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TRANSACTIONS
OF THE
ROYAL SOCIETY OF SOUTH AFRICA
VOLUME X.
1922.
WITH THIRTEEN PLATES, ONE FOLDING MAP,
AND ONE FOLDING TABLE.
269606
CAPE TOWN:
PUBLISHED BY THE SOCIETY.
1922.
Printed in Great Britain by
Neity & Co., Ltp., EDINBURGH.
CONTENTS
THe Action oF Hucomis undulata, Ait. By J. W.C. Gunn. (With
three Text-figs.)
THe WatER RELATIONS OF THE PINE (Pinus Pinaster) AND THE
SILVER TREE (Leucadendron argenteum). By R. D. AITKEN,
M.Sc. (With one Text-fig.)
Note on AXISYMMETRIC OrTHOGONANTS. By Sir THomas Mutr,
Bie. F.R.S:
OBSERVATIONS ON LIVING FISHES BROUGHT BY H.M.S. ‘*‘ CHALLENGER ”’
FROM TropicaL East Arrica TO CAPE Waters. By J. D. F.
GILCHRIST
Notes ON THE DEVELOPMENT OF THE OVULE, EmBRyo Sac, AND
Empryo oF Hydnora africana, THuns. By R. H. Dastur,
B.Sc. Bombay, F.L.S. (With thirteen Text-figs.)
(1) THz PERMANGANATE ABSORPTION-SPECTRUM: A CLAIM FOR
Priority. (2) A FoRMULA FOR CALCULATING THE URANIUM
Spectrum. By James Morr
CoLtouR AND CHEMICAL ConstTITUTION. Part XIII.—THE Catcv-
LATION OF THE CoLouR oF THE Monocyctic Dyess. By
JAMES Morr
Own Braula Caeca, Nitzscy, A DIPTEROUS PARASITE OF THE HonEY BEE.
By 8. H. Sxaire, M.A., M.Sc. (With eleven Text-figs.)
Note on A Frre-FLiInt oF STRANDLOOPER ORIGIN. By JOHN
Hewitt. (With Plate I)
A Note on THE USE oF THE SoutTH AFRICAN CLAWED TOAD IN THE
BroLtoaicaL ASSAY OF THE DigitTaLis Series. By J. W. C.
GUNN
SUNSPOTS AND HartH TEMPERATURES. By J. R. Sutron
RAINFALL AND THE PRESSURE GRADIENT. By J. R. Sutton
CoLOUR AND CHEMICAL CONSTITUTION. Part XIV.—THE CaLcvu-
LATION OF THE COLOUR OF THE DicycLtic Dyzrs. By JamEs
Morr
ii
PAGE
23
27
Bo
35
4]
49
65
iv Transactions of the Royal Society of South Africa.
PAGE
CRYSTALLINE STRUCTURE OF ANTIMONY AND BismutTH. By A.
Oaa. (With two Text-figs.) . ; ; mate)
MAP SHOWING THE DISTRIBUTION OF THE GENUS Agama. By G. A.
BouLenGER and J. H. Power. (Left over from the paper on
the subject published in Vol. IX, Part 3) : . facing page 80
On THE REPTILIAN GENERA Huparkeria Broom, and Mesosuchus
Watson. By 8. H. Haventon, B.A. (With Plates II and III) 81
NoTE ON THE PRODUCT OF ANY DETERMINANT AND ITS BORDERED
DERIVATIVE. By Sir THomas Muir, LL.D. . sees)
Comments oN Dr. PERinGuEY’s NotE on WuHates. By ArTHuR
F. Bearpark. (With one Text-fig.) . LP eoe
Note oN THE PEecToRAL FIN oF THE SOLE, Achirus capensis: Its
ORIGIN AND SIGNIFICANCE. By J. D. F. Gitcurist. (With
Plate IV) . : ee
On THE SENONIAN AMMONITE Fauna oF Ponpotanpd. By L. F.
SpatH, D.Sc., F.G.8. (With Plates V to TX) . ; Roelile,
A Funcus—Gibellula Haygarthi, sp. N.—oN A SPIDER OF THE
Famity Lycosipak. By Paut A. van DER Bist. (With four
Text-figs.) . ; ; . 149
Some SoutH Arrican StEREuMS. By Paut A. VAN DER BISL.
(With nine Text-figs.) ; 151
CoLouR AND CHEMICAL CONSTITUTION. Part XV.—A _ Sys-
TEMATIC StTuDY OF FLUORESCEIN AND RESORCIN - BENZEINE.
By James Morr . : ; ; ; . 159
Tur ““Account-Book” or JAN Haszine. By C. Prsprr, M.D.(Leiden),
Johannesburg, and H. ZwWARENSTEIN, M.B., Capetown os
I. On SoutH AFRICAN PARAMPHISTOMIDAE (Fiscu.). II. Some
TREMATODES IN SoutH AFRICAN ANURA, AND THE RELATIONSHIPS
AND DISTRIBUTION OF THEIR Hosts. By C.8.GRoBBELAAR, M.A.,
Lecturer and Demonstrator in the Department of Zoology, Uni-
versity of Stellenbosch, Stellenbosch, S.A. (With nine Text-figs.) 181
STUDIES IN THE MorpPHo.oey oF Selaginella pumila, Sprine. Parr I.
—THE VEGETATIVE ORGANS OF THE SPOROPHYTE. By A. V.
Durnie. (With twenty-six Text-figs.) . . Var
SOME OBSERVATIONS ON THE Errects oF a BusH Fire ON THE VEGE-
TATION OF SignaL Hitt. By Maraarst R. MicHeux, Lecturer
in Botany, University of Cape Town. (With Plates X to XJI
and one Text-fig.) . ‘ . F : : : : » (23
Contents.
CoLtourn “AND CHEMICAL ConstiITUTION. Part XVI.—FURTHER
MiscELLANEOUS OxsERvATIONS. By James Moir .
On Hyauite. By J.S. v. p. Lincen and A. R. EH. WALKER
CrrcUMCISION REGIMENTS AS A NativE CHronotoGcy. By the Rev.
Professor W. A. Norton, M.A., B.Litt. (With one folding
Table) .
SESUTO AND SECHWANA Praises. By the Rev. Professor W. A.
Norton, M.A., B.Litt.
NoTvTE ON A DETERMINANT WITH FACTORS LIKE THOSE OF THE DIFFER-
ENCE-PRopuct. By Sir THomas Murr, F.R.S.
CoLOUR AND CHEMICAL ConsTITUTION. Part XVII.—THE Azo DyYEs
AND OTHER Monocyciic CoLtours. By James Morr
FUNGI OF THE STELLENBOSCH DISTRICT AND IMMEDIATE VICINITY.
By Dr. P. A. vAN DER BrgL
OBSERVATIONS ON THE PROTECTIVE ACTION OF NORMAL SERUM IN
EXPERIMENTAL INFECTION wiITH Bacititus DIPHTHERIAE. By
ies MACKIE, M.D., Ch.B., D.P.H.
On somME Upper Braurort Tuerapsipa. By 8. H. Haueuton,
B.A., D.Sc. (With Plate XIII and two Text-figs.) .
245
253
267
273
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TRANSACTIONS
OF THE
ROYAL SOCIETY OF SOUTH
VOL Xe
THE ACTION OF EUCOMIS UNDULATA, Se Ry,
'@ alee) WY a es s 330
By J. W. C. Guwn. Jtional Ms <i
(From the Pharmacological Department, University of Cape Town.)
(With three Text-figures.)
Eucomis undulata, Ait, is a bulbous plant of the Natural Order of
Liliaceae, which grows in Bechuanaland and Basutoland. It has a local
reputation as a cure for galziekte and other diseases in stock. Nothing is
known as to its chemical character or pharmacological action. I was given
two bulbs, one about 3 in. in diameter and the other much smaller, in order
to investigate its constituents and action.
The total amount of material weighed under 200 grm. 100 grm. of the
bulb were dried, powdered and macerated in 70 per cent. alcohol for three
days, then filtered through three folds of linen. The filtrate was care-
fully dried over a water-bath. It weighed 10 erm.
ACTION ON THE FRoa.
The minimal lethal dose of the extract for frogs was found to be ‘005 germ.
per gramme of body-weight. The extract was dissolved in Ringer’s solution
and injected into the ventral lymph-sac. Death occurred in from 10 to
26 hours.
The only symptoms noted were slowing of the respiratory movements
and final paralysis. The heart was arrested in diastole. There was marked
inflammation at the site of injection.
ACTION ON MAMMALS.
(1) Oral Administration.—The extract dissolved in water was adminis-
tered to cats through a stomach-tube. The largest dose given was 6 grm.,
to a cat weighing 2050 erm. No general symptoms were seen, but evidence
li
2 Transactions of the Royal Society of South Africa.
of severe local inflammation, salivation, vomiting and purging were noted
In most cases.
(2) Subcutaneous Administration.—The extract had only slight toxicity
when injected subcutaneously. A dose of 1 grm. per kilo of body-weight
produced no general symptoms in rats, and all recovered. The injection was
made into the back, near the tail. In almost every case the sciatic nerve on
that side became paralysed.
As it was possible that the rat might have some congenital tolerance for
the active principle, a similar dose was injected into a cat and a rabbit, but
again the effects were purely local.
(3) Intravenous Injection.—A_ tincture, of which 5 c.c. were equivalent
to 1 grm. of the bulb, was made with part of the remainder of the material.
It!was used in the remaining experiments. The alcohol was driven off before
Fig. 1.
|
Bis ils Qe ae “ge titli fila ps ttt bo
it was used. The blood-pressure was recorded, and, asa rule, the respiratory
movements, the heart volume, or the intestinal volume as well.
Two main actions were seen: The heart was slowed and weakened. If
the injection were made very slowly, with a dilute solution, the cardiac
effect was lessened or absent. The blood-vessels as a rule were constricted.
As a result of these actions the effect on the blood-pressure varied. Usually
there was a sharp preliminary fall, which seemed to be due to the effect of
the substance on the heart. The blood-pressure then rose again, sometimes
not quite to its previous level, but sometimes considerably above it. If the
injection were made very slowly there was a slight fall or a slight rise.
These were the effects seen after injection of 0°5 c.c. of the alcohol-free
tincture into the jugular vein of a cat.
Three cubic centimetres of the tincture, if given in an undivided dose,
was fatal, death being due to respiratory failure.
Fig. 1 shows the effect of 0°5 c.c. of tincture of Hucomis wndulata on
the blood-pressure of a cat weighing 2340 grm.
The Action of Eucomis undulata, Art. 3
ACTION ON THE ISOLATED HEART.
The movements of the isolated mammalian heart were recorded by
perfusion through the coronary arteries. The heart of the rabbit, cat, rat
and guinea-pig were used. The effect of Hucomis undulata was to slow and
Fig. 2.
lt
ay
weaken the heart and arrest it in diastole. In Fig. 2 the effect of 0°5 per
cent. tincture is shown on the movements of the cat’s heart. a is the normal
heart tracing, B, c, D, E and F the movements after perfusing with the
experimental solution for 1, 3, 4, 6 and 8 minutes respectively.
The isolated frog’s heart was arrested in diastole, but sometimes there
was a slight increase of tone.
ACTION ON UNSTRIATED MUSCLE.
The tone of the uterus—pregnant and non-pregnant—of the cat and
rabbit is increased by E'ucomis undulata, as shown in Fig. 3.
4: Transactions of the Royal Society of South Africa.
THe Active PRINCIPLE.
It was evident that Hucomis undulata contained some saponin body, as. ©
a very persistent froth was seen when the tincture was shaken up with
water.
Forty grammes of the bulb were powdered and extracted with 90 per
cent. aleohol. The extract was dried and extracted with distilled water.
The resulting extract was dried, and weighed 2:015 erm. It was completely
soluble in water and in 90 per cent. alcohol, but insoluble in ether. It pro-
duced all the actions of the tincture and extract on animals and isolated
tissues.
On boiling with dilute hydrochloric acid a sugar was obtained. The
other product of hydrolysis is insoluble in water and is crystalline.
The active principle is, therefore, a glucoside. After the tests and
animal experiments were done with it.there was not enough left to determine
whether it is a pure principle or a mixture. If a small quantity were mixed
with water and the mixture shaken a persistent froth formed. Volatile
and fixed oils were emulsified with a little of the glucoside, and the emulsion
separated out only very slowly.
Harmo.uytic AcTIoN.
The glucoside is powerfully haemolytic. The following table shows its.
haemolytic action on washed corpuscles in one hour:
Concentration of
glucoside. bese
lin 2000 . a Caan . Complete haemolysis.
lin 4000 . : , : ‘ .
lin SSO00r y ”
lin 16,000 . : : ri a
1 in 32,000 . ' : . Partial haemolysis.
1 in 64,000 . é . . No haemolysis.
Control. ae . No haemolysis,
SuMMARY.
Eucomis undulata contains a sapo-glucoside which is a powerful haemo-
lytic agent. It does not seem to be absorbed from the alimentary tract, and
only slowly from the subcutaneous tissues. It is toxic when injected
intravenously, affecting the heart and the respiration. It is probably of no.
medicinal value.
(5)
THE WATER RELATIONS OF THE PINE (PINUS PINASTER)
AND THE SILVER TREE (LEUCADENDRON ARGENTEUM).*
By R. D. Arrrcen, M.Sc.
(Union Government Research Scholar).
Communicated by Prof. D. THopay.
(With one Text-figure.)
I. INTRODUCTION.
In two papers recently published in the ‘ Proceedings of the Royal
Society, Farmer (1) has given an account of experiments he has performed
with a view to ascertaining, among other matters, whether “the efficiency
of the wood can be usefully expressed in a quantitative form, and to what
extent separate species differ.” As a result of his experimeuts he concludes
that the efficiency of the wood can be thus expressed, and describes a
method of measuring it, and defines the “ specific conductivity” of the
wood as “the quantity of water transmitted through 1 sq. cm. of wood,
15 em. long, in 15 minutes under a pressure of 30cm. of mercury.”
The clearest result of his experiments is that evergreens possess wood of
markedly lower efficiency than that of the broad-leaved deciduous trees, and
that the absolute range of variation in individual species is far narrower
than in deciduous forms. A few examples from the tables in his papers will
illustrate this:
Evergreen. Deciduous.
Name. Spec. cond. Name. Spec. cond.
Scots pine . : » to 222) 4 “Commonjoals - ef Ondteelo
Euonymus japonica . 121 . Huonymus europaeus . 40-2
He further suggests that the xerophytic character of many of the ever-
greens may be due to the low conductivity of their wood, and says, “ It
becomes intelligible why a plant apparently xerophytic may yet be restricted
to localities in which it is never really subject to drought. The wood is
* The cost of this research was in part defrayed from a Union Government
Research Grant.—D. T.
2
6 Transactions of the Royal Society of South Africa.
capable of transmitting a limited supply of water, the leaves are fitted just
to utilise this amount with very little margin; in other words, though the
plant requires little, it can hardly do with less.”
In May of 1919 the present writer began, at the suggestion of
Prof. D. Thoday, a series of measurements of the specific conductivity of
the wood of certain South African trees with the object of ascertaining
whether this property of the wood played any part in deciding the usual
habitat of the various trees. It was soon realised, however, that this factor
could not be considered independently, but that the transpiring power of the
leaves must also be taken into account. For instance, a tree with wood of
high specific conductivity and mesophytic leaves, which use up nearly all
the water they can obtain from the stem, will probably be quite as much
restricted as to its habitat as a tree with wood of low conductivity and
xerophytic leaves. Accordingly it was decided to compare the transpiration
rates and the water-conducting power of the wood of two trees, and see
whether any indication could thus be obtained of the drought-resisting
power of the trees.
For the preliminary work the two trees selected were Pinus pinaster
and Leucadendron argenteum, both of which grow within practicable range
of the University of Cape Town. The leaves of each of these evergreen
trees are distinctly xerophytic. That of the Pine is so well known as to
need no description. The leaves of the Silver Tree are sessile, 25—53 in. long,
4-1} in. broad, and lanceolate. They are densely pubescent | on both
surfaces with adpressed, silvery, silky hairs. According to Marloth (2)
these hairs are closely pressed to the surface of the leaf in dry weather,
and stand out at an angle of about 30° from the surface during damp
weather. With the naked eye, however, the present writer could observe no
difference in the position of the hairs in dry and damp weather, though no
critical experiments were conducted to test the statement. As in the case
of other Proteaceae the stomata of Leucadendron argentewm are protected
by a small raised pore.
Il. Meruops.
A. Measurement of conductivity.—Lengths of 15 em. were cut under
water from the centre of the twigs chosen for the experiments, and placed
in jars of water, over which the pressure of air was reduced by means of a
suction pump. Half an hour to an hour was deemed sufficient to remove
all the air from the stem, and to inject it with water.
