PL batnensis Coquand from the Cenomanian of Tunis and Egypt.
The only difference between the African representatives and these speci-
mens from the Narbada Valley is that the latter have a slightly smaller
number of ribs, a variation which is not unexpected in a species occurring
in such widely separated regions.
Plicatula instabilis Stoliczka 23 occurring in the Ariyalur stage of South
India and in the Campanian of Tunis might at first sight be confused with
the present species ; but the former species has fewer and much weaker
ribs, which increase by division.
Ivike the previous species this is also quite common in the Chirakhan
area.
Occurrence.' -Deola-Chirakhan Marl and Coralline Iyimeston.es.
Family . . Mytillidse.
Genus . . Modiola Lamarck, 1758.
Modiola inflata sp. nov.
(Plate XIII, Fig. 2)
Description. -This species is roughly oblong and rather tumid at the
anterior third of the length. The unibones are low and near the anterior
side. The posterior region is slightly taller than the anterior one. The
posterior margin makes an angle of about 55 with the very feebly convex
ventral margin. The dorsal margin is almost straight. Anteriorly the shell
is well rounded. The surface of the shell is covered with concentric growth
lines. A ridge extends from the umbones postero-ventrally, with a feeble
narrow depression along its length on the anterior side. A few radial
striations are seen on this depression aiid on the anterior slope of the postero-
ventral ridge.
Comparison. As compared to M. vishnu Noetling 2 * from the msestridi-
tian of the Mari Hills, Baluchistan, this species has its growth striae of the
same strength throughout their course and the unibones are less anteriorly
situated.
Modiola typica Forbes 26 from the Trichinopoly stage of the Cretaceous
Series of South India and from the Alpine Gossan Deposits has, as compared
23 Stoliczka, 1871, p. 445, pi. 84, figs. 3-14 ; pi. 16, fig. 3 ; Quaas, 1902, p. 175,
pi. 20, figs. 16-22; Fourtau, 1917, p. 23.
24 Noetling, 1897, p. 44, pi. 11, fig. 3.
25 Forbes, 1846, p. 152, pi. 14, fig. 4; pi. 16, fig. 7; d'Orbigny, 1850, p. 247;
Stoliczka, 1871, p. 377, pi. 23, figs, 12-15 ; Zittel, 1866, p. 78, pi. 11, fig. 5,
262 Go W. Chiplonker
to the present species, its posterior side much more expanded and the
posterior margin more inclined to the dorsal margin.
Modiola lensi Quaas, 26 an upper Cretaceous species from the I,ybian
Desert can be distinguished from the present species by its antero-ventral
region less inflated and much smaller than the postcro-dorsal region, by the
absence of radial striations along the postero-ventral ridge and by the
greatest inflation situated sub-medianly.
As compared to M. reversa Sowerby, 27 a Cenomanian species in England
and Europe, this species has its posterior margin less sinuous.
This species shows the nearest relation to M. roquei Thomas and
Peron 28 from the Cenomanian of Tunis ; the latter has, however, its poste-
rior margin more oblique.
Occurrence. This species is recorded from the Deola-Chirakhan Marl.
Modiola minor sp. nov.
(Plate XII, Fig. 6)
Description. The shell is rectangular, rather high and tumid with
fairly prominent umbones. The anterior side is very short and rounded.
The ventral margin is almost straight. A ridge extends postero-veutrally
from the umbones and sets apart an attenuated dorsal area. Except for
the concentric growth lines the surface of the shell is smooth.
Comparison. From the associated M. inflata sp. nov., this species
differs in having a much taller shell, its umbones more prominent and no
radial striations on the postero-ventral ridge.
As compared to M. typica Forbes 29 from the Tricliinopoly stage of the
South Indian Cretaceous and the Alpine Gossau Deposits, the present
species has its umbones more anteriorly situated, its shell a little shorter
with the posterior margin less oblique and the posterior region less
expanded.
Modiola lensi Quaas 30 from the upper Cretaceous of the lyybiau Desert,
comes closest to the present species. The difference, however, lies in the
26 Quaas, 1902, p. 193, pi. 22, fig. 11.
27 d'Orbigny, 1844-48, p. 277, pi. 361, figs. 1-2 ; 1850, p. 105 ; Moms, 1854,
p. 211 ; Pictet and Campiche, 1867, p. 510 ; Geinitz, 1871-73, p. 210, pi. 48, lie 9
Woods, 1900-03, p. 94, pi. 15, figs. 15-18 ; pi. 16, figs. 1-3. '
26 Thomas and Peron, 1890, p. 247, pi. 27, fig. 18.
29 Forbes, 1846, p. 152, pi. 14, fig. 4; pi. 16, fig. 7; d'Orbigny 18^0 D 247-
Stoliczka, 1871, p. 377, pi. 23, figs. 12-15 ; Zittel, 1866, p! 78, pi [ 1^ % 5 '
30 Quaas, 1902, p. 193, pi. 22, fig. 11.
Lamellibranchs from the Bagh Beds 263
^former species having a little longer shell and the postero-dorsal region is
proportionally smaller than the antero-ventral region.
Occurrence. -Probably Upper Coralline Limestone.
Family . . Astartidse.
Genus . . Opis Defrance, 1828.
Opts corniformis sp. nov.
(Plate XII, Fig. 2)
Description. The shell is very high and trigonal with prolonged, acute,
slightly incurved beaks. The antero-ventral side is obtusely rounded and
the postero-dorsal side is almost straight. A rounded carination extends
from the umbo to the postero-ventral angle and cuts off a posterior area
which is divided into two parts by a rounded secondary ridge extending
to the postero-dorsal angle. The posterior area between the carinae is
concave. The lunule is large and deep.
Comparison. This species can be distinguished from Opissoma geinitziana
Stoliczka 31 from the Utatur stage of South India, by its beaks more prolong-
ed, ventral margin less convex and the antero-ventral margin more rounded.
To compare with the present species, Opis bicornis Geinitz 32 from the
upper Cenomanian of Bohemia, has its beaks shorter and less incurved,
its shell more oblique and its ventral margin more convex.
Opis elegans d'Orbigny 33 from the Turonian of Sarthe has, as compared
to the present species, its shell more oblique, beaks less produced and more
incurved.
The species described here shows a very close affinity to Opis haldo-
nensis Woods, 3 * an Upper Green Sand species from England. The only
observable differences are that the Indian specimens have slightly more
prolonged beaks and a little shorter shell,
Occurrence. Beola-Chirakhan Marl and Nodular Limestone.
Family . . Crassatellidae.
Genus . , Anthonya Gabb, 1864.
Anthony a tumida sp. nov.
(Plate XIII, Fig. 1)
Description. The shell is roughly trigonal, nearly twice as long as high
and strongly inflated at about one-fourth the height from the umbones.
31 Stoliczka, 1871, p. 288, pi. 10, fig, 11.
32 Geinitz, 1871-73, p. 227, pi. 50, figs. 1-3.
33 d'Orbigny, 1844-48, p. 35, pi. 254, figs. 4-9.
34 Woods, 1904-12, p. 119, pi. 18, fig. 1.
264 G. W. Chiplonker
The ventral margin is very feebly convex. A ridge extends from the
umbones to the postero-ventral extremity, setting apart a feebly concave
dorso-posterior area. The umbones are situated at about one-fourth the
length from the anterior side. Fine radial striations are observable only
on the postero-dorsal area, while the rest of the surface shows only taint
concentric growth lines.
Comparison. As compared to An. cantiana Woods, 35 a Gault species
from England, the species described here has a taller and more inflated
shell.
To compare with Anthony a sub-cantiana Nagao 36 from the Albian of
japan, the Indian species is more inflated and has a little higher shell.
Anthony* lineata Kitchin 37 from the Uitenhage Series has its shell
less tumid, more sub-equilateral and a little longer with its umbones placed
less anteriorly.
Occurrence. This species is recorded from the Deola-Chirakhan Marl.
Family . . Cardiidae.
Genus . . Protocardia Beyrich, 1845.
Protocardia pusilla sp. nov.
(Plate XII, Fig. 8)
Description. This species is a small-sized race, the largest of the indi-
viduals measuring nearly 21 mm. in length. The shell is nearly sub-circular
and slightly longer than high. The ventral and anterior margins are regu-
larly rounded and postero-ventrally there is a slight angulatiou. The shell
is thickest at a little above the middle of the shell. The ornamentation
consists of very fine, concentric rib-lets 9 to 11 per 5mm. in the middle
region. The posterior region of the shell is covered with line, smooth,
radial rib-lets ; the extent of this posterior region varies from one-fourth
to one-fifth of the shell surface.
Comparison. In comparison to Proto. hillana Sowerby/ 18 a species of
very wide destribution in Europe, Africa and South India, and ranging
from the Cenomanian to Senonian, the species described above is a much
35 Woods, 1904-12, p. 130, pi. 19, figs. 4-5.
36 Nagao, 1934, p. 222, pi. 25, fig. 11 ; pi. 30, figs. 6-7.
37 Kitchin, 1913, p. 137, pi. 7, figs. 7-8.
38 Sowerby, 1812, p. 41, pi. 14 ; d'Orbigny, 1844-48, p. 27, pi. 243 ; Forbes, 1810,
p. 146 ; d'Orbigny, 1850, p. 162 ; Pictet and Campiche, 1867, p. 268 ; Gemit//, 1871-73 ,
p. 230, pi. 1, figs. 11-12 ; Thomas and Peron, 1890, p. 276; Quaas, 1902, p. 218, pi. 1M- ,'
fig. 18 ; Fourtau, 1904, p. 331 ; Woods, 1906, p. 307 ? pL 37, fig. 6 ; 190-1-12, p. 11)7,
pi. 31, fig. 6 ; pi. 32, figs. 1-6 ; Pervinquiere, 1912, p. 264 ; Lehner, 1937, p. HO.
Lamellibranchs from the Bagh Beds 265
smaller race ; its shell is a little longer than high and its umbones are a
little more prominent,
Proto. parahillana Wade 39 from the'Ripley Formation of North America
as compared to the present species, has its outline more sub-circular and the
umbones less conspicuous ; also it has fine radial striations on the whole
of the surface.
Occurrence. Deola-Chirakhan Marl and the Nodular limestone.
Family . . Isocardiidae.
^ Genus .. Isocardia Klein, 1753.*
Isocardia aff. neglecta Coquand
(Plate XIII, Fig. 7)
Description. The shell is rather tumid, trigonal in outline and higher
than long, the length being nearly equal to the thickness. The anterior
side is short and rounded ; postero-dorsally the shell is slightly tapering.
The ventral margin is convex, being comparatively flatter in the middle
than at the two ends. The beaks are incurved and twisted anteriorly.
The maximum thickness of the shell is a little above the middle, while the
greatest length is situated medianly.
Comparison. -As compared to I so. similis Sowerby 40 from the Lower
Green Sand of England, France and Switzerland the present species is
taller, anteriorly more rounded and ventrally more convex.
In I so. ataxensis d'Orbigny, 41 a Turonian species from France, the
shell, as compared to that of the present species, is less inflated, slightly
longer and posteriorly more angulated.
1 did not have access to the original description and figures given by
Coquand, but Fourtau 42 gives Coquaud's diagnosis of Iso. neglecta Coquand
as " Coquille plus longue que large, oval, crochets legerment contourn^s,
etroits, ecartes, moule interieur montrant en arriere des crochets, une
impression transversale ". From this, it is obvious that the specimens
described here are closely related to this species from the Cenomanian of
Egypt, But since these specimens could not be compared with the original
39 Wade, 1926, p. 87, pi. 27, figs. 1, 5.
* Dall (in Zittel, 1927, p. 491) attributes the generic name Isocardia to Lamarck,
while Fourtau (1917, p. 66) uses it after Klein; since I myself had not had access to
the necessary literature for deciding the claims of priority, I use it here after Klein .
40 Sowerby, 1826, p. 27, pi. 66, fig. 1 ; d'Orbigny, 1850, p. 163; Morris, 1854,
p. 204 ; Pictet and Campiche, 1867, p. 240 ; Woods, 1904-12, p. 151, text-fig. 25.
41 d'Orbigny, 1844-48, p. 47, pi. 251, figs. 3-6,
42 Fourtau, 1917, p. 66.
266 G* W. Chi plonker
figures, their identity with Coquands species cannot be definitely estab-
lished.
Occurrence. Deol^Chmkh^n Marl and Nodular Limestone.
Family . . Veneridae.
Genus .. Callista Poli, 1791.
Callista sp. indet.
(Plate XIII, Fig. 6)
Description. This species is represented by a number of casts which
are longer than high, the maximum length lying a little below the middle.
The shell is thickest slightly above the middle. The anterior and posterior
margins are more strongly convex than the ventral margin. The umbones
are situated a little anterior to the middle.
Comparison.- Because of the general aspect of these casts they are
comparable with Callista plana Sowerby 43 from the Upper Green Sand of
England, France and Switzerland ; but the present state of preservation
of these specimens does not allow their identity being established.
Occurrence. Deola-Chirakhan Marl and Nodular Limestone.
Family . . Sportellidse.
Genus . . Anisodonta Deshayes, I860.
Anisodonta sp. indet.
(Plate XII, Fig. 4)
Description. These specimens are trapezoidal in outline. A strong
carination separates a more or less fiat dorsal area- from, the rest of the
surface ; it carries very indistinct radial striations while the rest of the
surface shows only faint growth lines. The ventral margin is feebly convex.
The anterior side is rather tapering. The umbones are turned slightly to
the anterior and are situated at a little less than one-third of the length
from the anterior end.
Comparison. The unfavourable state of preservation of the available
specimens does not permit of a close comparison with any of the known
species. But a reference might be made to Aniso. unzanbiensis Rennie 44 from
the upper Cretaceous of Pondoland, which, however, is a longer species.
Occurrence. Deola-Chirakhan Marl.
43 Sowerby, 1812, p. 58, pi. 20 ; Morris, 1854, p. 201 ; d'Orbigny, 1844-48, p. 447,
pi. 386, figs. 1-3 ; 1850, p. 159 ; Woods, 1904-12, p. 192, pi. 30, figs. 1-6 ; Pictet and
Campiche, 1867, p. 190.
44 Rennie, 1930, p. 194, pi. 21, fig. 11.
Lamellibranchs from the Bagk Beds
267
Table showing the Affinities and the Vertical Distribution of the
Lamellibranchia from the Bagh Beds
No,
Sp3cios from
Bagh Beds
Related species dth
geological horizon.
Nodular
Limestone
Lower
Coralline
Limestone
Dsola-
Ghirakhan
Marl
Upper
Coralline
Limestone
1
Pinna mathuri sp.
nov.
P. vanhc&peni Rennie ;
Upper Cretaceous of
Pondoland
X
X
2
Inoceramus pseudo-
lalus sp. nov.
Ino. latus Man toll ; Cfmo-
manian of England and
Central Europe
X
3
//io. lamarcki var.
indicus var. nov.
Ino. lamarcki var. cuvieri
Sow.; Upper Chalk of
England.
X
4
Ino. sp. A.
Ino. crispianus Stol.;
Ariyalur stage of South
India
X
X
5
Ino. sp. B.
Ino. striatus Mantell ;
Cenomanian and Seno-
nian of Bohemia
? X
x
6
Neithea morrisi Pic-
tet and Renevier
N. morrisi Pictet and
Renevier ; Aptian and
Albian of England,
Spain, Switzerland and
Japan
X
1
Plicatula spini-costata
sp. nov.
PI. aspera Sow.; Turonian
and Campani an of France
and Germany; and PL
aitressensi-s Coquand ;
Cenomani an of Tunis and
Egypt
? X
X
X
8
PI. batnensis Coquanc
P. batnensis Coqua-nd ;
Cenomanian of Tunis ami
Egypt
? X
X
X
9
Mod-tola inflata sp.
nov.
M. roquei Thomas and
Peron ; Cenomanian of
Tun i:s
X
"
10
M . minor sp. nov.
M . lensi Quaas ; Upper
Cretaceous of Lybian
Desert
? X
11
Opis corniformis sp.
nov.
0. haldonensis Woods ;
Upper Green Sand o
England
X
X
12
Anthonya tumida sp.
nov.
A. sub-cantiana Nagao
Albian of Japan
X
13
Protocardia pusilla
sp. nov.
Proto. parahillana Wade
Ripley Formation of
U.S.A.
X
X
268 G. W. Chiplonker
Table showing the Affinities and the Vertical Distribution of the
Lamellibranchia from the Bagh Beds (Contd.)
No.
Species from
Bagh Beds
Related speci es with
geolegical horison
Nodular
Limestone
Lower
Coralline
Limestone
Deola-
Chirakhan
Marl
Upper
Coralline
Limestone
14
Isocardia aff . neglecta
Coquand
Iso. negleda Coquand :
Cenomanian of Egypt
X
X
15
Callistasp. iridet.
O. piano, Sow.: Upper
Green Sand of England,
X
X
France and Switzerland
16
Anis-jdonta sp. indet.
An. unzanbiensiv Rennie :
Uppar Cretaceous of
Pondoland
*
X
Discussion of the Palceontological Resiilts and Conclusions
As can be seen from the above table of distribution of the lamel-
libranch species, we find that out of the sixteen species which are
recorded from this formation, all except one, namely Modiola minor sp. nov.,
are present in the Deola-Chirakhan Marl. The species which occur in the
Nodular Limestone include Pinna mathuri sp. nov., Inoceramus sp. A.,
Ino. sp. B., Opis comiformis sp. nov., Protocardia pusilla sp. nov., Isocardia
afL neglecta Coquand and Callista sp. indet. ; but all of them also occur in
the Marl bed. Similarly Plicatula spini-costata sp. nov., and PL batnensis
Coquand are recorded both from the Marl and the Coralline Iyirneston.es.
Thus, while most of the species known to occur in this formation, are
recorded in the Marl bed, there are some species which it shares with the
Nodular limestone, and others which it has in common with the Coralline
limestones. These lamellibranch species, therefore, while being parti-
cularly abundant in the Marl, are not characteristically confined to any of
the beds, so as to mark them out as distinct palseontological zones or stages
as was believed to be the case by Bose.
To discuss the faunal affinities and the age of the I/amellibranchia from
the Bagh Beds, described in the foregoing pages, we have in these strata two
species which are known also from outside India. Of these, Neithea morrisi
Pictet and Renevier, the commonest of the lamellibranch species in these
beds, is known from the Aptian and Albian horizons in England, Spain,
Switzerland and in Japan, while Plicatula batnensis Coquand, which also
is an abundant species in this formation, is recorded from the Cenomanian
of Tunis and Egypt. Another species, which is well represented in this
formation, of which, however, the identity could not be established for
Lamellibranchs from the Bagh Beds 269
certain for want of the necessary literature, is Isocardia aff. neglecta Coquand
a very near ally of /so. neglecta Coquand from the Cenomanian of North
Africa. Among the species which are described new, we have Plicatula
spini-costata, sp. nov., a species equally abundant as the last one; it is
related on the one hand to PL aspera Sowerby from the Turonian and
Companian of the Alpine region, and to PL auressensis Coquand from the
Cenomanian of Tunis and Egypt on the other. To take the less common
species, we have Modiola infiata sp. nov., with its nearest ally M. roquei
Thomas and Per on from the Cenomanian of Tunis ; Opis corniformis sp. nov.,
is allied to 0. haldonensis Woods, from the Upper Green Sand of England ;
and Inoccramiis pseudolatus sp. nov., is related to Ino. latus Mantell from
the Cenomanian of England and Central Europe.
From among the remaining species to consider those which are related
to upper Cretaceous species, we have Protocardia pusilla sp. nov., related
to Proto. parahillana Wade, from the Ripley Formation of the United
vStates of America. Modiola minor sp. nov., is allied to M. lensi Quaas from
the upper Cretaceous of the Lybian Desert ; Pinna mathuri sp. nov., has its
nearest ally P. vanhcepeni Rennie from the upper Cretaceous of Pondoland
and Inoceramus lamarcki var. indicus var. nov., is closely related to Ino.
lamarcki var. cuvieri (Sowerby) from the Upper Chalk of England.
It is thus clear that while these lamellibranch species from the Bagh
Beds show a mixture of affinities towards species ranging from the Aptian
to Senonian, a large proportion of them have Cenomanian affinities.
Mukerjee, 45 however, has lately tried to show that the Bagh Beds
range from the Cenomanian to the Upper Senonian because of the range of
affinities shown by his collection of Mollusca from the Jhabua and Ali
Raj pur States. In his report on these beds he records the presence of
Astarte, Mactra (?), Aucella (?), Nucula, Ostrea, Melania, some Auricullid
Pulmonata, Cidaris and Bryozoa, and claims to have specifically identified
Pr otocar dium pondicherriense d'Orbigny, Cardium (Trachycardium) incomptum
Sowerby, Macrocallista cf. sculpturata Stoliczka, Grotriana cf. jugosa Forbes,
Crassinella cf . planissima (Forbes), Cr. cf. trigonoides (Stoliczka) andTumtella
(Zaria] multistriata Reuss. Out of these, he mentions Proto. pondicherriense,
Card. (Track.} incomptum, Macrocall. cf. sculpturata and Turr. multistriata
as typical South Indian upper Cretaceous forms suggesting that the Bagh
Beds probably range up to the upper Senonian, while the presence of
characteristic Utatur forms Grotriana cf. jugosa and Crassinella cf.
planissima show that the Bagh Beds range down to the Cenomanian.
45 Mukerjee, 1934, pp. 71-73 ; 1935, pp. 80-81 ; 1938, pp. 196-98.
270 G. W. Chiplonker
From tliis list given by Mukerjee it is clear that the only species of
which he can be said to have definitely established the identity are only
three, namely: Protocardium pondichernense d'Orb., Cardium (Tmchy-
cardium] incomptum Sowerby and Turntella (Zaria) nwltistriata Rettss.
Of these, the first two are from the Trichinopoly stage, where only the
second species is reported to be common ; while Turritdla tnuUistriata is
recorded in South India from the Ariyalur and Trichinopoly stages
where it is not reported to have any particularly important position ; it is
also known from the Senonian and Turonian of the Lybian Desert, Germany
and the Alpine region. 46 The remaining four species which Mukerjee records
in the Bagh Beds of his area, are only comparable with some of the vSouth
Indian species which, moreover, are not of any special importance in those
deposits. It is unfortunate that Mukerjee has used the terms " character-
istic or typical species" in too loose a sense as can be seen from the above
remarks. None of the species which Mukerjee records in the Bagh Beds
of Jhabua and AH Rajpur States, are in any sense " characteristic " or
" typical " of any of the stages in the South Indian Cretaceous Series ; nor
are any of them particularly important in the Bagh Beds. They, therefore,
cannot be of any value in fixing the age of the Bagh Beds with any more
precision than, that they indicate an upper Cretaceous age for this forma-
tion.
It is thus clear from the foregoing discussion, that this series of strata
forms a single palseontological unit, in which Cenomanian affinities are the
predominant element of the fauna ; it is therefore, reasonable to assign to
this formation the Cenomanian age.
Mukerjee's collection is, however, of some interest in so far as it includes
the three South Indian species Protocardium pondicherriense d'Orb., Cardium
(Trachy.} incomptum Sowerby and Tunitella (Zaria) multistriata Reuss.
It brings in under discussion the possibility of faunal connections between
the Narbada Valley and the Trichinopoly District. Except Pinna mathuri
sp. nov., which is allied to P. arata Forbes from South India and also to
P. vanhcepeni Rennie from Pondoland, there are no Bagh species
related to those from South India. As is clear from the author's work on
other groups of Bagh fossils 47 also, the Bagh fauna have strong affinities
towards those from North Africa and Southern Europe. Since, of the three
South Indian species mentioned above, Turritella multistriata Reuss is also
46 Forbes, 1846, p. 124; Stoliczka, 1868, p. 224 ; Gteinitz, 1871-73, p. 101 ;
1902, p. 246.
47 Chiplonker, 1937, pp. 66-68 ; 1938, pp. 312-13 ; 1939, pp. 21-2-15 ; 1939 a,
pp. 105-07.
Lamellibranchs from the Bagh Beds 271
known to occur in the Lybian Desert, Germany and in the Alps, the affi-
nities of Pinna mat/hurt towards P. arata and P. vanhcepeni, as well as the
occurrence in the Narbada Valley, of Protocardium pondichemense and
C&rdium (Trachy.) incomptum in association with Tumtella (Zaria)
multistriata might be expected through the African and the Mediterranean
regions and not by any direct connection as was suggested by Bose.
While it is admitted that the explanation given above might account for
the occurrence of these South Indian species in the Narbada Valley, it is
also likely that a closer examination than hitherto done, of the molluscan
fauna of the Jhabua-Ali Rajpur area might show these species to be spe-
cifically different from the two above-mentioned South Indian species.
A cknowledgments
In conclusion, I have to express my great indebtedness to Dr. Raj Nath,
Head of the Department of Geology, for kindly going through the manu-
script of this paper and for his valuable criticism. My grateful thanks are
due to the Director, The Geological Survey of India, for permission to work
in the Survey Library and the Museum. I take this opportunity of thanking
also the Dhar, Indore and the Gwalior Durbars for the kind permission to
collect fossils from their territories and for the facilities accorded to me
during the field-work.
BIBLIOGRAPHY
1. Bose, P. N. . . " Geology of the Lower Narbada Valley between
Nimawar and Kawant/' Mem. G.S.I., 1883, 21,
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2. ChofEat, P. . . Recueil d'Etudes paleontologigues sur la faunc cretacique
du Portugal, 1901-02, 1, ser. 3 and 4.
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Sci., (B), 1937, 6, 60-71.
4. .. ' ' Rhynchonellids from the Bagh Beds," Ibid., 1938,
(B), 7, 300-16.
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(B), 9, 236-46.
6. . .. ' * Bryozoa from the Bagh Beds," Ibid., 1939 a, 10,
98-109.
7. Coquand, H. . . Monographic de V$tage Aptien de Espatfne, 1865.
8. Duncan, P. M. " Description of the Echinodermata from the Strata.
on the South-Eastern Coast of Arabia and at Bagh
on the Narbada in the Collection of the Geological
Society," Q.J.G.S., 1865, 21.
9, .. " Notes on the Echnoidea of the Cretaceous series of
the Lower Narbada Valley with remarks upon their
geological age," Rec. G.S.I., 1887, 20, 81-92.
272
10. Douville, H.
11. Forbes, E.
19
-i t~> "
13. Fourtati, R.
14.
G. W. Chiplonker
23.
24.
15. Geinitz, H. B.
i a ,
JLU. '
17. Kitchin, F. L.
18. Lartet, L.
19. Lehner Leonhard
20. Mantell
21. Morris, J.
22. Mukerjee, P. N.
25. Nagao, T.
26. Noetling, F.
27. d'Orbigny ; A.
28.
29. Pervinquiere, L.
30. Pictet, F. J. s and
Campiche, G.
*' Les terrains secondaires dans le Massif du Moghar
a Feast de Fisthixie de Suez," Mem. Acad. Sci.,
Paris, 1916, 54, ser. 2.
41 Catalogue of the Lower Green Sand Fossils in the
Mu seum of the Geological Society," Q.J.G.S., 1845, 1 .
'* Report on the fossil Invertebrata from South India
collected by M. Kaye and M. Cimliff-V Trans. Geo.-
Soc. Lo ndcw, 1840, (2), 7, pt. 3, 139-57.
" Contribution a 1' etude de la faune cretacique
d'Egypte," Bull. Inst. Egypt., (4), 1904, 4, 231-349.
Catalogues des Invertebres fossiles de TEgyple repre
sentes dans les collections dw MuseB de Geologic au
Caire. Geol. Surv. Egpt., pal. ser., terr. crefc., 1917,
3 S pt. 2.
"Des Elbthalgebirge Sachsen," Palceontographica,
1871-73, 20, pt. 1.
" Des Elbthalgebirge Sachsen," Ibid., 1872-73, 20,
pt. 2.
' ' The Invertebrate fauna and PaLnpontological relations
of the Uitenhage vSeries," Ann. 8. Afr. Mus., 1913,
7, 21-25.
4 * Exploration geologique de la Mer Morte, de la
Palestine et de ridume6," Paleontolor/ie, Paris,
1880 (?), 2.
* ' Fauna und Flora der frankischen Albiiberdukenden
Kreide I. Die Lanaellibranchiaten (ohne Inocera-
men)," Palccontogr aphica, 1937, 85.
Geology of Sussex, 1822.
C dialogue of British Fossils, 1854.
in Director, " General Report of the Geological Survey
of India for the year 1932," Rec. G.S.I., 1934, 67,
71-73.
in Director, " General Report of the Geological Survey
of India for the year 1934," Ibid., 1935, 69, 69-81.
in Gupta, B. C., and Mukerjee, P. N., " Geology of
Gujarat and Southern Rajputana," Ibid., 1938, 73,
pt. 2, 163-208.