To obtain the necessary pressure for the actual conductivity measure-
ments an aspirator was placed on top of a cupboard and filled with distilled
water. In order that the pressure might remain constant throughout the
series of experiments, a glass tube was inserted through a cork in the top
The Water Relations of the Pine and the Silver Tree. 7
of the aspirator. A long tube descended from the base of the aspirator,
and the twig was attached to the lower end of this tube. In this way a
head of water of 186 cm. was obtained. This is equivalent to a pressure
of 13°68 cm. of mercury.
The amount of water passing through the twig in 15 minutes under this
pressure was measured. ‘The values thus obtained are referred to as the
“standard transmission ” (i.e. the amount of water transmitted under the
standard conditions) of the twigs. In the early experiments four readings
were taken for each twig, and it was found that in nearly every case the
amount of water transmitted became less with each successive quarter of
an-hour, é.g. :
Pine, first quarter hour, 5°6 c.c. Silver Tree, 16 c.c.
,, second S 48 ,, f ame
Pee uhird AN = iso,
= fourth 4 otal eae : a ESOS
That this falling-off was due to clogging at the cut surface seems clear
from the facts that it was not nearly so marked in the Leucadendron as in
the Pine, and that it did not occur when the surfaces were shaved after each
reading. The first reading was therefore taken as most nearly representing
the true water-conductine power of the wood. In later experiments only one
or two readings were taken, and the first of these was used.
Having measured the standard transmission of the twig it was bisected,
and a cross-section cut from the centre. By means of a projection apparatus,
fitted up at the suggestion of Prof. Thoday from materials available in the
laboratory, a drawing of this section was made on squared paper, and the
area of wood in the section was measured.
All the data were then available for calculating the specific conductivity
of the wood.
B. Measurement of transpiration.—The twigs for these experiments were
cut from the trees in the evening and kept overnight with their cut ends in
water. They were then placed in small test-tubes of water, and the loss of
weight per half hour measured. In some cases the twigs were kept in the
laboratory throughout the experiment, in others they were exposed on the
roof of the laboratory on both dull and bright days.
©. Measurement of leaf area.—In the case of Leucadendron the method
was comparatively simple. The leaves were picked from the twig and
classified according to their size into about six groups. A representative
leaf was then chosen from each group. and placed under glass on a piece of
photographic paper, The leaf shape thus obtained was cut out and weighed,
and from this weight and the area and weight of the whole piece the area of
the leaf print was calculated. Knowing this, and the number of leaves in
each group, the leaf area was calculated for each group and then for the
whole twig.
8 Transactions of the Royal Society of South Africa.
The determination of the leaf area of the Pine presented greater diffi-
culties. The most hopeful method seemed to be to find an expression for
the area involving the length of the leaf and the diameter of its flat side.
As a first step towards this, sections were cut from various portions of a leaf
and camera lucida drawings made of these sections. Measurement of the
perimeter of the curved surface and of the diameter of the flat surface
showed that the relation between the two was fairly constant and averaged
1°8, e.g.—
Lear No. 1. Length, 16:7 cm.
Position of section. Diameter. Curved surface. C/D.
Base. ; 1:39 mm. 2:7 > am: 1:99
271 cm.from base. 126 On awe 3:14: \c, : 1:86
4:2, 5 169.73 BLU, os : 1°82
G3. & P. : TOO 2:30 : 1:78
SOD) = . ; 58 4 , 2S Oaee , 1°81
10°75 _,, 62:2 3028 ; 1:86
12°6 _,, = 150 eae) th Lev
14-7 —,, - : Le3or ; 2°42 ,, E 1:75
15-7, eli ar 2°05, 178
16:25) = - 107, : — , a
16°6 _,, - 0:54 ,, : O:57 wie ==
9)16°38
Average . : . ; 1:8(2)
Two other leaves, one 16:9 cm. long, the other 10:1 cm. long, gave as the |
average ratio C/D 1:77 and 1:84 respectively. In the case of nine other
leaves of varying length the diameter and circumference of the mid-section
were measured, and the average ratio of C/D in these cases proved to be
1:77. The value of C/D was taken as the average of these four values,
Ve aS
The next step was to measure as accurately as possible the area of the
flat surface of the leaf. This was effected by cutting sections of a leaf at
intervals from the base and plotting on squared paper the diameter of these
‘sections against their distance from the base. In this way one obtained a
magnified drawing of the flat surface of the leaf, and measured its area by
counting the number of squares it contained. Knowing the magnification,
the actual area of the flat surface was then readily calculated. The total
area of the leaf was then taken as —
Area of flat surface x (C/D +1)
i.e. Area of flat surface x 2°8,
The Water Relations of the Pine and the Silver Tree. )
Comparison of the leaf area measured in this way with the area calcu-
lated from —
Length of leaf x Total perimeter of mid-section
showed that the latter was always slightly greater than the former, e. g.—
a. °.
No. Area from graph. Mi ie eee cena a per cent. of b.
] 7°08 sq. em. 7°38 sq. em. ; 95°9 per cent.
2 Gio) ic, CA0E ,; Cosas
3 Boo 5; 348 ,, , De As ao
The actual leaf area was therefore taken to be 96 per cent. of the area
calculated from the expression, perimeter of mid-section x length.
To obtain the total leaf area of a twig, the leaves were arranged in
groups according to their length, and the diameter of the flat surface of
representative leaves from each group measured. The leaf area of each
group was then obtained from the expression, number of leaves x length x
diam. mid-section X 2°38 x 0:96. As already explained, 2°8 is the value of
(C/D + 1) and 0:96 represents the necessary reduction to 96 per cent., just
mentioned.
The total leaf area of the twig was then obtained by adding together the
areas of the groups.
IIl. ReEsvuts.
A. The water-conducting power of the wood.—The first experiments were
confined to measurements of the conductivity of the wood of the two trees,
and showed a considerable difference between the two. It should be noted
that the ‘conductivity’ in these experiments differs from Farmer’s
“specific conductivity.” The former is calculated for a pressure of 186 cm.
of water, the latter for a pressure of 80 cm. of mercury. At the outset a
few measurements were made of the conductivity of the wood of the Oak as
well as of that of the Pine and Silver Tree. The results obtained for these
three trees are in accordance with Farmer’s main conclusion that evergreens
possess wood of markedly lower efficiency than that of the broad-leaved
deciduous trees, and that the absolute range of variation in individual
species is far narrower than in deciduous forms.
In the few measurements made of the conductivity of the wood of the
Oak the values ranged from 18-9 to 107°8, the average of seven experiments
being 46:6.
Of the three trees the Pine showed least variation in its conductivity.
For the six preliminary measurements with Pine wood the average conduc-
tivity was 71, the values ranging from 5:9 to 82. These measurements
were made in July, 1919; in January, 1920, further measurements were
10 Transactions of the Royal. Society of South Africa.
made of both transpiration and conduction, the average conductivity then
obtained being 7:9. In the later set of experiments, however, the range of
values was greater, the lowest being 56 and the highest 9:3. The average
conductivity obtained from all the experiments was 7:5.
In the case of the Silver Tree, nine experiments in July gave values
ranging from 13:0 to 26:0, the average being 20°38. The January experi-
ments gave lower values ranging from 10-0 to 246 with an average of 16:2.
The average for all the experiments was 17°8.
These results are tabulated in Table I, in which some of Farmer’s
figures are included for comparison.
TapLE I.—Measurements of Conductivity.
Pinus Pinus
VES. pinaster. sylvestris.
Range. Se Ys ; : | GlSiS=10 7855) 5°6-9°3 =
Conductivity (average) (186 cm. water) | 46°6 75 -—
Specific conductivity (30 cm. mercury) : |
Aitken : : : ‘fl 102°5 165 —
Farmer ‘ ; ; : : : Of Scie dl) — io ate
. Euonymus Euonymus
salon ese : Japonica europaeus
ee : (evergreen). (deciduous),
Range . : : : : 10°0-26°0 — _
Conductivity (average) (186 cm. water) es - =
Specific conductivity (80 cm. mercury) :
Aitken : , b : ; ‘ 39:2 “= —
Farmer : : ; ‘ — 2:1 40:2
The present writer’s figures show clearly that the wood of the deciduous
Oak has a much higher conductivity than has that of the Pine or Silver Tree,
and it also gives a larger range of values. This is in accordance with
Farmer’s main conclusion already quoted.
Comparing the two sets of values for the specific conductivity, a con-
siderable divergence will be noticed in the case of the Oak. This is probably
due to the small number of experiments performed by the present writer,
for although the average is so much higher, the range of individual values
is similar. The specific conductivity of Pinus pinaster is only a little
ereater than that of the species investigated by Farmer. For an evergreen
tree Leucadendron argenteum has a remarkably high specific conductivity,
its value being very nearly equal to that of the deciduous Huwonymus
japonica.
The Water Relations of the Pine and the Silver Tree. 11
B. Rate of Transpiration.—In the experiments on transpiration two
objects were kept in view, viz. :
(1) A comparison of the rate of transpiration of similar twigs of the
Pine and Silver Tree under the same conditions.
(2) The relation, if any exists, between transpiration and water-conducting
power of-twigs of the same tree.
With regard to the first of these, all the experiments showed a much
higher rate of transpiration both per twig and per unit area of leaf surface
in the case of the Pine than of the Silver 'Tree. In view of the difference in
water-conducting power of the wood this result seemed very surprising, but
it was obtained under various sets of conditions, e.g. in the laboratory, and
on the roof of the laboratory, on both dull and bright days. Table II gives
the actual transpiration per twig and per unit area of leaf surface of those
twigs, the transpiration of which was measured under similar conditions.
TasueE II.—Transpiration in grammes per hour of Pine and Silver Tree
under similar conditions.
(The experiments bracketed together were concurrent with one another.)
Leucadendron. Pine.
Teamncpiration per Transpiration per SAHISti ; . Transpiration per
eee square metre) TOR. | ‘Slate metre
Lab. Roof. Lab. Roof. Lab. Roof. Lab. Roof.
1 1-0 as a ae 62; = eee
2 1°54 5:74, 595 | 22°25 462 | 14°88 31-9" |, 03°60
3 ool 6:54 as 87°5 as (eat — | -652
12 0-41 1-09 3-09 8:2 ey at Bae
13 0°36 0:90 3°64 971 — | — ae -
14 se an =a aa 296 | 677 16:25 | 48:67
15 = — a = 2:00: | > 95°81 478 | 189-0
16 | 0:94. 2-05 6°35 13°84 ube Ma aay
17 0°64 1:28 Heo 14°45 ae = ae a
18 ne eS: a = 1:60 4°37 15°49 4232
21) ay 162 — 15°43 fae | uel
yl — 1:20 _ 10:07 aut ee =
28 | e413 me 10:96 Tas aa car a
29 j = oS = — — 39 | 93535
30 ee = - ae a Hea | 38°72
In every case it will be seen that the actual transpiration per Pine twig
is greater than that per Leucadendron twig, and that the transpiration per
unit leaf area of Pine is greater than that per unit leaf area of Leucadendron.
12 Transactions of the Royal Society of South Africa.
It will also be noticed in the above table that in concurrent experiments
the results for the transpiration per unit leaf area are reasonably concordant
in the case of Leucadendron, e.g. Nos. 12 and 18, 16 and 17, 27 and 28.
With the Pine twigs fewer measurements were made, and Experiments 14
and 15 show a considerable divergence in respect of transpiration per unit
leaf area.
Two sets of experiments were performed to ascertain whether the high
rate of transpiration continued in the Pine as the twigs were allowed to dry.
Twigs of both Pine and Silver Tree were kept overnight in water and the
following morning the cut ends were blocked with paraffin wax. The loss
of weight was then followed in the one set for five hours and in the other
for two days. Both sets showed that the rate of loss of the Pine twig was
greater than that of the Leucadendron during the first two hours, but after
this it became less.
Experiments Nos. 20 and 26: Date, January 20, 1920.
No, 20: Leucadendron. No. 26. Pine
Time. Weight of twig. | Loss of weight. Time. Weight of twig. | Loss of weight.
| , pes
9.53 a.m 37°82 orm. — 9.57 a.m.| 66°54 grm. —
10.56 __,, 36°35 __s,, 1°47 germ. 10.59). 62°90 _,, 3°64 erm.
IBS 35°10 =, P25 5 11.58" 5: 59°52, 3°38 ss,
12.17 p.m 34°93 ,, Onin. 12-22 p.m. ||) 09:39 >. OS. 5;
116; 4; 34°19 ,, O74 .,; ee ae ao 1Ooe., 0-29 .;,
2.13 ,, 33°59 ,, 0°60 —z, 2 AGr es DS SOs. 024 ,,
3.16 5, 32°86 .,, 0°73 3.20 5; 98°59 on, O27
Experiment No. 33: Date, January 29, 30, 31, 1920.
No. 33: Leucadendron.
Date. Time. Weight of twig. Loss of weight.
January 29, 1920 11.6 a.m. 27°46 erm. ——
oo 2.4 p.m. 26°69 _,, O77 grm.
¥s SOen.. 9.20 a.m. Zoos is 3°54,
” 3 Le QroOb ae, INte10) 945 DIO)! G5
No. 33: Pine.
January 29, 1920. 11.13 a.m. 36°05 grm. —
— 2.8 p.m. 30°24 ,, 2°81 grm.
53 30 sé“; 9.24 a.m. 34) aT 1:67- ,;
» AON is
31,
9.0405...
30:17 1,
The Water Relations of the Pine and the Silver Tree. 13
These results are shown in graphical form in the figure on page 17.
In connection with Experiment No. 33 it is to be noted that at the end
of the experiment the Leucadendron leaves were very dried up, while the
Pine needles still appeared fairly fresh. In fact the Leucadendron leaves
were quite shrivelled up several days before the Pine needles began to die
and turn brown.
One or two determinations were made of the water content of fresh
leaves, in the hope that this would throw some light on the results of the
transpiration experiments. In Experiment No. 3 two twigs, one of Leuca-
dendron and one of Pine, were kept in water overnight, and left on the
roof of the laboratory with their cut ends in water for about an hour and a
half the following morning. At the end of this period the fresh weight of
the leaves was determined, and also their area; they were then dried and
the dry weight determined. In Experiments Nos. 22 and 24, two twigs were
kept overnight in water, and a few leaves were picked from each the
following morning. The fresh weight, area, and dry weight of each set were
then determined. The result of these experiments was to show that the
Pine leaves have a larger water content per square metre of leaf surface
than the Leucadendron leaves, but a lower water content per cent. of fresh
weight.
Heperiment No. 3.
No. 3: Leucadendron. No: .3:-Pine.
Leaf area. ; ‘ : .| 1742°8 square cm. 1090 square cm.
Fresh weight of leaves 33°82 grm. 36 45 grm.
Dry x ae : : 14°20 ,, 18°35 _,,
Water content. : : : : 19°62 ,, 1 ee
Water content per square metre leaf
surface. ; ; , 1126. - 5, 1661 sa,
Water content per cent. fresh weight 58°02 per cent. 49°66 per cent.
Experiments Nos. 22 and 24.
No. 22: Leucadendron. No. 24: Pine.
Leaf area. : ; : . | 173:48 square cm. 1152 square cm.
Fresh weight of leaves : ' | 2°995 grm. 3°145 grm.
Dry 5 re : al TOS. Bylo
Water content. . ‘ : <4 | 1-St5e.. ISTO 2
Water content per square metre leaf |
surface y ; : : , : 110°4 Be 157:9 Pe
Water content per cent. fresh weight . 63°93 per cent. 57°84 per cent.
A number of experiments were also performed with twigs of Leuca-
14 Transactions of the Royal Society of South Africa.
dendron of various thicknesses, but no very definite relation could be
discovered between the rate of transpiration, and either the conductivity of
the wood, or the area of the wood in cross section. The average of a number
of experiments showed, however, that twigs of Leucadendron usually bear
a larger leaf area per unit cross sectional area of wood than do those of
the Pine. ;
From the results of all the experiments described a table has been
compiled to show the relation between the actual transpiration per 15
minutes and the amount of water actually transmitted through 15 em. of
the twig in 15 minutes under the pressure of 186 cm. of water after
injection of the wood, 7. e. between the transpiration and the “standard
transmission.”
Tasxe ITI.—Relation between Transpiration and “ Standard Transmission.”
Leucadendron. Pine.
Nowe Se eo ony 7 Sia ape ia. ee ak? tai
Transpiration. | Transmission. | Transpiration. | Transpiration. Transmission. a ranspiration.
| Transmission. | Transmission.
|
2 1:44 39 | Ont 3°72 2°35 Nes
3 1°30 29 0°45 Fo 1680 18 1-0
S) ileal 2°3 | 0°60 ae — —
o- 2°50 4-0 | 0625 = = =
6) 1-28 15 O85 ¥ na =
10, O72 0:9 | 0-80 — — —
ay) 0:68 O'°6 al — — —
12 0:27 32 | 0:08 == — ae
13! 0:23 1:0 te 0123 “= ox a
14 / = a — 169 1°65 1:0
15] ee ue ae 1:45 12 aie
19, Bue oe = 1-12 05 Die
215 O41 1:9 0°22 — | - —
27 0:30 27 O11 | — | — —
28 0°53 30 OTS | —- —
297) as a = | 1:00 1-4 | O71
30 me = an 1°66 0-9 pls
(Concurrent experiments are bracketed together.)
In every case it will be noticed that the ratio Transpiration/Transmission
is far higher in the Pine than in the Silver Tree. This must mean that the
Pine leaves exert a much greater suction force than do the Silver Tree
leaves.
IV. Discussion.
For the purposes of discussion it will be convenient to summarise the
outstanding results of the above experiments at this point.
The Pine tree possesses wood of very low water-conducting power, but
twigs, kept with their cut ends in water, show a high rate of transpiration
The Water Relations of the Pine and the Silver Tree. 15
both per twig and per unit of leaf area. The Silver Tree, on the other
hand, possesses wood of fairly high water-conducting power, its conductivity
being rather more than double that 6f the wood of the Pine, but twigs with
their cut ends in water show a much lower rate of transpiration than do
similar twigs of the Pine.
When transpiration is followed in twigs, as they dry, it is found that
the Pine twigs at first transpire more rapidly than the Silver Tree twigs.
After a short time, however, the transpiration of the Pine twigs suddenly
drops and becomes less than that of the Silver Tree twigs.
Determinations of water content show that Silver Tree leaves have a
sreater water content expressed as a percentage of fresh weight than have
Pine needles.
Comparing the relation between transpiration and water transmission
in the two cases, we find that the ratio transpiration to transmission is far
higher in the Pine than in the Silver 'Tree.
Great caution is necessary in the interpretation of these results owing
to the absence of information as to how far experiments with isolated twigs
afford a true idea of processes taking place in the intact tree. For instance,
the fact that isolated Pine twigs transpire more rapidly than do Leuca-
dendron twigs does not enable us to draw any conclusions as to the rate of
transpiration of twigs attached to living trees. In all the experiments
comparatively short twigs were used, and these were saturated with water
overnight. Consequently the resistance to the flow of water to the leaves
during transpiration would be minimal, and far lower than when the supply
must be drawn from the roots of the tree through the trunk and branches.
The measurements of conductivity show that the resistance to the flow of
water to the leaves is very much greater in the Pine than in the Silver
Tree, and this in itself would tend to lower the rate of transpiration of
Pine twigs while attached to the tree. |
One conclusion which may safely be drawn is that under the experimental
conditions Pine needles exert a greater suction on the water in the stem
than do Silver 'Tree leaves. In experiment No. 2, for example, the “ standard
transmission ”’ of the twigs was-—
Leucadendron 3°9 c.c. Pine'2°5 c.c.
Therefore to force water through the Pine twig at the same rate as through
the Leucadendron twig would require a pressure or suction 1°56 times as
sreat as that apphed to the Leucadendron twig. The rate of transpiration
of the Pine, however, was 2°58 times as great as that of the Leucadendron.
Therefore the Pine leaves must have exerted a suction 2°58 x 1°56, i.e.
about four times as great as the Leucadendron leaves. In experiment No. 3
the Pine leaves must have exerted a suction twice as great as the Silver
Tree leaves, and in Nos. 13 and 14 four and a half times as great.
One is tempted to infer from this that the greater suction, which our
16 Transactions of the Royal Society of South Africa.
experiments show can be exerted by the Pine needles, is a response to the
structure of the wood, enabling them to obtain water sufficiently rapidly
for their requirements. At the same time the much greater tensions, which
must exist in Pine trees as compared with Silver Trees, probably neutralise
the greater suction force of the Pine needles, so that the transpiration of
twigs growing on the tree will probably be less in the case of the Pine than
of the Silver Tree. Here, however, it must be remembered that we have
no information as to the maximum tensions which the leaves of either tree
are capable of exerting or withstanding, and without such information
inferences as to the behaviour of twigs on the tree are of little value.
The experiments with drying twigs are suggestive in this connection.
The figures for these experiments are repeated in the following table, the
initial fresh weight of the twig being called in each case 100. Alongside is
given the loss of weight per hour expressed as a percentage of the fresh
weight, and the original water content per cent. of fresh weight, as
determined in Experiments 3, 22, 24:
TasLEe LV.—Loss of Weight of Drying Twigs.
Experiments Nos. 20 and 26.
No. 20: Leucadendron. No. 26: Pine.
ie | Loss of weight per hour oe ah Loss of weight per hour
ce Welght: | per cent. of fresh weight. PEN): Weight. per cent. of fresh weight.
9.53 a.m. 100 — 9.57 a.m. 100
10:06) 3; 96°12 Sul NOR 9E.; 94°54 a3
159, 92°81 374 ToS... 89°43 a2
12.17 p.m.| 92°31 — 12.22 p.m.| 89°28 —
Li6 ,, | 90:40 19 LOR: 88°82 | 0°46
213° 4; 88°82 ie Zalin vs, 88°45 0°38
SiO) os 86°90 18 3.205%, 88:06 0°43
EHzxperiment No. 38.
Leucadendron. Pine.
Date. |
Loss of weight per Loss of weight per
Time. Weight.| hour per cent. of | Time. Weight.| hour per cent. of
fresh weight. fresh weight.
January 29, 1920/}11.6 am.| 100 es 111.13 a.m.| 100 =
== 2.4 p.m. | 97°21 | 0:95 2.8 p.m.| 92:19 2
ie 304e 9.20 a.m. | 84°29 | 0-67 9.24 a.m. | 87°58 0°24.
5 Oy | wes 9:50 ,, | 64°82 0:80) 9.54 ,, | 83°70 0:16
The Water Relations of the Pine and the Silver Tree. 17
Water Content per cent. Fresh Weight.
Experiment No. 38, 58:02 percent. . Experiment No. 3, 49°66 per cent.
» , 22,0393 ,, ; 9 pedo Sas
These results are shown in graphical form in the accompanying figure :
5 /O 15 20 25 30 35 40 45 HOURS jog
LHDISM TWNIDIYO 3O 1N39D Y3d LHD5IIM
WEIGHT PER CENT OF ORIGINAL WEIGHT
/ 2 3 4 5 HOuRS
The striking difference in the behaviour of twigs of the two trees is that
in the Silver Tree the rate of loss does not markedly fall off as the water
content diminishes, whereas in the Pine the rate of loss falls after a few
hours to less than one-tenth of the rate in the first hour. Tension may be
taken to increase as the water content diminishes, unless at some point the
resistance of the leaves begins to break down. If this happens the water
columns in the vessels break, and the leaves lose water without being able
to obtain fresh supplies from the stem. ‘This may be the case with the
Silver Tree. In the case of the Pine the bend in the curve suggests either
stomatal or internal regulation of a far more drastic nature, and this brings
us face to face with another difficulty—the absence of information as to the
18 Transactions of the Royal Society of South Africa.
behaviour of the stomata, both on the tree and under the experimental
conditions.
Attempts were made to obtain this information by Lloyd’s method of
fixing strips of the epidermis in absolute alcohol, and measuring the size of
the stomata and the degree of opening with a micrometer eyepiece. The
method, however, proved unsatisfactory, and neither type of leaf is suitable
for observations with the porometer.
It is evident, therefore, that any interpretation of the experimental
results with reference to the behaviour of whole trees must necessarily be
highly speculative. One possible interpretation is that the Silver Tree keeps
its stomata only partially open; that it has relatively immobile stomata and
that if the conditions become too severe, and the tension exceeds a limited
value, the leaves dry back, as in the cut twig. In the Pine, on the other
hand, the stomata are more mobile, open wider under favourable conditions,
and close more effectively in dry. Another possible interpretation is that
there may be a greater power of internal regulation in Pine leaves. That is
there may be some way other than stomatal closure, or additional to it, of
reducing water loss, probably connected with lowered water content. Apart
from these purely speculative suggestions all that can definitely be said at
present is that the Pine leaves appear to be more resistant to unfavourable
conditions than the Silver Tree leaves.
Further research is needed to determine the effect of cutting on the
stomata and internal condition of the leaves. In this connection transpira-
tion measurements in the field are needed, (a) on twigs still attached to the
trees, and (b) on twigs immediately after detachment.
Another line of research is the measurement of the actual tensions
existing in trees, and hence of the resistances to be overcome by leaves
transpiring on the tree.
V. SUMMARY AND CONCLUSIONS.
1. Farmer’s experiments on the water-conducting power of the wood of
trees have been extended to certain trees growing in the Cape Peninsula,
and the results obtained are in general accordance with his conclusions.
2. Pinus pinaster has wood of low water-conducting power, its specific
conductivity being a little higher than that recorded by Farmer for
P. sylvestris. :
3. Leucadendron argentewm has wood of high water-conducting power
for an evergreen tree, its specific conductivity being rather more than
double that of the Pine.
4. A method of estimating the surface area of Pine leaves is described.
5. Pine twigs, kept with their cut ends in water, show a far higher rate
of transpiration, both per twig and per unit leaf area, than do similar twigs
of the Silver Tree.
The Water Relations of the Pine and the Silver Tree. 19
6. The ratio of transpiration to transmission is far higher in the Pine
than in the Silver Tree. This is shown to indicate that under the experi-
mental conditions Pine needles exert a greater suction on the water in the
stem than do Silver Tree leaves.
7. When transpiration is followed in twigs, as they dry, it is found that
the Pine twigs at first transpire more rapidly than the Silver Tree twigs.
After a short time, however, the transpiration of Pine twigs suddenly drops
and becomes less than that of the Silver Tree twigs.
8. Determinations of water content show that the Silver Tree leaves have
a greater water content expressed as a percentage of fresh weight than have
Pine needles.
9. The bearing of these results on the water relations of whole trees is
discussed, and possible interpretations suggested. It is concluded, however,
that all that can definitely be said at present is that the Pine leaves appear
to be more resistant to unfavourable conditions than the Silver Tree leaves.
10. Further lines of research are indicated.
In conclusion, I should like to take this opportunity of thanking
Prof. Thoday, at whose suggestion the above work was commenced, for his
invaluable help and advice throughout the course of the investigation.
I have also to acknowledge the assistance received from the Union
Government Research Grant Board, which awarded me a Research Scholar-
ship, thus enabling me to carry out a very large portion of the work
described in this paper.
REFERENCES.
1. Farmer, J. B.—‘ Proc. Roy. Soc.,’ 90 B, 218-31, 232-50, 1918.
2. Maruoru, R.— The Flora of South Africa,’ vol. i, p. 145.
(21)
NOTE ON AXISYMMETRIC OCRTHOGONANTS.
By Srr THomas Muir, LL.D., F.R.S.
(1) It will be recalled that Cayley’s rule for the construction of a
positive unit orthogonant is to take a unit-axial skew determinant, A say ;
replace it by its adjugate; multiply each element of the latter by 2/a ; and
then, lastly, from each diagonal element subtract 1. Since the original
n-line determinant, taken as it were for material of construction, involves
én(n—1) arbitrary constants, this is the number of such constants in the
orthogonant.
(2) There is an appreciable advantage to be gained by introducing
another constant, a say, the most natural place for it being that of each
unit of the original diagonal. The preceding rule then has to have its
multiplier 2/ A changed into 2a/A, or, what is the same thing, to give place
to the following theorem: If |a,,|, or A say, be a skew determinant with
ei =). S == Bn = a, then
|
A
2Ay) a i. Ado Masai Ain
i ON ere ep see:
Q
An va Wey eG, ee ae As ee _— =
is an orthogonant whose basic constant is (A/«)?. A short verificatory proof
is obtained by showing with the help of a result of Spottiswoode’s (‘ Hist.,’
li, pp. 289-291, 315) that
TOWpe x HOW, <==) ( At/on)r
and row, X row; = 0.
(3) Let us consider now the case of this where n is odd. A then, being
skew, can be expressed as a sum of terms in descending odd powers of a, so
that for A/a we have an expression beginning with a”~! and ending with a
sum of squares independent of a. If in the result as thus simplified we
put «= 0, A,, and A,, become equal, being conjugate minors of a zero-
axial skew determinant. We consequently obtain when » is odd an axisym-
3
22 Transactions of the Royal Society of South Africa.
metric orthogonant involving superficially the full number of arbitrary
constants. For example, when nis 3, and
Qa Vv — Pp
A=|;-—v a xX = a(a? + A2 + p? 4 7?)
p—A a
we have the axisymmetric orthogonant
N= pw — 2A pe 2r¥
2A —A2 + p? — rv? py
2A» 2pm —’M— wt Pv
with (A? + p? + v?)? for its basic-constant.
(4) The procedure fails when n is even, because then the expansion of
A has its last term independent of a and cannot be divided by a prepara-
tory to putting a = 0. Nevertheless if we introduce the condition that the
said last term vanishes, and we so, in effect, diminish the number of arbi-
trary constants by 1, a result worthy of note is arrived at when n is 4;
namely, If af + bg + ch = 0, the axysymmetric determinant
_— a? = be = c 9 Shes 92 paae 9 =
ay ee ee oe a Ds 2(ef — ah) (ag — Of)
2(vh— eg) ee et ys BUfg— ab) (fh — ae)
hy) 2
Q(ef—ah) A fg—ad) FAT Th 2(gh — be)
2(ag — bf) 2(fh — ac) 2(gh — be) a ae
g g —fi—-gth
is an orthogonant whose basic constant is (a® + b? + c? + f? + g? + h?)?.
What is equally interesting is the fact that this resolves into the product of
two orthogonants
f-a qg—b h—c , fta g+b h+e
a—f h—e b—g| ;-a—f . -h-—-c b+g
b—g c—h : f—a| —b—gq c+th : —f—a
c—h g—b a—f Sie oa aay Oey
which are both skew and have the same basic-constant
October 20, 1920.
(23 )
OBSERVATIONS ON LIVING FISHES BROUGHT BY H.M.S.
“CHALLENGER” FROM TROPICAL EAST AFRICA TO
CAPE WATERS.
By J. D. F. Giucurist.
The general aspect of the fauna of the warm waters of the southern
equatorial current and its branches is very different from that of the colder
waters of the circumpolar Antarctic current. ‘This difference is no doubt
ultimately due to the differences in the character of these waters as regards
temperature and salinity, nitrogen-contents, etc. Waters of such different
characters do not readily mix with each other, and, as many animals seem
highly sensitive to differences in salinity and temperature either directly or
indirectly (through the food supply), an effective barrier to migration is in
many cases established. We may therefore divide the marine fauna of the
southern hemisphere into two great regions, one which may be called the
Equatorial, or the Equatorial-Current Region, and the other the Antarctic,
South-Polar, or Antarctic-Current Region. The Equatorial Region is sub-
divided by the continents projecting into the southern seas, so that we may
recognise as divisions of this region the Pacific, Atlantic and Indian sub-
regions. These divisions are founded on fundamental physical differences
which are clearly seen in the southern hemisphere, but which probably apply
in a modified form to the northern hemisphere, where the Polar region is
broken up by land masses, which are not found in the South- Polar regions.
Three great regions may therefore be recognised, North Polar, Equatorial,
and South Polar, more or less sharply divided from each other, and one of
the most interesting problems of zoogeography is the relation of their
respective fauna to each other. There is the difficult problem of the
resemblance, in some cases identity, of species in the North-Polar and
South-Polar regions, on which a diversity of views still exists, and there is
the simpler one of the relation of the equatorial fauna to the adjacent polar
fauna. These two questions are, however, not unconnected, and it is
possible that the more easily ascertained facts in the case of the second may
have some bearing on the first.
The South African seas are well adapted for a study of the relation of
the Equatorial to the South-Polar regions, and I have already recorded
elsewhere some striking facts observed in South African waters in this
24, Transactions of the Royal Society of South Africa.
connection. These are, briefly, that, at a point on the South Coast, marking
the meeting place of the South-Polar and Equatorial waters, there 1s a
mortality amongst fish, and possibly other marine animals, which may be
sheht or well marked but occurs annually and at definite periods of the
year. A similar occurrence is found on the West Coast, at a point in the
branch of the South-Polar current which passes up the West Coast of
South Africa. The object of the present paper is to record some facts
bearing upon this subject from a different point of view.