"Cretaceous mollusca from the Miyako District, Honshu >
Japan," Journ. FacuL Sci. Hokkaido Imp. Univer.
1934, (4), 2, No. 3, 177-271.
4C Baluchistan Fossils," Pal. Ind., (14), 1897, 1, pt. 3,
Paleontologie Francaise. ter. cret., Lamellibrancb.es,
1844-48, 3.
Prodrome d'e Paleontologie Stratigraphie, etc., 1850, 2.
" Etudes Paleontologie Tunisienne. II. Gastropodes et
Lamellibranch.es des terr. cret.," 1912.
" Description des fossiles du terrain cretace des Environs
de Ste. Croix." Mater. Pal. Suisse, 1864-67, ser. 4.
Lamellibranchs from the Bagh Beds
273
31. Pictet, F.J., and
Campiche, G.
32. and Renevier, E.
33. Quaas, A.
34. Remiie, J V. L.
35. Sowerby, J.
36. Sowerby, J. de C.
37.
38. Stoliczka, F.
39.
40. Thomas, P., and
Peron, A.
41. Wade, B.
42. Wanner, J.
43. Woods, H.
44.
45.
46. Yabe, G. and Nagao, T.
47. Zittel, K. von.
48.
" Description des fossiles du terrain cretace des En-
virons de Ste. Croix.," Ibid., 1870, ser. 5.
" Description des fossiles du terrain Aptien de la Perte
da Rhone et des Environs deSte. Croix.," Ibid,., 1858,
ser. 1, 54-142.
' ' Beitrag zur Kenntniss der obersten Kreidebildungen
in der Hbyschen Wiiste, Overwigischichten und
Blatterthone.," Palczontographica, 1902, 30, pt. 2,
153-336.
" New Lamellibranchia and Gastropoda from the Upper
Cretaceous of Pondoland, (with an appendix on some
species from the Cretaceous of Zululand," Ann.
S. Afr. Mus., 1930, 28, 159-260.
Mineral Conchology of Great Britain, 1812, 1.
Ibid., 1826, 4.
Ibid., 1829, 6.
" The Cretaceous Fauna of Southern India.
Gastropoda." Pal. Ind., (5), 1868. 2.
" The Cretaceous Fauna of Southern India.
Pelecypoda," Ibid., 1871, (6), 3.
Description des Mollusyues fissiles des terrains cretaces
dela region sud des Hants-Plateaux de la Tunisie
recueilles in 1885 et 1886 par M.P. Thomis. 1890,
Paris .
' ' The Fauna of the Bipley Formation on the Coon
Creek, Tennesse." Prof. Papers U. S. G. S., 1926,
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* Die Fauna der obersten weissen Kreide in der lyb-
ischen Wiiste." Palccontographica, 1902, 30<, pt. 2,
91-152.
4 The Cretaceous Lamellibranchiata of England," Mon.
Palasontogr. Soc., 1900-03, 1.
** The Cretaceous Fauna of Pondoland," Ann. S. Afric.
Muss., 1906, 4, pt. 7.
)J The Cretaceous Lamellibranchiata of England," Mon.
Palmontogra. Soc., 1904-12, 2.
" New or little known Cretaceous fossils from North
Saghalin," Sci. Rept., Tohoku, Imp. Univ., (2),
1925,7, No. 4.
" Die Bivalven der Gossaugebilde in der Nordostlischen
Alpen. Ein Beitrag zur Character! stik der Kreide-
formation in Osterreich.," Denkschr. Reiser. Akad.
Wissenschaften., 1866, 25, pt., 2, 77-198.
Text-Book of Palaeontology, Edit. Eastman, C. R.,
2nd. Ed. ,1927,1.
EXPLANATION OF PLATES
PLATE XII
FIG. 1. Inoceramns latnarcM var. indicus var. nov. ; showing right valve (B. II. (J.
No. L/3).
FIG. 2. Opis cvrnifowiris sp. nov., showing right valve (B. IT. IT. No. L/ll).
FIG. 3. Inocermnus pseudo-latus, sp. nov., showing right valve (B. II. IT. No. L/2).
FIG. 4. Anisodonta sp. inclet., showing left valve (B.I.1.IT. No. L/k).
FIG. 5. Inoceranivs sp. A., showing left valve (B. IT. IT. No. L/-I).
FIG. G. Modiola minor sp. nov., showing loft valve (B. IT. IT. No. L/tO).
FIG. 7. Ncithca morrisi Pictet and Renevier : (a) showing left valve; (b) showing
right valve ; (B. II. U. No. L/0).
FIG. 8. Protocardia pus-ilia sp. nov., showing right valvo (B. II. IT. No. L/13).
PLA.TK X.III
FIG. 1. Anthony a tumida sp. nov., showing right valve (B.II.IT. No. LI 12).
FIG. 2. Modiola inflata sp. nov., showing right valve (B.II. IT. No. L/*))-
FIG. 3. PUcatula spini-costata. sp. nov. ; showing upper valve (B. II. IT. No. L/7).
Fi'G. 4. Pinna mathuri sp. nov., showing left valve; (B.II.IT. No. L/i).
FIG. 5. PUcatula batnensis Ooquand, showing upper valve (B, II. IT. No. Ij/8).
FiG. 6. CalHsta sp. indct., showing right valve (B.II.IT* L/1.5).
FIG. 7. Isocardia aft', nec/lccta Ooquand ; (a) sliowing left valve ; (/;) showing anterior
view (B.H.IT. ' No. L/ll).
N.B. All the figures are natural si7,e.
All the figures are photographs of the plates drawn from the Type specimens
under the author's supervision.
G. W. Chiplonker
Proc. Ind. Acad. Set., B, vol. X, PL XII
B
ON SOME STONE IMPLEMENTS FROM
HOSHANGABAD (CENTRAL PROVINCES)
BY MANOHAR I^AI, MISRA, M.Sc., 1,1,. B.
(From the Department of Geology, Benares Hindu University}
Received Saptember 12, 1939
(Communicated by Prof. L. Rama Rau, M.A., F.G.S.)
THE stone implements, which form the subject of the present contribution,
were collected by the author from the banks of the river JSTarbada and
neighbouring ravines, near Hoshangabad in the Central Provinces. So far the
implements collected from the Narbada Valley by various workers are most-
ly confined to collections made from certain areas round about Jubbulpore
and the literature available is mostly a record of the find. The very first
account of some discoveries of worked flints from near Jubbulpore was
given by Evans 1 in 1.853, followed by lyeMesuriar, 2 Sweney, 3 Carey, 4
Blanford 5 and Oakes. 6 In the year 1873, Medlicott 7 described a quartzite
Celt found in situ by Hacket from Bhutra in the Narsingpur District in the
Narbada Valley. This find is of exceptional importance as " Hacket him-
self dug it from where he found it lying flat and two-thirds buried, in a
steep face of the stiff, reddish, mottled, unstratified clay about six feet
above low water-level" 8 and which is about the same horizon at which
some of the implements, described here, have been found. Mention of
some palaeolithic and neolithic implements from near rock shelters about
three miles from Hoshangabad town has also been made by Ghosh. 9 The
present contribution contains the description of four palaeolithic and two
neolithic stone implements, which are unquestionably of human manu-
facture and some of them have been found in sitii in the same or about the
same horizon in which the vertebrate fossils have been known to occur.
1 Evans, 1853.
2 LeMesuriar, 1801, pp. 81-85.
3 Swiney, 18C9, pp. 17-18 ; 1865, pp. 77-79.
* Carey, 1866, pp. 135-36.
5 Blanford, 1866, p. 230.
6 Oakes, 1869, pp. 51-53.
7 Medlicott, 1873, pp. 49-53.
8 Ibid., p. 49.
9 Ghosh, 1932, pp. 21-22,
275
Some fossilised remains of vertebrate animals have also been collected by
the author along with the implements and these have been handed over
to Prof. D. K. Chakravarti of this University for study.
MAP or THE APEA
ABOUT
P/?OV//VC<fS
The area covered in collecting these implements is about 1.0 miles in
length along the banks of the river Narbada running from about a mile to
the east of Bandrabhan a locality at the confluence of the Narbada with
the Tawa, to about a mile to the west of the railway bridge over the
Narbada near Hoshangabad town. Two other localities, a small sandstone
hill containing some old red coloured paintings in the rock shelters, about
1| miles to the south-east of the Hoshangabad town and a laterite quarry,
at the village Tugaria about four miles to the south of the town, were also
investigated but no implement was found.
On Some Stone Implements from Hoshangabad (C.P.) 277
The general stratigraphical succession of beds at the banks of the river
in the area investigated is as follows in the descending order :
Beds Approximate
Thickness
Regur . . . . . . 8 feet
Yellow alluvium . . . . . . 12 ,,
Yellow alluvium with concretions . . 8 ,,
Gravels at places containing boulders . . 5 ,,
The thickness of the various beds is not constant. The gravelly bed is
conspicuous by constantly running along the banks forming a terrace.
Due to the meanders of the river, this gravelly bed is, often, overlain and
thus concealed by the recent deposits of sand. The yellow alluvium forms
the thickest bed of the series. Just below Saddar Bazar in the Hoshanga-
bad town all the beds are well exposed and attain a total thickness of
about 35 to 40 feet.
As far the age of these deposits, the topmost bed is, no doubt, of
recent time. The bottom bed (gravels) was considered, to be of Pliocene
age by Falconer, 10 while in the opinion of Medlicott 11 "these old ossiferous
alluvial deposits are not more ancient than the late Pleistocene ". Both
these views have been questioned by Das Gupta, 12 who is of opinion that
the Narbada ossiferous gravels are not older than middle Pleistocene.
The author takes this opportunity of conveying his grateful thanks
to Mr. K. Sripada Rao, Central College, Bangalore, for lending his personal
copies of the two papers of Prof. Sampat lyengar. The author's sincere
thanks are also due to Dr. Rajnath, Head of the Department of Geology,
Benares Hindu University and to Prof. D. K. Chakravarti, of the same
Department for facilities and help.
Specimen No. AT/19 (B. H. U.)
Plate XIV, Fig. 1
The specimen is fashioned out of brownish looking fine-grained
Vindhyan quartzite with a predominant yellowish tinge. It is almond
shaped, mostly symmetrical, and has not been retouched. The ' surface
d' accommodation' is like an inverted U, with diverging arms. The two
points, at which these diverging arms end, mark the greatest width of the
specimen. From beyond these two points, the specimen begins to narrow
10 Medlicott, 1873, Pb. 3, p. 49.
11 IHc?., p. 54.
!2 Das Gupta, 1923, p. 22.
B3a IT
down towards the ' surface d'utilisation ', where it finally ends in a small
semicircular edge about |-" in width. A somewhat blunt edge all round the
specimen forms the periphery which is undulating and contains a number
of marks of chipping. One of the two main faces of the implement is made
convex by three main bold strokes, the axis of convexity running along its
greatest width. Two strokes, one on each side of the axis of convexity,
have produced two surfaces. One of these two surfaces slopes towards
the ' surface d'utilisation ' and the other towards the ' surface d'accommocla-
tion '. The line along which these two surfaces meet forms a ridge which
runs more or less along the greatest width of the specimen and coincides
with the axis of convexity on this face. The third stroke not as bold as
the other two is near about the middle of the right edge of the specimen
and it has produced a third surface, small and triangular in shape, sloping
towards the edge. This side shows some signs of retouching. The surface
on the reverse side is also convex, but the axis of convexity here is along
the length of the specimen and runs from the ' surface d'accommodation '
to the ' surface d'utilisation '. From along this axis the specimen slopes
mainly in two opposite directions towards the right and towards the left.
Only one, roughly pentagonal surface, produced perhaps by only one bold
stroke, forms the whole of the right-hand face. It is bounded by two edges
and three ridges which are formed by conjunction of differently slotting sur-
faces. Two four-sided surfaces, comprise the left-hand face. These faces
slope towards the edge from the main ridge. There is a small triangular
area at the ' surface d'accommodation' bound by two ridges and one edge.
The main ridge of the specimen coincides with the axis of the convexity
and slightly bulges towards the left.
The specimen weighs 7 oz. Its greatest length from ' surface d'ac-
commodation ' to the ' surface d'utilisation ' is 3 - 9", the greatest width
3- 2" and the greatest thickness 1-05".
The specimen bears a very close resemblance, in practically all the
details, to a boucher from the Cuddapah District of the Madras Presidency
figured by Coggin Brown. 13 It also resembles K. Sripada Rao's specimen
No. Z 6/441. u The only main point of difference between these two is
that the present specimen is marked by the absence of notches, and this
negatives the idea of the present specimen having ever been used with a
handle. On the other hand, the smooth and convenient grasp at the
' surface d'accommodation ' and its close resemblance to Coggin Brown's
13 Coggin Brown, 1917, pi. I, fig. 1.
14 Sripada Eao, 1930, pi. VI, figs. 1, 2, 3.
On Some Stone Implements from Hoshangabad (C.P.) 279
boucher point strongly towards this specimen being designed for a hand-
grasp, and used as a hand axe.
Tlie specimen was found in situ and was extracted from the steep face
on the left bank of the river Narbada near Hoshangabad. It was found
embedded just below a gravelly bed containing vertebrate fossils, about
6 to 8 feet above the lowest water-level.
The bold and rough design of the implement and a smooth and com-
fortable hand-grasp indicate that the implement probably belongs to
Cheleaii Culture, but appears to be more primitive either to Racket's find 15
or to that of Wynne. 16
Specimen No. NJ20 (B. H. U.)
Plate XIV, Fig. 2
The specimen is fashioned out of hard, compact, brownish looking
ferruginous Vindhyan sandstone, which is found in abundance round about
the locality where it was found. It is evidently an almond-shaped side-
scraper with semicircular ' surface d'utilisation '. The ' surface d'accom-
; modation ' is thick and convenient for a hand-grasp. The bulb of percus-
] sion is well developed by a bold stroke at the ' surface d'accommodation '
; on one of the two main faces. Two deep scars or ' eraillures ' can be very
clearly seen on the bulb of percussion. One of these is along the edge of
i the ' surface d'accommodation ' and the other one, which is slightly curved,
} is at right angles to the first. Excepting these two scars and the well-
developed bulb of percussion, the specimen does not show any other major
signs of chipping. Few small chips have been taken out along some por-
i tions of the ' surface d'utilisation '. The other corresponding opposite face
pi of the specimen presents enough marks to show that the implement has
; been fashioned mostly on this side. This face shows three main surfaces
sloping in different directions towards the edges from a common point.
These faces have been produced in the attempt to get a sharp edge and
convenient hand-grasp. The semicircular working edge shows all along a
number of marks of chipping. On both the sides the specimen has a general
slope from the ' surface d'accommodation ' to the ' surface d'utilisation '.
The implement measures 3-75* along its greatest length and 3-0" along
its greatest width. Its greatest thickness at the ' surface d'accommodation '
is - 75". The specimen weighs 4 7 oz. It bears a very close resemblance
to one side-scraper from Pilt-down gravels so prominently figured by
15 Medlicott, 1873, p. 49.
16 Oldham, 1868, p. 65,
Boule, 17 Osborn 18 and Sollas. 19 The specimen was found embedded, rather
loosely, in a sort of boulder or gravelly bed which projects into the bed of
the river near Hoshangabad town. The spot from where it was extracted
lies about 6 to 8 feet above the low water-level of the river.
As for the age of the implement nothing very definite can be said
as the specimen has not been found truly in situ. The fact that it is worked
on one side only and that it is a side-scraper with a bulb of percussion which
is a characteristic of the Mousterian industry, leads one to believe that the
implement belongs to the Mousterian age. But the consideration that the
implement was found embedded, though loosely, near about the same
horizon as that of N/19, described in this paper, and which is of Chellean
Culture, negatives the idea of its being of Mousterian age. The view that
the implement belongs to Mousterian time further loses support when it
is considered that " although flints worked on one side only were formerly
regarded as characteristic of Mousterian civilisation, they are very often,
sometimes to the exclusion of the other forms, found also in the Chellean/' 20
and also that " the form of the chipped implements is not always a sure
criterion of the particular cultural stage which is represented by it." 21
Therefore, the present author feels inclined to believe that the age of this
implement is Chellean rather than Mousterian.
Specimen No. N/21 (B. H. U.)
Plate XV, Fig. 3
The specimen is made out of hard, fine-grained, compact, brownish
looking dark Vindhyan sandstone. It is roughly almond-shaped, crudely
fashioned and unretouched. One side the bulb of percussion is sufficiently
well developed with shallow, thin, " eraillure ". The face containing the
bulb of percussion, further shows a few other marks of chipping at the
' surface d' accommodation ' as well as at the ' surface d'utilisation '. These
chips were evidently taken out in order to provide a comfortable hand-
grasp and a sharp working edge. The opposite corresponding face shows
one prominently triangular surface, and two other surfaces the boundaries
of which are not clearly defined. The periphery is undulating and all
along the edge marks of chippiiigs are evident, but towards the working
face they become more prominent.
17 Boule, 1923, fig. 94.
18 Osborn, 1927, p. 127, fig. 60, 1 and 1 a.
19 Sollas, 1924, p. 191, fig. 81.
20 Boule, 1923, p. 161.
21 Das Gupta, 1923, p. 4.
On Some Stone Implements from Hoskangabad (C.P.) 281
Tlie implement measures along its greatest length, from ' surface d'
utilisation ' to ' surface d* accommodation ' 4-25" and 3-2* along its greatest
width. It assumes greatest thickness of 1" at the bulb of percussion. The
specimen weighs 7 oz.
In appearance and by the presence of the bulb of percussion the imple-
ment appears to be a side-scraper. But the edge just opposite to the bulb
of percussion is very blunt and thick. This side is 5" thick and conse-
quently does not provide in anyway, a working edge for the implement,
so that it may be used as a side-scraper. The real ' surface d'utilisation '
is provided by a a V-shaped edge. One arm of this V ends in the bulb
of percussion and the other in the blunt edge, opposite to the bulb of per-
cussion. The number of marks of chippings at this V edge proves that this
obviously has been done only to provide a sharp working edge more con-
venient for chopping. This being the ' surface d'utilisation ', the surface
opposite to this automatically becomes the ' surface d'accominodation '
and this provides a very convenient and comfortable hand-grasp along
with two dents for the fingers at the side. Hence this implement is more
likely to be a coupe-de-poing than a side-scraper. It may have been ori-
ginally designed to serve as a side-scraper but afterwards probably, either
intentionally or accidentally was transformed into a hand axe.
The specimen was found lying loose at a distance of about 50 yards
from where the specimen No. N/20 was extracted. It seems to belong to
earlier Palaeolithic Culture more probably Chellean.
Specimen No. N/22 (B. H. U.)
Plate XV, Fig. 4
The implement is fashioned out of hard, brownish, ferruginous, com-
pact Vindhyan sandstone. It is massive and heavier than the other
implements described here. The implement is roughly circular in shape and
double convex in appearance. It is thick at the ' surface d'accommodation '
and slopes gradually towards other directions till it thins out at the peri-
phery and forms a semicircular edge. The edge is quite sharp and the
periphery is undulating specially along the working edge. The working
face shows more marks of chipping on one side of the implement than 011 the
other. Out of the two main faces of the implement, one shows gradual
bulging from the ' surface d'utilisation ' and becomes very prominent to-
wards the ' surface d'accommodation ' where the specimen .becomes very
thick. This face is marked by having a few radial striae which make this
still more undulating towards the ' surface d'utilisation '. Towards the
'surface d'accommodation' this face has one more surface which is in-
clined to it. This surface is triangular in shape with rounded corners and
sides bulging out. The whole side shows marks of prominent chippings
along the edge. The corresponding face on the reverse side shows a tend-
ency towards flatness, but no doubt, it too shows a very gradual rise from
the edges towards the ' surface d'accommodatiou ', where the specimen
possesses a prominent but depressed bulb of percussion with two very clear
eraillures on each side of the origin of the bulb. The bulb of percussion
has lost its prominence due to the thickness and the massiveness of the
specimen. The face shows some concentric striae which of course start from
the bulb of percussion and are seen even upto the ' surface d'utilisation '.
They are shallow and flat and not at all prominent. I^ike the reverse face,
this face too, possesses a triangular surface, with rounded corners and the
sides bulging outside, at the ' surface d' accommodation '. This face also
contains marks of chippings which are very prominent at the ' surface
d'utilisation '.
The specimen weighs 17 oz. It measures 5-2" along its greatest
length, and 4-45" along its greatest width. It is 1-5" thick at the butt
end. The implement was found near the confluence of the river Narbada
with the Tawa about six miles at the east of the Hoshangabad town. It
was picked up from the bed of a ravine in the land between the two rivers,
which shows no outcrop of any sandstone.
This implement, without doubt, was intended to be a scraper with a
semicircular face. But looking to its size and weight it becomes doubtful
if it was used as such. The hand which wielded it must have been power-
ful and big to have a convenient grasp to use it as a scraper. With an
average modern hand it can very well be used as a hand axe, for the purpose
of chopping or cutting. The specimen, with its size and weight must have
afforded an unique implement for cutting the branches of the trees. As
for the age, the implement seems to belong to Mousterian Culture. It
bears a very close resemblance to a Mousterian side-scraper figured by
Schmucker. 22 It also resembles in shape to a flake No. 2436 figured by
R. B. Foote. 23
Specimen No. JV/23 (B. H. U.)
Plate XVI, Fig 5.
Fashioned out of fine-grained, brownish looking ferruginous Vindhyan
sandstone, elongated with pointed butt and curved edges, this celt or
hatchet falls in groups 4 and 1.0 given by Coggin Brown. 24 The specimen
251 Schmucker, 1925, p. 79, fig. 25.
23 Foote, 1916, pi. XII.
24 Coggin Brown, 1917, p. 6.
On Some Stone Implements from floshangabad (C.P.) 283
is tapering towards the butt end along its length. The butt end is bluntly
pointed. The specimen shows three faces, two main ones, opposite to each
other and the third at one of the sides. The side face is more or less an
elongated hexagonal area, the elongation being along the length of the speci-
men. There is a small bulb of percussion on this surface with a prominent,
elongated " eraillure " and few marks of mild strokes for retouching the
bulb of percussion. Out of the two main faces, one shows a convexity, the
axis of the 'curve being along the width of the specimen and near about its
central part. This face is devoid of any marks of fashioning, except a
slight dent towards the left. The whole face looks like a trapezium with
rounded corners and bevelled edges, the butt end being its shortest side.
The corresponding opposite face, includes one small triangular area at the
butt end. This area is inclined to the main face. At the union of these
two faces there is a ridge. This triangular area seems to be the result of
chipping off a small flake with one bold stroke in order to provide a con-
venient accommodation surface. The main face, though flat, slopes gently
towards the left till, at the union with the corresponding opposite faces, it
forms an edge. Towards the ' surface d'utilisation J the specimen shows a
number of marks of chipping. These marks are of mild strokes used evi-
dently to obtain a sharp edge. The utilisation edge is adze-shaped, curved
and sharp. The specimen seen as a whole is smooth and it appears that
this has been achieved by grinding. Though the ' surface d'accommoda-
tion ' gives a very convenient hand-grasp, it is very likely that the imple-
ment may have been mainly used with a haft.
The specimen measures 5-1" from the 'surface d'utilisation' to the
butt end, 3 4" along its utilisation edge and 1 80" near the butt end. The
thickness of the specimen at one of the lateral sides is 1. -1" while the other
side is a mere edge. It weighs 12-7 oz.
The specimen is a neolithic celt. It closely resembles the celt from
Burma described and figured by Theobald 25 and also to one figured by
Schmucker. 20
The specimen was found loosely embedded, about half out, in yellow
alluvium about 2 ft. below its junction with black cotton soil. The spot
from where it was extracted is near the mouth of a nala near "the Railway
bridge on the right bank of the river. It probably belongs to Campignian
Culture.
25 Theobald, 1874, pt. 2, pi. Ill, figs. 1 a and 1 6.
26 Schmucker, 1923, p. 90, fig. 32.
284 Manohar Lai Misra
Specimen No. N/M (B. H. U.)
Plate XVI, Fig. 6
This implement is obviously a crudely made hatchet and is fashioned
out of olivine Basalt. It is thickest at the butt end and thinnest at the
' surface d'utilisatioii ' with the result that it has got a tapering appear-
ance. The two side-faces are triangular in shape, while the other three
remaining faces are oblong. Except for one bold stroke at the butt end
and one near an edge which shows a slight depression, the specimen shows
no signs of chipping. All the different planes of the faces are tilted. This
causes the specimen to appear like a solid trapezium. The edge, utilised
for the cutting purposes, is slightly curved and the two end corners have
been rounded. One side of this edge shows some indications of its having
been ground just like a chisel.
The specimen shows uniformity in length and breadth but not in
thickness. The length from the ' surface (Tutilisation ' to the butt end is
4-65" and the width is 2-2". The greatest thickness is 1-5* at the butt
end and reduces itself merely to an edge at the ' surface d'utilisation '.
The specimen weighs 14 oz.
It was found near the Railway bridge about three miles from the
Hoshangabad town lying loose in the bed of a small nala on the left bank
of the river Narbada. The specimen is fashioned out of olivine Basalt
which does not occur near about the locality. The interesting feature of
the specimen is in the presence of three marks of slightly brownish colour
which appear to be originally due to grease or fat, and which are yet pre-
served in the implement. These marks point definitely that this imple-
ment has been fitted into a cloven handle and lashed with gut or stripes of
wet hide or strong vegetable, the fat or grease of which has left these marks.
If a haft is attached to this implement it will appear like a stone axe figured
by W. J. Sollas. 27 Though this implement has not been found in situ, yet
from the mode of its fashioning and the way in which it was used with a
haft indicate clearly to this celt belonging to later Neolithic age perhaps
Campignian or even Housian.
REFERENCES
1. Blanford, H. F. . . Proc. As. Soc.. Beng., 1866, pp. 230-31.
2. Boule, M. .. Fossil Man, 1923.
3. Carey, V. J. . . Proc. As. Soc. Beng., I860, pp. 135-38.
4. Coggin Brown, J. . . Catalogue of Prehistoric Antiquities in the Indian Museum
1917.
Sollas, 1924, p. 269.
Manohar Lai Misra
Proc* hid. A cad. Set., B % vol. X\ PL XIV
a b
FIG. 1. Specimen No. N/19 X i
FIG. 2. Specimen No. N/20 X J
Manohar Lai Misra
Proc. Ind. A cad. ScL, /?, vol. X, PL XV
a b
FIG. 3. Specimen No. N/21 X i-
a b
FIG, 4. Specimen No. N/22 x -|
Manohar Lai Misra Proc. Ind, Acad. Sa, B, vol. X, PL XVI
a
FIG. 5. Specimen No. N/23 X -
a
FIG. 6. Specimen No. N/24 X
On Some Stone Implements from Hoshaugabad (C./ J .)
285
5. Das Gupta, H. 0.
6.
7. Evans, J.
8. Ghosh, Manoranjan
9. Foote, R. B.
10. LeMesuriar, H. P.
11. Medlicott, H. B.
12. Oakes, R. E.
13. Oldham, T.
14. Osborn, H. F.
15. Schmucker, S. C.
16. Sollas, W. J.
17. Sripada Rao, K.
18. Swiney, J.D.
19. Theobald, W.
" Indian Prehistory ", Journ. Dept. Sci. CaL Univ.,
1923, 5, 1-29.
''Bibliography of Prehistoric Indian Antiquities, 5 *
Journ. As. Soc. Beng., (N. S.), 1931, 27, 1-90.
Proc. Soc. Antiquaries, 1853, 8.
'" Rock paintings and other antiquities of Prehistoric
and later times," Mem. Arch. Surv.. Ind., 1932 >
24, 1-22.
Prehistoric and Protohistoric Antiquities, Notes on Ages
and Distributions, 1916.
Proc. As. Soc. Beng., 1861, pp. 81-85.
' * Note on a celt found by Mr. Hacket in the ossiferous
deposits of the Narbada Valley (Pliocene of
Falconer)," Rec. Geo. Surv. Ind., 1873, 6, 49-54.
Proc. As. Soc. Beng., 1869, pp. 51-54.
" On the agate flake found by Mr. Wynne in the
Pliocene (?) deposits of the Upper Godavary," Rec
Geo. Surv. Ind., 1868, 1, 65-69.
Man of the Old Stone Age, 1927.
Man's Life on Earth, 1925.
Ancient Hunters, 1924.
" On some stone implements from South India," HalJ-
Y early Jour. Mysore Univ., 1930, 4, No. 2.
Journ. Roy. As. Soc. Bom., 1864, 8, 17-18.