The H.M.S. “ Challenger,” namesake of the ‘‘ Challenger” of zoological
fame, arrived in Simons Bay in January, 1919, from British East Africa,
Zanzibar and other Eastern ports. She had on board a number of live fish,
which had been collected at various places, and kept alive in a tank of circu-
lating water. Dr. N. Spencer Nairne, Surgeon, R.N., who, along with the
officers of the ship, were instrumental in securing and keeping the fish alive,
readily assented to transfer the fish to the tanks of the Government Marine
Laboratory at St. James, near Cape Town, thus affording a unique oppor-
tunity of testing the effects of the colder waters of the Cape seas on
tropical fish.
The following is a list of the fish, with their localities ;
Carnax djeddaba . Four from Dar es Salaam Harbour.
Carnax affinis . One from Dares Salaam Harbour.
Lutianus gohnia . QOne from Port Amelia, Portuguese E. Africa.
Holocentrum rubrum . One from Port Amelha, Portuguese E. Africa.
Balistes aculeatus . One from Mnazi Bay, Portuguese E. Africa.
E'pinephelus sp. Two from Zanzibar.
The fish were safely frametarred in barrels by steam launch and placed
together ina large tank (14 x 5 ft.). Balistes had, however, to be removed to
a separate tank on account of its aggressive habits. The others adapted them-
selves almost at once to their new surroundings, the two species of Carnaz
swimming about together, the Epinephelus (rock-cod) remaining, as is the
habit of these fish, under the shelter of the rock-work in the tank.
A marked feature of all the fish was the apparent ease with which they
adapted themselves to their new surroundings, and more especially their
oreater activity as compared with that of the Cape fishes in captivity. The
latter feature was more conspicuous in the case of the Carnaz. Their
movements were very rapid and sustained, and it was soon found that no
Cape fish of a small size were immune from their attacks. This was
specially so in the case of the common Cape mullet (Mugil capito), which
happened to be in the tank with them.
All of the fish throve well; they became plumper, and some abrasions,
caused doubtless by their confinement in the smaller tank on board the
‘‘Challenger,” healed up rapidly, and one might wonder why those warm-
water fish had not naturally spread to the apparently more congenial con-
Observations on Living Fishes brought from Tropical East Africa. 25
ditions of the Cape seas. The reason, however, became apparent at a later
date. Up to the 25th of April the four Carnax continued active and fed
well, but on the 26th, after a strong N.W. wind, the temperature of the
water rather suddenly fell to 13° C., and on the 26th three of the fish were
found dead in the tank, though apparently quite as usual the day before.
The dead fish were carefully examined, and beyond the fact that the skin was
covered with an abnormal secretion of mucus all the tissues seemed normal.
On the 27th, when the temperature had fallen to 12°5° C., the remaining
Carnax was found dead under similar conditions.
On the same date the Lutianus appeared to be very slow in its move-
ments and it was transferred to another and smaller tank, but soon died.
The most interesting and striking of the fishes was the Balistes. It is a
gaudily-coloured fish, with body of a salmon-pink colour; three bright blue
bands pass from the eye to the base of the pectoral, a yellow stripe from the
angle of the mouth, and conspicuous white bands in an oblique direction
over the body above the anal fin. The fish had the pecuhar habit of retiring
at sundown to a particular nook between two rocks, where it lay down on
one side and apparently fell asleep, with fins extended and perfectly motion-
less. If disturbed at night it was aroused with difficulty and usually turned
over on the other side and went to sleep again. In the morning it was early
awake and swimming about, feeding as actively as before. It was specially
fond of crabs. On the onset of the cold weather the fish was found
apparently asleep in its corner during parts of the day, and finally, during
the whole day through. When aroused it was fairly active and took crabs
readily enough. This fish, however, and the Holocentrum died shortly after
the Carnaz.
The Rock-cods did not seem to be affected to the same extent. They
lived for about a month longer. One was killed by leaping out of the tank
and the other died some days afterwards.
There seems little doubt but that the fish, though otherwise well
adapted to live in the colder waters, are unable to stand the low temperature
of the winter season or perhaps the sudden change of temperature.
It is true that we are here dealing with fish in the artificial surroundings
of an aquarium, and this must not be lost sight of in drawing conclusious
from this experiment.
Another aspect of the distribution of marine animals on the South
African coast is connected with the variation of the strength of the two great
conflicting currents which meet in this region, and this, again, is connected
with the relative force of the winds which primarily cause the currents.
The strong southerly winds of the Hast Coast have a tendency to check the
force of the Mozambique current which flows southward along the Hast
Coast. The absence or diminution of these winds will therefore tend to
accelerate the flow of the current of warm water, which will, in that case,
26 Transactions of the Royal Society of South Africa.
extend further along the South Coast. The Cape fishermen have a great
belief in the effect of the prevailing winds (8.E.\ and N.W.) on the
relative abundance and occurrences of fish, and it has been stated that in
the marked absence of south-easters in the summer time strange fish are
to be found in False Bay and are often cast up dead on the beach. I have
examined some of these fish, and they proved to belong to species usually
found in the warm waters of the East Coast. <A fisherman of many years’
experience, on seeing the Balistes, stated that he had seen a similar fish cast
up on the beach of False Bay.
The fact thus demonstraned that such tropical fish of the Hast Coast live
and thrive for a time in Cape waters would seem to indicate the possibility
of their being able to pass the barrier of the Cape and reach the warm
waters on the western side of the continent. In these waters, similar in
character to those in which they normally live and propagate, they would
probably survive and increase, so that we could have identical species on the
east and west side of the continent, with an apparent absence of such
species in the intervening parts of the ocean.
If some tropical fish can thus run the gauntlet of the colder waters of the
Cape and ultimately reach waters in which they can thrive and propagate, it
is reasonable to suppose that fish of the northern hemisphere may be able
in the same way to pass, in limited numbers and on specially favourable
occasions, through warm equatorial waters to the southern hemisphere and
vice versé. The supposed absence of bi-polar species in the equatorial
regions may be only apparent, and they may at special times and on special
occasions occur in these regions. The same, of course, holds good for other
forms of marine life, more especially, perhaps, of planktonic forms, which
may be carried by exceptional currents in limited numbers from their usual
haunts, through regions not suited to their increase, to conditions of life not
different from their normal environment. Alleged cases of strict bi-polarity .
in marine animals may, therefore, be more apparent than real.
(27 )
NOTES ON THE DEVELOPMENT OF THE OVULE, EMBRYO
SAC AND EMBRYO OF HYDNORA AFRICANA, THUNB.
By R. H. Dastur, B.Sc.Bombay, F.L.S.
(With thirteen Text-figures.)
INTRODUCTORY.
The genus Hydnora is confined to South Africa, Bourbon and Madagascar,
and comprises several species, out of which three only occur in South Africa.
They are all parasites, and the species which forms the subject of these notes,
viz. Hydnora Africana, uses mainly the common milkbush of the karoo,
Euphorbia mauritanica, L., as its host, and is of fairly common occurrence in
South Africa (3). The flowers appear as buds on the underground stem, but
soon come above the surface of the ground, and open when mature by means
of three longitudinal slits.
The material for this investigation was collected in the neighbourhood
of Worcester, Cape Province, mostly by Prof. Saxton in December, 1912,
and in part by Mr. Izak Meiring in May, 1912, and fixed in chromacetic acid
with or without osmic acid. A short series of sections was prepared by
Prof. Saxton, but pressure of other work prevented him from completing the
series, and in November, 1919, he handed over the whole material to me.
The fixed material consisted mainly of small pieces of the extensive placentas
cut out from a series of about a dozen plants ranging from the youngest to
the oldest that could be found.
INVESTIGATION.
The placentas are extensive and much branched, and bear radially
_ enormous numbers of orthotropous ovules. The sections were cut, in the
main, transverse to the longitudinal axis of the placentas, thus giving
longitudinal sections of the ovules in great numbers. A very young ovule
before the differentiation of the integument is shown in Fig. 1. Very soon
a massive integument makes its appearance, at the top of which the cells
divide by periclinal walls giving rise to four layers of cells, while it is only
two cells thick on the sides. The very young ovules are nearly spherical
but elongate afterwards. No definite archesporial cell or cells are clearly
28 Transactions of the Royal Society of South Africa.
EXPLANATION OF FIGURES.
All sections were cut with a Cambridge rocking microtome, the thickness varying
from 4-7. The figures were drawn with the help of a Leitz microscope and a
Zeiss camera lucida. The sections were cut transverse to the placenta, giving longi-
tudinal sections of the ovules.
In all figures: 1. Integument; m.m.c. Megaspore mother-cell; n. Nucellus; £:s.
Embryo sac; £.c. Ege cell; s.n. Synergid nuclei; p. Polar nuclei; p.t. Pollen tube.
mM. Male nuclei; a. Antipodal nuclei; &. Embryo; s. Suspensor; p.T.N. Pollen tube
nucleus.
Fig. 3
Fias. 1-7.—1. Young ovule about the archesporial stage. x 196. 2. Young ovule
showing m.m.c. which directly functions as the embryo sac. x 196. 3. Embryo sac
nucleus dividing. x 196. 4. Equatorial arrangement of the chromosomes at the
heterotypic division. x 196. 5. The embryo sac with two nuclei, with the curious
thickening above. x 196. 6. Embryo sac showing two nuclei with the vacuole.
x 196. 7. Division of two nuclei into four. x 196.
Development of the Ovule, Embryo Sac and Embryo of Hydnora africana. 29
recognisable, but a hypodermal cell larger than the rest becomes at once the
megaspore mother-cell (Fig. 2). At this stage a curious thickening is seen
at the top of the megaspore mother-cell and persists till a later stage.
The megaspore mother-cell functions as an embryo sac directly, without
first dividing into four megaspores. The nucleus divides into two by the
heterotypic division (Figs. 8 and 4). The sporophyte number of chromo-
somes is believed to be twenty-four,* and in the reduction division twelve
were counted. Vacuoles appear in the embryo sac after the first division
and the two nuclei arrange themselves at the poles (Fig. 6). The stages
showing the homotypic division of the two nuclei into four were also seen
(Fig. 7). Fig. 8 shows the embryo sac with an ege cell nucleus, two
synergids, two fusing polars, three antipodals and a pollen tube, with the
pollen tube nucleus and two curved male nuclei, Stages were seen in the
progress of the polar nuclei towards the centre of the sac, and they fuse
before fertilisation (Fig. 8). The antipodals may also disintegrate and
disappear before fertilisation. Cases believed to show actual fertilisation
were observed, but they have not been figured as the evidence was inconclusive.
The embryo divides into two cells by a transverse wall and transverse
divisions continue till a very long chain of cells is formed. As many as
fifteen cells were counted in a chain in one proembryo. Then longitudinal
divisions occur in about the fifth, sixth, seventh, eighth and ninth cells
from the distal end and these cells alone give rise to the embryo proper.
The longitudinal divisions begin even before the divisions in the distal part
of the filament are complete. The cells of the suspensor slowly shrink and
at least the proximal cells disappear gradually, while the distal cells persist
in the most mature embryo observed. Fig. 12 shows two longitudinal rows
of four and three cells each and Fig. 13 shows two longitudinal rows of five
cells each.
DISscUSSION.
The massive integument seems to be an unusual feature in this species,
as generally two integuments prevail among Archychlamydeae, except in
some species of Ranunculus and Leguminosae (4).
Among Archychlamydeae nearly all the species investigated have three
or four megaspores (except in three genera of Piperaceae), and a row of four
megaspores seems to be less common than a row of three (4). In this plant
the megaspore mother-cell does not divide at all but directly functions as
the embryo sac.
The curious thickening which is seen at the top of the megaspore mother-
cell, and which persists tilla later stage, was thought at first to be the remains
of a parietal cell cut off from the archesporial cell, and a careful examina-
* Twenty-three were actually counted, but it is presumed that twenty-four were
present.
30 Transactions of the Royal Society of South Africa.
Fig. 13.
Fig. 11.
Fias. 8-13. Embryo sac showing egg cell nucleus with two synergids, two fusing
polars, three antipodals and the pollen tube with a pollen tube nucleus and two male
nuclei. x 252. 9. Two-celled embryo. x 196. 10. Older embryo consisting of a
chain of cells. x 196. 11 and 12. Beginning of the formation of a massive embryo.
x 196. 138. The massive embryo. x 196.
Development of the Ovule, Embryo Sac and Embryo of Hydnora africana. 31
tion of earlier stages was undertaken to test the point. So far as can be
ascertained, however, it seems clear that no parietal cell is formed at all.
The long filament of cells produced by transverse divisions from the
fertilised egg is unusual among Dicotyledons though paralleled in certain
Leguminosae (2). Generally the formation of the embryo proper takes
place in the distal cell of the proembryo amongst flowering plants, but in
this plant it is approximately in the fifth, sixth, seventh, eighth and ninth
cells from the distal cell that the formation of the embryo proceeds. So
there are three regions in the proembryo: (1) the suspensor, (2) the embryo
proper, and (3) the cells beyond the embryo. The most mature embryo
observed (Fig. 13) is still undifferentiated, and the material available does
not admit of the investigation of later stages, but field observations indicate
that probably these would only be met with in germinating seeds. A small
embryo in which neither radicle nor cotyledons are differentiated has been
described for a number of parasitic and saprophytic Dicotyledons, but none
of them shows an embryo of precisely the same type observed and figured
for Hydnora. In Engler and Prantl’s ‘ Pflanzenfamilien’ (1) there is figured
a mature embryo of Prosopanche Burmeisteri, De Bary (the only genus
allied to Hydnora), but the latter does not appear to show the peculiar
features described here for Hydnora, so far as such features are shown in
Solms’ figure cited.
In conclusion I have to thank Prof. Saxton for advice, suggestions and
criticisms on various points in this investigation. Thanks are also due to
Mr. Izak Meiring for his help in the collection of material.
SuMMARY.
(1) The ovule of Hydnora is orthotropous with a single integument.
(2) The megaspore mother-cell is hypodermal and becomes the embryo sac.
(3) The proembryo consists of a row of about fifteen cells.
(4) The embryo is produced from the middle region of the proembryo,
and no differentiation had taken place in the latest stages seen (probably
from nearly ripe seeds).
Note.—This investigation was carried out in the Botanical Laboratory
of the Madhavlal Ranchhodlal Science Institute, Ahmedabad.
LITERATURE CITED.
(1) Soums, H. Grar zu.— Rafflesiaceae”’ and “ Hydnoraceae” in Engler and Prantl,
‘Pflanzenfamilien,’ III Teil. Ab. 1, 1889.
(2) GuianarpD, L.—* Recherches d’embryogénie végétale Comparée. I: Legumineuses,”
1881. (In Coulter and Chamberlain’s ‘Morphology of Angiosperms.’)
(3) Maruoru, R.—-‘‘ Notes on the Morphology and Biology of Hydnora africana,”
‘Trans. 8. Afr. Philos. Soc.,’ vol. xvi, Part v, 1907.
(4) CoULTER AND CHAMBERLAIN.— Morphology of Angiosperms,’ 1912.
act
_
ce
-
4
‘tame ¢
*
by i
=
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1)
—=2
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i
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—
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e Cyne
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(33 )
(1) THE PERMANGANATE ABSORPTION-SPECTRUM: A CLAIM
FOR PRIORITY. (2) A FORMULA FOR CALCULATING
THE URANIUM SPECTRUM.
By James Morr.
(1) In Part VII, Section III, p. 47 of “Colour and Chemical Constitu-
tion,” published in these Transactions two years ago, I gave the formula—
== : = 1750 + 78 N (in which N represents the 7 natural numbers
from 0 to 6) by means of which the positions of the 7 absorption-bands of
the permanganates in dilute water solution could be calculated.
I now find that an Italian investigator, E. Adinolfi, has practically redis-
covered this formula. He states his result as n = n, + 75°35 N, but his n
refers to the third (viz. the strongest) band, whereas my figure 1750 refers to
the first band. Subtracting therefore 2 x 75:3 from his value of ns (viz. 1902°8)
pee aig 2 2s Acimolf’'s formulacthua reads: : = 1752-2 + 75:3 N and
is practically the same as mine, since the decimal points mean nothing in
observations of this sort, which are not capable of a higher accuracy than
O-l per cent. The following table exhibits the results of the calculations :
, Adinolfi’s formula. Moir’s formula. Moir’s observations.
LineI . 1752 . ~ hZ50 . 1750
Line II . 182734 . 1828 ; . 1830
Tame 1. 1903 . ; . 1906 ; . 1906
Tine LV. 1978. : ; . 1984 ; . 1990
Line V . 2058 . : ; . 2062 ; . 2060
Fane Vi. 2129 . : . 2140 ; : . 2188
Line VII 2204 . : . 2218 ; : . 2204
Adinolfi’s observations are in the Journal of the Academy dei Lincei,
1920, and are therefore inaccessible to me, but they apparently do not
differ from mine by more than 0:1 per cent. It is probable that the 7 bands
are really 10, 3 of which overlap and thus cause the great intensity of II
and III, and also the small irregularity of the spacing.