" Geology of Pegu," Mem. Geo. Surv. Ind., 1873, 10,
1-171.
1621-39 Printed at The Bangalore Press, Mysore Road, Bangalore City, by G. Srinivasa R
and Published by The Indian Academy of Sciences, Bangalore
:ao, Superintendent,
ON SOME NEMATODE PARASITES FROM
AFGHANISTAN
BY S. A. AKIITAR
(Prow the Department of tiloloyu, Faculty of Medicine, Kabul}
Received October (5, 191*9
(Communicated by Dr. M. B. Mirza)
Ix the following paper certain parasitic nematodes have been dealt with,
which were recovered from different animals of this place, and a new species
of the germs Thubunea Seurat, 1914, has been described. The writer is
greatly indebted to Dr. H. A. Baylis of the British Museum (Natural History),
I v ondoii, for the useful suggestions on Syphaciella and Spirocerca. He
expresses his thanks to Dr. G. D. Bhalerao, Helminthologist, Veterinary
Institute, Mukteswer, for kindly providing him with the necessary literature.
He is also thankful to Dr. M. B. Mirza, Director, Zoological laboratories,
Aligarh, for kindly going through the paper.
.1. Family Strongylidte Baird, 1.853. Sub-family Strongylina Railliet,
1.893. Genus Strongylus Mueller, 1780, or Goeze, 1782. Strongylus equimts
Mueller, 1780..
Host. Horse. Location. - -Caecum.
"2. Family Oxyurid& Cobbold, 1 8 (VI. Sub-family Qxyurince Hall, 19JO.
Genus Passalums Duj., 1.845. Passalurus ambiguits (Rudolphi, 1819).
Most of the rabbits which were dissected in the laboratory, were found
infected with the parasite.
HOvSt. --Rabbit. Location. Colon.
Genus Enterobius I v each, 1853. Enlcrobius vermicular is (Ijnn., 1758).
Host , Child. location. Rectum (I'tucesj .
Genus Dennatoxys Schneider, 18(>(>. Demuitoxys sj>.
Among many (about 50) rabbits dissected, only two were found infected.
The specimens were, unfortunately, lost in the course of study, and hence
their specific determination not clone. The examination, however, confirmed
Mirza's observations : " the cervical alee in certain specimens extend beyond
the (Bsophageal bulb, while in others they are shorter and above it or
287
B.1 F
288
S. A. Akhtar
terminate at the level of the cesophageal bulb, i.e., the length of the cervical
ate may differ in male and females of the same species."
Host. Rabbit. Location. Colon.
Sub-family Cosmocercina Railliet, 1 91 G. Genus Syphaciella Monnig,
H'iiM. SvphacieJla indica Maplestone, 1931.
PIG. I. tiyphacicUa indica. Tail of male, ventral view
These worms are the same as those of Syphaciella indica, but the writer
linds five instead of four pairs of caudal papillae in the male. Maplestone
seems to have overlooked one pair of comparatively large and conical papilla?
which is situated immediately behind the cloacal aperture. The tips of the
papillae of this pair are often bent towards each other. As the specimens of
the writer were not quite mature, it was very difficult to see the most
posterior pair but one, and so this pair of the caudal papilla? is not. shown,
in the diagram.
Host. Pterodes sp. (Local name : Burnaqara). Location. Geeum.
3. Family Spirurida Oerley, 1885. Sub-family Spirurince Railliet,
1916. Genus Spirocerca Railliet and Henry, 1911. Spirocerca lupi (Rud,
1809) ( = S. sanguinolenta) .
A pair of the worms of this species was recovered from the aorta of a
domestic cat of Kabul. They were quite mature, but of much smaller size
than that given in literature. The species occtirs in the dog, wolf, jackal
and fox, and has also been transmitted to cat experimentally. Therefore it
appears that they have occurred there accidentally and their smaller size is
connected with the fact that they were in an unusual host.
On Souie Neuiatode Parasites from Afghanistan- 289
The male measures 8 -902 mm. in length and 0-540 mm. in thickness,
the female 31 -421mm. and 0-654 mm. respectively. The buccal cavity is
about 0-070 mm. long. The oesophagus is 5-247 mm. long in the male and
0-558 mm. in the female. The left spicule is 2-305 mm. long and the right
0-600 mm. in the female the tail is 0-283 mm. long and the vulva is
0-083 mm. from the posterior end of the oesophagus. The eggs measure
0-02(5 x 0-010 rnm.
Host. Cat. Location. Aorta.
4. Family Physalopteridce Leiper, 1809. Sub-family Physalopterincp
Railliet, 1893. Genus Thubuwaa Seurat, 1914. Thubunaa baylisi N. Sp.*
The cuticle is coarsely striated and lateral alas are absent. The head
bears a pair of submedian cephalic papillae. The mouth is with two lateral
rounded lips, the internal surfaces of which are armed with three blunt and
forwardly directed tooth-like structures. The pharynx-, is. ^short and with
delicate walls. The oesophagus is clearly divided into two parts, an
anterior muscular and a posterior glandular. The cervical papillae are
situated near and behind the nerve-ring and the posterior part of oesophagus
begins in front of the cervical papillae. The excretory aperture is behind
the cervical papilla and the intestine is straight.
The male measures 14-334 mm. or more in length and 0-286 mm. in
thickness near the middle of the body. The head is 0-043 mm. in diam.,
and the length of pharynx is about 0-049 mm. The first part of the
oesophagus is 0-190 mm. in length while the second part is. 1 393 rnm.
The nerve-ring is situated at about 0-167 mm., the excretory aperture
at 0-300 mm. and cervical papillse at 0-207 mm. from the anterior end of the
body. :
The caudal alse are well developed, quite symmetrical, finely striated
transversely and meeting ventrally in front of cloacal aperture. The tail
is - 246 mm. long simply conical and digitif orm and slightly curved to the
dorsal side. The ventral surface of the tail with papilliform elevations
exhibiting a verrucose appearance. There are altogether thirty caudal
papillae 1.6 on the left and 1.4 on the right side. The spicules are subequal,
lightly chitinised, and about 150 mm. long, their proximal ends nearly
truncate and distal pointed.
The female measures 22 -01 6 mm. or more in length and 0-400 mm.
in diam. near the middle of the body. The head is 0-063 mm. in diain.
* The species is named after Dr. II. A. Baylis of the British Musftmn, London
(England).
290
S. A. Akhtar
and the pharynx is 0-050 mm, in length. The iirst part of the oesophagus
is 0-193 mm. long while the second part is 1 -667 mm.
The nerve-ring is situated at about 0-217 mm., the excretory aperture
at 0' 313 mm, and cervical papillae at 0-273 mm. and vulva at 3 -083 nun.
(or at about 1/7 of the total length) from the anterior end. The vulva is a
little salient and the vagina is 0-134 mm. long, is nearly right-angle to the
bod} r surface. The eggs are with thick shells and contain larvte when
deposited. The larva is 0-007 mm. thick and the size of eggs is
0-050 x 0-037 mm.
The tail is 0-400 mm. in length and terminates into a small conical
process.
Six species of the genus Thubunece- have been so far described : T. p-uclica
by Seurat in 1914 ; T. parkeri and T. asymmetrica by Baylis in 1920 and
1938 ; T. fitzsamonsi by Ortlepp in 1.931 ; and T. greyicola and T. agama by
vSandground in 1933. The new species T. baylisi differs from all the above-
named species in most of the measurements and is differentiated from all
of them by its lips armed internally with only three blunt, forwardly directed
teeth. The posterior part of the oesophagus begins near and behind the
nerve-ring. In spite of the peculiar arrangement of its caudal papillae, the
caudal alse are symmetrical and join with each other in front of the cloacal
aperture. The vulva is situated at about the anterior 1/7 of the body
length.
Host. Agama sp. Location. Stomach.
FIG.
a h
Thubunecr bay Us i . a-Anterior extremity of female
extremity of male, ventral viow : c-Egg
lateral view ; ^-Posterior
On Some Nematode Parasites from Afghanistan
291
The type specimens of the new species have been deposited in the
Museum of the Zoological laboratories, Muslim University, Aligarh.
Baylis, H. A.
Hall, M. 0.
Mirza, M. B.
Ortlepp, K. J.
Sandground, J. H.
Seurat, I,. G.
Yorke, W., and Maplestone,
P. A.
''On a New Species; of the Nematode Genus Thubunece,"
Ann. ct- Mag. Nat. Hist., 1926, 9, 18, 361-64.
A Manual of Heltnintholoffy, Medical and Veterinary,
London, 1929.
" A third species of the Nematode Genus ThuhunecB,"
Ann. & Mag. Nat. Hist., 1930, 10, 5, 240-19.
The Fauna, of British India, Nematoda, London, 1 93t>, 1.
l ' Neinatodo Parasites of Mammals of the Orders
Rodentla Lagomorpha, etc.," Proc. 27. *S r . Nat. Mu$. 9
WasJt., 3910, 50, 258.
" A New Species of the .Nematode Genus Dennatoxys
from Lepus riifi-raitdatus," Proc. Ind. A cad. ScL,
1936, 3, Sec. T> r 234.
** A New Species of Thubuneaa, T. fitsslmonsl from
a Kalahari lizard.," Jour. S. Afr. Yet. Med. Assn.,
1931, 2, 328-31.
" Parasitic Nematodes from East Africa and Southern
Rhodesia," Bull. Mus. Comp. Zool. Harv., 1933.
75, 6.
k< Sur mi nouveau Nematode parasite des Reptiles,"
C. R. Soc. BioL, 1914, 76, 721.
C. R. Soc. ftiol., 1914, 76, 714.
The Nematode Parasites of Vertebrates^ London, 1926.
1723-39 Printed at The Bangalore Press, Mysore Road, Bangalore C'ty, by G. Srinivasa Rao, Superintendent,
and Published by The Indian Academy of Sciences, Bangalore
A SYSTEMATIC ACCOUNT OF SOME
SOUTH INDIAN DIATOMS*
BY G. VENKATARAMAN, B.Sc. (HONS.), M.Sc.
University Botany Laboratory, Madras
Received October 16, 1939
(Communicated by Dr. M. O. P. lyengar)
Tim number of papers dealing with systematic accounts of Diatoms from
the different parts of India has been so far very few. The first important
account of the diatom flora of the Indian region was given by Grunow (1865)
in a paper on the Diatomacese and Desmidiacese of the Island of Banka near
Singapore. In this paper he gives an account of 31 species of diatoms from
the island. Previous to this Wallich in 1.860 recorded 2 diatoms from the
digestive cavities of the Salpse. Zeller in 1873 recorded one diatom from
Burma. The following foot-note in his paper states, "The diatoms from
Burma (about 60 or more species) are not yet described ; Dr. I,. Rabenhorst
of Dresden has, however, been kind enough to undertake the determination
of them (S. Kurz)." I am not able to trace if any account of these diatoms
has been published anywhere. In 1.882 George Dickie in a paper on some
algse from the Himalayas gives an account of 28 diatoms. i,euduger-
Fortmorel (1879) has given an account of a number of diatoms from Ceylon.
He (1.893) has also recorded nine diatoms from Malaya and the neighbouring
areas. Schaarschmidt (1.88G) in a paper on Afghanistan algae, has given an
account of 22 diatoms. W. W.est and G. S.,West (1.902) has recorded 49
diatoms from Ceylon. In 1907, they published a paper on the Fresh Water
Algae from Burma including a few from Bengal and Madras, wherein they have
given an account of 59 species, of which two were from Vizagapatam in South
India. Prain (1.905) recorded three forms from Hughli-Howrah districts.
Carter (1.926) in her account of some algae from North India recorded 49
diatoms. Skvortzow (1930) recorded 99 forms from Ceylon, and 56 forms
from Calcutta (1.935). Biswas (1932) has given an account of the records of
diatoms from India by various workers upto 1932. He (1.932, 1935)
records 3 diatoms from Upper India and 9 diatoms (1936) from the Loktak
I^ake, Manipur, Assam. In 1935 Abdul Majeed published a paper on the
Panjab diatoms wherein he has given a detailed account of 62 forms.
* From the University Botany Laboratory, Madras. Thesis (in part) submitted
for the degree of Master of Science in the University of Madras.
293
Bl F
294 G. Venkataraman
It may be seen from the above that all the records have been from
Upper India, Burma, Malaya and Ceylon, and that, with the exception of the
two diatoms recorded by West and West (1907) from Vizagapatam, no work
has been so far done on the Diatomacese of South India. I have therefore
attempted to give a detailed account of the Diatomaceee collected in this
region.
This work was taken tip at the suggestion of Prof. M. O. P. lyengar.
The diatoms dealt with in this paper comprise the collections of several people
including some made by myself. Prof. M. O. P. lyengar was kind enough
to place Ms valuable collections from different parts of South India at my
disposal. A good collection of diatoms from the Nilgiris was kindly handed
over to me for examination by Dr. T. Ekambaram. The stomach contents
of a fish caught in the river Adyar at Madras by Prof. R. Gopala Aiyar and
of another caught by Dr. B. Sundararaj in the Periyar I^ake, Travancore,
were examined, and quite a number of diatoms were found in them.f
A large number of brackish-water diatoms was collected by me in
connection with my paper on the ecology and seasonal succession of the
diatom flora of the river Cooum which runs through Madras. A systematic
account of these diatoms also is included in the present paper.
I take this opportunity of expressing my sincere thanks to all the above
people for their kindness in placing their diatom material at my disposal.
I also wish to thank my several friends who were kind enough to hand over
at various times stnall samples for examinations.
All the drawings were made with the aid of a Zeiss Camera lucida with a
Zeiss oil-immersion objective 90 (ap. 1 -25 ; m Iris) or 120 apochromat
(ap. 1. 3) and with compensation ocular No. 1.0 or 1.5. The drawings were made
from specimens carefully cleaned, dehydrated and mounted in canada balsam.
The material is first treated with concentrated hydrochloric acid. After
an hour, it is washed in water two or three times with the help of a centrifuge.
It is then treated with an equal quantity of concentrated sulphuric acid to
which a few crystals of potassium dichromate is added. The mixture is then
Diatoms found in the. stomach, contents of the fish from. Adyar :
Pleurosigma salinarum Nitzschla obtusa var. scalp elliforniis
Amphora coffewformis Nitzschla dosterium
Navicula digitoradiata
Diatoms found in the stomach contents of the fish from Periyar :
M elosira granulata Pinnularia inter rupta f. yenuina
Cyclotella Meneghiniana Gynibellaturgida
Eunotia pseudolunaris sp. nov. Cocconeis placentula var, euglypta
A Systematic Account of Some South Indian Diatoms 295
allowed to stand for two to four hours. After thorough washing in water with
the help of a centrifuge, the frustules are preserved in 6% formalin. In the
case of diatoms with weakly silicified walls, dilute hydrochloric acid alone is
used.
On a slide previously smeared with a thin coating of Mayer's albumen,
a drop of the cleaned material is placed. This drop is either allowed to dry
up of its own accord or gently heated over a flame to drive off all the water.
The material on the slide is then dehydrated in 95% alcohol and then in
absolute alcohol, and then given two changes in xylol and finally mounted
in canada balsam.
Where the specimens are stray, the slide with the material on it is gently
heated over a flame so that all the water evaporates. Then the material
sticks to the slide. After cooling, the slide is placed in a jar containing
concentrated sulphuric acid to which a few crystals of potassium dichromate
is added. The slide is kept in the mixture from about 1.0 minutes to hours
depending upon the nature of the silicification of the diatoms. It is then
washed in running water for about an hour and then passed through the
alcohol and xylol and finally mounted in Canada balsam.
On the whole 98 forms are described representing 33 genera ; of these
9.8 forms, 67 are new records for India, 3 are new species, 6 new varieties and
6 new forms.
The classification of Hustedt (1930) was followed in the arrangement of
the several forms. As far as possible, only those references that I was
actually able to consult are given under the several species.
Bacillariophyta (Diatomese)
A. Order CENTRAI V ES
I. Suborder DISCINE^
(1) Family COSCINODISCACK/IC
(a) Subfamily Melosiroidese
Genus Melosim Agardh, 1824
vSubgenus Eumelosira
\. Melosira dubia Kiitz.
(Kg- 8)
Hustedt, Fr., Uabenhoi-siAs Kryptoymnen-Flora, Bd. VII, Toil 1, 1930 ; p. 234,
fig. 1)7.
Frustules in long chains united by gelatinous cushions. Frustules with
short mantle portions and arched discs. Sulcus and neck absent. Pseudo-
sulcus broad. Cell walls strong.
296 G 8 Venkataraman
Dimensions.*
Diameter . . . . . . 17-38jx
Height of the half cell . . . . 8-1.4 /u-
Habitat. Brackish water. Ennore back-waters, Madras (!)
The discs are more arched than in the type and look elongated in the
specimens observed. The faint punctae on the valve surface were not
observed.
2. Melosira grantdata (Ehr.) Ralfs.
(Fig. I)
Schonfeldt, Pasclier's Susswaaser-Flora, Heft 10, 1913, p. 1(5, tig. 9 ; Boyer, Syji.
N. Am. Dial., 1927, p. 30 ; Husteclt, Fr., Pascher's Sustncastfcr-Flora, Heft 10, 1930,
p. 87, fig. -14 ; Hustcdt, Fr., Rabcnhorst's Kryptoyamen-Flora, Bd. VII, Toil 1, 1930,
p. 248, fig. 104, a, ft, *, e.
Mdosira polymorphic subsp. granulata (Ralfs) Bethge ; Bethge, Kolkwitz Planzcn-
forschung, Heft 3, 1925, p. 30, Tafel l r fig. 1.
Frustules cylindrical, robust and stiff in detached filaments. Mantle
portions cylindrical, discs flat. Small pseudo-sulcus present. Stilcus some-
what shallow. Neck fairly big. Mantle line straight, parallel. Mantle
surface punctate, puncta coarse in more or less spiral rows. The outer shell
always coarsely punctate, their puncta rows being parallel. The same cells
have spines projecting outside as well as inside the cells.
Dimensions.
Diameter . . . . . . 8-12 ^
Height of the half cell . . . . 8-1.2 /z
Rows of puuctse in the upper cell . . 8-9 in 10 p
No. of punctse in the upper cell . . 8-10 in 10 p
Rows of puncte in the lower cell 10-1.4 in 10 \L
No. of puiictse in the lower cell . . 10-12 in 10 ju,
Habitat. Fresh water. Plankton, Red Hills I/ake, Madras, leg.,
S. V. Ganapati. Periyar Lake, Travancore, from the stomach contents of a
fish, leg., B. Sundararaj.
This form agrees with the type.
* The dimensions given under the various forms in this papor aro those actually
found in the material examined by me.
A Systematic Account of Some Soitth Indian Diatoms 297
3. Melosim gramilata (Ehr.) Ralfs var.
ang'itstissima Mull.
(Fig. 2)
Hustedt, Fr., Pascher's Susswaaser -Flora, Heft 10, 1930, p. 88, fig. 45 ; Hustedt,
Fr., Babenhorst's Kryptog amen- Flora* Bd. VII, Teil 1, 1930, p. 250, fig. 104, d.
Filaments long with narrow and long cells, the height of the cells being
several times the diameter.
Dimensions.
Diameter . . . . . . 3-5 /x ; mostly 4 /t
Height of the half cell . . . . 11-1.4 p
Rows of punctse in the upper cell . . 8-10 in 10 /*
No. of ptincte in the Upper cell . . 8-10 in 10 ^
Rows of punctae in the lower cell . . 10-12 in 10 /*,
No. of punctae in the lower cell . . 10-12 in 10 \L
Habitat. Fresh water. Red Hills Lake, Madras, leg., S. V. Ganapati.
This form agrees with the type.
(b) Subfamily Sceletoneinoidese
Genus Sceletonema Greville, 1865
4. Sceletonema costatum (Grev.) Cleve
(Fig. 6)
Van Heurck, Traite* des Diatomees, 1899, p. 437, pi. 33, fig. 889-890 ; Gran, H. H.,
Nordisches plankton, Botanisher Teil, Bd. VIII, 1908, p. XIX. 15, fig. 7 ; Hustedt, Fr.,
llabenhorst's Kryptogamen- Flora, Bd. VII, Teil 1, 1930, p. 311, fig. 149; Lebour,
Mari<-\ V., Planldonic. Diatoms of Northern Seas, 1930, p. 70, fig. 43.
Friistnles weakly silicified. Cells lens-shaped. Ends of cells rounded.
IvOng spines connect the cells to form usually straight chains. The spaces
between the cells are longer than the cells themselves.
Dimensions.
Diameter . . . . . . 6-5-14 ^
Habitat. Marine. Plankton of the river mouth Cooum, Madras (!)
This form agrees with the type.
Genus Thalassiosira Cleve, 1873
5. Thalassiosira marginata sp. nov.
(Figs. 1,2, 13)
Frustules drum-shaped, quadrangular in girdle view, single, occasionally
united in twos. Valves round with two chromatophores more or less lobed
298 G. Venkataraman
between which lies eccentrically a very small nucleus. Along the
margin of the valve are present 18-22 small punctae. The pttncte are seen
in the girdle view also.
Dimensions.
Diameter . . . . . . 4-6 JJL
Marginal punctse . . . . 18-22
Habitat. Brackish water. River Coottm, Madras (!)
This form occurred as plankton of the river Cooum during the North-
East Monsoon Season (Oct.-Dec., 1.936). Sometimes it was found in such
enormous numbers as to impart a brownish yellow colour to the water.
A similar thing was noticed by Kolbe and Tiegs in lower Werra because of
the presence of a small species of Thalassiosira akin to the present one
(Kolbe and Tiegs, Ber. d. Deut. Bot. Ges. } Bd. XI, VII, 1.929, p. 418, Abb. 2).
In girdle view the height of the diatom as compared with Thalassiosira
nana lyohmann is much more in the present form than in I v ohniaim's figure
of Th. nana (Hustedt, Rabenhorst's Kryptogamen-Flora, Bd. VII, Teil 1,
1930, p. 331, Fig. 1.67 a, b). In girdle view the outline of the valve side is
wavy in Th. nana whereas in this form it is quite straight and the sides
parallel. In the case of Th. nana there are no striations or punctse at the
periphery of the valve view ; but here, there are 1.8-22 pttnctse clearly visible.
Again, I^ohmann's form has 4 chromatophores whereas there are only two in
the present form.
This form agrees with Kolbe and Tiegs' form (Ber. d. Dent, Bot. Ges.,
Bd. XL, VII, 1.929, p. 418, Abb. 2) in certain respects, viz., the two chromato-
phores, the absence of the unpaired process and in the presence of the
marginal punctse. The marginal punctse are 8-1.0 (constantly 9) in their
form whereas in the present one they vary from 1.8-22. In the girdle view
also the punctse are seen along the margin.
(c) Subfamily Coscinodiscoideae
Genus Cyclotella Kiitzing, F.T., 1.834
6. Cyclotella stelligera Cleve and Grunow
(Fig. 10)
Do Toni, SylL Alg., Vol. II, part 2, 1894, p. 1355 ; Schonfeldt, Pascher's Suss-
wasser-Flora, Heft 10, 1913, p. 18, fig. 13 ; Hustedt, Fr. 5 Pascher's Susswasser-Flora*
Heft 10, 1930, p. 100, fig. 65 ; Hustedt, Fr., Rabenhorst's Kryptogamen-Flora, Bd. VII,
Teil 1, 1930, p. 339, fig. 172.
Valves discoid, margins narrow, striae coarse and distinct. The middle
field has a star-like structure with a central puncta and radiating thick and
short lines.
A Systematic Account of Some South Indian Diatoms 299
Dimensions.
Diameter . . . . . . 1.1-13 /x
Striai . . . . . . 10 in 10 p,
Habitat. Fresh water. As stray specimens in the plankton of an
irrigation tank at Vandalur, Madras, leg., K. G. Veeraraghavan.
The radiating short lines in the middle field as given in the descriptions
are elliptic lanceolate in the specimens observed.
7. Cyclotella Meneghiniana Kiitz.
(Figs. 11, 14)
Do Toni, Syll. Alff., Vol. II, part 2, 1834, p. 1354 ; Van Hexirck, Traite' des Diato-
IH&S, 1899, p. 447, pi. 22, fig. 656 ; Boyer, Syn. N. Am. Diat. 9 1926, p. 38 ; Hustedt'
Ki*., Paseher's Susswasser-Flora, Heft 10, 1930, p. 100, fig. 67; Hustedt, Fr., Raben-
horst's Kryptog amen- Flora, Bd. VII, Teil 1, 1930, p. 341, fig. 174.
Frtistules discoid in valve view, rectangular and Undulated in girdle
view. Margin well defined, coarsely striated and the striae wedge-shaped.
Dimensions.
Diameter . . . . . . 11-30 ju,
Strise . . . . . . 8-10 in 10 p.
Habitat. Fresh and brackish waters. Common.
The central portion at first sight appears to be quite smooth, but under
very high magnifications show extremely fine radially arranged punctse as
figured by Van Hettrck (op. cit,, pi. 22, fig. 656). This form agrees in all
respects with the type. It is found as a very common form in the ponds and
pools and is also capable of accommodating itself to salinity. This form
occurs fairly in abundance in the plankton of the river Coourn and its
development is stimulated through an increase in the salinity. Therefore
this form is grouped under " Halophilous forms" according to the classifica-
tion of Kolbe.
8. Cyclotella Kutzingiana Thwaites
(Fig. 9)
I)e Toni, Syll. Alg., Vol. II, part 2, 1894, p. 1358 ; Schonfeldt, Pascher's Suss-
wd$ser-Plora, Heft 10, 1913, p. 10, fig. 18 ; Boyer, Syn. N. Am. Diat., 1926, p. 38 ;
Hustedt, Fr., Pascher's Susswasser- Flora, Heft 10, 1930, p. 98, fig. 32 ; Hustedt, Fr.,
Kabcnhorsti's Kryptogamen-Flora, Bd. VII, Teil 1, 1930, p. 338, fig. 171 a.
Frtistules single, free floating. Valves disc-shaped. Central portion
of the valve punctate, puncta scattered. Striations symmetrically radial,
somewhat extending to the middle. Near the margins they appear broken
through their sparkling nature.
300 G* Venkataraman
Dimensions.
Diameter . . . . .. 18-24 /z
Striae . . . . . . 12 in 10 /i
Habitat. Brackish water. River Cooum, Madras ( ! )
This form agrees quite well with the type.
Genus Coscinodiscus Ehrenberg, C.G., 1838
9. Coscinodiscus Granii Gougli
(PL XVII, Fig. 2 ; Figs. 16, 1.7)
Gmn, H. H., Nordisches plankton, BotaniscJicr Tell, 1908, p. XIX 34 fig. 35 ;
Husteclt, Fr., Raberhorst's Kryptog amen- Flora, Bd. VII, Teil 1, 1930, p. 43t>, fig. 237;
Lebour Marie, V., Planldon Diatoms of Northern Seas, 1930, p. 44, fig. 20.
Valves rounded, aeriolated, aeriolations bigger in the centre. The highest
point in the valve is eccentric and therefore wedge-shaped in girdle view.
Dimensions.
Diameter . . . . . . 229-240 jit
Habitat. Marine. From the plankton of the river month Coonrn,
Madras (!)
Under a high magnification there appears to be a thickening like a
nodule in the central portion of one of the valve surfaces. The other valve
has got a different pattern in the central portion.
II. Suborder
(2) Family
Genus Chatoceros Ehrenberg, 1844
Section Brevicatenata*
10. Ch&toceros orientalis Schiller
(Figs. 3, 4, 5, 7)
Haste dt, Fr., Rabenhorst's Kryptogamcn-Flora, Bd. VII, Teil 1, 1930, p. 721,
fig. 412.
Chains long, straight and stiff. Valves elliptical. Frustules connected
to each other by a short process arising fr>m the slightly gibbous middle
portion of the cell. Bristles long, thin and smooth. End bristles stronger
and longer.
Dimensions.
Length of the valve (apical axis) . . 10-15-5 \L
Habitat. Brackish water. Plankton of the river Cooum, Madras ( ! )
Abundant in the month of September 1936.
A Systematic Account of Some South Indian Diatoms 301
A solitary specimen with a resting spore (Fig. 16) was found in the
collection. The resting spore of this diatom has according to Hustedt
(op. cit., p. 722) not been known previous!} 7 . The resting spore has a smooth
thick wall and in general shape resembles that of Chatoceros Muelleri I/emm-
(op. cit., p. 750, fig. 439) and Chcctoceros subsalsum I/emm. (Kolbe, Pflanzen-
forschung, Heft 7, 1927, Taf. II, Fig. 28-31) which Hustedt considers it the
same as Chcetoceros Muelleri.