34 Transactions of the Royal Society of South Africa.
(2) The spectrum of the Uranyl Ion has already been given (1. c., p. 48).
The following formula explains it in mathematical terms, viz. en:
2113 + 66 N (in which N is 0, 1, 2, 3, and 4):
Calculated: 21132179; 2245, 2311 and 23877.
Observed : 2114, 2180, 2243, 2310 and 2381.
As shown in the same paper the bands of the unionised solid salts can
be calculated fairly closely from these figures by dividing by the sixth-root
of the ratio of the molecular weight of the salt to that of UO,+*+. Thus
the bands of autunite are given by—
_1_ 2113 + 66N
A 6/ 914
N/ 2 x 270
Calculated : 1937, 1997, 2058, 2118 and 2179.
Observed : 1945, 2000, 2062, 2128 and 2190.
I wish to thank Mr. T. P. Waites for the fine specimen of autunite in
crystals large enough to permit observing the absorption-spectrum.
— 1937 + 604N
( 35 )
COLOUR AND CHEMICAL CONSTITUTION.
Part XIII.—TuHE CaAaLcuLATION OF THE COLOUR OF THE MonocycLic
Dyess.
By James Morr.
Readers of Part X of this work will remember that by assuming
triphenylcarbinol (or its anhydride) to have a potential colour with 593
for the position of its absorption-band, I was able to show that the colour
of any dye derived from it could be calculated by successive multiplying
of 593 by factors characteristic of every substituting atom (or group) and
position contained in the dye.
Among the simpler members of the family, however, several exceptions
were met with. Thus the salts of fuchsimonium or amino-triphenylcarbinol
had a calculated \ = 593 x 0:'972= 576, whereas observation gave the much
lower value of about 4425 (in water). When Part X was written I
suggested, as many other writers on colour have also done, that certain
substances only exhibit harmonics of their calculated colour. Further
experiment has now shown me that this hypothesis is not only unnecessary,
but probably quite untrue.
The true explanation is that the scheme described in Part X does not
apply when the dye is “ monocyclic,” 7. e. when it contains only one active
or ionisable colour-ring. This means that the high colours (between 470
and 4760) of the commercial dyes are in practically every case due to a
‘“‘dicyclic”” constitution, namely, one in which two ionisable colour-rings
co-operate. .
To make my meaning perfectly clear it will be sufficient to give a list
of “ionisable colour-rings,” viz.
Vera, DF eee end
< NH —C Ou, A >—NuMe, -—K NM,
and the other four in which the loose linkage is ortho, not para, to the
ionised group. There are no doubt others in which thiophene, furfurane
and pyrrhole are the rings.
An example of a monocyclic dye is dimethylaminobenzhydrol Ph .CH
(OH) .C,H,NMe,. Two analogous dicyclic dyes are:
36 Transactions of the Royal Society of South Africa.
Me,.N —— __/NMe,
aes Vey,
CHi(OE) 4
I.—Bis-dimethylaminobenzhydrol. ‘“ Michler’s hydrol.”
Me.N__ NMe,
SEZ,
\ CH(OH)”
II.—Bis-dimethylamino-xanthhydrol. The dye pyronine.
I have now found that the colours of the monocyclic dyes (which, by the
way, are all low, viz. below 4500) can be calculated by a factorial scheme
similar to that of Part X, but that the factors are all different from the
corresponding factors of the dicyclic scheme.
The parent substance is paraoxybenzylalcohol
Ye ~
HO CH,0H
isan aes
(oxyphenylearbinol), which has a calculated » of 290 for faintly alkaline
water solution. This substance is interesting since its anhydride is the
hypothetical quinomethane,
A SS.
Org >: CH,
which is the parent-substance of the common ‘aniline dyes” according to
the upholders of the quinonoid theory of colour.
The factors required for colour-calculations are :
A. Phenyl-factor, for replacement of H in carbinol by Ph = 1:135.
B. Orthocarboxyl-factor, for conversion of a phenylcarbinol
into a phthalein = = 1060:
C. Phthalein-factor, or factor A x cco B, Sr ene H
by C,H,CO,H = 1-208.
ip. AenibeoMipoene factor, for renineine OH i in ring o NH, = 110,
HK. N-methyl-factor, for replacing H in NH, by CH, . == d0so,
EF. Methylaminocompound-factor, for replacnmg OH by
INE Mie=— ex Gh = . = iso:
G. Dimethylaminocompound-factor, for ies achive OH by
INDNie; == aD ex Gh : : == a2
Colour and Chemical Constitution. OY
Armed with these data we proceed to calculate the colours of the known
monocyclic coloured substances, and to predict the colours of the unknown
or unobserved members of the series.
(1) Fuchsone.—This is the diphenyl] derivative of the parent-substance.
Its calculated \ is therefore 290A2 = 290 x 1135 x 1:185 = 374. Meyer
and Fischer observed \ 380 in alcohol (‘ Berichte,’ 19138, p. 70). Observations
in alcohol generally give results about 6 units higher than in water.
(2) Paraoxybenzhydrol.—This is the mono-phenyl derivative, and its
calculated A is 329. It has not been observed, but this value is predicted
for its colour in alkali. Ortho-oxybenzhydrol is known, and is yellow
with x about 360 (‘ Beilstein,’ ii supplement, p. 694). This is about the
usual difference in colour between ortho- and para-compounds, so that the
prediction is probably correct.
(3) Fuchsimine (fuchsone-imine).—This is fuchsone with NH, in place
of OH. The calculated \ is—
{ 290 x 1:135? x factor D
or 374 x 114
Observations by Meyer and Fischer (/. c.) gave , 430 in alcohol. My own
observation, in water, gives d 420.
(4) Fuchsonedimethylimonium salts.—The calculated value is that of
fuchsone multiplied by the factor G, viz. 8374 x 122 = 2456. Meyer and
Fischer give \ 460 in alcohol. My observation in water is A 455.
(5) Mono-orydiphenylphthalide (phenylphenolphthalein).—This is the
ortho-carboxylic acid of fuchsone. Its calculated colour corresponds
to .374 x 1:060 = 1397. My observation gives \ 395 in water.
In Parts I and III of this work I stated that I had observed a pink
(instead of a yellow) colour in this substance (A560). This work I now
find to be erroneous, for if oxydiphenylphthalide is made from a specimen
of benzoylbenzoic acid which has been purified by frequent crystallisation of
its ammonium salt (whereby ammonium phthalate is completely eliminated),
the product gives a pure yellow in alkali without a trace of the band near
560. The similar compound from salicylic acid (second diagram on p. 125
of Part IIT) is also wrong: 4562 should be 420.
(6) Phenyl-dimethylaniline phthalein.—As will be seen by writing out
its formula, this 1s the ortho-carboxylic acid of No. 4 mentioned above.
Its calculated colour is therefore that of No. 4 multiplied by 1:06 = \ 483.
The substance is made from benzoylbenzoic acid and dimethylaniline, and my
observation of its colour gives 4 480 for the centre of its absorption band.
The unknown phenylanilinephthalein should have d 452.
(7) Para-dimethylaminobenzhydrol.—This is most simply calculated by
multiplying the colour-figure for oxybenzhydrol (No. 2) by factor G. The
calculated X is 3829 x 122 —=—2401. Observation in a specimen from
benzaldehyd and dimethylaniline gave A 400.
+
= d 4263.
38 Transactions of the Royal Society of South Africa.
(8) Mono-phenolphthalein.—
Wis Nae ee
(Ee a Ge
from phenol and phthalaldehydic acid is yellowish and has the calculated
\ = 290 x factor C = 1349. This agrees with my observation so far as such
can be made, as the centre of the band is just below the limit of visibility,
using sunhght.
The number of substances in the monocyclic class is limited, and the
remainder are, I believe, all coloured in or near the ultra-violet, and therefore
beyond my observation. I therefore predict their colours so that some other
observer possessing an adequate photographic apparatus can verify them.
(9) Para-aninobenzhydrol.—Calculated 4 = 290 x 1:1385 x 1:14 = A875.
(10) Para-aminobenzyl alcohol.--Calculated A = 290 x 1:14 = 28305.
(11) Para-dimethylaminobenzyl alcohol.—Caleulated A = 290 x 122 =
rN dd4.
(12) Para-oxyphthalide.—
HOS
ae
< >
\co —¥
probably has a = 290 x 1:06= 307 and so on for the corresponding
amino- and dimethylamino-compounds.
It is probably an important poimt that the parent-substance (whether
regarded as quinomethane or as oxybenzylaleohol) has its absorption-band
close to those observed for sodium phenate and paracresolate (Baly and
Ewbank, ‘J.C.S.,’ 1905, p. 1352), the former being ~290, and the latter,
calculated to water from alcohol, about 2285. The difference is similar to
that caused by a methyl-group,* so we can infer that in alkalne water
HO-< CH; and HO-C »—e H.OH
have very nearly the same colour (absorption, speaking more strictly), viz.
a 290.
It would therefore seem that colour cannot be traced to any lower terms
than this, namely, the presence of one benzene ring with one ionisable group,
and (usually) another active atom. All the classes of colours except the
azo-dyes and certain nitroso-derivatives are therefore accounted for in these
terms, so it is probable that the conception of the physical cause of colour
as the periodic orbit of an electron rownd a molecule is the correct one.
The sinuous orbit of Part XII is not very likely, but the tautomeric pauses
assumed therein are quite a probable feature of the orbit.
It is to be noted that in the dicyclic compounds two rings, each carrying
* Baly and Ewbank’s diagram is too small to show the difference.
Colour and Chemical Constitution. 39
an ionisable group, always co-operate: consequently the electron-orbit is very
much larger and the colours much higher. Part X of this work might
therefore be re-written without the hypothesis of fuchsene by making
dioxybenzhydrol the parent-substance, whereby on using the dicyclic colour-
factors given in Part X, all the colours are calculated without exception,
because the exceptions given there wre all monocyclic. As an example,
dioxybenzhydrol
HO-< > -cmon< ou
of 4539 in alkali can be treated with the dicyclic factor 1-059 for NMe,/OH
to give Michler’s hydrol (factor applied twice), and so on. This is elaborated
in Part XIV.
Cai)
ON BRAULA CAECA, NITZSCH, A DIPTEROUS PARASITE OF
THE HONEY BEE.
By S. H. Sxairz, M.A., M.Sc.
(With eleven Text-figures.)
INTRODUCTION.
Braula caeca is an aberrant dipteron that is a common parasite of the
honey bee in this country and other parts of the world. It is a small,
active insect, reddish-brown in colour, and about 2 mm. in length
(Fig.1). Instrong hives these so-called bee lice rarely become abundant, but
in weak colonies they are sometimes so numerous that hardly a bee is without
one or more of the parasites. According to Sharp (1) there is only one
_ species known, and this species is placed in a separate family by itself, the
Braulidae. This family, together with the Hippoboscidae and two small and
little-known families, the Streblidae and the Nycteribiidae, make up the
series Pupipara. Braula caeca is linked with the Pupipara, mainly because
of the statements of Boise, Packard and others concerning its life-history,
but, as will be shown later, these statements are erroneous.
Boise states that a pupa is deposited in the cell in the hive by the side
of the young larva of the bee, and appears as the perfect insect in about
twenty-one days. Packard says that on the day the larva hatches from the
egg it sheds its skin and turns to an oval puparium of a dark brown colour.
According to Comstock (2) the mode of reproduction of Braula is similar to
that of the Hippoboscidae. Cowan (3) asserts that the “‘eggs hatch inside
the insect and the larvae are nourished by the secretions from a gland.
The pupa is extruded on to the floor-board of the hive, and fourteen days
later a perfect insect emerges. The young lice remain on the floor-board
until they have the opportunity of climbing on to a passing bee.”
THE Eaas or BRAULA CAECA.
Some time ago Mr. R. H. Harris, Assistant Entomologist. called the
present writer's attention to the fact that certain white specks which are
often found on the brood combs in the hives are the eggs of some insect or
42 Transactions of the Royal Society of South Africa.
other. The writer had often noticed these specks in the hives at Cedara, but
had failed to detect their true nature until Mr. Harris suggested that they
should be mounted in xylol for examination under the microscope. The
xylol dissolves the adherent wax from the white specks, clears them, and
renders their examination a simple matter.
The eggs measure about ‘65 mm. xX ‘56 mm. and are dead white in
colour (Fig. 2). They are oval in shape, and have on either side a flattened
wing-like expansion, marked with a delicate, raised reticulation. They seem
to be deposited in a haphazard manner over the brood combs, some being
Sikes
Sb
i
i
Fic. 1.—Braula caeca, Nitzsch. x 265.
placed on the caps of the cells containing sealed brood, others inside empty
cells glued to the sides and bottoms of the cells, and still others on the tops
of the partitions between the cells. Some of the eggs were found to
contain fully developed embryos (Fig. 2) and their dipterous nature was
unmistakably revealed. Thus the writer’s attention was turned at once to
Braula caeca, the only dipterous parasite present in the hives. A number
of the lice were caught, and carefully dissected under a binocular dissecting
microscope. In three or four of the females examined a fully formed egg
was found in the common ovarian duct (Fig. 9, cod), and these eggs were
obviously identical with those found on the brood combs. The appearance,
size, Shape and markings were exactly identical, so that there could be no
possible doubt as to their identity. The characteristic wings with the
On Braulu caeca, Nitesch, a Dipterous Parasite of the Honey Bee. 43
reticulations and peculiar notches at either end make the recognition of the
egos easy and certain.
THe Larva or BRAULA CARCA.
Having thus established the fact that the bee louse is oviparous and not
pupiparous as is so often stated, an endeavour was next made to trace out
its life-history. For some time no traces of the immature stages of the
parasite could be found in the hives, but finally a neglected hive was found
in which the lice were very numerous. This colony had evidently been
queenless for some time and was very weak; many laying workers were
Fie. 2.—Egeg of Braula caeca. x 70.
present, and these had given rise to large numbers of drones. Two of the
drones from this hive were very badly infested with the parasites, one
carrying no less than eighteen of them and the other twelve. Both of the
drones were undersized and both had obviously emerged recently.
A number of the drone larvae were removed from their cells and
examined under the lens. In several cases one, and in a few cases two,
small dipterous larvae were found lying side by side with the bee larvae.
The largest of these dipterous larvae measured about 2 mm. in length
(Fig. 3). The buccopharyngeal armature, the tracheal system and the
sensory papillae of these larvae were exactly similar to those of the embryos
found in the eggs of Braula caeca (cf. Figs. 2 and 3), hence there could be
no doubt but that these were the larvae of the bee louse. The bucco-
pharyngeal armature is of typical muscoidean form, with well-developed
4A, Transactions of the Royal Society of South Africa.
lateral hooks and a large pharyngeal sclerite, but there is no separate inter-
mediate sclerite (Fig. 6). The tracheal system is well developed, meta-
pneustic, but without the hard, horny spiracular plates usually present in
muscid larvae. The larval antennae (Fig. 7, a) are small and inconspicuous
and of the usual dipterous form. At each end of the larva there are a
number of peculiar sensory papillae. Those at the anterior end are long and
conspicuous and are armed at the tip with a number of minute blunt cones
OSH?
a
CHO
fb See Be
Si
ersaeete aul (e
Fic. 3.—Larva of Braula caeca.—sp. Sensory papillae. ba. Buccopharyngeal armature.
tr. Trachea. fb. Fat body. mt. Malpighian tube. st. Spiracle. x 30.
(Fig. 7,c), whilst those at the posterior end are shorter and armed each with
a single sensory hair (Fig. 7, b)., -
The drone larvae from the cells containing the parasites seemed to be
quite normal and healthy. The contents of the alimentary canal of the
dipterous larvae were carefully examined and found to consist mainly
of pollen, the characteristic pollen grains from the black wattle being
especially abundant and easily recognised. The drone larva’s stomach was
found to contain exactly similar food. Hence there seems little doubt but
On Braula caeca, Nitasch, a Dipterous Parasite of the Honey Bee. 40
that the newly-hatched Braula larva makes its way into a cell containing a
bee larva, and feeds side by side with it on the food supplied by the nurse
bees. Their minute size, white colouring and comparative scarcity in normal
hives make their detection very difficult, and would account for their having
remained unnoticed hitherto.
THE Pupa or BRAULA CAECA.