In this connection may be mentioned I v ebour's view (Planktonic Diatoms
of Northern Seas, 1930, p. 105). With regard to the resting spores of ChcBto-
ceros I v ebour says that " the young resting spores are often smooth, the
armature coming on later". It is not clear whether the spore observed by
me is only in a young condition which will develop armature later on. But
the resting spores of Chcetoceros Muelleri and Chatoceros subsalsum also
show smooth walls. So it is just probable that the smooth-walled resting
spore in my specimen represents the fully developed condition.
Subfamily Anattlese
Genus Terpsince Ehrenberg, 1843
H. Terpsince musica Elir.
(Figs. 15, 18, 19, 20, 21)
Do Toni, Syll. Ale/., Vol. II, 1804, p. 891; Van Ileurok, Tralte* tics Diatoniees,
1890, p. 452, fig. 170 ; Boyer, Syn. N. Am. Died., 1.927, p. 1 11 ; Hustedt, PP., Kaben-
horst-'s Kr-yptogamcn-Flora, Bel. VI I, Toil 1, 1030, p. 898, fig. 510.
Frustules in girdle view quadrangular, United in long zig-zag chains.
The several septa of the frustule have their inner margins curved and slightly
thickened resembling the musical notes. Valves linear, elliptical with undu-
lating sides and slightly knobbed at the ends, divided by the septa into 5 to 7
parts. vStirface coarsely punctate and finer at the ends. In some cases the
surface is both punctate and reticulate, the reticulations being irregular.
A central big pore is seen in the middle of the valve.
Dimensions.
Length of the valve . . . . 95-148 p,
Breadth of the valve in the central
part . . . . . . 35-40 p,
Rows of puncte in radial rows . . 10 in 1.0 ju,
Habitat. Fresh water. Agri.-Hort. Gardens, Madras ( ! )
This is an epiphyte on Pithophom and Cladophora occurring in plenty
adorning the algal filaments like festoons. It is found almost throughout
the year in one particular pond.
It agrees with the type quite well.
1 ""
.
<?,'
7o v&
*
DQ c2 ,. ._,.... .. w ^^
f^fl^
o^
=^o
15
19
A Systematic Account of Some South Indian Diatoms 303
B. Order
I. Suborder
(1) Family
(a) Subfamily Tabellarioidese
Genus Tabellaria Ehrenberg, C.G., 1840
12. Tabellaria fenestrata (I/yngbye) Kiitz.
(Figs. 29, 35, 41)
Smith, W., Syn. Brit. Dial., Vol. II, 1856, p. 46, pi. XLIII, fig. 317 ; Be Toni,
S-j/ll. Alg., Vol. II, part 1, 1891, p. 743 ; Van Heurck, Traite' des Diatom 4es, 1899,
p. 356, pi. 11, fig. 477 Boyer, Syn. N. Am. DM., 1926, p. 151 ; Hustedt, PP., Pascher's
Susswasser-Flora, Heft 10, 1930, p. 122, fig. 99 ; Hustedt, Pp., Rabenhoret's Krypto-
yamen-Flora, Bd. VII, Teil 2, Lief 1-4, 1931-32, p. 26, fig. 554.
Frustules rectangular in girdle view, united at the corners by gelatinous
cushions to form zig-zag chains. Septa straight, two at each end, extending
nearly to the centre. Valves linear, elongated, inflated in the middle and at
the ends.
Dimensions.
Length .. .. .. 7 1-85 p
Breadth . . . . . . 7-8 ^
Habitat. Fresh water stream, Pykara, leg., T. Ekarnbaram.
FIG. 1. Melosira(/ranulata($hv.)Il&\ts. x 1000.
FIG. 2. Melosira granulata (Ehr.) Ralfs var. anguslissima Miill. x 1000.
FIG. S.Chrctocwos orienta Us Schiller. Cell in valve view. x 700.
FIG. 4. Chattoceros orienta I is Schiller. A portion of the chain. x 700.
FIG. Z.ChMtoceros orientaUs Schiller. Cell with a, resting spore. x 700.
FIG. 6. -Sceletonerna cosiatnm (Grev.) Oleve. x 600.
FIG. T.C'hfletoceros orlentalis Schiller. Beginning cell of *i chain. x 800.
FIG. S.'M.e.losira dulna Kutz. x 700.
FIG. 9. CydoteUa Kutzingiana Thwaites. X 1000.
FIG. 10. CydoteUa &telli(/era 01. u. Grun. X .1600.
FIG. 11. CydoteUa Meneyliiniana Kiitz. Valve view. X 1000.
FIG. 12. Thalassiosira marginata sp. nov. Valve view. X 1600.
FCG. 13. Thalassiosira maryi-nata sp. nov. Girdle view showing the chloroplasts and
the small nucleus. X 1600.
FIG. 14. Cyclotella Meneghiniana Ktitz. Girdle view. X 1000.
FIG. 15. Terpsincc muslca Ehr. Cell with chloroplasts and nucleus, x 500.
FIG. 16. Coscinodiscus Gran-ii Gough. Central portion on one side of the valve
showing a nodule-like thickening, x 1600.
FIG. 17. Coscinod'iseus Granil Gough. Central portion on the other side of the valve.
X 1600.
FIG. 18. Terpsince musica, Ehr. Cells united in a zig-zag chain. X 85.
FIG. 19. Terpsinoe musica Ehr. Cell in girdle view. X 800.
FIG. 20. Terpsinoe musica Ehr. Cell in valve view. x 350.
FIG. 21. Terpsinoe musica Ehr. A part of the girdle view magnified. X 1600.
304 G, Venkataraman
The frustules of this diatom were found at the mouth of the Adyar
estuary in brackish water. No living specimens were found in the region.
Since this is a fresh-water form, it is evidently brought down from the
upper reaches of the river where the water is fresh and not saltish at all.
It agrees with the type in all respects.
13. Tabellaria flocculosa (Roth) Ktitz.
(Figs. 23, 24)
Smith, W., Syn. Brit. Dial., Vol. II, 1856, pi. 45, pi. XLIII, fig. 310 ; T)e Toni,
%//. Alt/., ' Vol. II, part 1, 1891, p. 714; Van Ileurck, Traite* des Diatomees, 1809,
p. 357, pi. 11, fig. -178; Schonfeldt, Pa-scher's Sussicasser- Flora, Heft 10, 1913, p. 27,
fisi. 32 ; Boyer, Si/n. N. Am. Dial., 1920, p. 152 ; Hustedt, Fr., Pascher's Susswasser-
Flora, Heft 10, 1930, p. 123, fig. 101 ; Hustedt, Fr., Rabeiihorst's Kryptoyamen-Flora,
Bd. VII, Teil 2, Lief 1-4, 1931-32, p. 28, fig. 558.
Frustules quadrangular with from 4 to 8 septa, incurved at each
and alternating with those of the opposite end. Valves linear with median
inflation larger than the terminal.
Dimensions.
length . . . . . . 20-24 /*
Breadth .. .. .. 8-8-3^
Stride . . . . . . 15 in 1.0 p,
Habitat. Fresh water. Road side ditch, Kodaikanal, Pulneys, leg.,
K. S. Srinivasan. Fresh-water stream, Masingudi, Ootacamttnd, leg,,
T. Ekambaram. Fresh- water stream, Pykara, leg., T. Ekambaram.
The specimens observed are wider in the middle of the valve agreeing
with the figure given by Van Hettrck (op. cit., pi. 11, fig. 478).
(6) Subfamily Fragilarioidese
Genus Fragilaria Lyngbye, 1.819
14. Fragilaria intermedia Grun. var. robusta var. nov.
(Figs. 27, 42)
Frustules in girdle view linear, rectangular, united together to form
long bands. Valves linear with parallel sides and gradually tapering ends.
Ends slightly capitate. Striae coarse and distinct and on one side absent
in the middle region and therefore with a unilateral central area.
Dimensions.
length . . . . . . 72-1.40 p
Breadth . . . . 5-8 p
Stride , . . . . . U-12 in 10 p,
A Systematic Account of Some South Indian Diatoms 305
Habitat. Fresh water stream, Vaiyampatti, Tricliinopoly, leg., M. O. P.
lyeugar.
This form shows some resemblance to Fragilaria virescens Ralfs (Hustedt,
Fr., Rabeiihorst's Kryptogamen-Flora, Bd. VII, Teil 2, Lief 1-4, 1931-32,
p. 162, lug. 672 in its size and to some extent to Fragilaria capucina Desma-
zieres (op. cit., p. 144, fig. 659, a-e) in the nature of the central area. But in
the girdle view the frustules do not show the white hyaline band in the middle
characteristic of F. capucina. The striae are coarser and far apart in the
Vaiyampatti specimens than in jp. virescens. This form resembles Fragilaria
intermedia Grim. (op. cit., p. 152, fig. 666) somewhat in the slightly capitate
poles, the unilateral central area and in the nature and number of the strke.
The Vaiyampatti form is by far larger than F. intermedia and unlike the
latter has parallel sides.
15. Fragillaria brevistriata Grttn., forma elongata f. nov.
(Figs. 25, 26)
Frttstules in girdle view linear, rectangular, forming small bands.
Valves linear lanceolate with rounded ends. Striae very short and marginal.
Pseildoraphe broad.
Dimensions.
Length . . . . . . . 30-41 p
Breadth . . . . . . 3-4 - 5 p
StriiXi . . . . . . 12-14 in 10 JJL
Habitat. Fresh water pond, Agri.-Hort. Gardens, Madras (!)
The form agrees in all respects with the type but differs from it in being
very much longer. The Madras form has a length of 30-41 ^ while the type
is only 12-28 ju, long.
Genus Synedra Ehrenberg, C.G., 1830
16. Synedra ulna (Nitzsch) Ehr.
(Figs. 37, 39, 43)
De Toni, Syll. Alg., Vol. II, part 1, 189.1, p. Oo3 ; Van Ifeurck, Truite* des Diuto-
mees, 1890, p. 310, pi. 10, fig. 409 ; Boyer,tyn. N. Am. Died., 1926, p. 198 ; Hustedt.,
Fr., Paschev's Sussivasser-Flora, Heft 10, 1930, p. 151, figs. 158-59; Hustedt, Fr.,
Rabcnhorst's Kryptocjamen-Flora, Bd. VII, Teil 2, Lief 1-4, 1931-32, p. 195, fig. 091 A,
a-c.
Frustules in girdle view linear, somewhat broadened at the ends. Valves
linear to linear lanceolate gradually tapering or tapering only near the ends ;
36
V
37 38
39
27
31
32
40
42
43
A Systematic Account of .Some South Indian Diatoms 307
ends rounded. Pseudoraplie narrow, linear. Central area rounded or rect-
angular. vStrise coarse.
Dimensions.
Length . . . . . . 80-137 /x,
Breadth . . . . . . 3 5-7 p
Stride . . . . . . 10-1.2 in 10 ^
Habitat. Fresh water stream, Vaiyampatti, near Trichinopoly, leg.,
M. O. P. lyengar. Red Hills I v ake, Madras, leg., S. V. Ganapati. Fresh
water pond, Presidency College Garden, Madras (!) Fresh water ponds,
Agri.-Hort. Gardens, Madras (!)
The specimens observed agree with the type quite well.
17. Synedra ulna (Nitzsch) Ehr., var. Oxyrhynchus (Ktitz.) Van Heurck
(Mg. 38)
Van Hourck, Traite 1 des Diatomees, 1809, p. 311, pi. 10, %. 418; Jlustedt, Fr.,
Piischcr's Susswasser-Flora, Heft, 10, 1930, p. 152. fig. 100 ; Hustedt, Fr., Rabenhorst's
Kri/pto</a men-Flora, Bd. VII, Teil'2, Lief 1-4, 1931-32. p. 198, fig. fHU B.
Frustules below 100 p. in length, delicately striated and the striae closer.
Dimensions.
Length . . . . . . 60-90 //,
Breadth . . . . . . 5 p
vStrise . . . . , . 14-16 in 10 /x
FIG. 22. Eunotia pectinalls (Dillw. ? Kiitz.) Rabh. var. yibbulosus var. nov. x 800.
FIG. 23. Tabellariafloccidosa(T&ot}\)KTjLt<z. Oils in valve view. X 1100.
FIG. 24. Tabellarlaflocfndosa (Roth] KiiiK. Cells in a., chain. X 1100.
FIGS. 25-26. Fragilaria brevistriata Grun. f. clone/ at a f . nov. X 1000.
FIG. 27. Fragilar-ia intermedia Grun. var. robusta var. nov. Cells in a band. X 800.
FIG. 28. Synedra ulna (Nitzsch) Ehr. var. amphlrhynchus (Ehr.) G-run. End portion
showing the nmscihige poi.'e. x 1000.
FIG. 29. Tabellar-iafenetstmta (Lyng.) Kul-ss. .V coll in valve view showing the septa.
X 325.
FIG. 30. Synedra ulna (Nit/..) Ehr. vtu\ ampJiirhynchus (Ehr.) Grun. x 500.
FIGS. 31-32. -Synedra ulna (Nifcz.) Ebr. var. amphirliynclius (Ehr.) Grun. Middle
portion of the valve showing the variations in the striae. X 1600.
FIG. 34. Asterionella japonica Cleve spiral colony ; cells with chloroplasts. X 250.
FIG. 35. Tabellariafenestrata (Lyng.) Kiitz. A cell in girdle view. X 325.
FIG. 36. Synedra ulna (Nitz.) Ehr. var. constricta var. nov. x 1000.
FIG. 37. Synedra ulna (Nitz.) Ehr. x 700.
FIG. 38. Synedra ulna (Nitz.) Ehr. var. oxyrhynchus (Kiitz.) Van Heurck. x 700.
FIG. 39. Synedra ulna (Nitz.) Ehr. x 700.
FIG. 40. EunotiamonodonJShr. x 1500.
FIG. 41. Tabellarla fenestrata (Lyngb.) Kiitz. (Jells connected in a zig-zag chain.
x 1GO.
FIG. 42. Frayilaria intermedia Grun. var. ro6 usta var. nov. x 1000.
FIG. 43. Synedra ulna (Nitz.) Ehr. x 800.
308 G. Venkataraman
Habitat. Fresh water pond, Presidency College Garden, Madras (!)
This form was found attached to some debris near the edge of the pond.
1.8. Synedra ulna (Nitzsch) Ehr., var. amphiyhynchus (Ehr.) Grun.
(Pigs. 28, 30, 31, 32)
Schonfeldt. Paselier's Suxsica-nser-Floru* Ilet't 10, HHrj, p. :>0 ; Ilustedt, Fr .
Pasu'her's Suxswasser-Flora, 11 eft 10, 19tfO, p. 15-1, fig. 107 ; Hustedt, Fr., Rabenhorst \s
Krypto(jcu)ien,-Flora, Bd. VII, Toil 2, Lief 1-4, lim-;*2. p. 200, fig-. o9I A, e.
Valves lanceolate, broad nearly to the end where it suddenly constricts
to form capitate ends.
Dimensions.
Length . . . . . . 130-240 p,
Breadth . . . . . . 5-7 fi
Striae . . .. .. 10-12 in 10 /x
Habitat. Fresh water stream, Vaiyampatti, near Trichinopoly, leg.,
M. O. P. lyengar. Pond inside Botanical Gardens, Ootacamund, leg.,
T. Ekambaram. Fresh water stream, Glen Morgan, Pykara, leg. , T. Ekain-
baram. Fresh water pond, Presidency College Garden, Madras (!)
The central area varies widely. The striae are completely absent in the
central area or only one striation projecting in the central area or the striae
shorter on one side and absent on the other.
19. Synedra ulna (Nitzsch) Ehr., var. constricta var. nov.
(Fig. 36)
Valves linear lanceolate, strongly constricted in the middle, the segments
gradually tapering towards the ends. Ends rounded. Pseudoraphe
narrow, linear. Central area broad, striae absent. vStriations distinct.
Dimensions.
Length .. .. .. 84-96^
Breadth in the middle . . . . 3-3 . 5 ^
Breadth at the broadest portion
of the segment . . . . 4-5 ^
Stride .. . . .. 10-12 in 10 /*
Habitat. Fresh water. Red Hills Lake, Madras, leg., S. V. Ganapati.
This form was found along with Synedra ulna (Nitzsch) Ehr. and
Cymbella Hustedtii Krasske.
In S. ulna var. impressa Hust. (Hustedt, Fr., Rabenhorst's Krypto-
gamen-Flora, Bd. VII, Teil 2, Lief 1-4, 1.931-32, p. 199, fig. 691, A i) sides
A Systematic Account of Some South Indian Diatoms 309
are slightly concave and the ends wedge-shaped and the form also seems to
be smaller whereas the present form has a deep constriction in the middle
with the segments having tapering sides and the form is also larger.
Genus Astenonella Hassal, 1855
20. Astenonella ja-ponica Cleve
(Fig- 34)
Gran, II. II. , Nordisch-cs Plankton, Botanlscher Teil, JBd. VIII, 1008, p. XIX IIS,
litX. 100 ; Lebour Mario, V., Planktonw Diatoms of Northern Seas, 1930, p. 195, fig. .155 ;
Hustedt, Fr., Rabcnhorsfc's Krtjptogmnen'Flora, Bel. VII, Teil 2, Lief 1-1, 1931-32.
p. 254, fig. 73-1.
Frttstules forming spiral colonies, linear, narrow with parallel sides and
broadened at the base. Chromatopliores plate-like, two in the broadened
base.
Dimensions.
Length . . . . . . 74-100 ^
Breadth at the base . . . . 8-12 p
Habitat.-- -Marine. River mouth Coo urn, Madras (!) The fine delicate
transverse striae recorded by other workers could not be oberved in my
material even with a very high magnification.
II. vSuborder RAPHIDIOIDINE^
(1.) Family EUNOTIACE^
(a) Subfamily Eunotioidese
Genus Eunotia Ehrenberg, C.G., 1837
21. Eunotia pectinalis (Dillw. ? Kiitz.)
Rabenhorst var. gibbulosus var. nov.
(I'ig. 22)
Valves linear. Dorsal side slightly tumid in the middle. Ventral side
slightly concave and gibbous in the middle. Near the end slightly con-
stricted on the dorsal side but not capitate. Ends rounded. Striations
coarse and clear.
Dimensions.
Length .. . . .. 42-120 \i
Breadth . . . . . . 5-8 /A
Strke .. . . .. 7-11 in 1.0 p
Habitat. Fresh water. Pools near Elliot beach, Madras, leg., M. 0. P.
lyengar.
310 G. Venkataraman
This form was found as a brownish scum inside the water on the dissected
leaves of Limnophila grateoloides. They are united to each other by thin
valves to form long bands.
This form is in all respects identical with the figure given by Hustedt
in Pascher's Susswasser-Flora, Heft, 10, 1930, p. 182, fig. 241. In his later
work (Rabenhorst ; s Kryptogamen-Flom, Bd. VII, Teil 2, Lief 1-4, 1.931-32,
p. 297), be gives two entirely different figures (figs. 763, & and c) for the
variety ventralis (Ehr.) Htist. and his fig. 241 of the earlier work does not
find a place in his later work. The present alga, in possessing a dorsal
swelling in the middle, does not agree with the figure of E. ventricosa Ehr.
var. ? elongata Grunow (Rabenhorst's Beiir. Kenntn., Alg. 2, 1865, p. 4, Tafel
1, fig. 4) which Hustedt considers as a synonym of Eunotia pectinalis var.
ventralis.
This appears in a way somewhat distantly similar to var. undulata. In
a w r ay this could be considered as var. undulata with only a single undulation
in the middle of the dorsal region. It forms a sort of a transition between
var. ventralis and var. undulata. Moreover, this form does not show any
constriction on either side of the middle swelling on the ventral side of the
valve nor the club-shaped halves as described and figured by Hustedt
(Rabenhorst's Kryptogamen-Flora, Bd. VII, Teil, 2, p. 297, fig. 763 b and c).
22. Eunotia monodon Ehr.
(Kg. 40)
Smith, W., Syn. Brit. Died., Vol. I, 1853, p. 16, pi. II, fig. 16 ; Schonfeldt, Pascher's
Susswasser-Flora, Heft 10, 1913, p. 45, fig. 73 ; Boyer, Syn. N. Am. Died., 1027, p. 221 ;
Hustedt, Fr., Pascher's Susswasser-Flora, Heft 10, 1930, p. 185, fig. 254. ; Hustedt, Fr.,
Kabenhorst's Kryptogamen-Flora, Bd. VII, Teil 2, Lief 1-4, 1931-32, p. 305, fig. 772
"> k-
Valves arcuate with the dorsal side well bent and gradually narrowing
towards the ends, ends rounded. Striations coarse, slightly narrower near
the ends.
Dimensions.
Length . . . . . . 26-70 /x
Breadth . . . . . . 7-1.1 /*
Striae .. .. .. 8-10 in 10 p
Habitat. Fresh water. Pools near Elliot beach, Madras, leg., M. O. P.
lyeiigar. Presidency College Garden pond, Madras, leg., Miss John ; Agri.-
Hort. Gardens pond, Madras (!) Fresh water pond, Adyar, Madras (!)
This seems to be a narrower form, the maximum breadth reached being
equal to the minimum of the type.
A Systematic Account of Some South Indian Diatoms 311
23. Eunotia pseudolunaris sp. nov.
(Figs. 53, 61)
Valves linear, .slender, arcuate with parallel sides. Ends rounded.
End nodules small. Raphe in the valvular plane with a line-like appendage
beginning from the end of the raphe and going backwards parallel to the
apical axis. Raphe very small, comma-shaped. Strife fine and clear.
Dimensions.
Length . . . . .. 68-102 /A
Breadth .. .. .. 3 -5-4^
Striae .. .. .. 14-15 in 10 /x
Habitat. Fresh water. Periyar Lake, Travancore, leg., B. Sundararaj
from the stomach contents of a fish.
This form at lirst sight looks like Eun. lunayis (Ehr.) Grun., the dimen-
sions agreeing very well with the type. But on closer examination, with a
higher magnification there is a line-like appendage beginning from the end
of the raphe and going backwards parallel to the apical axis, which resembles
that of Eun. flexuosa (Breb.) Kiitz. and Eun. pseudopectinalis Hust. The
present form has got neither a capitate end characteristic of Eun. flexuosa
nor the sudden tapering near the end, characteristic of Eun. pseudopectinalis.
But for the presence of the line-like appendage at the end of the raphe which
can be observed only under a very high magnification, and the smooth nature
of the ends of the valve without even the slightest depression just below the
poles, this form resembles Eun. lunaris (Ehr.) Grun. in all other respects.
Hustedt in Rabenhorst's Kryptogamen-Flora, Bd. VII, Teil, 2, Lief 1-1, 1931-
32, p. 266, classifies liunokia according to the presence or absence of the line-
like appendage. Only two forms have been recorded so far having this
appendage, viz., Eun. flexuosa and Eun. pseudolunaris. The present form
does not resemble either of these.
III. Suborder
(1) Family ACHNANTHACE^E
(a) Subfamily Cocconeoidese
Genus Cocconeis Ehrenberg, 1838
24. Cocconeis placent-ula Ehr. var. euglypta (Ehr.) Cle^e
(Figs.. 04, 65, 66)
Clevo, K. 8v. VeL-AJtad. HandL, part 2, Bd. 27, No. 3, 1895, p. 170 ; Hustedt,
Fi\, Pascher's Susswasser-Plora, Heft 10, 1930, p. 190, fig. 261 ; Hustedt, Fr., Ttabon-
horst's Kryptogamen-Ftora, Bd. VII, Teil 2, Lief: 1-4, 1931-32, p. 349, fig. 802 c.
58
47
60
A Systematic Account of Some South Indian Diatoms 313
Valves broadly elliptical, the two valves being similar in outline but
dissimilar in punctation. Hypotheca with a raphe and a distinct central
nodule. PitnctcC fine, in radial rows. The margin of the valve separated
from the central part by a hyaline narrow portion. Epitheca with the
pseudoraphe with longitudinal hyaline wavy lines as a result of the transverse
striations being composed of short thick lines.
Dim-en sions.
Length . . . . . . 14-28 ft, '
Breadth . . . . . . 9-15 /*
Stride on the epitheca . . . . 1.8-19 in 10 p.
Habitat. Fresh and slightly brackish waters. Common. Fresh water
stream, Vaiyampatti, near Trichinopcly, leg., M. O. P. lyengar. Stream,
Kodaikanal, Pulneys, as scum near the water edge along with other diatoms,
leg., K. V S. Srinivasan. Fresh water ponds and pools, Madras (!) as epiphyte
on Cladophora and Rhizoclonium.
The forms are variable in outline from more or less rounded to fairly
elongated.
(b) Subfamily Achnanthoidese
Genus Achnanthes Bory, 1.822
25. Achnanthes Hauckiana Grun.
(Figs. 50, 51, 56)
Do Toni, 8yll. Aly., Vol. II, part 1, 1.891, p. 481 ; Boyer. Syn. N. Am. Dial., 1927,
p. 237 : Ilustedt, Fr., Pascher's Suaftwa-saer-Flora, Heft 10, 1980, p. 202, fig. 290;
Hnstodt., Fr., Ra,benlmr*t's Kryptorjamen- Flora, Bd. VII, Teil 2, Lief 1-4, 1931-32,
p. :*fiS, %. SIM.
FIGS. 11-15. Mastof/loia cxitjua Lewis f. Inwiroslris f. nov. X 1GOO.
FIGS. 4^ -i^.- Achnanthes coamtdta Brob. var. parallella var. nov.
FIG. W.Mafttogloia dolosa, sp. nov. x 1000.
FIGS. SQ-rA.AchnantheaHaiicldanaGvwn. X 1000.
FIG. 52. AchnanthesinflataKuto. >< 800.
FIG. frA.~Eunotia pseudolunaris sp. nov. End portion showing the raphe and the
lino -like appendage. X 1600.
FIG. M.Achnantties inflata Kiitz. The valve with the pseudoraphe. x 800.
1^0. 55. Achnanthes brevipes Ag. vav. intermedia (Kutz.) Cleve. A terratological
form. X 800.
I^ta. 50. Achnanthes Hauckiana Grun. X 1600.
Fin. 57. -Mastogloia Hrauni Grvin. Showing loculi. X 1600.
FKW 58-50. -Ac/manMes 'brevlpes Ag. var. intermedia (Kiitz.) Cleve. X 1100.
Fro. (to. Achnanthes brevipes Ag. var. intermedia (Kutz.) Cleve. A terratological
form. X 1100.
FIG. 01. Eu-notia psc.udolunaris sp. nov. X 800.
FlG< 02.- Mastogloia Brauni Grun. x 1 600.
FIG, 03. Afi7wn#/iefiZ)rei?^eAt<. var. intermedia (Kiitz.) Cleve. Girdle view, x 1100
FIOP. 64-00. Cocconei*pZaceH.wte (Ehr.) var. euglypta (Blir.) Cleve. X 1100.
FIG. C)l.~- Achnanthes inflata Kiitz. Girdle view. X 1100,
314 G, Venkataraman
Valves elliptic lanceolate with slightly truncate ends. Pseudoraplie
linear lanceolate. Raplie thin, thread-like, axial area narrow, central area
somewhat broadened. Hypotheca with the raphe having clearly radial
stride ; the stride on the epitheca only slightly radial.
Dimensions.
Length .. .. . . 18-20 /x
Breadth . . . . . . 5-7 p.
Striae .. .. .. 12-15 in 10 /*
Habitat. Brackish water. Plankton of river Cooum, Madras (!)
This form was found here and there in the plankton of the river Cooum.
It never occurred in abundance. Boyer states that this occurs in the hot
springs as well as in brackish waters in Canada. The specimens agree with
the type quite well.
Subgenus Achnanthidium (Kiitz.) Heiberg, 1863
26. Achnanthes coarctata Breb. var. parallelled var. nov.
(Pi. XVII, Fig. I ; Figs. 46, 47, 48)
Valves linear elliptic with rounded ends and almost parallel middle
portion. Raphe straight and coarse in the middle. Axial area narrow but
clear, central area transversely widened, rectangular. Striae slightly radial
delicately punctate. PseUdoraphe narrow, excentric. Frustules in girdle
view slightly bent.
Dimensions.
Length . - . . . . 44-75 /*
Breadth . . . . . . 8-11 p
Strise . . . . . . 1.2-14 in 10 p
Habitat. Brackish water. Cooum estuary, Madras (!), epiphytic on
the filaments of Lyngbia.
This form has got some resemblance to A. coarctata (Breb.) Grun. var.
elliptica Krasske (Hustedt, Fr., Rabenhorst's Kryptogamen-Flora, Bd. VII,
Teil 2, I^ief 1-4, p. 420, fig. 872 d, e}. But the present form has got more
parallel sides and the form itself is narrow and many times longer than broad.