Six puparia were brought to light after a prolonged search in the hive
mentioned above, and all of these were found in sealed cells containing
Fies. 4-8.—4. Antenna of adult. 5. Tarsus of adult. 6. Buccopharyngeal
armature of larva. 7. a. Antenna of larva; b. Sensory papilla of larva,
anal segments; c. Sensory papilla of larva, anterior segments. 8. Mouth-
parts of adult.
drone pupae. They were white, oval, about one and a-half millimetres in
length, and attached to the sides of the cells. The puparium was found to
consist of the last larval skin, not modified or thickened in any way, and the
Braula pupa could be easily discerned inside. The peculiar sensory papillae
(Fig. 7,6 and c) and the moulted buccopharyngeal armature (Fig. 6) proved
the identity of the puparia and the larvae described above. The adults had
already emerged from four of the puparia when found, and these adults, pale
yellowish white in colour, had made their way on to the drone pupae inside
the cells. The remaining two puparia were kept in order to watch the
emergence of the adults, but unfortunately both died soon after their
A6 Transactions of the Royal Society of South Africa.
removal from the hive. The exit hole in the puparium consists of a rough
tear and is apparently made by a ptilinum, for a well-marked ptilinal suture
can be seen on the head of the adult.
THe ApuLT BRAULA CAECA.
The antennae of the adult recall both the muscid and the hippoboscid
types (Fig. 4). They are lodged in deep cavities in the front of the head as
in true hippoboscids, and remind one strongly of the antennae of Melophagus
Fras. 9-10.—9. Reproductive organs, female. 10. Reproductive organs, male.
ot. Ovarian tubes. sp. Spermatheca. cod. Common ovarian duct. ov. Ovi-
positor. te. Testis. ag. Accessory gland. cd. Common ejaculatory duct.
pe. Penis. x 40.
ovinus. The mouth-parts (Fig. 8) are pecuhar, and do not resemble closely
those of the muscid or the hippoboscid flies; the task of making out the
different parts 1s too involved for the present writer to attempt it. The foot
also seems to be unique among the Diptera, for the tarsus 1s armed with two
well-developed chitinous combs instead of the two simple claws usual among
the Diptera (Fig. 5).
The alimentary canal is of the usual type found among the higher
Diptera. A narrow oesophagus leads to a small proventriculus (Fig. 11),
behind which the canal branches, one branch leading into a large, thin-walled
On Braula caeca, Nitzsch, a Dipterous Parasite of the Honey Bee. 47
crop, and the other to the chylific ventricle. There are four Malpighian
tubes, uniting to form a single pair just before entering the small intestine.
The rectal ampulla is large, conspicuous, and furnished with four typical
rectal glands, well supplied with trachea. This part of the alimentary canal
is strongly reminiscent of that of Hippobosca.
The reproductive organs are shown in Figs. 9 and 10 and need little
description. There are two ovarian tubules on either side closely applied
one to the other. The follicles consist of groups of nurse-cells with a
developing ovum posterior to them; the nurse-cells get smaller and the ovum
larger as the egg approaches maturity. There is only one spermatheca,
Fig. 11.—Alimentary canal of adult. pv. Proventriculus. cv. Crop. cv. Chylific
ventricle. mt. Malpighian tubes. rg. Rectal glands. ra. Rectal ampulla.
tr. Trachea. x 40.
closely applied to the side of the common ovarian duct—not three as in the
true muscids.
The testes are comparatively large sac-like bodies, and there is one pair
of accessory glands filled with a white granular fluid. The penis is large
and conspicuous and is armed with a pair of strongly chitinised curved
spines.
Our knowledge of the feeding habits of the adult is very scanty, and the
following, quoted from A. I. Root (4), is perhaps the most authoritative
statement so far put forward:
‘When the louse wishes to feed it goes to the bee’s mouth, where the
4.8 Transactions of the Royal Society of South Africa.
motions of its feet, armed with bent claws, produce a tickling sensation
perhaps disagreeable to its host, but at least provoking some movement of
the buccal organs, which slightly open and release a small drop of honey,
which the louse at once licks up.”
Although the present writer has never witnessed the feeding of these lice,
he has been able to prove that their food consists of honey. In the study of
the anatomy of this insect several individuals were dissected which had the
crop distended with a colourless fluid, and when this fluid was apphed to
the tongue it was proved unmistakably to be honey. No sold matter was
ever found in the intestine.
SUMMARY AND CONCLUSIONS.
1. Braula caeca is oviparous, not pupiparous as was hitherto supposed.
2. The eggs are deposited on the brood combs in the hives, hatch out
into typical muscid larvae which make their way into cells containing young
bee larvae.
3. The larvae feed on food supplied to the brood by the nurse bees, and
beyond robbing the bee larvae of a little of their food do no harm.
4. The larvae pupate inside the cells beside the bee pupae; they emerge
before the bees do and make their way at once on to the bodies of their
hosts.
5. The adults feed on honey, probably supplied to them by their hosts.
6. Although Braula is not pupiparous, it shows many marked resem-
blances to the Hippoboscidae, and the family Braulidae should probably be
placed between the Muscidae and the Hippoboscidae, not between the latter
and the Streblidae as at present.
BIBLIOGRAPHY.
(1) SHarp, D.—‘ Cambridge Natural History,’ Pt. IT, p. 520.
(2) Comstock, J. H.—‘ Manual for the Study of Insects,’ 1917, p. 489.
(3) Cowan, 'T'.—‘ British Bee Keepers’ Guide Book.’
(4) Root, A. I.—‘A.B.C. and X.Y.Z. of Bee Keeping,’ 1919, p. 256.
( 49 )
NOTE ON A FIRE-FLINT OF STRANDLOOPER ORIGIN.
By Joun Hewitt.
(With Plate I.)
In his important work on the Stone Ages of South Africa, Dr.
Péringuey described a remarkable hafted stone implement, found in one of
the Outeniqua Caves. He placed the type in his group of South African
Neolithic implements, for, having been found along with the skeleton of one
of the aboriginal cave-dwellers, it evidently belonged to that race of people
variously known as Strandloopers, Coastal Bushmen or Primitive Hottentots.
It is the object of these few notes to give further records of the same
type of implement, and to suggest a mode of use.
The earliest reference to the above-mentioned specimen is found in the
record of the monthly meeting of the South African Philosophical Society for
September, 1892, which reads as follows: “ Mr. Péringuey exhibited a stone
implement with wooden handle, which was found in a Bushman or
Hottentot grave near George. The handle had been attached with some
kind of cement, and the implement had been used as a hand weapon.” At
the December meeting of the Society another note on the same specimen
was contributed by Dr. Marloth. His report was as follows: “He had
examined the stone implement, and found that the cement used in con-
necting the handle to the head consisted of resin—probably fine resin and
chalk. He had also found starch grains from wheat and rice, which led him
to conclude that the natives must have used wheat and rice, and therefore
this cement must have been made after the arrival of white men in South
Africa.”
The account of the discovery by Mr. R. E. Dumbleton, as given in Dr.
Péringuey’s monograph, contains the following: ‘‘On coming to the head
(of a human skeleton) I discovered immediately in front of the face two
tortoise shells, etc. ete. With these there was the lumbar vertebra of a
large ruminant, several flint scrapers, and also a peculiar instrument con-
sisting of a piece of flint fixed in gum-cement, in which was inserted
a piece of wood about 4 in. long, serving as handle. The latter
unfortunately was perfectly rotten and broke off short.”
Another example of this implement but in better state of preservation,
has been in the Albany Museum for some years, and a short description was
50 Transactions of the Royal Society of South Africa.
published in the ‘Records of the Albany Museum,’ vol. ii, p. 282, and in
the Guide Book to the Albany Museum. This specimen also was found in
a cave near Plettenberg Bay. The essential features of the implement are—
a small piece of chalcedony, roughly trimmed, presenting a concave cutting
edge; this is mounted in a large lump of resin, a horn handle with slender
and tapering distal end fitting into a comparatively short socket in the
resin. The total length of the specimen is 95 in. There is nothing to
suggest a great antiquity for the specimen. The horn handle is well
preserved, and, in shaping it, several sharp cuts have been made, which
clearly point to the use of a steel or iron blade; it is possible, however, that
the present shape of the handle is not as left by the original maker, for
whilst the resinous socket is rounded, the socketing portion of the horn has
more or less squared surfaces.
Now, what is the function of this unique implement? Dr. Péringuey
rightly remarks that “as a serviceable cutting or gravine tool it could not
prove of much service.” He then made the following suggestion: ‘My
first impression was that this hafted tool was that of a ruler or medicine
man, somewhat on the lines of the baton de commandement, and I am
inclined still to look upon it in that lght.” This explanation seems to me
quite untenable, in view of the weak attachment of the handle in our
specimen, which would be useless as a baton, or as a weapon, for on
moderately rough treatment the lump of resin would break away from the
handle. As a cutting implement, on the other hand, it could only be
effectively employed when grasped firmly by the resinous lump, the handle
being quite free. !
The solution of the problem, in my judgment, lies in a suggestion made
to me by Mr. Frank Brownlee, who remarked on the strong resemblance in
shape between the piece of chalcedony and the imported flints commonly
used by the natives of the Eastern Province for striking fire. It could
certainly be used as a fire-flint, and, I think, has actually been thus
employed, for the cutting edge is worn down in the middle as such flints
commonly are. From the fine illustration that accompanies Dr. Péringuey’s
account I infer that his specimen also has been used in the same way;
indeed he says—‘ The very sinuous edge had been broken before burial
probably, as indicated by the comparatively fresh fracture.” It may be
noted, too, that the stone was actually described by its discoverer as a flint.
Further, the implement as a whole could be utilised as a combined
firelight and torch, for the resin is very inflammable, and the horny haft
would make quite a serviceable handle when thus employed. I cannot
positively assert that the resin has actually been burnt since the implement
was made, but one point may be stated in favour of this view. It is
suggested that the resin originally covered more of the surface of the stone
than at present, for the basal portion imbedded in the resinous socket seems
Note on a Fire-Flint of Strandlooper Origin. 51
hardly sufficient to afford firm attachment to the stone, however it be used ;
in its present condition the stone is imperfectly mounted.
The mounting of small chips of quartz or other hard stones in resin
or gum-cement seems to have been a practice of Bushmen, and there are
several specimens of stone-tipped arrows thus prepared in the South African
Museum. Iron, again, has been known to all the South African tribes for
centuries, although as a comparative rarity to the Hottentots and Bushmen ;
its scarcity may perhaps explain the very late adoption of the flint and steel
amonest the South African tribes as a whole.
Tinder would have to be used in conjunction with such an implement,
but this was no novelty to the aborigines, and indeed is constantly used by
the Kalahari Bushmen of the present day along with their fire-sticks. It is
true that owing to the brittle nature of the resin a moderate blow on steel
would weaken the attachment, and for use in this way the implement would
be quite impracticable. To produce fire there is no need to strike a blow:
it is sufficient to draw the steel rapidly over the edge of the flint, the latter
and its mount of resin being held firmly in the hand.
It may be urged that an easier and more direct way of producing fire
was well known to the aborigines, and that Bushmen produce fire by means
of fire-sticks with astonishing rapidity. Nevertheless, fire-sticks have their
limitations, and in the dreary wet seasons of the Southern Cape Coast the
flint and steel may be more rehable. This is indeed just the kind of
implement that the coastal cave-dwelling people might be expected to evolve.
Moreover, Dr. Péringuey has actually received from the Coldstream Cave
what appears to be the steel striker of a tinder-box, and this same cave
contained Strandlooper remains.
It is also suggested that the tortoise shells found by Mr. Dumbleton
along with the implement were improvised tinder- boxes.
According to the original report, three such shells accompanied the
skeleton, but there were no records of tortoise shells from other Strandlooper
burial places. In prosecuting his researches, Dr. Péringuey obtained very
detailed records of circumstances and accessories from a number of burial
sites ; indeed, one of his investigators was specially asked to search for
more examples of the above-mentioned stone implement and for tortoise
shells, but failed to find either, though beads and sundry other objects were
taken. Therefore the association of tortoise shells with this unique type of
stone implement is probably not accidental. But on this point I do not
lay great stress, for at the present day tortoise shells are much used by
natives throughout the sub-continent, and for a variety of purposes; they
are included amongst the numerous items in the outfit of a Kaffir witch-
doctor, and are commonly used by Bush women in Kalahari as powder
boxes.
Another specimen of essentially the same type, though differing greatly
52 Transactions of the Royal Society of South Africa.
in details, has been in our collection for some years, and was referred to by
me in the ‘Records of Albany Museum,’ vol. 11, p. 283. It consists of a
limb bone of some bird with a lump of resin at one end. The bone has
been marked with incised crosses and transverse lines which perhaps
originally served as a tally. The resinous lump expands a little distally,
and on one side presents a moderately deep depression, at the base of which
is a fragment of hard stone, firmly attached to the resin. This stone cannot
be removed for examination without injury to the specimen, but is evidently
of flinty nature. It is presumably the broken base of a much larger stone
which was formerly mounted in the resin.
The total length of this specimen is 4 in. Thus it is much too small for
use aS a weapon or baton. It came from the same cave as the first-
mentioned specimen, and was no doubt used for a similar purpose, It may
have functioned as the striker of a fire-making apparatus and thus can be
termed an aboriginal match. Lastly, a piece of resin mounted on a stick was
found many years ago in a rock shelter at Woest Hill, Grahamstown, along
with a number of bones. The resin is a cylindric-ovoid lump about 2 in.
long and #? in. thick; it has a deep socket which still contains decayed
wood. There is, however, no indication that this was used as a mount for a
stone. It is of interest as additional evidence that the cave-dwelling
aborigines made use of resin. Mrs. H. M. Barberton informs me that such
resin was quite familiar to the European settlers under the name of “ Bush-
man resin.” It was generally believed that the Bushmen employed this
material, when available, in the attachment of arrow-heads to the shaft.
The chief source of the resin seems to be the roots of Pterocelastrus
variabilis. Thus obtained, it is very brittle, and for cementing purposes
must be mixed with other substances.
Again, in association with typical implements of the Bushman type,
there have been found at several inland localities a few worked stones of
striking resemblance to fire-flints, though made of local rock. One such
specimen, of agate, is in the Albany Museum from Barrow Hill, O.F.S.
(Miss Joan Whitworth), and others from Kimberley have been collected
by Mr. Jas. Swan and Mr. J. H. Power. But, so far as I know, nothing of
this kind has been found under circumstances pointing to great antiquity.
Here it may be added that Sir John Evans, in his work on the ‘ Ancient
Stone Implements of Great Britain,’ commented on the resemblance between
the modern “ strike-a-lights ” and the ancient ‘scrapers,’ and came to the
conclusion that a certain proportion of these latter were in use not for
scraping hides but for scraping iron pyrites, and not improbably in later
days even iron or steel for procuring fire. He also cited various instances of
the occurrence in ancient graves of flint implements in association with
nodules of iron pyrites—which for the purpose of producing sparks seems to
be as effective as iron,
Note on a Fire-Flint of Strandlooper Origin. 53
There is much probability that fire-flints were introduced to the
Strandloopers by Europeans. They are mentioned by various writers of past
centuries as suitable objects for barter with the natives. John Barrow,
referring to some Kaffirs whom he met on the Kareiga River (1797), says:
“They received each a small present, consisting of tobacco, knives, flints and
steels, tinder-boxes, and a few glass beads. These are the sort of articles
which the farmers have been in the habit of exchanging for their valuable
breed of cattle.”
Me oy lint ;
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Trans. Roy. Soc. S. Afr. Vol. X. Plate I.
Strandlooper implements from cave in Plettenberg Bay. x 2. (In B, only a
fragment of the stone remains at the base of the socket of resin. ]
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(55)
A NOTE ON THE USE OF THE SOUTH AFRICAN CLAWED
TOAD IN THE BIOLOGICAL ASSAY OF THE DIGITALIS
SERIES.
By‘ J. W. C. GUNN.
The clawed toad, Xenopus laevis, is the most common batrachian in
laboratory use in South Africa. Morphologically it is most closely related
to the toad, but it is active in its movements like the frog, and there is some
evidence that its electrical responses are at least as quick as those of Rana.
Vulpian (1) was the first to state, and his observations have been
repeatedly confirmed, that the European toad has a considerable degree of
natural tolerance for poisons of the digitalis series, the minimal lethal dose
for the toad being several times greater than for the frog. Abel and
Macht (2) have shown the same to hold true for the tropical toad,
Bufo agua.
It is therefore not only of scientific interest to determine whether
Xenopus reacts to digitahs bodies hke Bufo or Rana, but also of practical
importance, since the most convenient method of standardising preparations
of digitalis is to determine their toxicity for the frog.
METHOD.
The method employed was that in common use in biological assay of
digitalis preparations. A measured quantity of the drug was made up to
‘0 ¢.c. with Ringer’s solution and injected through the floor of the mouth into
the ventral lymph-sac of the toad. Freshly caught animals were used.
Hither the minimal lethal dose or the amount required to arrest the
ventricle in systole in one hour (the minimal systolic dose) was estimated.
: The room temperature varied between 21° and 23°C.
Minima LetHat Dost or STROPHANTHIN.