This is by far a bigger form than the type whose length varies only from
28-40 p whereas the present one has a length of 44-75/u,.
27. Achnanthes inflata (Kiitz.) Grun.
(Figs. 52, 54, 67)
De Toni, Syll. Alcj., Vol. II, part 1, 1891, p. 475 ; Boyer, Syn. N. Am. Diat. 9 .1.927,
p. 233 ; Hustedt, Fr., Pascher's Sftssioasser-Flora, Heft 10, 1930, p. 209, fig. 307,
A Systematic Account of Some South Indian Diatoms 315
Hustedt, Fr., Rabenhorst's .Kri/ptor/a men-Flora, Bd. VII, Teil 2, Lief 1-i, 1931-32,
1>. 421, fig. 873.
Frtlstules epiphytic. Valves linear, bulged in the middle with broadly
rounded poles. The pseudoraphe is excentric, stride raidal punctate, puncta
coarse and clear.
Dimensions.
Length . . . . . . 44-50 \L
Breadth . . . . . . 1.0-14 ^
Stride . . . . . . 10-12 in 10 ^
Habitat. Fresh water. Agri.-Hort. Gardens pond, Madras (!)
This is an epiphytic form found along with Ach. brevipes Ag. var. inter-
media (Kiitz.) Cleve on Cladophora and Pithophora. It agrees well with the
type.
28. Achnanthes brevipes Agardh var.
intermedia (Kiitz.) Cleve
(Figs. 55, 58, 59, 60, 63)
Schonfeldt, Pascher's Susswasser-Flora, Heft 10, 1913, p. 58 ; Boyer, Syn. N. Am.
DiaL, 1927, p. 232 ; Hustedt, Fr., Pascher's Susswasser-Flora, Heft 10, 1930, p. 210,
fig. 310 ; Hustedt, Fr., Rabenhorst's .Kryptogamen-Flora, Bd. VII, Teil 2, Lief 1-4,
1931-32, p. 425, figs. 877 d, c.
Valves linear with rounded poles, constricted in the middle in the
hypotlieca and the constriction not seen in the epitheca. Central area
stattroid. The pseudoraphe on the epitheca side, very near the margin.
Rows of punctse more radial towards the poles ; punctae large and distinct.
Dimensions.
length . . . . . . 35-44 /*
Breadth . . . . . . 8-12 /*
Rows of punctse . . . . 9-10 in 10 /L
Habitat. Fresh and brackish water. Presidency College Garden pond,
Madras (!). Agri.-Hort. Gardens ponds, Madras (!). Fresh water stream,
Vaiyampatti, near Trichinopoly, leg., M. O. P. lyengar. River mouth, Cooum,
Madras (!). Adyar estuary, Madras (!)
This form is found as an epiphyte on the filaments of Rhizodonium,
Cladophora, Pithophora, Oedogonium, Lyngbia and Oscillatoria in the fresh
water ponds. They are attached by their valve faces often forming groups
containing 2 to 4 individuals. This form is often associated with other
epiphytic diatoms such as Cocconeis placentula Ehr. var. euglypta (Ehr.)
Cleve and Gomphonema lanceolatum Ehr. Near the mouth of the river
316 G. Venkataraman
Cooum this form is found throughout the year in plenty as a brownish coating
on rocks immersed in the water. Many of the filaments of Ch&tomorpha
litorea Harv. growing in this place were often fully covered with this diatom.
This was found as a stray epiphyte 011 some Enteromorpha plants also in the
same place.
Two teratological forms of this diatom were collected from this place
which possess normal shape but the punctae in some portions are irregu-
larly disposed (Figs. 55, 60).
IV. Suborder BIRAPHIDINE^:
(I) Family NAVICUI/ACEJ^
(a) Subfamily Naviculoidese
Genus Mastogloia Thwaites, 1856
29. Mastogloia dolosa sp. nov.
(Fig. 49)
Valves elliptic lanceolate, subrostrate. Axial area narrow. Central
area big, rectangular, in conjunction with the two longitudinal hyaline
furrows form a H-shaped figure. The furrows on either side of the raphe
converge and seem to meet at the top. Striations slightly radial and very
finely punctate. I/oculi of equal size, ending at a distance from the ends.
Dimensions.
Length . . . . . . 44-62 //,
Breadth . . . . . . 14-20 //,
vStriae . . . . .. 24-26 in 10 ^
Breadth of the loculi . . . . 2 /^
No. of loculi . . . . . . 4|-5 in 1,0 p
Habitat. Brackish water. Adyar estuary, Madras (!) Occurs on the
moist soil as a brownish film along with Mastogloia Brauni Gritn.
This form at first sight looks like Mast. Smithi Thwaites which it resem-
bles in its elliptic lanceolate shape, the rostrate ends and the equal size of its
loculi. But it differs from it in having two longitudinal hyaline furrows
which together with the central area forms definite H-shaped figure. This
H-shaped figure owing to the extremery delicate structure of the stride, is not
seen at first but can be seen very clearly under higher magnifications. In
having this H-shaped figure it resembles Mcist.pumila (Grun.) Cleve (Hustedt,
Rabenhorst's Kryptogamen-Flora, Bd. VII, Teil, 2, p. 553, fig. 983) but the
loculi of Mast, pumila are of different sizes, the middle 1 or 2 being bigger and
the rest smaller, where as in the Madras diatom the loculi are more or less
A Systematic Account of Some South Indian Diatoms 317
equal in size. Again the two longitudinal hyaline furrows in M. pumila are
quite parallel throughout their length whereas in the Madras diatom the
two furrows on either side of the raphe converge and appear to meet towards
the ends. In this latter respect it resembles Mast, exilis Hust. (op. cit.,
p. 553, fig. 985) where the two furrows converge towards each other at the
end, but Mast, exilis is a much smaller diatom (15-20/x) long and the loculi
much smaller in number and confined only to the middle portion of the valve.
The present form may therefore be considered as a new species which can be
placed between Mast, pumila and Mast, exilis.
30. Mastogloia Brauni Grttn.
(Figs. 57, 62)
Ciovr., K. #*. VcL-Akad. HandL, part 2, Bd. 27, No. 3, 1895, p. 158 ; Van Ileurck.
TraUtf des Diatomees, .1899, p. 15(5, pi. 2, fig. 56 : Skvortzow, Jot.tr. Bot., Vol. 05, 1927,
p. 101, %s. ;!, .1, ; Hustodt, Fi-., Fischer's Susswasser- Flora, Heft 10, 1930. p. 218,
fig. 320; llustedt, Fr., Rabenhorst's Kryptogamcn- Flora, Bd. VII, Teil 2, Lief 1-4,
I.9:U-32, p. 551, fig. 982.
Valves elliptic lanceolate. Axial area narrow. Central area big,
rectangular. The two narrow longitudinal horns together with the central
area form a H-shaped figure. Striations transverse in the middle, radial
towards the ends, punctate, punctre fine but distinct. Loculi bigger in the
middle, smaller at the ends.
Dimen sions.
Length . . . . .. 55-76 //,
Breadth .. .. .. 15-18- 5 /x
Ivoculi . . . . . . 4-6 in LO /x.
vStriai . . .. . . 18-20 in 10 //,
Breadth of the loculi in the
middle . . . . . . about 2-5/x
Breadth of the loculi near the poles about 1. -5 /x,
Habitat. Brackish water. Adyar estuary, Madras (!) Occurs as a
brown film on the moist soil along with Mastogloia dolosa sp. nov.
The punctse were fine but distinctly seen under high magnification
conforming with the figures given by Van Heurck and Skvortzow. Hustedt's
form seems to have bigger puncte.
3 1 . Mastogloia exigua I v ewis forma brevirostris f . nov.
(Figs. 4-4, 45)
Valves elliptic. Raphe straight. Axial area narrow. Central area
square. Striations radial. lyoculi mostly 5 in number, bigger in the middle
318 G. Venkataraman
and smaller at the ends, bending suddenly inwards and out again before
joining with, each other at the end of the poles.
Dimensions.
Length . . . . . . 30-33 p
Breadth . . .. . . 9-9 -5 ft
Stride . . . . . . 21-23 in 10 ft
Breadth of the bigger loctili . . 2 2-2 5 p
Breadth of the smaller loctili . . 1. 2 ft
Habitat. Brackish water. Adyar estuary, Madras (!) Found as a
greenish film on the mud inside water along with other diatoms.
This form agrees in all respects to Mastogloia exigtia Lewis (Hustedt,
Fr., Rabenhorst's Kryptogamen-Flora, Bd. VII, Teil 2, Leif 1-4, 1931-32,
p. 569, fig. 1003) excepting for the characteristic end portion of the longi-
tudinal septa. The two longitudinal septa before joining together near the
ends of the poles bend suddenly inwards and out again, the outline formed
by the two septa appearing very much constricted near the poles and then
becoming rounded at the ends. This feature was seen in all the specimens
observed. Therefore this form may be considered as a new form of Masto-
gloia exigua Lewis.
Genus Gyrosigma Hassal, 1.845
32. Gyrosigma balticum (Ehr.) Rabh.
(Figs. 71,72)
Cleve, K. Sv. Vet.-Akad. Handl. 9 paxt 1, Bd. 26, No. 2, 1894, p. 118; Boyer, Syn.
N. Am. Diat.* 1927, p. 456 ; Hustedt, Fr., Pascher's Siisswasser- Flora, Heft 10, 1930,
p. 224, fig. 331.
Valves broad, linear, slightly sigmoid. Raphe eccentric and sigmoid.
Central area small, oblique. Transverse and longitudinal striations equally
distant.
Dimensions.
length . . . . . . 250-300 p
Breadth .. .. .. 26-32 /x
longitudinal and transverse stride . . 11-1.3 in 1.0 /x
Habitat. Brackish water. River mouth Cooum, Madras (!)
This form agrees well with the figure and description given by Hustedt.
33. Gyrosigma distortum (W. Smith) Cleve
var. Parkeri Harrison
(Fig. 69)
Cleve, K. Sv. Vet.-Akad. HandL, part 1, Bd. 26, No. 2, 1894, p. 11C ; Hustedt,
Fr., Pascher's Susswasser-Flora, Heft- 10, 1930, p. 224, fig. 335.
A Systematic Account of Some South Indian Diatoms 319
Valves lanceolate, slightly sigmoid, with obtuse protracted ends curved
in opposite directions. Raphe central, sigmoid. Transverse stride more
distant than longitudinal.
Dimensions.
Length .. .. .. 91-U5/X
Breadth .. .. .. 19-2 1 p,
Long. vStrise . . . . . . 23 in 10 /z
Trans. vStrte . . . . . . 20-21 in 10 /x
Habitat. Slightly brackish water. Plankton of the upper portion of
the river Cooitm, Madras (!) Only stray specimens were found here and
there.
This form agrees with the type quite well.
34. Gyrosigma scalproides (Rabh.) Cleve
var. eximia (Thwaites) Cleve
(Fig. 76)
Cleve, K. Sv. Vet.-Akad. HandL, part 1, Bel. 26, No. 2, 1801, p. 118 ; Htistedt,
Fr., Pascher's Susswasser-Flora, Heft 10, 1930, p. 226, fig. 339.
Valves linear with parallel sides and obliquely rounded ends. Raphe
straight, nearly central, slightly sigmoid at the ends. Transverse strife
finely punctate. longitudinal stria? very faint.
Dimensions.
Length . . . . . . 50-55 ^
Breadth . . . . . . 9-10 ^
Trans. vStriae . . . . 24 in 10 /x
Habitat. Brackish water. Adyar estuary, Madras (!)
This form agrees with the description of Cleve and Hustedt. The
longitudinal striae were very faint and were observed only with difficulty
under a very high magnification.
Genus Pleurosigma W. vSmith, 1852
35. Pleurosigma angulatum (Qtiekett) W. Smith
(Figs. 68, 73)
Smith, W., Syn. Brit. Died., Vol. I, 1853, p. 05, pi. XXI, fig. 205 ; Cleve, K. Sv.
Vet.-Akad, HandL, part .1, Bel. 26, No. 2, 1894, p. 40 ; Hnstedt, Fr., Pascher's Suss-
wasser-Flora, Heft 10, 1930, p. 228, fig. 34.2.
Valves slightly sigmoid with acute ends. Raphe central, sigmoid.
Central area small, rhombic. Oblique striae at an angle of 60 equidistant
with the transverse stride,
\>
68
78
79
A Systematic Account of Some South Indian Diatoms 321
Dimensions.
length . . . . . . 151-100 //,
Breadth . . . . . . 29-32 /A
Oblique and transverse striae . . 20 in 10 \L
Habitat. Brackish water. Adyar estuary, Madras (!) Mouth of the
river Cooitm, Madras (!)
This form agrees with the type quite well.
30. Pleurosigma salinarum Grun.
(Figs. 78, 79)
DeToiii, fii/ll.Ah/., Vol. U, part 1, p. 1891, p. 247; Clove, K. 8v. Vet.-Akad.
HandL, part 1, Bd. 26, No. 2, 1894, p. 39 ; Hustedt, Fr., Pascher's Susswasser-Flora.
Iloft 10, 1030, p. 228, lig. 344.
Valves linear lanceolate, slightly sigmoid with rounded ends. Raphe
central less sigmoid. Central nodule elongated.
Dimensions.
length .. .. .. 126-140 /A
Breadth .. . . .. 17-20 /x,
Trans, striae . . . - 23 in 10 ^
Oblique striae . . . . .' 25-27 in !()/*
Habitat. Brackish water. River mouth, Cootim, Madras (!) Adyar
estuary, Madras (!)
FIG. 08. PLeurosi(jma ancjLilatuni (<^iiekiM) W. Smith. X 700.
FIG. (59. Gyrosiytnadistortum (W. Smith) Oleve var. Parkerl UaiTison. X 000.
FIG. 70. Anomceoneit* sphcarophora (Kiitz.) Ptitzer var. sculpta (Ehr.) Midi. f. iadica f.
nov. Showing the two kinds o pimcttc. X 1000.
FIG. 71. Gyrosiynia lalticum (Ehr.) Babli. Showing the longitudinal and transverse
VtrifiB. X 1100.
FIG. 72. Gywsifjma balticum (Ehr.) llabh. X 250.
FIG. 73. Pleuroaigma anyulalum (Quekkott) W. Smith. Showing the stride and the
rhombic central area. X 1600.
FIG. 7-1. Di'ploneis subovalis Oleve. X 1600.
FIG. 75. Anomcjeoneis sphoerophora (Kiitz.) Pfttzer. X 1000.
FIG. 76. Gyrosigma scalproides (Rabh.) Cleve vav. cxlmia ( Tlwvail.es) Cleve. X cSOO.
j^ IG> 77. Colonels Schurnanniana (Grun.) Cleve var. biconstricta Grun. X 1100.
FIG. 78. Pleuroslgma salinar um Grun. x 700.
p IG . 79. Pleuroslgma salinarum Grun. Showing the st>rie. x 1100.
jn IG . 8(). Anomoeoneis aplimrophora (Kiitz.) Pfitzer var. sculpta (Ehr.) Mull. f. indica f .
nov. X 1100.
FIG. 81. Staur one-Is anceps Ehr. X 1500.
FIG. 82. Diploncis interrupta. x 1600.
322 G. Venkataraman
This form occurs in plenty on moist soil near the water edge at the mouth
of the river Coouni. Sometimes it is found in the plankton also as stray
specimens.
Genus Caloneis Cleve, 1894
37. Caloneis Schumanniana (Grim.) Cleve
var. biconstricta Grim.
(Fig. 77)
liustedt, Fr., Pascher's Susswasser-Flora, Heft 10, 1980, p. 21-0, fig. .>70.
Frttstules constricted near the poles and the middle portion swollen as
broad as the polar regions. Ends more or less wedge-shaped, rounded.
Axial area narrow, lanceolate. Central area big, elliptic with a lunate
marking on each side of the central nodule. Striations slightly radial.
Dimensions.
Length . . . . . . 37-43 \L
Breadth .. .. .. 9-11-3/1.
Stride .. .. .. 1.7 in 10 /x
Habitat. Fresh water. As plankton in an irrigation tank at Vandalur,
Madras, leg., K. G. Veeraraghavan.
In the specimens observed the middle portion was as broad as the polar
swellings. In no case was the middle portion broader than the polar
portions. The form agrees with the type.
Genus Diploneis Ehrenberg, 1840
38. Diploneis subovalis Cleve
(PI. XVII, Figs. 3 & 4 ; Fig. 74)
Cleve, K. Sv. Vet.-Akad. HandL, part 1, Bd. 26, No. 2, 1894, p. 00, pi. I, fig. 27 ;
Kicli, P., Trans. Roy. Soc., S. Africa, Vol. XXIV, part III, 193H, p. 2.11, pi. X. I.
Valves elliptical. Central nodule large, rounded. Furrows narrow,
elosely following the central nodule and its horns. Coste strong and far
apart, alternating with double rows of alveoli.
Dimensions.
Length . . . . . . 35-40 p.
Breadth . . . . . . 22-24 /*
Costse .. .. .. 8-1 Oin It)/*
Rows of alveoli . . . . . . 1.8 in 10 ^
Habitat. "Fresh water pond, Pykara. leg., T. Ekambaram. vStream,
Coonoor Park, leg., T. Ekambaram.
A Systematic Account of Some South Indian Diatoms 323
The costse of this diatom are very characteristic. They are seen clearly
below the outer surface of the valve when examined at a slightly lower focus.
The costae are very strong and prominent. They have each a capitate end
towards the raphe side. This aspect of the costse has not been referred to by
Cleve (1894, p. 96) in his description of this diatom, nor was I able to find
any reference to this structure in any of the available literature here.
Plate XVII, Fig. 3, is a photomicrograph of the diatom taken at a higher
foctts where the double rows of alveoli are clearly seen. But the costse with
the capitate ends are seen only very hazily. But PI. XVII, Kg. 4, is a
photomicrograph of the diatom taken at a slightly lower focus. Here the
costse have come out very well and each of the costse is seen possessing a
prominent capitate end towards raphe side.
The specimens agree with the type in all other respects. It was only a
rare form in the collections.
39. Diploneis intermpta (Kiite.) Cleve
(Fig. 82)
Clove, K. So. Vet.-Akad. Handl, part 1, Bd. 26, No. 2, 1894, p. 84 ; Boyer, Syn.
N. Am. Dial., 1927, p. 348 ; Carter, N. } Jour. EcoL, Vol. XXI, No. 1, 1933, p*. 175, figs.
7, 19 ; Hustedt, Fr., Pascher's Susswasser-Flora, Heft 10, 1930, p. 252, fig. 400.
Valves deeply constricted, the segments elliptical, rounded at the ends.
Central nodule elongated, quadrate with parallel horns. Furrows linear,
narrow. Coste strong, usually interrupted in the middle of the valve.
Dimensions.
Length . . . . . . 44-60 p.
Breadth in the middle . . . . 1.0-14 - 5 p
Breadth at the segments . . 15-17 ^
Coste ' .. .. .. 1.0-12 in 10 /A
Habitat. Brackish water. Adyar estuary, Madras (I)
When examined carefully Under very high magnification, the furrow
region shows a large number of broad transverse ridges. These transverse
ridges are absent in Hustedt's figure but are shown very well in Carter's
figure. In the figure given by Hustedt, a row of big punctse is seen in the
furrow region. I was unable to see any such puncte in my specimens.
Genus Stauroneis Ehrenberg, 1843
40. Stauroneis anceps Ehr.
(Fig- 81)
Smith, W., Syn. Brit. Dial., Vol. 1, 1853, p. GO, pi. XIX, fig. 190 ; Cleve, K. 8v.
Vet.-Akad. HandL, part 1, Bd. 26, No. 2, 1894, p. 147 ; Van Heurck, Traite' des Diato-
mees, 1899, p. 100, pi. 1, fig. 55 ; Schonfeldt, Pascher's Susswasser-Flora, Heft 10, 1913,
324 G. Venkataraman
p. 114, tig. 249 ; Buyer, tifjH.X.Am. DM., 11)27, p. 422; Peterson, J. !>.. /Jot. of
Iceland, Vol. II, part 2, 1928, p. 380 ; Hustetlt, Fr., Paschor's Sussicatntcr- Flora, Heft
10, 1930, p. 250, fig. 405.
Valves linear lanceolate with produced sttbcapitate cuds. Stauros
reaching the margin. Axial area narrow. vStriae radial, distinctly but finely
punctate.
Dimensions.
Length . . . . . . 44-6:2 p
Breadth . . . . . . 12-14 /*
Striae .. .. .. 20-22 in 10 p
Habitat. Fresh water. Pools near Elicit beach, Madras, log., M. (), P.
lyengar (!) Stream, Masingiidi, Ootacamuncl, Nilgiris, leg., T. Kkambanim.
This agrees very well with the type in all respects.
Genus Anomceoneis Pfitzer, 1.871 l
41. Anomceoneis sphcerophom (Kiitz.) Piitzer
(Fig. 75)
Schonfeldt, Pa-scher's Siisswasser-Flora, Heft 10, 19KJ, p. 87, %. 171 ; Boyer, Mi/n.
N. Am. Diat., 1927, p. 324 ; Hustetlt, Fi. 9 Pascher's Sussicasser-Flom, Heft- H), IHtfl),
p. 262, fig. 422.
Valves elliptic lanceolate with rostrate, capitate ends. Axial area broad,
linear. Central area big, asymmetrical. Striations radial, punctate, puncta
interrupted by longitudinal, wavy, blank lines.
Dimensions.
Length . . . . . . 44-58 ^
Breadth .. .. .. 16-20/1
vStrise - . . . . . . 15-17 in !()/*
Habitat. Fresh water stream, Vaiyampatti, near Trichinopoly, leg.,
M. O. P. lyeugar, Pools near Elliot beach, Madras, leg., M. O. P. lyengar (!)
Slightly brackish water, upper portion of the river Cooum, Madras (!)
This form agrees with the type.
42. Anomceoneis sphczrophom (Kiitz.) Pfitzer
var. sculpta (Ehr.) Miiller forma indica f. iiov.
(Figs. 80, 70)
Valves elliptic lanceolate with only slightly produced but not capitate
ends. Axial area broad. Central area big. Strke radial, punctate, punela>.
interrupted. There are two sets of striations on the same valve surface.
One set is composed of bigger punctse with an asymmetrical central area and
A Systematic Account of Some South Indian Diatoms 325
the other set is composed of finer punctse which fill up the whole valve
surface leaving only a small central area.
Dimensions.
Ivength . . . . . . 62-76 //,
Breadth .. .. .. 2 1-26 /x
Striae .. .. .. 11-1511110^
Habitat. Fresh water pond, Adyar, Madras (!)
This form agrees with A. sphterophora var. sculpta in all respects, but the
central area is asymmetric and on one side the axial area is completely free
from the punctse. But when carefully examined under a very high magni-
fication most of the spaces which appear to be empty show numerous very
fine punctse. These fine punctae are, however, absent in the axial area.
Genus Navicula Bory, 1.822
Section Navicula orthosticha Cleve
43. Navicttla cuspidata Kiitz. var. conspicua var. uov.
(Figs. 83, 88)
Valves rhombic to elliptic lanceolate with rounded and slightly constrict-
ed ends. Axial area narrow, central area slightly broadened. The raphe bent
like hooks in the central area. Transverse striations parallel, slightly
convergent near the poles. Longitudinal striations coarse, clear and promi-
nent closer towards the margins and wider towards the middle field. Some-
times the longitudinal striations are broken in the middle portion near the
central area. The craticular plates characteristic of Navicula cuspidata
Kiitz. are also present.
Dimension s .
Length .. .. .. 120-144 /A
Breadth . . . . . . 28-35 /*
Trans. Stride . . . . . . 12-15 in 10 p
IvOiig. Stria: . . . . . . 8-14 in 10 ju,
Habitat. Fresh water, Masingudi brook, OotacamUnd, Nilgiris, leg.,
T. Ekambaram. Fresh water pond inside Museum compound, Madras,
leg., M. T. Philipose.
This form differs from Navicula, cuspidata Kiitz. in having the longi-
tudinal striae closer towards the margins than towards the middle line. These
longitudinal striae appear to be very strong. They are only 8-1.4 in 1.0 ^
whereas in the type the longitudinal striations are closer (25 in 1.0 p.) than the
transverse striations. Besides this, the present form has slight constriction
133 *
\
17
84
TOffiti-:
t tj::h
ms
m.
88
1
A Systematic Account of Some South Indian Diatom3 327
near the poles. On these grounds, this form has been created as a new variety
of Navicula cuspidata Kiitz.
44. Navicula cuspidata Kiitz. var. ambigua (Ehr.) Cleve
(Fig. 94)
Olevo, K. Sv. Vet.-Akad. HandL, part 1, Bd. 26, No. 2, 1894, p. 110 ; Boyer, Syn.
N. Am. Dial., 1927, p. 336 ; Hustedt, Fr., Pascher's Sussicasser -Flora, Heft 10, 1930,
p. 268, fig. 434.
Valves elliptic lanceolate with produced rostrate ends. Axial area
narrow. Central area slightly widened in the middle. The transverse
striae are slightly radial. The longitudinal stride are equally placed.
Dimensions.
length . . .. . .. 65-77^
Breadth .. .. . . 18- 5-20 fc"
Trans. Stride ; Long. Stride . . 17-19 in 1.0 //,
Habitat. Fresh water stream, Vaiyampatti, near Trichinopoly, leg.,
M. O. P. lyengar.
The produced ends of this form are not so much pronounced as given by
Hustedt in Pascher's Susswasser-Flora, Heft 10, 1.930, p. 268, fig. 434.
45. Navicula halophila (Gnin.) Cleve
f. subcapitata Ostrup
(Kg. 91)
Kolbe, R. W., Plawsenforschung, Heft 7, 1927, p. 68, Tafel 1, fig. (5 ; Hustedt, Fr.,
'Paseher's Sussivasser-Flora, Heft 10, 1930, p. 209.
FIG. 83. Navicula cuspidata Kiitz. var. couspicua var. nov. x 700.
FIG. 84. Navicula gracilis Ehr. x 1000.
FIG. 85. NaV'icula peregrina (Ehr.) Kiitz. var. Kefvingensis (Ehr.) Cleve. x 800.
FIG. 86. Navicula lacustris Greg. var. major var. nov. x 1100.
FIG. 87. Navicula digitoradiata (Greg.) A. Schmidt. X 1100.
FIG. 88. Navicula cuspidata Kiitz. var. major var. nov. Showing the prominent
longitudinal striae and the punctate transverse striae, x 1600.
FIG. 89. Navicula cincta (Ehr.) Ktitz. var. Heufleri Grun. x 1100.
FIG. 90. Navicula rostellata Kiitz. X 1000.
FIG. 91. Navicula halophila (Grun.) Cleve f. subcapitata Ostrup. x 1500.
FIG. 92. Navicula later ostrata Hust. x 1100.
FIG. 93. Navicula protracta Grun. x 1100.
FIG. 94. Navicula cuspidata Kutz. var. ambigua (Ehr.) Oleve. X 1000.
FIG. 95. Navicula salinarum Grun. X 1100.
FIG. 96. Navicula lyra Ehr. x 1000.
FIG. 97. Navicula pigmcea Kiitz. x 1600.
FIG. 98. -Navicula ha$tal>&nt. X 800.
328 G. Venkataraman
Valves lanceolate with slightly produced and capitate ends. Axial area
narrow, linear, central area slightly widened in the middle. Striations
parallel and slightly convergent at the ends.
Dimensions.
Length . . - . - 32-36 ^
Breadth . . . . - - 8-9 ^
Strise . . . . - - 14-15 in 1,0 p
Habitat. Brackish water. River Cooum, Madras (!)
' This form was collected from the littoral region in the upper portion o1
the river Cooum, occurring in plenty as pure forms without any other diatoms.
This occurs in the plankton also sometimes. Kolbe (op. cil.> p. 08) has
collected these forms as pure individuals. Mesohalobic form.
This form slightly differs from the figure' given by Kolbe in its having
a somewhat rounded central portion and a less pronounced capitate end.
Section Navicultz decipientes Cleve
46. Nevicula protracta Grun.
(Fig. 93)
Cleve, K. Sv. Vet.-Akad. HandL, part 1, Bd. 2(5, No. 2, 1894, p. 140 ; lluatedl.,
Fr., Paseher's Susswasser- Flora, Heft 10, 1930, p. 284, fig. 472.
Valves linear with rostrate, truncate ends. Axial area very narrow.
Central area small, slightly widened. Striatioii slightly radial and at the ends
parallel, in the middle somewhat coarse.
Dimensions.