The strophanthin used had been tested on Rana temporaria in the
Wellcome Physiological Research Laboratories and the minimal lethal dose
found to be about 0:0008 mgrm. per gramme weight of frog.
56 Transactions of the Royal Society of South Africa.
Experiments on Xenopus laevis :
Ext, mat eat Time. Result.
1. ‘001 mg. : 25 min. =. Death.
2: ‘0009 _,, ; 54, : es
a “0008 _,, : 1OP : vA
4. ‘0007 ,, ; 150. 5
5. ‘0006, ; — ; Recovery.
6. "0005 _,, : _ ; 3
de 70007 : SO min. : Death.
8. 0007, 130 |; 2
9. O07. : Overnight. : >
10. 0007 oon — Recovery.
11. ‘0006. ,, ae "
12. ‘0006 _,, } o x
This would show that the minimal lethal dose of this strophanthin for
the toad is ‘0007 mgrm. per gramme, practically the same as for Rana
temporaria.
Minimat Systoric Dose or Tinctures (B.P.).
Two freshly prepared tinctures of digitalis and one of squills were
tested by the same method. At the end of one hour the toads were pithed
and the condition of the heart examined. ‘The tinctures were known to be
active, in two cases by estimating the minimal lethal intravenous dose for
cats, and in the other by subsequent clinical trial.
The minimal systolic doses of the tinctures of digitalis were ‘0055 c.c.
and ‘005 ¢.c. per gramme of toad, and of the tincture of squills ‘006 ¢.c. per
eramme. ‘These figures are in close agreement with what would have been
expected with Rana temporaria, in which an accepted standard of activity
of these tinctures is a minimal systolic dose of 0:006 c.c. per gramme.
From these experiments it is concluded that Xenopus laevis is as
susceptible as Rana to the action of the digitalis group, and that it may be
used like Rana for the biological assay of digitalis preparations, the doses
for both being the same.
REFERENCES.
(1) Vutpran.—‘ Compt. Rend. Soe. Biol.,’ ser. 2, 1, 1854, p. 133.
(2) AspgL and Macutr.—‘ Journ. Pharm. Exper. Therap..,’ iii, 1912, p. 319.
(57)
SUNSPOTS AND EARTH TEMPERATURES.
By J. R. Sutton.
Table I gives the results of a comparison between the six-foot earth
temperatures observed at Kenilworth (Kimberley) for the nineteen years
1900 to 1918, and the Wolfer’s observed sunspot numbers as published in
the ‘Monthly Weather Review.’ The comparison has been made by taking
the mean earth temperature of any month and the sunspot number of the
previous month: this in order to allow something for the lagging of the
earth temperature wave with depth.* The means have then been grouped
according to spot numbers below or above thirty. The work was begun more
out of curiosity than with the expectation of getting any tangible result ;
and the direct comparison of curves was not very encouraging. The
grouping, on the contrary, shows a somewhat higher temperature for a lower
spot number, and that most definitely in the summer half of the year. If
the one could be regarded as the direct result of the other this would
indicate more intense radiation when the sun is free from spots than at other
times. Direct observations of the solar radiation, however, are interpreted to
mean less intensity when the sun is free from spots.
The 1-7-metre earth temperatures observed at Cérdoba for the fourteen
years 1887 to 1900 have also been compared with the spot numbers in the same
way. The results appear in Table II. They are of the same tendency, 7. e.
higher temperatures with fewer spots, though much more pronounced. So
also the summer half is the more definite.
Table III refers to the five-foot earth temperatures observed at Adelaide
during the thirty years 1878 to 1907. Proximity to the sea may have
influenced the temperatures. Only in the summer months is any difference
of temperature shown, though that is in the same direction as it is at the
other stations.
In all three cases the results, so far as they go, are in agreement with
previous ones.
* But no allowance has been made for the lagging in the annual values at the
foot of the ‘Table.
+ “It seems likely that increased values of the solar constant attend increased
sunspot numbers.” McAdie, ‘The Principles of Aérography,’ 1917, p. 276.
58 Transactions of the Royal Society of South Africa.
Taste I—Sia-foot Earth Temperatures at Kimberley and corresponding
Sunspot Numbers.
Sunspot numbers below 30. Sunspot numbers above 30.
BUEpe ‘ | igteoneee ]
; | month, Mean of | pjmes, | Earth tem-| Meanof | mines | Earth tem-
numbers. peratures. numbers. * | peratures.
December. January .| 5°8 2 |-foce Boe 10 |) F2zo sue
January . February.| 8-4 10 | 745 57:9 o Wo gers
February . March 8:0 a ae 56°3 10 | 73°6
March . | April 10°5 10 720 60°6 9 72:0
April . |May . 10-0 9 69:4 03°6 10 69:0
May June. 9°3 9 | 65°6 56°] 10 | 65-2
June . | July . 8:8 10 62°3 62°1 ° 62°6
July August 73 OG 62° 7 10° | 645
August . September; 83 10 | 62°5 722 a o2425
September October 9-0 9 | 65:0 63°3 10 | 65:0
October November 8:6 8 67:9 576 11 67°6
November | December 577 8 Ale Aad 58°9 11” 4) 703
Summer half . 76 53 70°9 58°2 61 70°4
Winter half 9:0 ov” 65°6 62°4 57 65°5
Year . 9-0 -— 68:2 59:1 — 67:9
TasLE I].—1:7-metre Earth Temperatures at Cordoba and corresponding
Sunspot Numbers.
soa Sunspot numbers below 30. Sunspot numbers above 30,
cheats pene ee
mon | eee on. | Times, | Cevsbaee | mmcibers, | "PSs eee
December . | January IIE) (| 20-0850.) 030 7 |18-93°C.
January .|February.| 12°6 8 206 60:2 6 | 19°d5
February .| March 13-0 8120772 65:7 6 11966
March April . 11-3 8 | 19-94 53°2 6: | 13°s9
April. May). 10-4 8 | 18-56 65:2 6 | 18-00
May .| June . 14°5 Slices) 72 74:8 5 | toe
June. .|July . 12:1 8 | 15°55 720 6 | 14-95
July .| August 13:3 8 | 14°97 70°5 6 | 14-47
August September | 13:7 8 | 15:34 72:4 6 | 1412
September | October 9-4 Gye 1e;:07 57°38 8 | 15-10
October .| November 11:3 8- | e238 63:3 6. | lok
November | December 69 i 11883 50°7 7 | 17-44
Summer half . 10-7 44, | 18:98 59°3 40) | 17°73
Winter half 12°5 48 | 16°89 68-0 30 | 16:10
Year . 13°5 — |17:82 62°9 — {16-91
Sunspots and Earth Temperatures.
59
Tasie III.—Five-foot Earth Temperatures at Adelaide and corresponding
Sunspot Numbers.
Sunspot Aehaeee :
month. te ee 4
December. | January
January February .
February . | March
March __. | April
April . May .
May . .|June.
June. ./July .
July? .| August
August September |
September | October .|
October .| November
November | December
Summer half .
Winter half
Year .
Sunspot numbers below 30.
Mean of
numbers.
ee ed pe
ee ee ee ee ee ee
q&> oO
ANN RH OWN OHO Or
— ee
pd
~
Oo
Sunspot numbers above 30.
Times. | Deratures, | numbers. | THM | Heratures.
15. 566-9°R) |) -55°9) |e Bony | e6o2b"
15 | 682 564 | 15 | 676
16 | 67°8 677 | 14 | 67:8
14 | 66:4 559 | 16 | 66:8
14 | 63-9 616 | 16 | 63-9
i sieoies 581 | 15 | 613
14 | 59-1 61-1 jp edGeneno
15 | 579 682 | 15 | 580
14 | 58-2 p95 |e lous) ood
13 | 59:4 stop wll a sees
14 | 61:8 596 | 16 | 61's
12 | 642 539 | 18 | 641
85 | 647 586 | 95 | 644
86 | 611 60-7 | 94 | 611
= 630 578 | — | 62:7
(61)
RAINFALL AND THE PRESSURE GRADIENT.
By J. R. Surron.
The seasonal rainfall over the greater part of South Africa may be
regarded as due mainly to a planetary rain belt following the sun to and fro
across the equator. It is this that gives us the summer rains. As the belt
moves north it is followed at a distance by the winter rains of the south-west,
which, however, do not penetrate often so far inland as the middle reaches
of the Orange River, and which recede as the belt returns south. Over the
central table-land of South Africa the seasonal rains are very unequal ; and
the climate is to be regarded in general as a permanently semi-arid one
punctuated at occasional intervals by wet years. Why the rainfall varies
so much from year to year may be suspected from a consideration of the
questions raised by Colonel Rawson in his notable papers on the movements
of anticyclones. It appears to depend upon the variations of barometric
pressure, or perhaps, rather, upon the causes which produce these. Father
Goetz has made an important advance in this aspect of the matter by his
discovery of a relationship between the fluctuations of rainfall at Bulawayo
and the variations of the second harmonic term in the annual pressure
curves.
The present paper is a brief discussion of the possibility that the pressure
gradient between Kimberley and Cape Town may have some sort of connec-
tion with the rainfall of the two places. For our purpose the monthly means
of pressure and rainfall of the two places have been compared for the twelve
years 1897 to 1908. The pressure gradient is supposed to be represented
month by month by the simple arithmetic difference between the monthly
means of pressure at the two places.*
Table I gives the mean gradient, and the mean rainfall of each place for
each month of the period. On the whole the gradient is least in autumn
(March to May), and greatest in spring (October and November). There is
a maximum of rainfall at Cape Town near midwinter, and a secondary maxi-
mum near midsummer. The maximum rate of fall at Kimberley occurs in
February, the minimum in the late winter, with some tendency to a
secondary maximum in July.
Table II shows the mean monthly rainfall of the two places for gradients
less than the median (‘‘ Small Gradients”), and for those greater (“ Large
Gradients’’).
(1) Cape Town.—At Cape Town, speaking at large, the rainfall when the
gradient is small exceeds that when the gradient is large by about 18 per
cent., both summer and winter. Eight months show the excess, four do not.
* T.e. at the Royal Observatory, Cape Town, and the Observatory at Kenilworth
(Kimberley).
62 Transactions of the Royal Society of South Africa.
Considering individual months, it happens that of the 88 months having a
fall less than the mean, 38 occur with a small gradient and 50 with a large
one ; whereas of the 56 months having a rainfall greater than the mean, 34
occur with a small gradient and 22 with a large one. That is to say,
deficient rainfall tends towards occurrence with months of large, while
abundant rainfall tends towards occurrence with months of small gradient.
(2) Kimberley.—At Kimberley, on the whole, we have the opposite. The
rainfall when the gradient is large exceeds that when the gradient is small
by about 15 per cent , the effect being most marked in the winter. Eight
months agree, three do not, and one (October) is about neutral. By a
coincidence the number of months showing excess or defect is the same as
at Cape Town though in the opposite direction like the quantities, 7.e. of the
88 months having a fall less than the mean, 38 occur with a large gradient
and 50 with a small one: whereas of the 56 months having a fall greater
than the mean, 34 occur with a large gradient and 22 with a small one.
These facts may be summed up in—
Rule 1.—An excess of rain inclines to occur either at Cape Town or
at Kimberley according as the synchronal gradient is small or large
respectively.
The question now arises whether this tendency is something more than
synchronal: e.g. whether it persists at all into the following months?
Obviously, by the above Rule, it can only persist at the best to a partial
extent; for if the months of small and large: gradient were to alternate
continually, any persistence would necessarily be masked by the current
conditions. But the length of a spell of gradient, large or small, may be
anything from one to six months, the average duration being about three.
It seemed worth while, therefore, to compare the gradient of each month
with the rainfall of the following month. This gives Table IIT.
The unexpected result shown by Table III is a considerable reversal of
the conditions shown by Table II: more rain going to Cape Town and less
to Kimberley when the gradient of the previous month was large. With
this qualification, that the effect is a winter one at Cape Town, and a
summer oneat Kimberley. In the summer half at Cape Town and the winter
half at Kimberley the normal condition persists into the following month.
Hence we have—
Rule 2.—An excess of rain inclines to occur either at Cape Town in the
winter or at Kimberley in the summer according as the gradient of the
- previous month is large or small respectively.
Rule 3.—At Cape 'Town in the summer and at Kimberley in the winter
the conditions under Rule | incline to continue unaltered into the following
month.
These results, based on a comparatively short period of observation, can,
of course, only be regarded as provisional. As they stand they can only
Rainfall and the Pressure Gradient.
have a limited use in long-range forecasting. It is to be hoped that the
investigation may be carried further, seeing that it promises a certain
measure of information beyond the mere requirements of weather prediction.
As Lord Rayleigh, thirty years ago, truly said, “In order to introduce
greater precision into our ideas respecting the behaviour of the earth’s
atmosphere, it seems advisable to solve any problems that may present
themselves.”’
TasLe [.—Monthly mean Pressure Gradients and Rainfall.
Rainfall.
Cape Town.
Gradient.
in.
January 3864
February. 3°848
March 3°828
April 3°838
May 3°852
June 3°843
July 3°845
August ; 3°866
September . 3°895
October : 3°914
November. 3°918
December. 3°889
Year 3°865
TasLe Il.—Monthly mean Rainfall corresponding to Synchronal Gradients
in.
111
‘46
“7A
2°29
3°98
4°33
3°60
3°22
27
2°27
‘90
94,
26°11
Greater or Less than the Median.
Rainfall.
Cape Town.
Small
gradient.
in.
January. ‘87
February. 68
March ‘81
April 2°20
May ar 3°45
June ; 567
July 4°55
August : 3 30
September . 2-11
October 2°36
November . oe)
December . 1-11
Summer Half 6°82
Winter Half 21:26
Year 28:08
Large
gradient.
in.
1-34
"25
67
2°39
4-5]
Rainfall.
Kimberley.
in.
3°09
2°85
3°06
157
“06
‘26
30
‘Ll
88
117
1°55
2°06
17°44
Rainfall.
Kimberley.
Small Large
gradient. gradient.
in. in.
3°90 2°28
Zane, 2°98
2°26 3°87
1-47 1-67
32 (9
10 “42
“AS 14
‘07 ‘14
25 151
1-16 113s
1:63 1:43
1:87 2°26
13°54 13°95
2°66 467
16°20 18°62
64 Transactions of the Royal Society of South Africa.
Taste I1Il.—Monthly mean Rainfall corresponding to. Gradients of the
Previous Month Greater or Less than the Median.
Rainfall. Rainfall.
Cape Town. Kimberley.
—_— eee 5 rE
Rain Small Large Small Large
Month. gradient. gradient. gradient. gradient.
in. in. in. in.
January. 1:24 ; iS . 988 2°49
February. “42 : OO me Ss: . 324 2°47
March wi) (oy - 73230) ; 2°83
April ; 2°47 ; ra ree ‘90 2°24,
May 3°43 A522 62 “49
June ; 2°82 SE ee : “42 09
July 341 ; Son = : 03 56
August 3°79 ; 204 12 ; 1)
September . 1:94: ; 259 ’ "62 ; 113
October 2°38 Qe 2 06 1:27
November . 1-01 aay ee ey 47h 1-42
December . 1:10 eo 8 . 2°38 1:80
Summer Half 6°90 ; 0°92 15:08 12:28
Winter Half 17°86 . 21°42 201 461
Vent 24°76 °°, OF 34 . | qelvg alee
( 65 )
COLOUR AND CHEMICAL CONSTITUTION.
Part XIV.—THE CALCULATION OF THE COLOUR OF THE Dicyciic DyYEs.
By James Morr.
The remarkable discovery published last year in the Addendum to Part X,
whereby I was enabled to calculate the colour of methylene-blue from that
of a dye containing neither nitrogen nor sulphur, has in the meantime proved
very fertile. I have now worked out all the factors whereby the colour of
any dye made up of two ionisable rings and either one or two linkages can
be calculated. This means calculating quite two-thirds of the known dyes.
Dioxybenzhydrol, HO: C,H, CH(OH) - C,H,OH is an example of a
singly-linked dicyclic dye, and dioxy-xanthhydrol
te
a
is an example of a doubly-linked dicyclic dye. Mono-oxybenzhydrol on the
other hand is “ monocyclic,” having only one ionisable ring.
I find if the assumption is made that the wave-length of an electron-
orbit going round two phenol rings is about A 1380, that this orbit is
contracted by linkage of the rings always in the same proportion for the same
linkage, whatever other groups are present. Hach linkage thus has a colour-
factor (less than unity) which, when multiphed by 1380, gives the colour of
the corresponding di-oxyphenyl-dye.
Table of Linkage Factors.
A. Methylene or “CH. . . Value 0°65.
B. Imino or 7 NL , . Value 0°78.
C. Oxo- ee OR . Value 0°89.
D. Thio- oe : - Value.0°92,
66 Transactions of the Royal Society of South Africa.
If two of the linkages are present in a phenolic dye, the figure 1380 is to
be muliplied by both factors.