Length .. .. .. 20-26 /LC
Breadth . . . . . . 7 - 5-8 - (5 ^
Stride . . . . . . 18-19 in 10 p
Habitat. Brackish water. Stray in the plankton of river Cooum,
Madras. (!)
This form agrees with the type quite well.
Section Name-nice lincolatcv Cleve
47. Navictda salinarum Grun.
(Fig. 95)
Cleve, K. Sv. Vet.-Akad. HandL, part 2, Bd. 27, No. 3, 1805, p. 19 ; Van Hourck,
Traite 9 des Diatomees, 1899, p. 178, pi. 3, fig. 108 ; Boyer, Syn. N. Am. DM., 1927,
p. 383 ; Hustedt, Fr., Pasclier's Sussicasser-Flora, Heft 10, 1930, p. 295, fig. 498.
A Systematic Account of Some South Indian Diatoms 329
Valves elliptic lanceolate with produced subcapitate ends. Axial area
narrow ; central area big, rounded. vStrise strongly radiate, in the middle
longer and shorter, and in the end transverse.
Dimensions.
length . . . . .. 35-41 ju,
Breadth . . . . . . 1.0-1.2 p
vStrue .. .. .. 1.3-14 in 10ft
Habitat. Brackish water. Plankton of the river Cooum, Madras (!)
This form never occurred in plenty but only as stray individuals. The
specimens agree best with the figure given by Van Heurck in the nature of
the slightly capitate ends.
48. Navicula rostellata Ktitz.
(Fig. 90)
Klitzing, Species Algarum, 1840, p. 75 ; B,abenliorst, Flora Europcea Alg., 1,
1864, p. 200 ; Hustedt, Fr. 5 Pascher's Susswasser- Flora, Heft 10, 1930, p. 297, fig. 502.
Valves narrow, elliptic lanceolate with short, thin, narrowly produced,
rostrate ends. Axial area narrow, central area big, rounded. Striations
delicate radial, and at the ends convergent.
Dimensions.
Length ... .. .. 38-45 p,
Breadth .. .. .. 9-1 0/x
Stride .. .. .. 10-11 in 10 ju,
Habitat. Brackish water. River Cooum, Madras (!)
This form was often met with in the plankton of the river Cooum.
Though it did not occur in plenty, it was found fairly in good number. It
agrees well with the type. The striations are extremely fine and were
observed only with much difficulty.
49. Navicula cincta (Ehr.) Kiitz.
var. Heufleri Grun.
(Fig. 89)
Cleve, K. Sv. Vet.-AJcad. HcmdL, part 2, Bel. 27, No. 3, 1895, p. 16 ; Van Heurck,
Traite' des Diatomees, 1899, p. 178, pi. 3, %. 106 ; Schonfeldt, Pascher's Susswasser-
Flora, Heft 10, 1913, p. 92 ; Hnstedt, Fr., Pascher's Susswasser- Flora, Heft 10, 1930,
p. 298, fig. 511.
Valves linear lanceolate with obtuse ends. Axial area narrow. Central
area small, broadened. Striations radial in the middle and at the end conver-
gent, coarse and distinct. The stride are wider apart than in the species.
330 G, Venkataraman
Dimensions.
Length .. - 26-32 /*
Breadth. .. . - -- 5~ 6 P-
Strias .- 10 in 10 /t
Habitat. Brackish water. River Cootim, Madras (!)
This form was found in the littoral region, on moist soil. This form
agrees with the type in all respects.
50. Navicula grades Ehr.
(Fig. 84)
Oleve, K. Sr. Vet.-Akad. Handl., part 2, Bd. 27, No. 3, 1895, p. 17 ; Schonfeldf ,
Pascher's Susswasser-Flora, Heft 10, 1913, p. 90, fig. 182 ; Boycr, 8yn. N. Am. Dial. ,
1927. p. 385 : Hustedt, Fr., Pasoher's Stosswasser-Flora, Heft 10, 1930, p. 299, fig. 514.
Valves linear, the middle portion with parallel margins and with obtuse
ends. Axial area narrow ; central area rectangular. Striatioiis transverse
in the middle and slightly radial at the ends.
Dimensions.
Length . . . . 36-42 ^
Breadth .. .. - - 6-7-5/A
Striae .. - U-12inl.O^
Habitat. Slightly brackish water. River Coottm, Madras (!)
This form occurs in the littoral region. Not a common form.
5L Navicula peregrina (Ehr.) Ktitz.
var. Kefvingensis (Ehr.) Cleve
(Fig. 85)
Oleve, K. Sv. Vet.-Akad., HandL, part 2, Bd. 27, No. 3, 1895, p, 18 ; Hustodt-, Fr.,
Pascher's Sussicasser-Flora, Heft 10. 1930, p. 300.
Valves lanceolate with obtuse ends. Axial area distinct, narrow, central
area broadened, elliptical. Striatioiis coarse, radial and at the ends conver-
gent.
Dimensions.
Length .. .. .. 40-80 ft
Breadth .. .. .. 1.2-1.6 ft
Strise . . . . . . 7-8 in 10 ft
Habitat. Brackish water. River mouth, CooUm, Madras (!)
The ends are clearly rotlnded and not slightly truncated as given in the
description by Hustedt. This form occurred as pure individuals forming a
thin yellow film over the moist soil near the river mouth, Coottm.
A Systematic Account of Some South Indian Diatoms 331
52. Navicula digitoradiata (Greg.) A. Schmidt
(Kg. 87)
ITnstedt, Fr., Pasehcr's Susswasser-Flora, Heft 10, 1930, p. 301, fig. 518.
Valves lanceolate with obtuse ends. Axial area narrow, central area
transversely widened and irregular. vStriations radial, in the middle alter-
nately shorter and longer and at the end parallel.
Dimensions.
length . . .. .. 51-60 ^
Breadth . . . . . . 1.3-16 ^
Striae . . . . . . 8-9 in 1.0 ^
Habitat. Brackish water. River mouth, Cooum, Madras (!)
This form was collected once from the littoral region among other
diatoms. The specimens show not much alternating shorter and longer
striations in the middle region. It agrees with the type in all other respects.
53. Navicula lot ero strata Hust.
(Fig. 92)
Hiistedt, Fr., Pascher's Susswasscr-Florct, Heft 10, 1930, p. 301. fig. 521.
Valves elliptic lanceolate with bioadly rounded and with more or less
capitate ends. Axial area very narrow. Central area big, rounded.
vStriations delicate, slightly radial and closer towards the ends.
Dimensions.
Length . . . . . . 18-22 p
Breadth . . . . . . 6-9 p
Striae in the middle . . . . 15 in 10 ju,
Stride towards the ends . . . . 21-22 in 1.0 /x
Habitat. Brackish water. Adyar estuary, Madras (!)
This form is slightly smaller than the type described by Htlstedt. The
striations are very delicate and were seen only with difficulty.
54. Navicula hasta Pantocsek
(Fig. 98)
Cleve, K. Sv. Vet.-Akad. HandL, part 2, Bd. 27, No. 3, 1895, p. 25 ; Boyer, Syn.
N. Am. Dial., 1927, p. 399 ; Hustedt, Fr., Pascher's Sussivasser-Flora, Heft 10, 1930.
p. 306, fig. 541.
Valves lanceolate, gradually tapering from the middle to the subacute
ends. Axial area narrow, widened slightly in the middle. Striations
strongly radial, middle few are coarse and slightly wide part.
332
G. Venkataraman
Dimensions.
Length
Breadth
Striae
Habitat. Brackish water.
. . f>0-80 /A
. . 15 p
9-1.0 in 1.0
River Coottm, Madras (!)
This form occurs on moist soil near the water edge along with other
littoral diatoms. Forms that are shorter in length by 10 //, than the type
have been met with.
Section Naviculce punctate- Cleve
55. Namcula lacustris Greg. var. major var. nov.
(Fig. 86)
Valves broadly elliptical with obtuse rounded ends. Axial area narrow,
linear, central area large,' transversely widened, more or less rectangular.
Striations radial, punctate, puncta large and round.
Dimensions.
Length .. .. . . 70-84 p
Breadth . . . . . . 26-30 p.
Stride .. .. .. 9- 1.0 in 10^
Habitat. Brackish water. Adyar estuary, Madras, leg., 8. Dorai-
swami (!)
This form differs from the type (Gregory, Quart. Jour. Micr. Sci. , Vol. IV,
1856, p. 6, pi. I, fig. 23) in several respects. The present form is a bigger
one than the type which is only 35-60 /x long and 1.6-20 ju, broad. The
type has a linear lanceolate shape and with subrostrate or acute ends
(Hustedt, Fr., Pascher's Susswasser-Flom, Heft 10, 1930, p. 310, fig. 555;
Cleve, K. Sv. Vet.-Akad. Handl., Part 2, Bd. 27, No. 3, 1895, p. 4.1) whereas
the Madras form is broadly elliptical with obtuse or nearly rounded ends.
The central area is orbicular in the type (cf. Cleve and Hustedt) whereas in
the present form it is transversely expanded and more or less transversely
elliptic. Again in the type, the punctse limiting the axial area are bigger
than the other punctse, but in the present form these are more or less of the
same size as the other punctse.
Section Naviculcz lyratcz Cleve
56. Navicula lyra Ehr.
(Fig. 96)
Cleve, K. Sv. VeL-Akad. Handl., part 2, Bd. 27, No. 3, 1895, p. (53 ; Van Hcurck,
Traiie* des Diatomees, 1899. p. 202, pi. 4, fig. 161 ; Boyer, Syn. N, Am, Dint., 1927,'
p. 411,
A Systematic Account of Some Soidh Indian Diatoms 333
Valve elliptical with rounded ends. lateral areas linear, narrow,
constricted in the middle and slightly divergent at the ends. Striae
punctate ; ptincta coarse and clear.
Dimensions.
Length . . . . . . 52-60 //,
Breadth , . . . . . 37 /x
Stria; .. .. .. 10- Urn 1.0 /z
Habitat. Brackish water. Adyar estuary, Madras (!)
This form was found as a stray one in a collection of the brown film over
the mud containing a number of other brackish water diatoms. This agrees
very well with the type.
57. Navicula pygmcea Kiitz.
(Fig. 97)
Kiitzing, Species Alftarum, 1849, p. 77 ; Gleve, K. Sv. Vet.-Akad. HandL, part 2,
Bd. 27, No. 3, 1895, p. 05 ; Van Heurclc, Traite des Diatomees, 1899, p. 200, pi. 4,
fig. 164 ; Boyer, Syn. N. Am. Diat., 1927, p. 416 : Hustedt, Fr., Pascher's Susswasser-
Flora, Heft 10, 1930, p. 312, fig. 50.1.
Valves hyaline, elliptical with broadly rounded ends. Axial area
indistinct. Lateral areas constricted in the middle and convergent at the
ends. Striations very delicate.
Dimensions.
Length . . . . . . 23-27 ^
Breadth . . . . . . 1.0-U. /*
Striae about . . . . 25 in 1.0 \L
Habitat. Brackish water. Plankton of river Cooum, Madras (!)
This is a form that is often met with in the plankton of the river Cooum.
The structure is very delicate so that a very high magnification is required
for deciphering it. It agrees with the type. ,
Genus Pinnularm Ehrenberg, 1843
Section Parallelistriatce
58. Pinnularia fasciata (Ivagerstedt) Hust.
(Fig. 107)
Hustedt., Fr., Pascher's Susswasser- Flora, Heft 10, 1930, p. 316, fig. 569.
Valves linear lanceolate with rounded ends. Axial area narrow.
Lanceolate central area widened into a broad fascia reaching the margin.
Striations parallel in the middle and slightly radial towards the ends ; deli-
cate T
G, Venkataraman
A Systematic Account of Some South Indian Diatoms 335
D imensions .
Length .. .. .. 16-21 /u
Breadth . . . . . . 4 5-7 p
. . . . . . 23-25 in 1.0 p
Habitat. Fresh water stream, Vaiyampatti, near Trichinopoly, leg.,
M. O. P. lyengar.
This differs from Hustedt's species in having slightly radial striations at
the ends rather than slightly convergent at the ends. He feels that this
species is doubtful of identification since there is a similar species in
Caloneis bacillum (Grttn.) Meresch. (op. cit., p. 236, fig. 360 a). The present
form in the nature of the striations and the dimensions agrees with Caloneis
bad-Hum (Grttn.) Meresch. But the longitudinal line close to the margin which
is characteristic of the genus Caloneis is not clearly seen. Under very high
magnification, however, it looks as though there is a faint longitudinal line
running close to the edge of the valve, but the line-like appearance may
be a false one due to the edge of the valve and is therefore not quite
convincing.
Section Capitattz
Pinnularia interrupta W. Smith
Smith, W., Sijn. Brit. Diat., Vol. 1, 1853, p. 59, pi. XIX, fig. 184 ; Clevo, K. Sv.
Vel.-Akad., Handl., part 2, Bd. 27, No. 3, 1895, p. 76 ; Fritsch, F. B., Ann. S. Afr. Mus.,
Vol. IX, part VII, 1918, p. 590, figs. 40 a-d ; Fritsch, F. E., and Rich, F., Trans. Roy.
Soc. S. Africa, Vol. XI, 1925, p. 381- ; Hustedt, Fr., Pascher's Susswasser- Flora, Heft
10, 1930, p. 317, fig. 573 a. &.
This is a very variable species. All the forms present in the various
collections showed interruption of the striae at the middle of valve.
FIG. 99. Pinnularia gibbet, Ehr. f . undulata f. nov. x 1600.
Fia. 100. Amphora holsatica H\ist. X 1100.
FIG. 101. Tropiodoneis lepidoptera. x 800.
FIG. 102. Amphora coffeceformis Ag. var. africana Fritsch and Bicli. X 1600.
FIG. 103. AmphiprorapaludosaW. Smith var. subsalina, Cleve. x 1000.
FIGS. 104-105. Amphora coffeccjormis Ag. x 1100.
FIG. 106. Tropiodoneis lepidoptera (Greg.) Cleve. Valve view, x 800.
FIG. 107. Pinnularia fasciata (Lagerst.) Husfc. x 1600.
FIG. 108. Pinnularia giVba Ehr. f. subundulata Mayer. X 800.
FIG. 109. Pinnularia 'borealis Ehr. X 1100.
FIG. 110. Pinnularia interrupta W. Smith f. subcapitata Fritsch. X 1100.
FIG. 111. Pinnularia acrosphceria Breb. x 800.
FIG. 112. Pinnularia interrupta W. Smith f. genuina Fritsch. X 800.
FIG. 113. Pinnularia Braunii (Gran.) Cleve var. amphieephala (A. Mayer) Htist. f.
siibconica f. nov. x 1600.
FIG. 114. Pinnularia viridis (Nitzsch) Ehr. X 800.
336 G. Venkataraman
Fritscli in Ann. S. Afr. Mus., Vol. IX, Part VII, 1918, p. 590, lias grouped
the different forms according to the nature of the margins, the polar regions
and the interruptions of the striae in the middle.
59. Pinnulana intemtpta W. Smith f. gemtina Fritscli
(Fig. 112)
Pinmdarin intcrrupta W. Smith; W. Smith, Syn. Brit-Diat., Vol. I, 1853, p. 50,
pi. XIX, fig. 184.
vSides of the valves straight and parallel, striations interrupted at the
centre of the valve. Ends markedly capitate.
Dimensions.
Length .. .. .. 51-70 /x
Breadth . . .. .. 9-10 -5^
Stride . . .. . . 1 2 in 10 /A
Habitat. Fresh water pond, Adyar, Madras (!) Very slightly
brackish water, river Coottm, Madras (!) From the stomach contents of a
fish in Periyar lake, Travancore, leg., B. Sttndararaj.
This form agrees with the description given by Fritscli and the figure
given by W. Smith and Hustedt (Pascher's Susswasser-Flora, Heft 10,
1930, p. 317, fig. 573 i).
60. Pinnidaria interrupts W. Smith f. subcapitata Fritscli
(Fig. HO)
Syn. : P. subcapitata, Gregory, Quart. Jour. Micr. Sci., Vol. IV, 1850, p. 0, pi. 1,
fig. 30.
Sides of the valve straight and parallel. Striations interrupted at the
centre of the valve. Bnds not very markedly capitate.
Dimensions.
Length .. .. .. '26-7-36 /z
Breadth . . . . . . 5-7 p
Stride .. . . .. IO-UinlO/A
Habitat. Fresh water. Pools near Elliot Beach, Madras, leg., M. O. P.
lyengar. Agri.-Hort. Gardens pond, Madras (!)
This form is slightly smaller than f . genuina Fritsch from the specimens
observed. The striations are far apart and it agrees with the figure given by
Gregory (op. cit.).
A Systematic Account of Some South Indian Diatoms 337
61. Pinnularia- Bmunii (Griin.) Cleve
var. amphicephala (A. Mayer) Httst., forma subconica f. nov.
(Fig. US)
Valves linear lanceolate with more or less markedly convex sides.
vSHghtly constricted below the poles and the ends slightly capitate and more
or less wedge-shaped. Axial area narrow. Central area widened. Strise
interrupted in the middle portion of the valve, radial in the middle and
convergent at the ends.
Dimensions.
Length . . . . . . 40-13 p
Breadth .. e .. . . 5-5-6 p
Stride .. .. .. 14 in 10 p.
Habitat. Fresh water. Pools near Elliot Beach, Madras, leg., M. O. P.
lyengar (!)
This form comes very near P. Braunii var. amphicephala but differs from
it in its ends being not so capitate as in the type variety, but in having
only slightly capitate ends and also in the ends being not rounded as in the
type but slightly wedge-shaped. The form resembles in all respects the
figure P. inter rupta f. Braunii Fritsch (Fritsch, F. E., Ann. S. Afr. Mus.,
Vol. IX, Part VII, 1918, p. 592, fig. 40 a, b).
vSection Distances
62. Pinnularia borealis Ehr.
(Fig- 109)
Clove, K. Sv. Vet.-Akad. HandL, part 2, Bd. 27, No. 3, 1895, p. 80 ; Van ileiuck,
TraUe' des Di.atomees, 1899, p. 170, pi. 2, fig. 77 ; Sclionfeldt, Pasclier's Sussicasser
Flora, Heft 10, 1918, p. 106, fig. 227 ; Boyer, Syn. N. Am. Diat., 1027, p. 437 ; Peterseii,
J. B., Botany of Iceland, Vol. II, part 2, 1928, p. 401 ; Hustedt, Fr., Pascher's Suss-
weaver-Flora, Heft 10, 1930, p. 326, fig. 597.
Valves linear elliptical with broadly rounded ends. Axial area somewhat
widened in the middle. Raphe thread-like, with strongly hooked terminal
fissures. Striatioiis coarse, slightly radial in the middle.
Dimensions.
Length . . . . . . 40-55 ^
Breadth .. .. .. 9-U>
Strise . . . . . . 4-5 in 10 p
Habitat. On a decaying fern leaf in water. Fresh water stream,
Bearshola, Kodaikanal, Pulneys, leg., K. S. Srinivasan.
338 G. Venkataramaii
This form agrees with the type ; only the maximum length in the
material is much below that of the species given by Hustedt (28-lJLO/x,
long and 8-18/4 broad).
It was a rare form in the collection.
Section Tabellarice
63. Pinnulana gibba Ehr. f. sitbunchihita Mayer
(Fig. 108)
Hustedt, Fr., Pascher's Siteswavser- Flora, Heft 10, 1030, p. 827, fig. 001.
Valves linear with slightly wavy margins and gradually tapering from
the middle to the subcapitate ends. Ends rounded or slightly wedge-shaped.
Axial area of variable breadth, dilated in the middle. Striations radial in
the middle, parallel above and convergent at the ends. Striae interrupted at
the middle portion.
Dimensions.
Length .. ., .. 1 00-127 p,
Breadth . . . . . . 12-14 /*
Striae . . .. . . 9-11. in 1.0 //,
Habitat.' Fresh water. As plankton of ail irrigation tank at Vandalur
near Madras, leg., K. G. Veeraraghavan. Only few 7 specimens were found.
64. Pinnularia gibba Ehr. f. undulata f. iiov.
(Fig. 99)
The undulations of the valve margins are more pronounced. There is
only a very slight decrease in the breadth of the valve towards the ends.
The ends are strongly wedge-shaped.
Dimensions.
Length . . . . . . 58-70 p
Breadth .. .. .. 9-10- 5 /*
Stride . . .. . . 9-10 in 10 /*
Habitat. Fresh water. As plankton of an irrigation tank at Vandalur,
Madras, leg., K. O. Veeraraghavan.
This form in the wavy outline resembles P. gibba f . sufamdulata but differs
from it in the undulations being more pronounced and the ends of the valves
being not rottnded but more or less wedge-shaped. Again it is a very much
smaller form than P. gibba f. subundulata both in length and in breadth.
P. gibba f, subundulata is 1 00-127 /x, long and 12-14 /.c broad whereas the
present form is only 58-70 p long and 9-10- 5 /z, broad.
A Systematic Account of Some South Indian Diatoms 339
Section Brevistriata
65. Pinnularia acrosphceria Brebisson
(Fig. HI)
Oleve, K. Sv. Vet.-Altad. HandL, part 2, Bd. 27, No. 3, 1895, p. 80 ; Schonfeldf-,
Pasoher's Susswasser-Flora, Heft 10, 1913, p. 106, fig. 228 ; Boyer, Syn. N. Am. .DM.,
1927, p. 441 ; Hustedt, Fr., Pascher's Susswasser-Flora, Heft 10, 1930, p. 330, fig. 610.
Valves linear, gibbous in the middle and at the ends. Axial area broad,
linear. Median area punctate. Striations nearly parallel and slightly radial
at the ends.
Dimensions.
Length . . . . . . 38-62 ^
Breadth .. .. .. 8-12/1,
Striae . . .. .. 9-12 in 10 /u,
Habited. Fresh water brook, Masingudi, Ootacamitnd, Nilgiris, leg.,
T. Ekambaram, Agri.-Hort. Gardens pond, Madras (!) Presidency College
garden pond, Madras (!)
In a few specimens the punctse in the axial area were a bit difficult to
decipher, though in all the other cases they were quite clear. In the present
form the middle portion of the Valve is always rather more swollen than
the form figured in Hustedt. This form was found here and there among
other diatoms in the collections.
S ection C omplexce
66. Pinmilaria viridis (Nitzsch) Ehr.
(Fig. U4)
Boyer, Syn. N. Am. Diat., 1927, p. 446 ; Hustedt, Fr., Puncher's Susswasser-Flora,
Heft 10, 1930, p. 331, fig. 617 a.
Valves linear with slightly convex margins and rounded ends. Raplie
complex. Axial area narrow, slightly widened in the middle. The central
knot is clearly seen and is slightly shifted to one side. vStrise coarse, slightly
radial in the middle and convergent at the ends. The longitudinal band is
present.
Dimensions. . .
Length .. . . .. 93-13G p
Breadth .. .. .. 18-6-20/a,
Stride . . . . 8-9 in 10 JLG
jfffl&itoJ. Fresh water. Pools near Elliot Beach, Madras, leg., M. O. P.
lyengar (!)
340 G. Venkataraman
In the specimens observed the two margins of the valve appear slightly
more parallel in the middle region than in the type. In this respect it
resembles P. viridis var. sitdeticd ; but the longitudinal bands arc quite well
seen instead of its being not clear as in var. sudetica. The Madras ionii has
also got a resemblance to P. streptoraphe Cleve (Hustedt, Pascher's Suss-
wasser-Flora, Heft 10, 1.930, p. 337, fig. 620). But it is a bigger form (KSO
260 ju, long) and also the raphe is more complex.
(b) Subfamily Amphiproroideae
Genus Amphiprora Ehrenberg, 1843
67. Amphiprora paludosa W. vSmitli
var. subsalina Cleve
(Fig. 1.03)
Cleve, A'. Sv. Vet.-Akad. Handl., part 1, Bd. 20, No. 2, 1891, p. 1 1 ; Hustedt, Fr.,
Pascher's Sussicasser-Flora, Heft 10, 1930, p. 340.
PrUstules membranaceous. In girdle view broad with truncate ends.
Median line sigmoid. Junction line with a large sinus. Frustule strongly
constricted in the middle.
Dimensions.
Length . . . . . . 68-84 ^
Stride . . . . . . 23 in 10//,
Habitat. Brackish water. On moist soil at the banks of the river
Cooum (!)
The extremely delicate striae were seen under very high magnification.
The striae are finely punctate.
Genus Tropidoneis Cleve, 1891
68. Tropidoneis lepidoptem (Greg.) Cleve
(Figs. 101, J.C6)
Cleve, K. Sv. Vel.-Akad. HandL, part 1, Bd. 20, No. 2, 189-1. p. *:, ; Hover M /ti
N. Am. Dial., 1927, p. 480. '
- A ^ o P n h r! Pr01 n WMotoovbtepMoptom (Greg.) Cleve]; Van Heurck, TruiM to D-tult,-
mees. 1899, p. 263, p. 5, %. 287.
Frustules elongated, more or less rectangular, strongly constricted in
the middle. Valves linear lanceolate with acute ends, central area indistinct,
small. Wing Unilateral, projecting above the central nodule. Transverse
striations finely punctate.
A Systematic Account of Some South Indian Diatoms 341
Dimensions.
I v ength . . . . . . 85-120 /z
Breadth .. .. .. 15-2 5 p
Stride . . . . . . about 23 in 10 /x,
Habitat. Brackish water. Coottm estuary, Madras (!) Adyar estuary,
Madras (!)
This form occurs in the littoral regions, on the moist soil among other
diatoms, forming a thin brown film.
(c) Subfamily Gomphocymbelloidese
Genus Amphora Ehrenberg, 1840
Section Halamphora Cleve
69. Amphora holsatica Hust.
(Fig. 100)
Hustedt, Fr., Pascher's Susswasser -Flora, Heft 10, 1930, p. 345, tig. 633.
Frustules lunate with slightly rounded poles. The dorsal side is convex
and the ventral side straight. Raphe straight, parallel to the ventral margin.
Axial area on the dorsal side very narrow. Transverse striations radial,
punctate, puncta big.
Dimensions.
Length .. .. .. 38-5-50^
Breadth .. .. .. 10- 7-18 /A
Stride .. .. . . 15 in 10 jit
Punctse . . . . . . 13-15 in 10 //,
Habitat. Brackish water. CooUm estuary, Madras (!) Adyar estuary,
Madras (!)
The form occurs in the littoral region, on the moist soil near the river
mouth, CooUm and in Adyar estuary. This is found also in the plankton
as stray specimens. It agrees very well with the type.
70. Amphora coffeaformis Agardh
(Figs. 104, 105)
Uleve, K. Sv. Vet.-Akad. Handl., part 2, Bd. 27, No. 3, 1895, p. 120 ; Van Heurck,
TraiW des Diatomees, 1899, p. 13*, pi. 24, fig. 681 ; Schonfeldt, Pascher's Susswasscr-
Flora, Heft 10, 1913, p. M3, fig. 314 ; Boyer, Syn. N. Am. Diat., 1927, p. 260 ; Hustedt,
FP. S Pascher's Susswasser-Flora, Heft 10, 1930, p. 345, fig. 634.
Frustules in girdle view elliptic lanceolate, truncate. Valves arcuate
011 the dorsal margin and straight or slightly concave on the ventral margin.
Ends of the valves slightly protracted and capitate. Striae delicate.
B-t JT
342 O. Venkataraman
Dimensions.
Length .. .. .. 35-49/1
Breadth . . .. - - 11-16 -5 /A
Stride .. 21 in 10 p,
Habitat. Brackish water. Cootim estuary, Madras (!) Adyar estuary,
Madras (!) Bnnore backwaters, Madras (!)
The figures given by the various authors differ very much. The Madras
form agrees best with the figure of W. Smith, Syn. Brit. Dial., Vol. I, 185:*,
p. 19, PL XXX, fig. 251. (Syn : Amphora salina W. Smith).
The diatom is found in abundance on rocks immersed under water near
the mouth of the river Cooum and on moist soil on the banks of the; river.
This is one of the forms which is present throughout the year.
71. Amphora coffeceformis Agardh var. africana
Fritzsch and Rich
(Kg. 102)
Fritsch and Rich, Trans. Roy. Soc. S. Africa, Vol. XVIII, part 2, p. 112, fig. <S a cL
Valves arcuate on the dorsal margin and straight on the ventral margin.
The ends are pronouncedly capitate and slightly bent outwards. The dorsal
side bears slightly divergent punctate striae ; the ventral side structureless.
Dimensions.
Length . . . . . . 25-37 //,
Breadth of the valve . . . . 4 5-7 p,
Stride . . . . . . 17-19 in 10 /u,
Habitat. Brackish water. River Cooum, Madras (!)
This form was collected from the upper portion of the river Cooum
where the water is fairly brackish.
Genus Cymbdla Agardh, 1830
72. Cymbdla alpina Grun.
(Fig. 124)
Cleve, K. $i\ Vet.-Akad. Handl.* part 1, Bel. 20, No. 2, 1891, p. 161 ; Muslrdt, Kr.,
Pascher's Sussicasser-Flora, Heft 10, 1930, p. 352, fig. 641.
Frustules slightly asymmetrical. Valves linear elliptical with convex
sides and slightly obtuse ends. Raphe straight, slightly eccentric. Axial
area narrow, slightly dilated in the middle. Strise slightly radial and finely
transversely lineate.
A Systematic Account of Some South Indian Diatoms 34o
Dimensions.
length . . . . . . 33-38 p.
Breadth . . . . . . 8-1.0 p.
Striae . . . . . . 6 8 in 10 /*
Habitat. Fresh water. Hill stream, Kodaikanal, Pttlneys, leg., K. S.
Srinivasan.
This form was found among other diatoms as a brown scum on decaying
fern leaves. It agrees very well with the type.
73. Cymbella turgida (Greg.) Cleve
(Fig. 125)
Cleve, K. Sv. Vet.-Akad. HandL, part 1, Ed. 26, No. 2, 1894, p. 108 ; Hustedt,, "Fr.,
Pascher's Susswasser-Flora, Heft 10, 1930, p. 358, fig. 660.
Valves lunate with highly convex dorsal side and gibbous ventral margin
and acute ends. Raphe straight, the terminal fissures turned downwards.
Axial area somewhat broad. Striations radial in the middle and slightly
convergent at the ends on the ventral side, punctate, puncta coarse.
Dimensions.
length .. .. 26 -3-53 -4 ft
Breadth .. .. .. 8-5-12 -5/4
Striae .. .. .. 8-1] in 10 /*
Habitat. Fresh water. Agri.-Hort. Gardens ponds, Madras (!) Presi-
dency College garden pond, Madras (!) Masingudi brook, Ootacamund,
Nilgiris, leg., T. Ekambaram. Vaiyampatti stream, near Trichinopoly, leg.,
M. O. P. lyeiigar. Kodaikanal stream, Pulneys, leg., K. S. Srinivasan.
Hustedt gives the range of the length of the diatom as 30-100 p, but
states that most of the individuals are on the smaller side. The South
Indian forms range from 26 3-53 - 4 p. The minimum length recorded for
the South Indian form, viz., 26-3, is slightly lower than the minimum
recorded previously for the type.
74. Cymbella Hustedtii Krasske
(Fig. 123)
Hustedt, Fr., Pascher's Suss wasser- Flora, Heft 10, 1930, p. 363, fig. 674.
Valves asymmetrical, elliptic lanceolate with convex sides and rounded
ends. Raphe less eccentric. Axial area narrow, not widened in the middle.
Striations somewhat coarse and slightly radial.
344 G a Venkataraman
Dimensions.
Length 1.8- 6-20 /,
Breadth . . 7 P
Striae .. 12-13 in 10 /z
Habitat. Fresh water stream, Vaiyampatti, near Trichinopoly, leg.,
M. O. P. lyengar. Red Hills Lake, Madras, leg., S. V. Ganapati.
This form agrees with the dimensions and figure given by Hitstedt.
75. Cymbella cistula (Hemp.) Gruii. var.
maculata (Kiitz.) Van Hetirck
(Fig. 136)
Van Heurck, Trade' des Diatomees, 1899, p. 1 17, pi. 1, fig. 11 ; Hustocll, Fr.,
Pascher's Sussicasser-Flora, Heft 10, 1930, p. 303, fig. 676 6.
Frustules asymmetrical, boat-shaped. Valves with ventral gibbous
margin and truncate rounded ends. Terminal fissures reflexed. Striations
radial, punctate. No isolated puncta on the ventral side.
Dimensions.
Length . . . . . . 74-140 //,
Breadth . . . . .. 18-31 ju
Stride . . . . . . 7-9 in 10 //,
Puncte . . . . . . 18-22 in 10 p
Habitat. Fresh water stream, Vaiyampatti, near Trichinopoly, leg.,
M. O. P. lyengar.
This form agrees with the type quite well.
76. Cymbella naviculiformis Auerswald
(Fig. 119)
Cleve, K. Si\ VeL-Akad. Handl., part 1, Bd. 26, No. 2, 1894, p. 166 ; Boycr, Ryu.
N. Am. DiaL, 1927, p. 281 ; Hustedt, Fr.. Pascher's Susswasser-Flora, Heft 10, 1930,
p. 356, fig. 653.
Valves elliptic lanceolate with rostrate capitate ends. Raphe less
eccentric. Axial area narrow, linear, suddenly dilated in the middle.
Striations radial, closer towards the ends, punctate, puncta very line.
Dimensions.
Length . . . . . . 30 /x
Breadth .. .. . . 10/A
Stride in the middle . . . . 12-14 in 10 p
Stride at the ends .. .. lSinl.0/^
A Systematic Account of Some South Indian Diatoms 345
Habitat. Fresh water stream, Bearshola, Kodaikanal, Pulneys, leg.,
K. S. Sritdvasan.
This form agrees with the type quite well.
Genus Gomphonema Agardh, 1824
77. Gomphonema sphtzrophomm Ehr. forma subcapitata f. nov.
(Figs. 126, 127, 128)
Valves elliptic, club-shaped with capitate round apex and base. The
capitate base is either smaller, equal or bigger than the capitate apex. The
constriction at the apex is only slight. Axial area narrow, linear, central
area small, unilateral with one stigma. Striations slightly radial.
Dimensions.
Length . . . . . . 25-38 ju,
Breadth .. .. .. 6-8-5^
Stride .. .. .. 13-15 in 10 /x
Habitat. Fresh water. Presidency College pond, Madras, leg., Miss
John. Fresh water stream, Masingudi, Ootacamund, Nilgiris, leg., T.
Ekambaram.
The form agrees in most respects with the type but differs from it in the
nature of its apical portion which is not so capitate as in the type. The
constriction below the capitate portion is not so very pronounced as in
Hustedt's figure (Hustedt, Fr., Pascher's Susswasser-Flora, Heft 10, 1930,
p. 372, fig. 695). Again in shape the diatom varies from thin and long to
short and stout. In all other respects it agrees with the type.
78. Gomphonema parvulum (Kutz.) Grun.
(Fig. 121)
Cleve, K. Sv. Vet.-Alcad. HandL, part 1, Bd. 26, No. 2, 1894, p. 180 ; Van Heurck.,
Traite' des Diatomees, 1899, p. 272, pi. 7, fig. 306 ; Schonfeldt, Pascher's Susswasser
Flora, Heft 10, 1913, p. 124, fig. 270 ; Boyer, Syn. N. Am. Diat., 1927, p. 294 ; Petersen,
J. B. 9 Botany of Iceland, Vol. II, part 2, 1928, p. 411 ; Hustedt, Fr., Pasclier's Suss-
wasser- Flora, Heft 10, 1930, p. 372, fig. 713 a.
Valves lanceolate, club-shaped gradually tapering from the middle to
the ends. Apex rostrate. Axial area very narrow, central area Unilateral
with an isolated stigma. Striations radial, very finely punctate.
Dimensions.
Length . . . . . . 24-32 p
Breadth . . . . . . 5-7/1.
Striae . , . , . . . 14-16 in 10 p.
si
fe i
125
116
126
127
130
A
119
120
124
128
132
133
A Systematic Account of Some South Indian Diatoms 347
Habitat. Fresh water. Vaiyampatti stream near Trichinopoly, leg.,
M. O. P. lyengar.
This form agrees with the type quite well.
79. Gomphonema parvulum (Kiitz.) Grttn.
var. subelliptica Cleve
(Fig. 122)
Cleve, 1C. Sv. VeL-Akad. Handl., part 1, Bd. 26, No. 2, 1894, p. 180 ; Hustedt. Fr.,
Pascher's Suawoasser-Flora, Heft 10, 1930, p. 373, fig. 713 b.
Frastules more symmetrically elliptic lanceolate with scarcely rostrate
apex. Smaller than the species.
Dimensions.
Length . . . . . . 14 /*
Breadth . . . . . . 5 /x,
Striae . . . . . . U in 10 /*
Habitat. Fresh water. Vaiyampatti stream near Trichinopoly, leg.,
M. O. P. lyengar.
It was a very rare form in the collection. The specimens agree with
the type,
80. Gomphonema intricatum (Kiitz.) var.
vibrio (Ehr.) Cleve
(Fig. 116)
Cleve, K. Sv. Vet.-Akad. HandL, parb 1, Bd. 20, No. 2, 1894, p. 182 ; Schdnfeldt,
Pascher's Susswasser-Plora<> Heft 10, 1913, p. 122 ; Hustedt, Fr., Pascher's Susswasaer-
Flora, Heft 10, 1930, p. 376, fig. 698.
FIG. 115. Rhopalodia gibbet (Ehr.) O. Mull. X 1000.
FIG. 116. Gomphonema intricatum (Kiitz.) var. vibrio (Ehr.) Cleve. x 700.
FIG. 117. GomphonemalanceolatumJ&hv. x 1100.
FIG. 118. Gomphonema lanceolatum Ehr. var. insignis (Greg.) Cleve. x 1100.
FIG. 119. Cymbella naviculiformis Auerswald. X 1600.
FIG. 120. Rhopalodia gibberula (Ehr.) O. Mull. var. protracta Gran. X 1100.
FIG. 121. Gomphonema parvulum (Kiitz.) Grun. X 1500.
FIG. 122. Gomphonema parvulum (Kiitz.) Grun. var. subelliptica Cleve. x 1500.
FIG. 123. Cymbella Hustedtii Krasske. X 1100.
FIG. 124. Cymbella alpina Grun. X 1100.
FIG. 125. Cymbella turgida- (Greg.) Cleve. X 1100.
FIGS. 126-28. Gomphonema sphcerophorum Ehr. f . subcapitata f . nov. x 1000.
FIG. 129. Rhopalodia gibba (Ehr.) O. Mull. var. ventricosa (Ehr.) Grun. x 1000.
FIG. 130. Rhopalodia gibberula (Ehr.) O. Mull. Showing rows of seriolse between
two ribs. X 1600.
FIG. 131. Epithemia zebra (Ehr.) Kiitz. var. saxonica (Ktitz.) Grun. X 1100.
FIGS. 132-33. Nitrschia closterium (Ehr.) W. Smith. Showing chloroplasts. X 700.
FIG. 134. Gomphonema laceolaium Ehr. x 1100.
FIG. 135. Rhopalodia gibberula (Ehr.) O. Miill. X 1100.
FIG. 136. Cymbella cistula (Hemp.) Grun. var. maculata (Kiitz.) Van Heurck. x 1100 ?
348 ^ Venkataraman
Valves very slender, narrow and linear, slightly gibbous in the middle
with rounded truncate ends. Axial area fairly broad. Central area broad,
unilateral with an isolated stigma. Striations slightly radial, clearly
punctate.
Dimensions.
Length .. .. .. 100 /*
Breadth . . . . . . 10 /*
Striae .. . . . . 10 in 10 /x
Habitat. Fresh water. Pond inside Agri.-Hort. Gardens, Madras (!)
This form was found as a rare one in the collection. It agrees very well
with the description given by Cleve and Hustedt and with the figure given
by Hustedt.
81. Gomphonema lanceolatiwn Khr.
(Figs. 117, 1.34)
Cieve, K. Sv. Vet.-Akad. Hcmdl., part 1, Bd. 26, No. 2, 1804, p. 183 ; Schdnfoldt,
Pascher's Silsswasser-Flora, Heft 10, 1913, p. 123, fig. 268 ; Boyer, y/n. A 7 . Am. Diat>,
1927, p.2Q5 : Hustedt, Fi\, Pascher's Sus$wasser-Flora t Heft 10, 1930, p. 376, fig. 700.
Valves club-shaped to lanceolate club-shaped with obtuse apex and
somewhat tapering basis. Axial area broad. Central area small, unilateral
with an isolated stigma. Striations coarse, slightly radial, punctate, pttncta
big.
Dimensions.
Length . . . . . . 40-54 p
Breadth .. .. ... 9-1 0^
Striae . . .. . . 12-13 in 10 /x
Habitat. Fresh, water. Ponds in the Agri.-Hort. Gardens, Madras (!)
This form agrees with the type.
82. Gomphonema lanceolatum Ehr. var.
insignis (Greg.) Cleve
(Fig. 118)
Cleve, K. Sv. Vet.-Akad. Handl., part 1, Bd. 26, No. 2, 1894, p. 188 ; Boyer, Syn.
N. Am. Diat., 1927, p. 295 ; Hustedt, Fr., Pascher's Susswasser- Flora, Heft 10, 1930,
p. 876, fig. 700.
This is distinguished from the species through the slightly distant strise,
8-10 in 10 p,. The apex is slightly more tapering than the type.
Habitat. Fresh water. Ponds in the Agri.-Hort. Gardens, Madras (!)
This form agrees very well with the description and figure given by
Hustedt.
A Systematic Account of Some South Indian Diatoms 349
(2) Family
(a) Subfamily Epithemioidese
Genus Epithemia Brebisson, 1838
83. Epithemia zebra (Ehr.) Kiitz. var. saxonica (Kiitz.) Grun.
(Fig. 131)
Schonfeldt, Pascher's Siisswasser- Flora, Heft 10, 1913, p. 146 ; Hustedt, Fr.,
Pn-scher's Sussioasser-Flora, Heft 10, 1930, p. 385, fig. 730.
Valves arcuate with rounded and slightly produced ends. Dorsal side
rather highly convex and the ventral side slightly concave. Costse robust,
alternating with more than two rows of aeriolse.
Dimensions.
length . . . . . . 44-46 ft
Breadth . . .. .. 9-5-10^
Coste .. .. .. 2-4 in 10 ^
Rows of aerioles . . . . 1.2 in 10 ft
Habitat. Fresh water. On decaying fern leaves in a stream in
Bearshola, Kodaikanal, Pulneys, leg., K. S. Srinivasan.
It was a very rare form in the collection. This specimen agrees with
the type.
(b) Subfamily Rhopalodioidese
Genus Rhopalodia O. Muller, 1895
84. Rhopalodia. gibba (Ehr.) O. Mull.
(Fig. 115)
O. Muller, Engler's Botan. Jahrbucher, Bd. 22, 1895, p. 05, Taf. 1, figs. 15-17;
Scbonfeldt, Pascher's Susswasser-Flora, Heft 10, 1913, p. 147, fig. 324 ; Boyer, Syn.
N. Am. Diat., 1927, p. 491 ; Hustedt, Fr., Paschec's Susswasser-Flora, Heft 10, 1930,
p. 390, figs. 74.0, 740 a.
Frustules in girdle view linear, gibbous in the middle, broad, gradually
slightly tapering to the ends. Valves linear, arcuate on the dorsal, straight
011 the ventral side, reflexed at the ends. Costse strong, two to three aeriole
rows between two costse.
Dimensions.
length .. . . .. 83-104 p
Breadth . . . . . . 21-25 p
Costse , . , , , . 6-8 in 10 p.
350 G. Venkataraman
Habitat. Fresh water brook, Masingudi, Ootacamund, Nilgiris, leg.,
T. Ekambaram. Red Hills Lake, Madras, leg,, S. V. Ganapati.
The rows of aeriolse were not observed even under a very high magnifica-
tion. The specimens agree with the type.
85. Rhopalodia gibba (Ehr.) O. Mull,
var. v&ntricosa (Ehr.) Grun.
(Fig. 129)
Hustedt, Fr., Pascher's Sussivasser- Flora, Heft. 10, 1930, p. 391, fig. 741.
This variety is distinguished from the species by its strongly gibbous
nature in the middle, tapering strongly towards the ends, thus making the
frustule compact.
Dimensions.
Length . . . . 66 /x
Breadth .. .. . . 21 p.
Costae . . . . . . 6-8 in 10 ft
Habitat. Fresh water brook, Masingudi, Ootacamund, Nilgiris, leg.,
T. Ekambaram.
This form was found only as a stray specimen among the species.
Hustedt (op', cit., p. 391) has observed such an occurrence. The few speci-
mens that were noticed are similar to the figure given here. The tapering
from the middle to the ends is intermediate between the species gibba
(Ehr.) O. Mull, and the var. ventricosa (Ehr.) O. Mull.
86. Rhopalodia gibberula (Ehr.) O. Mull.
(Figs. 130, 135)
Schonfeldt, Pascher's Susswasser- Flora, Ileffc 10, 1913, p. 118, fig. 320 ; Hustedt.,
Fr., Pascher's Susswasser-Flora, Heft 10, 1930, p. 391, fig. 742.
Frustules in girdle view long, elliptical with rounded ends. Dorsal
side highly convex ; ventral side more or less straight in the middle portion.
The aeriole rows distinctly seen between two costse.
Dimensions.
Length . . . . . . 35-42 ^
Breadth . . . . . . 20-24- p
Coste .. .. .. 3-4 in 10 /*
Rows of aerioles between two costse 2-8
Habitat. Brackish water. Adyar estuary, Madras (!)
This form agrees with the figure and description given by Hustedt,
A Systematic Account of Some South Indian Diatoms 351
87. Rhopalodia gibberula (Ehr.) O. Miill.
var. protmcta Grun.
(Fig. 120)
Hustcdt, Fr., Pascher's Susswas&er-Flora, Heft 10, 1930, p. 391, fig. 743.
This slightly differs from the species in having a slight depression on the
dorsal margin near the ends. Ends taper and are pointed.
Dimensions.
Length .. .. .. 53-4/1
Costse . . . . . . 3-4 in 10 ju,
Rows of aerioles between two costse 2-8
Habitat. Brackish water. Adyar estuay, Madras (!)
This form was found along with other brackish water diatoms on niud
inside water as a greenish film. It agrees with the type.
(3) Family NITZSCHIACE.E
Subfamily Nitzschioideae
Genus Hantzschia Grun. 1.880
88. Hantzschia amphioxys (Ehr.) Grun. var. vivax (Hantz.) Grun.
(Fig. 148)
Van Heurck, Traite' des Diatomees, 1899, p. 381* pi. 15, fig. 486 b ; Schonfeldt,
Pascher's Susswasaer- Flora, Heft 10, 1913, p. 161 ; Boyer, Syn. N. Am. Dial., 1927,
p. 528 ; Hustedt, Fr., Pascher's Susswasser-Flora, Heft 10, 1930, p. 394, fig. 750.
Valves linear, slender and with longer ends than the species. Keel
puucta short, the median distant.
Dimensions.
Length . . . . . . 61-75 p
Breadth .. . . .. 6-5-8jx
Keel puucta .. . . . . 6-8inlO/i
Striae .. -. . . 13-18 in 10 JLL
Habitat. Fresh water. Pools near Elliot Beach, Madras, leg., M. O. P.
lyengar. An irrigation tank at Vandalur, Madras, leg., K. G. Veeraraghavaii.
This is a rare form in the collections. Only a few stray specimens were
found. This form agrees with var. vivax in all respects.
Genus Bacillaria Gmelin, 1788
89. Bacillaria paradoxa Gmelin
(Figs. 144, 145)
De Toni, SylL Aly., Vol. II, part 1, 1891, p. 493 ; Schonfeldt, Pascher's Susswasser-
Flora, Heft lo! 1913, p. 149 ? fig. 328 ; Karsten, G-., Bngler and Prantl's Nat. Pflazen-
favnilien, Bd/2, 1928, p. 294, figs. 399, 100, 190. Lebcmr, M. V., Planktonic
352 G. Venkatarainan
Diatoms of Northern Seas, 1930, p. 211. fig. 175 ; Hustcdt, Fr., Pascher's Sussicawcr-
Flora, Heft 10, 1930, p. ?>96 ? fig, 755 a. 1).
Frustules in girdle view linear rectangular united in bundles to form
plates. Valves linear, spindle-shaped. Carinal dots form a row in the
middle portion of the valve. The striee are very fine.
D im en sions.
Length .. .. .. 80-105 /*
Breadth . . . . . . 5-6 /i
Carinal dots . . . . . . 6-8 in 1.0 //,
Strise .. .. .. 20-25 in 10 /z
Habitat. Brackish water. Adyar estuary, Madras (!)
This form occurred among other diatoms on the surface of the moist soil
as a brownish scum. They were showing the characteristic gliding move-
ments when collected.
Genus Nitzschia Hassal, 1845
Section Tryblionella (W. Smith, Grtinow) Hustedt
90. Nitzschia tryblionella Hantzsch var. levidensis (W. Smith) Grim.
(Fig. Ul)
Van Heurck, Traite' des Diatomees, 1899, p. 385, pi. 15, fig. 494 ; Rchonfeldt,
Pascher's Siissivasser-Flora, Heffc 10, 1913, p. 151 ; Boyer, Syn. N. Am. Diat., 1927,
p. 495; Hustedt, Fr., Pascher's Susswasser -Flora, Heffc 10, 1930, p. 399, fig. 700.
Valves linear with slightly concave margins. Ends wedge-shaped atid
rounded. A clear longitudinal fold in the middle of the valve is present.
Keel very eccentric. Striae clear.
Dimensions.
I^ength . . . . 40 fjL
Breadth . . . . . . 9 ju
Keel punctse . . . . . . 12 in 10 /^
Stride .. .. .. 1 1-1.3 in 10 /x
Habitat. Fresh water stream, Vaiyampatti, near Trichinopoly, leg.,
M. O. P. lyengar. Fresh water pond, Presidency College Garden, Madras (!)
This is a rare form in both the collections. The ends are a bit more
produced than in the var. levidensis as figured by Hustedt.
Section Linear es (Grun.) Hust.
91. Nitzschia vitrea Norman
(Fig. 143)
Van Heurck, Traite' des Diatomees, 1899, p. 399, pi. 16, fig. 544 ; Boyer, Syn. N.
Am. DiaL, 1927, p. 519; Hiistedt, Fr., Pascher's Susswasser-.Flora, Heft- 10, 1030,
p. 411, fig. 787,
A Systematic Account of Some South Indian Diatoms 353
Frustules in girdle view linear, rectangular, with parallel sides and
truncate rounded ends. Valves narrow, with parallel margins. Keel puncta
somewhat rounded. Striations fine but distinct.
Dim ensions.
Length . . . . . . 76-101 ^
Breadth ... .. . . 7- 6-1 1ft
Keel puncte . . . . . . 5-7 in 10 /x
Stride . . . . . . 20-22 in 10 p
Habitat. Brackish water. Plankton of river Cootlm, Madras (!)
This was only a stray form in the plankton. It agrees with the type
quite well.
Section Lanceolate Grun.
92. Nitzschia amphibia Grun.
(Fig. 149)
Van lleurck, Trailc"* ties Diatomees, 1899, p. 10o, pi. 17, fig. 5Go ; Sclionleldl,
Pascher's Susswasser-Flora. Heft 10, 1913, p. 158, fig. 349 ; Boyer, ?/??. A r . Am. Biat.,
1927, p. 521 ; Hustedt, Fr., Pascher's Susswasser-Flora, Heft 10, 1930, p. 414, fig. 793.
Valves linear to linear lanceolate with the ends slightly produced some-
times and rounded. Striations coarse and punctate, puncta big.
Dimensions.
Length .. .. .. H-30ju,
Breadth . . . . . . 4-5 p
Keel puncte . . . . . . 6-8 in 10 /x
Stride .. .. .. 14-17 in 10 /x
Habitat. -Fresh water. Common and widely distributed. Ponds, Agri-.
Hort. Garden, Madras (!) Adyar fresh water pond, Madras (!) Roadside
ditch, Guindy, Madras (!) Stream, Vaiyampatti, near Trichinopoly, leg.,
M. 0. P. lyengar. On the moist wall of a house, Triplicane, Madras, leg.,
M. O. P. lyengar, wherein only this species was found as a pure species in
abundance.
This form agrees very well with the type.
93. Nitzschia palea (Kiitz.) W. Smith
(Fig. 146)
Smith, W., Syn. Brit. Diat,, Vol. II, 1853, p. 89 ; Van Henvck, Traite' des Dialo-
mees, 1899, p. 40.1, pi. 17, fig. 554 ; Schonfeldt, Pascher's Susswasser-Flora, Heft 10,
1918, p. 159, iig. 352 ; Boyer, Syn. N. Am. Died., 1927, p. 521 ; Hn.stedt, Fr., Pascher's
Su8*wa88er-I*lora, Ilcft 10, 1930, p. 416, fig. 801.
334
G. Venkataraman
144
145
m
V
141
142
147
Vi/
149
FIG. 137. Surirella tenera Greg. . x TOO.
FIG. 138. Surirella elegans Ehr. Grirdle view, x 300.
FIG. 139. Surirella ovalis Breb. x 1000.
FIG. 140. Surirella elegans Ehr. Valve view, x 350.
FIG. Ul.NitzscJiia tryWonella Hantz. var. levidensis (W. Smith) Grun. x 800.
A Systematic Account of Some South Indian Diatoms 355
Valves linear to linear lanceolate with short wedge-shaped tapering ends,
Striations very delicate.
Dimensions.
Length .. .. .. 40-54^
Breadth .. .. .. 3-4 -5 /*
Keel punctse . . . . . . 10-15 in 10 p
Strise . . . . about 40 in 10 y,
Habitat. Fresh and brackish water. This is a common form
found in pools, ponds, streams, ditches, moist soil, etc. This forms the
plankton of the river Cooum, during the North-East Monsoon season. It
occurs in abundance on the moist soil of the beach during this season.
It agrees well with the type.
Section Obtusce (Grun.) Hust.
94. Nitzschia obtusa W. Smith var.
scalpelliformis Grun.
(Figs. 142, 147)
Van Heurck, Traite* des Diatom ees, 1899, p. 397, pi. 16, fig. 538 ; Boyer, Syn. N.
Am. Diat., 1927, p. 517 ; Hustedt, Fr., Pascher's Susswasser-Flora, Heft 10, 1930,
p. 422, fig. 817 d.
Prustules broad, linear with ends obliquely truncate. Keel fairly large,
inilexed in the middle region. The two median keel puncta distant. Stria-
tions punctate, punctae fine and linear.
Dimensions.
Length .. .. .. 89-1 12 /A
Breadth .. .. .. 10-12 p,
Keel punctai . . . . . . 6-8 ii) 10 ^
vStriac .. .. .. 26-27 in 10 /x
Habitat. Brackish water. River mouth, Cooum, Madras (!) Adyar
estuary, Madras (!)
FIG. 142. Nitzschia ohtusa W. Smith var. scalpelliformis Grim. Showing the carinal
clots and the transverse punctate strite. x 1600.
FIG. 143. Nitzschia vitrea Norman. Girdle view, x 800.
FIG. 141. Baelllaria paradoxa Gmeliii. Cells in a band, x 350.
FIG. 145. Bac.illaria paradoxa Gmeliu. Valve view, x 700.
FIG . 1 4 ( > . Nitzschia pale a ( Ktitz . ) VV . Smith . x 1 1 00 .
FIG. 147. Nitzschia ohtusa W. Smith var. scalpelliformis Grun. x 700.
FIG. 1.48.- Hantzsch-ia aniphioxijs (Ehr.) Grun. var. vivax (Hantzscb) Grim. X 800.
FIG. 1-19.- Nilzschid ampMW-a Gv\m. x 1500.
356 G* Venkataraman
This form is found in the littoral region and also in the plankton of the
river Cootim. It agrees with the type. The striations when seen under
very high magnification are composed of fine linear punctee.
Section Nitzschiella (Rabh.) Grun.
95. Nitzschia dosteriwn (Ehr.) W. Smith
(Figs. 132, 133)
Smith, \V. 5 Si/n. Brit. JJiat., Vol. I, 1853, p. 12, pi. XV, fig. 120 ; Gran, IT. 11.,
NorcUsc/ies Plankton, Sotanischer Tell B. VIII, 19, 1908, p. 129 %. 172 ; Boyer, tit/n.
N. Am. Diat., 1927, p. 520 ; Lebour, M. V., The Planldonic Diatoms of Northern, ftcua,
1930, p. 212. fig. 176 ; Hustedt, Fr., Pasclior's Sussica-sser-Flora, Ueffc 10, 1030, p. 42 1,
fig. 822.
Valves spindle-shaped in the middle. The ends extended into long
beaks usually slightly bent in a semi-circle or sometimes curved in opposite
directions. Striations not recognizable.
Dimensions.
Length .. .. .. 7 0-1 K> 5 /x
Breadth .. .. .. 3 -5-6 /A
Habitat. Brackish water. Plankton of the river Cooum, Madras.
This was almost constantly present in the plankton throughout the year.
The maximum development was during the month of January when they
were found as 52,000 per c.c. of the water, colouring the water yellowish
brown. Kolbe (Planzenforschung, Heft 7, 1927, p. 103) remarks about the
occurrence of this in abundance during spring.
This form agrees with the type.
(4) Family vS
(a) Subfamily Surirelloideae
Genus Sunrella Turpin, 1828
90. Surirella tenera Gregory
(Fig. 137)
Gregory, Quart. Jour. Mlcr, 8ci. 9 Vol. IV, 1850, p. 11. ; J>oyor, S f ///<.. A r . ,l?w. Dial,.,
1927, p.*540 ; Hustedt, Fr., Pascher's Susswasser-Flora, Heft 10, 1930, p. -I.'JS, fig. S5U.
Valves narrowl} 7 elliptical, rounded at one end and acute at the other.
Central space narrow. Costae rather indistinct and faint.
Dimensions.
Ivength .< .. .. 110-140 /A
Breadth .. .. .. 31-37 /u
Coste .. .. .. 25-40 in 1 00 x
A Systematic Account of Some South Indian Diatoms 357
Habitat. Fresh water stream, Bearshola, Kodaikanal, Pulneys, leg.,
K. S. Srinivasan.
This form agrees with the description and figures given by Gregory and
Hustedt.
97. Surirella elegans Ehr.
(Figs. 138, UO)
Schonfeldt, Pascher's Susswasscr-Flora, Heft 10, 1913, p. 167, fig. 372 ; Boye.r,
Syn. N. Am. Dial., 1927, p. 537 ; Hustedt, Fr., Pascher's Susswasser-Flora, Heft 10,
1930, p. 440, figs. 858, 859.
Frttstules in girdle view wedge-shaped. Valves narrowly or broadly
ovate, rounded at one end and acute ai; the other end. Central space narrow,
lanceolate. Costse broad.
Dimensions.
Length .. .. .. 146-200^
Breadth . . , . . . 52-70 /*
Costae .. .. .. 1 4-16 in 100 /*
Habitat. Fresh and slightly brackish water. Common. Fresh water
ponds, Agri.-Hort. Gardens, Madras (!) Stream, Vaiyampatti, near
Trichinopoly, leg., M. O. P. lyengar. An irrigation tank at Vandalur,
Madras, leg., K. G. Veer aragha van. River Cooum, from the upper
portion (!)
This fresh water form was also able to grow in portions of the river
where the salinity of the water ranged upto 0-075%. Kolbe groups
this under " indiffernt forms ".
98. Surirella ovalis Brebisson
(Fig- 139)
Smith, W., Syn. Brit. Dial., Vol. I, 1853, p. 33, pi. IX, fig. 68 ; Schonfeldt,
Pascher's Susswasscr-Flora, Heft 10, 1913, p. 170, %. 371. ; Beyer, Syn. N. Am. Diat.,
1927, p. 541 ; Hustedt, Fr., Pascher's Susswasser-Flora, Heft 10, 1930, p. 441, figs.
360, 861.
Valves ovate, acuminate, costse short, marginal, radial and unequal.
Central space rather indistinct. Striations radial.
Dimensions.
Length . . . . . . 42-80 p.
Breadth . . . . . . 28-43 ^
Costse . . . . . . 44-48 in 10 \L
Stride , . T , . 15 in 10 \L
358
G. Venkataraman
Habitat. Fresh water. Brook, Masingudi, Nilgiris, leg., T. Ekambaram.
This form agrees with the type. The inner limit of the costae are clearer
tinder a higher focus.
The author wishes to express his great indebtedness to Prof. M. O. P.
lyengar, M.A., Ph.D. (Lond.), FX.S., for his constant help and guidance during
the whole course of this investigation and in the preparation of this paper.
My sincere thanks are also due to the authorities of the University of Madras
for the award of a Research Scholarship during the tenure of which the
present investigation was carried out.
Abdul Majeed, M.
Bethge, H.
Biswas, K.
Boyer, C. S.
Bristol, B. M.
Carter, N.
Cleve, P. T.
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Pflanzenforschung, Heft 3, Jena, 1925.
" Notes on the organisms in the Filtered water of
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Series, 1930, 26, No. 4.
. " Census of Indian Algae. Scope of Algological studies
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cule 2, 197-219.
" Association of some of the common algno with animals
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" A comparative study of the algal flora of two
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, " Synopsis of the Naviculoid Diatoms," A". Sv. Vet.-
Akad. HandL, 1894, Part 1, Bd. 26, No. 2 ; Part 2,
1895, Bd. 27, No. 3,
A Systematic Account of Some South Indian Diatoms
359
Be-Toni, J. B.
Dickie, George
Fortmorel, G. L.
Fritsch, F. E.
llich, F.
Gopala Aiyar, "K., Sankara
Menon, K., and Mcnon,
M. G. K.
Gran, H. II.
Gregory, W .
G i'u now, A.
Hustodt, Fr.
Karsten, G.
Kolbe, H. W.
, and Tiegs, B.
Kutzing, F. T.
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S. Afr. Mus., 3918, 9, Part VII, 183-011.
" Contribution to our knowledge of the fresh water
algse of Africa. 4. Fresh water and subaerial algse
from Natal," Trans. Roy. Soc. S. Africa, 1925, 11,
297-398.
" Plankton Records for the years 1929 and 1930,"
Journ.. Madras Univ., S s No. 1, 97-139.
" Diatomeen, Nordisches Plankton von Brandt und
Apstein," Botanischer Tell, B. 8, 19, Keil und
Leipzig, 1908.
" Notice of some new species of British fresh water
Dlatomacece," Quart. Journ,. Micr. 8ci. 9 1S5G, 4, 1-14,
" Siisswasser-Diatoniacceu und Deaniidiaccen von der
Insel Baiika," Rabh. Beitr. Zur. Kenn. u. Verb.
Algcn, 1805, Heft II.
Bacillariopyhta (Diatomcaj) in A. Pascher's Die
Sussioasser-Flora, Miltclcuropas, Jena, 1930.
Die Kieselalgen in Dr. L. Ilabenhorst's Kryptogamen-
Flnra von Deutschlands, Osterreichs und der
Schweiz, 1930, 1931, 1932, Bd. 7, Teil 1, Teil 2,
Lief. 1-4- .
Bacillariophyta (Diatomcse) in Engler and Prantls'
Die Naturlic'hcn, PJlanzenfamilien, Leipzig, 1928,
Bd. 2.
"Zur Okologie, Morphologic und Systematik der Brack-
wasHcr-Diatomeeix. Die Ivieselalgen des Speren-
borgor Salxgebiets," PJlanzenforschung, 1927, Heft 7,
1-14:3.
" Zur mosohaloben Diatomeen-Flora des Werrage-
bictes," Ber. B. Deut. ~Bot. Gest., 1929, Bd. 47,
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Species Algarum, Lipsicr., 1849.
360
Labour, Marie, V.
Leuduger- Fort-morel, G-. M. D.
MtLller, Otto
Petersen, J. B.
Prain, D.
Rabenhorst, L.
Rich, Florence
Bchaarschrnidt, J.
Schonf eldt
Skvortzow, B. W.
Smith, W.
Van Heurck
Wallich, G-. C.
West, W., and West, G-. S.
Zeller, GL
G. Venkataraman
The plankstonic Diatoms of Northern Seas, Bay Society,
London, 1930.
From De Toni, Sylloge Air/arum. 1879.
Rhopalodia, ein neues Genus dec Bacillariaceen,''
'Bugler's Botanische Jahrbucher, 1895, Bd. 21, Heft
4, 54-71.
" The aerial al^ffi of Iceland," The Botany of Iceland.
1928, 2, Part II, No. 8, 325-447.
" The vegetation of the districts of Hughli-Howrah and
the 24 Pergunnahs," Rec. Bot. Surv. India, 1905,
3, No. 2.
Flora Europcea Alyarum Lipsicc, 1864.
'* Contributions to our knowledge of the fresh water
algae oi Africa. 12. Some diatoms from the
Victoria Falls," Travis. Roy. Soc. S. Africa, 1930 ?
24, Part III, 207-20.
" Notes on Afghanistan Algae," Jour. Linn. Soc.,
London, 1830, 21.
Eacillariales (Diatomese) in A. Pasclier's Die flti.s-
swasser-Flora Deutschlands, Oster-reichs und der
Schweiz, 1913. Heft 10.
t4 Diatoms from. Calcutta, India," Philippine Journ.
Sri., 1935, 58, No. 2, 179-92.
, " Notes on Ceylon Diatoms 1," Ann. Roy. Bot.
Gard., Ceylon, 1930, 11, Part III, 251-00.
. A synopsis of the British Diatornacea?, Vols. I aixd IE,
London, 1853-50.
. Traite' des Diatomees, Anvers, 1899.
. " On the >Siliceous organisms found in the digestive
cavities of Salpae," Trans. Micro. Soc., Neiv Series,
1860, 8.
. " A Contribution to the Fresh water Alga3 of Ceylon,"
Trans. Linn. Soc. Lond., 1902, 6, Ser. 2, Part III.
. " Fresh water algae from Burma, including a few
from. Bengal and Madras," Ann. Roy. Bot. Gard.,
Calcutta, 1907, 6, part II.
. " Algse collected by Mr. S. Kurz in Arracan and British
Burma," Journ. Asiatic Soc. Bengal, 1873, 13
part II.
A Systematic Account of Some South Indian Diatoms 361
EXPLANATION OF MICROPHOTO GRAPHS
PLATE XVII
FIG. 1. AcTinanthes coarctata Breb. var. parallella var. nov. Showing their epiphytic
nature. Cells are united both by their valves and by their ends. X 180.
FIG. 2. Coscinodiscus Granii Gough. Valve showing the seriolations. X 130.
FIG. 3. Diploneis subovalis Cleve. Showing the double rows of alveoli. x 2000.
FIG. 4. Diploneis subovalis Cleve. The same frustule (as in fig. 3) taken at a lower
focus. Showing the prominent costae with their capitate ends almost
touching one another. X 2000.
362
G. Venkataraman
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Systematic Account of Some. South Indian Diatoms 363
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364
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FIG. i
FIG. 3
FIG. -t
STUDY IN DISEASES OF FISH: PIN-ROT
A BACTERIAL DISEASE OF FINS OF FISH
BY DR. HAMID KHAN, M.Sc., I^L.B. (PUNJAB), PH.D. (CANTAB.)
(Fisheries Research Officer, Punjab, Lyallpur)
Received October 13, 1939
(Communicated by Dr. B. L. Bhatia, D.SC., F.N.I.)
IN November 1938 heavy mortality occurred among Gourami fiiigerlings
(Osphormenus goummy Lacepede) when these were being transported by
canal from Madras Fisheries Farm, Sunkesula to Kurnool Town a distance
of 17 miles in a "live-car". 1 The fingerlings, before despatch, had been
" conditioned" for a day or two, i.e., had been confined in a " conditioning-
car" 2 a wooden framed box with wire netting on all sides kept in water
and starved.
In all the dead fry it was observed that the caudal fin was frayed, and
observations made on the dying fingerlings, en route from Kurnool Town
to Delhi, showed that the epidermis from the caudal fin was gradually being
eaten off and the caudal rays were consequently getting exposed. In some
cases the dorsal and anal fins had also been affected (Plate XVIII, Figs. 1-4).
The sick fry swam near the surface of the water in the container, lost their
balance on the caudal fin becoming frayed upto its base, turned upside down
and died, the dead ones sinking to the bottom.
The author had noticed similar symptoms among Rainbow trout fry
when they were attacked by fin-rot. 3 In order to investigate the matter
in detail, freshly dead Gourami fry were taken to the Imperial Agricultural
Institute, New Delhi, and examined by the Assistant Bacteriologist. His
preparation from affected caudal fin (Plate XVIII, Fig. 5) showed exactly
similar rod-shaped bacteria as had previously been observed by the author in
Rainbow trout suffering from an attack of fin-rot.
1 " Live-car " is a wooden barrel, conical at one end and round at the other, with
a small opening at one side. It is made to float in water and has been in xise for trans,
port of alive fish by Madras Fisheries Department for a number of years.
2 " Conditioning-car " is similar to " live-car " used in the Punjab for keeping
fry under observation. Its photograph was given by the author in Journ. Bomb.
Nat. Hist. Soc., 1924, 29, No. 4.
3 Hamid Khan, " Notes on the diseases of Trout at the Mahili Hatchery Kulu
Punjab", ibid., 1939, 40, 4, 653-56.
369
B6 F
370 Hamid Khan
Inoculation of Rainbow Trout Fry
Some dead Gourami fingerlings were brought in water to I.yallpur and
examined by the Agricultural Bacteriologist. v Sapropliytic bacteria^ in
addition to pathological ones, hcd also mcde their appearance by that time.
To obtain pure culture of pathological bacteria, healthy Rainbow tiout fry
measuring fioni 52 mm. to 66 mm. in length, bi ought fiom rmiji b Govern-
ment Fish Farm at Madhopur, were inoculated on the 9th November
1938, at 4-30 p.m. with bacteria from the affected parts of the dead Gourami
as follows:-
(i) The caudal fins of three healthy Rainbow trout fry were scratched
with sterilized needle and rubbed with the caudal fin of the
diseased Gourami fingerling, and
(ii) three Rainbow trout fry, with scratched caudal fins, were simply
placed in contaminated water in which the diseased fish had. been
lying.
Four Rainbow trout fry with scratched caudal fins were kept as control
in uncontaminatLd water. The experiment was conducted in aquaria in
Fisheries Research Laboratory at I/yallpur. Temperature of water ranged
from 62 F.-66 F. and source of water supply was ordinary tap water.
Aeration of water in the aquaria was maintained by Semper J s Aerating
Apparatus.
Progress of the Disease
Rainbow Trout Fry No. 1. On the 10th November, at 8 a.m., fry
No. 1 (Plate XIX, Fig. 2) showed slightly whitish area on the free extremity
of its caudal fin. On the Hth, it was seen swimming near the sin face of
water and its caudal fin was slightly exposed at its extremity. On the 1 2th,
the rays of its caudal fin were almost totally naked and the disease had
spread with great rapidity to its caudal peduncle and anal fins. The caudal
fin rays got disintegrated. and soon after the fry lost its balances and while
it was dying its caudal fin was cut for culture of bacteria. The fry died on
the 12th November at 2-30 p.m., i.e., after about 70 hours after
inoculation and 54 hours after the symptoms of the disease became visible
externally.
Rainbow Trout Fry No. 2 (Plate XIX, Fig. 3). Showed external symptoms
of the disease on the Hth November, at 8 a.m. On the 1.2th the fin
rays of the caudal fin had become exposed and on the 1 3th, the caudal fin, as
well as the caudal peduncle, were seriously affected. The colour of the fry
had become dark and it swam near the surface of the water. The fry died
St^tdy in Diseases of Fish : Fin- Rot 371
at II a.m., on the 13th November, i.e., 90 hours after inoculation and
5.1 hours after the disease had become visible externally.
Rainbow Trout Fry No. 3. In Rainbow trout fry No. 3 (Plate XIX,
Fig. 4), external symptoms were visible on the 12th November and the fish
died early in the morning on the 14th.
Rainbow Trout Fry No. 4, Out #f the second lot, in which the caudal fin
had been scratched but not inoculated and the fry had been put in contami-
nated water, Rainbow trout fry No. 4 (Plate XIX, Fig. 5), showed external
symptoms on the llth November. On the 12th, its caudal fin rays
were slightly exposed and on the 13th it died at 11 a.m.
Rainbow Trout Fry Nos. 5 and 6. Rainbow trout fry Nos. 5 and 6
showed external symptoms of the disease on the 12th November and were
found dying on the 14th. Upper lobe of the caudal fin (Plate XIX, Figs. 6
and 7), had partially disintegrated and caudal peduncle was affected upto the
base of the adipose fin. The infection had spread to the anal fin too. Fry
No. 5 died on the 12th, at 12 a.m., and No. 6 at 3 p.m., on the 14th
November.
Rainbow trout fry kept as control did not show any symptoms of the
disease.
Rainbow Trout Fry No. 7. On the 14th November, when all inocu-
lated fry, as well as those which had been put in contaminated water, had
died, one fry from the control was transferred to the contaminated water
and its caudal fin, without being scratched, was tubbed against the caudal
fin of recently dead Rainbow tiout fry at 12 a.m. On the 1.5th, the disease
appeared on its adipose fin and the tissues at the base of this fin were
seriously affected (Plate XIX, Fig. 8). The fry died on the morning of the
16th November.
Rainbow Trout Fry No. 5. On the 16th November, at 9-30 a.m.,
another fry from the control was inoculated on its caudal fin after it had
been scratched, by rubbing it against the caudal fin of recently dead Rain-
bow trout fry. On the 17th the fry showed disease at the base of the caudal
fin. and the caudal peduncle was seriously affected (Plate XIX, Fig. 9). On
the 1,8th attempts were made to cure the fry by putting it in copper sulphate
solution (1 : 5,000), but the disease had advanced too much, caudal fin rays
had disintegrated and the tissues at the base of the caudal fin had been
invaded. Consequently the fry died on the morning of the 19th November.
The Rainbow trout fry infected with the disease, thus, died within 70-118
hours after inoculation and within 50-55 hours after the external appearance
of the disease.
372
Hamid Khan
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Study in Diseases of Fish . Fin-Rot
373
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374 Hamid Khan
The dead fry in all the cases sank to the bottom.
Immunity of Indigenous Species
Six months old fry of Roliu (Labeo roJiita H.B.) and Morakha (Cirrhina
mrigala H.B.), well-known indigenous food fish oftlie Punjab, were inoculated
at the same time and in a similar manner as those of the Rainbow trout.
But these fish were found to be totally immune to the disease.
Etiology and Pathology
Preparations made from Gourami (Plate XVIII, Fig. 6), and from Rain
bow trout fry (Plate XVIII, Fig. 5), show similar rod-shaped bacteria, which
are found in large numbers in the infected tissues. It is, therefore, apparent
that the causative organism of the disease, both in Gourami and Rainbow
trout, is the same, as it is so uniformly present in the diseased tissue. Davis
(1937) 4 described the causative organism of fin-rot to be " a rod-shaped
bacterium, which can usually be found in large numbers in the infected
fins' \ Though no figures are given by him, it is, however obvious from the
etiology and pathology of the fin-rot disease given by Davis, that the
causative organism 5 of the disease in Gourami and Rainbow trout is the same
as causes the fin-rot. The infection usually starts on the outer margin of the
fin and rapidly spreads to the underlying tissues. The fins which have
been injured or scratched are the first to be attacked.
Davis (1937) is of the opinion that "in small fmgerlings the pectoral
fins usually are affected first, the dorsal fins being attacked next, and the
infection may spread eventually to the other fins, although they are rarely
as badly affected". In the Gourami and the Rainbow trout fmgerlings, in
the present case, however, the sequence in the order of infection was different
from that described by Davis. In almost all cases in the Gourami, caudal
fins were affected before the dorsals and pectorals. In the Rainbow trout
fry, too, the inoculated parts of the caudal fin developed the infection first.
According to Davis (1937), "In young fingerliiigs death usually occurs
before the infection extends beyond the fins, so that the occurrence of lesions
on the body is rare/' The present investigations show that the disease in
the case of Rainbow trout fry had spread with great rapidity to the under-
lying tissues at the caudal peduncle where lesions were formed before the
death occurred, while in Gourami, the fingerlings died before the infection
extended beyond the fins.
4 Davis, H. B., " Care and Diseases of Trout," Bureau of Fisheries, U.S.A.,
Investigational Report, 1937, No. 35, 58-60.
5 Culture of causative organism, on some of the commonly used Laboratory media
at Lyallpur was not successful. '
Stiidy in Diseases of Fish : Pin- Rot 375
Control
Attempts to save the diseased Rainbow trout fry, under experiment,
by giving them bath of copper sulphate solution (1 : 5,000) for 2 minutes
were not successful as the disease had spread to the underlying tissues.
Control measures can only be effective in early stages of the disease
before the bacteria have penetrated the tissues. Davis (1937) recom-
mends " dipping the fish for 1 or 2 minutes in a 1-2,000 solution of
copper sulphate''. The copper sulphate solution of the strength recom-
mended by Davis has been found to be too strong for young fry.
It has, however, been possible to treat the diseased Rainbow trout fry
successfully by putting them in copper sulphate solution of 1,~20,000 strength
for 10 to 15 minutes. It is impossible to cure fish in advanced stages of the
disease and all such fish should be destroyed before any control measures are
adopted as otherwise they will contaminate the water and form a permanent
source of infection. Even healthy Rainbow trout fry, as the investigations
have shown, when placed in contaminated water, develop the disease with
great rapidity. It is, therefore, very essential to disinfect thoroughly all
troughs, hatching and rearing trays, " live-cars " and "conditioning-cars"
before putting healthy fiy in them.
Rainbow trout fry have been found to suffer from the fin-rot at the hat-
cheries, when they are overcrowded in a live-car. It has also been found that
any injury, whether mechanical or otherwise, caused by external parasite,
makes the fin more susceptible to the disease. It has also been noticed that
dirty, muddy rain water not only aggravates the disease, but has been found
to be directly responsible for an outbreak of the disease among Rainbow
trout fry. In the present case too, it seems, that transport of Gourami
fingerlings by canal, where water was to some extent dirty, aggravated the
disease and the fry died very rapidly.
Conclusion
Fin-rot has been found to be a serious disease of Rainbow trout finger-
lings at the Hatchery. The mortality is very high and it has been observed
that within one week from the outbreak of the disease thousands of fry have
died. The present investigations indicate that if once this disease enters
a Gourami Farm, and if due precautions are not taken, it is likely to play a
great havoc with the fry as well as with the adults. With due precautions
it is possible to safeguard an outbreak of the disease. It is expected, there-
fore, that the present investigations, which have established the suscepti-
bility of Gourami to fin-rot, will have far-reaching effect on cultivation of
Gourami in inland waters in India.
376 Hamid Khan
A cknowledgments
My thanks are due to the Director of Fisheries, Madras, for providing
me with facilities during my visit to Inland Fish Farms in Madras. I am
indebted to Mr. N. V. Joshi, Assistant Bacteriologist, Ptisa Imperial Agricul-
tural Research Institute, New Delhi, and Dr. M. R. Madhok, Agricultural
Bacteriologist, Lyallpur, for examination of the diseased fish and for prepara-
tions of bacteria. I am thankful to Mr. S. M. Nasir, Punjab Agricultural
Research Institute, Lyallpur, for the photographs and microphotographs
appearing in this paper.
Summary
1. Study of fin-rot, a bacterial disease of the fins of fish, has been made
by inoculation of healthy Rainbow trout fry with the disease from diseased
Gourami (Qsphromenus gouramy Lacepede) fingerlings.
2. The Rainbow trout fry infected with the disease died within 70 H8
hours after inoculation and 50 to 55 hours after the first externally visible
symptoms of the disease.
3. Preparations from the diseased Gourami and infected Rainbow
trout fry show exactly similar causative organism, which is a rod-shaped
bacterium, present in large number in the infected tissues.
4. Overcrowding, dirty and muddy water aggravates the disease.
5. Copper sulphate solution (1-20,000) bath for 10 to 15 minutes
has been found to cure the fry in early stages of the disease. It is impossible
to cure fish in advanced stages of the disease, and all such fish should be
destroyed before adopting any control measures.
6. Fry of indigenous fish such as Rohu (Labeo rohita H.B.) and Morakha
(Cirrhina mrigala H.B.) have been found to be immune to fin-rot.
1881-39-Printed at The Bangalore Press. Mysore Road, Bangalore City, by G. Srinivasa Rao, Superintendent
and Published by The Indian Academy of Sciences. Bangalore
Dr. Hamid Khan
Proc. Ind.Acad. Set., B, voL.X, PL XVIII
Gourami fingerlings (Osphro menus fjoiiramy Lacepede), showing fin-rot on caudal,
dorsal and anal 15ns
Microphotograph-of bacteria from caudal
fin of Gourami fingerling suffering from fin-
rot. X 1,200. (Oil immersion.) Acid f-itch-
sin.
Microphotograph of bacteria from caudal
fin of Rainbow trout fry inoculated with fm-
rot "disease. X 1,200. (Oil immersion.)
G. Violet.
Dr. Ham-id Khan
Proc. hid. Acad. Sri., J3, vol. X, PI- XIX
FIG. 1. Rainbow trout fry with healthy caudal fin.
FIGS. 2 to 4. Rainbow trout fry inoculated with fin-rot disease Jrom dead
Gourami fry, showing fin-rot on their caudal fins.
FIGS. 5 to 7. Rainbow trout fry put in contaminated water showing (in-
rot on their caudal fins,
FIG. 8. Rainbow trout fry with fin-rot on its adipose fin.
FIG. 9. Rainbow trout fry with fin-rot on its caudal fin.
Acharya, B. N., and
Devadat.tn, 8. 0.
Akhtar, S. A.
INDEX TO VOL. X (B)
AUTHORS' INDEX
Compounds of phosphorus in milk, I, 1*2.1.
Phosphorus, calcium and magnesium in milk, 'U,
Ayyangar, <*. N. liau<>'a-
swanri, ami Ponnaiya,
B. W. X.
Ohiplonker, 0. \V.
Dcvadatta, fc*. ( -.
Ganapathy, 0. V., and
Sastri, B. .N.
Manuel Kliau
Krishnaswami, ( 1 . S.
Lai, M. B.
Mello, I. l\ do
Misra, M. L.
Poiiuaiya, >>. \V. X.
K)aina.s\vaini, L. S.
Sadasivan, T. S.
Mastri, B. N.
S(\sha.char, B. U.
On some nematode parasites from Afghanistan,
287.
Studies in tiorghum tiuda-nense, Stapf The Sudan
grass, 237.
Bryozoa from the Bagh beds, 98.
Lamellibraachs from the Bagh beds, 255.
tfec Aeharya and Devadatta.
Oxidation of thiols and ascorbic acid in the
latex ol' papaya, 81.
Study in diseases of fish: tin-rot a bacterali
disease of fins of lish, 369.
*SVf* Thomas and Krislinaswami.
Studitvs i.u Helminth ology trematode i)ara.sites
of; birds. 111.
Two .now additions to the list of the Indian
aspergilli, 21.8.
On some stone implements from Hoshangabad
(C( k ntral Provinces), 275.
Nw Ayyaag'ar and Pormaiya.
Some aspects of the anatomy of amira (am])hibia)
a review, 41.
(om petition in fungi, I. A study of the growth
reactions of non-parasitic fungi in. associated
culture, J.
>SVff Ganapathy and Sastri.
T( 4 stic.ular ova in T'rffiotyp'hl'U8 narayani
char, 213.
Singh, B. .N'., and Hiiigh, Effectiveness of chemical fertilisers oil the growth
J. E. and water requirement of wheat, 27.
Singh, J. ~R. . . See Singh and Singh.
Srinivasau, K. S. . . On tho developmental morphology of andro-
gynous receptacles in Martfiantia
es, 88.
Thomas, K. M., and Little leaf a. transmissiblo disease of brinjal, 20.1.
Krislinaswaini, C. 8.
Venkataraman, G. . . A systematic account of some South Indian
diatoms, 293.
TITLE INDEX
Anura (amphibia), some aspects oi! the anatomy a review (Ramaswaxni), 4.1.
Aspergilli, Indian, two new additions to the list (Mello), 218.
Bagh Beds, bryozoa (Ohiplonker), 98.
Bagh Beds, lamellibranehs (Chiplonker), 255.
Diatoms, South Indian, some, a systematic account (Venkataraman), 29.'*.
Fish, study in diseases : Fin -rot a bacterial disease of fins of fish (II amid
Khan), 369.
Fungi, Competition, I (Sadasivan), 1.
Ilelminthology, studies trematode parasites of birds (Lai), 1.1 JL
Little leaf a transmissible disease of brinjal (Thomas and Krishnaswami), 20.1.
Marchantia palmata N ees, androgynous receptacles in, on the deve opmental
morphology (Srinivasan), 88.
Milk, compounds of phosphorus I (Acharya and Devadatta), 221.
Milk, phosphorus, calcium and magnesium II (Acharya and Devadatta),
229.
"Nenuitode parasites, some, from Afghanistan (Akhtar), 287.
Papaya, latex, oxidation of thiols and ascorbic acid (Gana.pat.by and Wastri),
81.
Morghum tiudanvn&e,, Stapf the Sudan grass, studies (Ayyaogar and
Ponnaiya), 237.
Stone implements, some, from ICoshangabad (('-(Mitral ProviiuMja) (Misra), 275.
Testicular ova in Urasotyphhut narw/ani Scshachar (Seshachar), 213.
Whc^at, the growth and water requirement, elTectiveness of cheinica,! fert.ilisorB
and Hingh), 27.
379
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