I have also discovered that the effect of oxidising the linkage is always
the same whatever the linkage is. The effect is to make the factor of the
oxidised linkage exactly 2 of the factor of the unoxidised linkage.* This
may be due, as explained in Part XII, to changing from five tautomerisms
to three only.
Table of Factors for Oxidised Linkages.
E. Oxymethylene or carbinol linkage, ,CHOH\ Value 0:39
F. Dioxymethylene or keto linkage, 7 CO \ Value 0-234
G. Oximino or hydroxylamine linkage, 7 NOH \ Value 0°47
O
H. Peroxide linkage, 79 \ . . Value 0°535
I. Sulphoxide linkage, /SO\ 2. s,s Wale 0°55
J. Sulphone linkage, “SO. . . . . Value 0:33 (?)
[K. Trebly-linked carbinol,=COH _. . Value 0°386]
These are all the new data required for the calculations, but in addition
all of the 27 ring-colour-factors published in Part X may be used so as to
extend the theory from the phenolic dyes to all sorts of classes of dicyclic
dyes. Thus, for bis-dimethylamino dyes the figure for the corresponding
phenolic dye is to be multiphed by the square of 1:059, the factor for
replacement of OH by NMe, (Part X, p. 306, middle), or alternatively,
instead of A 1380, use A 1380 x (1:059)? = »A 1548 for the fundamental
vibration round two dimethylaniline rings. Similarly the fundamental
9 9
Toe) or 4 1400 (Part X,
vibration for two aniline rings is A 1380 x (
p. 304).
According to this theory the colour of the majority of the singly-linked
dicyclic substances, such as dioxydiphenylmethane, is in the infra-red : those
which are observable to the eye are calculated below.
(1) Dioxybenzhydrol (formula on p. 65).—The calculated A is 1380 x
factor E = 13880 x 0:39 = A538. Observed A539.
(2) Benzaurine (di-para-variety ).—This is the C-phenyl derivative of the
foregoing. The C-phenyl factor is 1:026 (Part X, p. 310). Calculated A»
= 1380 x 0°39 x 1:026 = 2552. Observed A 553.
(3) Michler’s hydrol—This is two dimethylaniline rings joined by
CHOH. Calculated X = 1548 x 0°39 = X604. Observed 2 603.
— (4) Malachite green.—This is the C-phenyl derivative of No.3. Calculated
AX = 1548 x 0°39 x 1:026 = A620. Observed A619.
* Watson and Meek first showed this to be the case for the CO group.
Colour and Chemical Constitution. 67
(5) Doebner’s violet.—This is No. 4 less four methyl groups. Calculated
X = 1400 (see 15 lines above) x 0°39 x 1:026 = 2561. Observed A 562.
(6) Aurine.—Here the linkage K is used instead of linkage H, although
they are nearly the same. As shown in Part XII, p. 209, only two of the three
rings act in producing the colour. The calculated is 1380 x 0°386 = 2d 538.
Observed A 534.
(7) Fuchsine (para).—Two aniline rings and the factor K are used.
Calculated A = 1400 x 0°386 =A 541. Observed 4543. Commercial fuchsine
has an extra C-methyl group and has A 549. Calculated, using the C-methyl
factor 1-012 = » 547.
(8) Crystal violet.—Two dimethylaniline rings and factor K to be used.
Calculated 1548 x 0°386 = 2597. Observed A 595.
(9) Aniline blue.-—Two diphenylamine groups and the factor K to be
used. The N-phenyl factor has twice the effect of the C-phenyl factor
(cf. ortho- and meta- groups in Part IV, p. 188). Its value is therefore
(1:026)? = 1:054. Hence two diphenylamine groups have the value A 1400
x (1:054)? = 1554, and the calculated value for aniline blue is 1554 x 0°386
= 599. Observed A 600.
(10) Phenolphthalein.—This is No. 1 with a benzoic acid group attached.
The factor for the latter is C-phenyl multiphed by o-carboxyl (see Part X,
p. 306) = 1:026 x 1:002 = 1:028. Calculated X for phenolphthalein = 1380
x 0°39 x 1:028 = X1553. Observed 2 554.
(11) Phenolsulphonephthalein.—Yhis is No. 1 plus a benzenesulphonic
acid group. The latter’s factor is 1026 x 1-018 (Part X, p. 308) = 1:045.
The calculated » is 1880 x 0°39 x 1:045 = 562. Observed A 563.
(12) Michler’s ketone.—Two dimethylaniline rings and the factor F are to
be used. Calculated A 1548 x 0°284 = A 362. Observed d 364.
(13) Indophenol.—This is the anhydride of dioxy-diphenylhydroxylamine.*
Two phenol rings and the factor G are therefore to be used. Calculated
X = 1380 x 0:47 = X 649. Observed A 650.
(14) Indamine.—This in the same way is derived from two aniline rings
and the factor G. Calculated A = 1400 x 0°47 = 2658. Observed A 660.
(15) Bindschedler’s green.—This is tetramethylindamine, and the calcula-
tion requires two dimethylaniline rings and the factor G = 11548 x 0:47
= 728. Observed A 730.
Doubly-linked Dicyclic Colowrs.
Of these there is an enormous number in use, varying from eosine to
safranine and methylene-blue. If all the ten factors were used two at a
time there would be 100 colours of each of the three classes, viz. phenols,
anilines and dimethylanilines. I therefore limit myself to those I have
observed, or for which data exist in the literature.
* Commercial “indophenol” is a naphthol compound.
68 Transactions of the Royal Society of South Africa.
(16) Pyronine—This has the formula :
O..
NMer7 No NNes
eS el
It is calculated by multiplying 11548 (for the dimethylaniline rings) by
factors C and H, viz. 1548 x 0°89 x 0:39 = 5386. Observed A 542 in |
alcohol.
(17) Rhodamine.—This is the benzoic-acid derivative of pyronine (cf.
phenolphthalein). The benzoic factor is 1:028 (see p. 67). The calculated
Nis 586 x 1:028 = 4552. Observed A 553.
(18) Dioryxanthhydrol.—This is pyronine with 20H instead of 2N Me,.
Calculated 1380 x 0°89 x 0°39 = 2478. Observed A479. This substance
is the parent of fluorescein.
(19) Fluorescein.—The above plus benzoic acid is fluorescein. Calculated
A478 x 1:028 = rX492. Observed A 494.
(20) EHosine.—This is fluorescein plus four ortho-bromines. Calculated
A = 492 x (1:0128)* (see Part IX) =A519. Observed A 521.
(21) Resorufine.--This has the formula :
BANC es? Sir
\Y ion ae
1880 for the phenol rings is therefore multiplied by factors C and G.
The result is 1880 x 0°89 x 0°47 = 2577. Observed A 576.
(22) Thionol.—This is the foregoing with S in place of O. The calculated
value is 1880 x 0:92 x 0°47 =2597. Observed A = 593,
(23) Methylene-blue.— This is the example given in the Addendum to
Part X. It is thionol with two dimethylaniline rings instead of two phenol
rings. Calculated 1548 x 0°92 x 0-47 =A 668. Observed 1665. Formanek
gives \ 6673 in what was probably a better specimen.
(24) Thionine-—This has the formula:
ny
ee
Calculated value = 1400 (for two aniline rings) x 0°92 x 0°47 = A 604.
Observed A 608.
(25) Gentianine.—This is halfway between Nos. 23 and 24 in constitution.
Since the scheme is factorial its calculated A is the geometric mean (not the
arithmetical mean) of the AA of Nos. 23 and 24 = 1635. Formanek observed
d 638.
Colour and Chemical Constitution. 69
(26) Dioxyacridine.—This can be written in four ways and calculated
from either of the hydrated forms, viz. :
nS poo
|
Neu NO \7\cHH
(a) (b)
pNE
HON SO) Nise Fi SW Nat
a bo te
\7 NcHOH, SZ NN
(c) (d)
According to formula (c) it is dioxybenzhydrol with the NH linkage:
calculated 1380 x 0°39 x 0°78 = dX 420.
According to formula (d) the calculation is 1880 x 0°65 x 0-47 = A 422.
The difference is due merely to the factors not yet being accurately known
to three places of decimals. Observed A 421.
(27) Benzoflavine-—This has the formula :
xo \/\wu,
-60=
Ph
The calculation involves 4 1400 for the two aniline rings, multiplied by
(a) the CHOH factor, (b) the NH factor, (c) the phenyl factor, (d) the
square of the orthomethyl factor (see example No. 7). The calculated value
is 1400 x 0°39 x 0°78 x 1:026 x (1:012)?=A449. The observed value was
about 2» 454.
(28) Dioxyphenazine —This, being symmetrical, has only two formule,
Wiz. :
UN ok FN
Soe et
OO NY
Its calculated » is 1880 x 0°78 x 0°47 =2A506. Observed d 510.
(29) Neutral red.-—This is dimethyldiamino-toluphenazine. 'The parent
rings are one aniline and one dimethylaniline ring, the value of which
(see No. 25) is 1/1400 x 1548 = 1473. This is to be multiplied by the
NH and NOH factors and by 1:012 for the ortho-methyl group (see No. 27).
Calculated 4 546. Observed A 540 (commercial specimen).
(30) Safranine.—This has the formula (when hydrated) :
RO SNe,
CH,
|
Cee
70 Transactions of the Royal Society of South Africa.
Calculated X= 1400 x 0:78 x 0-47 x 1:026 (for phenyl) x (1-012)? (= benzo-
flavine x 0°47 + 0°39) = 2541. Observations in commercial specimens vary
between d 530 and A540; probably some do not contain the methyl groups.
(31) Methylene violet.—This is isomeric with safranine, but the two
methyls are on nitrogen not in the ring. Calculated 1473 (see No. 29) x
0°39 x 0°78 x 1:026 = 554. Commercial specimens vary in observation
between A550 and 565, being mixtures. Another way of calculating this
isomeric substance is to note that the N-methyl group has twice the effect
of the C-methyl group, the latter having a factor of 1:012; the former has
a factor of (1:012)? = 1:0245 (see Part X, p. 805). Consequently No. 31
calculated from No. 30 is A541 x (1:0245)2 + (1:012)2 = \541 x (1:012)?
= A554. This doubling of effect on changing from C-substitution to N-
substitution has already been mentioned under substance No. 9, but it is to
be noted that phenyl (and probably methyl also) on a linkage has always
the value 1:026 whether on C or on N.
(32) Magdala red.—This is safranine with two naph idea
attached, and a third in the linkage instead of phenyl. The calculated
colour of phenyldiaminophenazonium salts is 527, whereas Magdala red
shows A559. Assuming that the linkage-naphthyl has the same effect as
phenyl, this gives a factor of 1:030 for the naphthalene-ring residues, each
C,H, This is a much smaller value than that found from the naphthol-
phthaleins (1-092), and remains an anomaly until further work can be done
on naphthalene colours in general.*
(33) Azine green.—This is about the most complicated member of the
series. It has the formula:
NPh
NMe,“ \“
V7 SER
WA SNE, SO
As shown under No. 32, phenyldiaminophenazonium has \ 527. Azine-green
has, in addition to the latter, a naphthalene-ring, two N-methyls and one
N-pheny] (value 1-054). Assuming naphthalene to have the normal value
of 1:09, because the amino group is outside it in this case, the calculated
colour of azine-green is 527 x (1:0245)? x 1:054 x 1:09 = A636. I have
not observed it, but the description in the literature indicates that the
observed value lies between A 620 and A 640.
(34) Dioxyanthracene-—The ordinary formula of this substance when
written hydrated becomes :
NAECVA. SOA
NAN i NZ
* Since this was written it has been discovered that naphtholphthalein has ortho
linkages instead of para as hitherto supposed. There is thus no anomaly after all.
COD aa
Colour and Chemical Constitution. "al
The calculated colour is 1380 x 0°65 x 0°39 =2350. It is described in
‘Beilstein’ as pale yellow, which agrees with the calculation. Dioxydihydro-
anthracene should have a deeper colour corresponding to 1380 x (0°65)? =
d 583.
Dioxyanthraquinone is an exception: the two CO groups cause a new
kind of vibration, and the colour is much higher—eight times as high in
fact as theory predicts. No doubt the theoretical colour (A 76) is there also,
but is too low to observe.
These nineteen doubly-linked dicyclic colours have been selected so as to
cover the whole field, so far as known (the peroxide and sulphoxide dyes
are not yet known): it would merely be taking up space, for example, to
calculate erythrosine when eosine is calculated, or to calculate the fourteen
derivatives of thionine observed by Formanek.
There are also dicyclic dyes in which the rings are joined by chains
longer than one element: indigo and the azo-dyes are examples, but
the published data are insufficient—indeed almost useless—as regards a
quantitative explanation.
Indigo may be calculable from the hydrated formula :
NG? AN
oa
\_/-©0—C(OH) : ((OH)—Co-\/,
and the thiazole and primuline dyes similarly, viz. :
aes
\ J-s-00-< Dam
but the laws of the co-operation of two or more factors in a linkage are not
yet known. The azo-dyes do not come into the scheme for the reason that
they are coloured whether they contain ionisable groups or not; but it is
worth noting that the three which do contain two ionisable rings (azophenol,
azoaniline, etc.) have colours fairly similar to those of the corresponding
phenazines, from which they differ by two hydrogen atoms only. This
suggests that the azo group can sometimes be written —NH—NOH— and
the colour calculated accordingly, allowing about 0°73 instead of 0-78 for
the “NHQ\. factor when not replacing two hydrogen atoms of the benzene
rings but inserted in a previously existing linkage.
The most of the azo-dyes, however, owe their colour to the azo-linkage
independently of all rings, since diazomethane shows colour.
It is probably desirable to show the way in which the linkage-factors
were ascertained from the observations, particularly as this also exhibits the
degree of variation of each factor, and enables the reader to see that each
factor is really a constant within the observational error.
(1) The oxo-linkage.—(a) Resorcin-benzene: observed \ 492. The corre-
6§
(2 Transactions of the Royal Society of South Africa.
sponding substance without the oxo-linkage is benzaurine, which has A 553.
Oxo-linkage-factor from (a) = 492 + 553 = 0°890.
(b) Fluorescein = 494: corresponding substance = phenolphthalein of
554. Oxo-factor = 494+ 554 = 0°892.
(c) Eosine = A 521, whereas tetrabromophenolphthalein has A 584. Oxo-
factor = 521 + 584 = 0892.
(d) Resorcin-sulphophthalein = 4498, compared with phenolsulpho-
phthalein of A565 gives the oxo-factor = 0°885. :
(e) The tetrabromo-derivatives (3—3’-5-5’) of the foregoing two sub-
stances have 4527 for the resorcin compound and 594 for the phenol
compound. The oxo-factor in this case 1s 0°887.
(f) Pyronine has A 542 in alcohol, and Michler’s hydrol has » 610 in the
same solvent. ‘The oxo-factor in this case is 0°889.
(g) Dioxyxanthhydrol has A479 in water, and dioxybenzhydrol 2 539.
The oxo-factor for this pair is thus 0°888.
(h) 'Tetramethyl-rhodamine has 4555 (same as benzaurine). ‘The corre-
sponding substance without the oxo-linkage is “ dimethylanilinphthalein,”’
which I have not observed. It is related to malachite green just as
phenolphthalein A 554 is to benzaurine A 553, therefore its ’ must be that
of malachite green plus 1 = A620. Hence the oxo-factor for this pair is
553 + 620 = 0-892.
(‘) Resorufine has 4576: the substance without the oxo-linkage is
indophenol of A650. The oxo-factor here is 0°887.
(7) Hexabromo-derivatives of fluorescein and phenolphthalein substituted
in the same six places (acfgjk of Part XI) have AA 535 and 602 respectively.
The oxo-factor from this pair is 0-889.
(k) Isomeric hexabromo-derivatives derived from 3-6 dibromophthalic
acid (adfgjk of Part XT) have 4A 538 and 606 respectively. This oxo-ratio
is 0888.
The average of these eleven observations gives 0°889 for the oxo-ratio,
but I am of opinion that much more weight should be given to those
depending on common substances, and that consequently the true value is
0-891.
(2) The thio-linkage.-—This could be derived from as many substances as
the oxo-factor, but the substances, thiofluorescein, thiopyronine, etc., are not
obtainable or capable of being made here, I have therefore calculated it
from the methylene-blue and thionol classes alone :
(a) Methylene-blue and Bindschedler’s green have AA 665 and 730
respectively. The ratio is 0:912.
(b) Thionine and indamine have AA 603 and 660 respectively. The ratio
is 0-915.
(c) Thionol and indophenol have AA 593 and 650 respectively. ‘The
ratio is 0-915,
Colour and Chemical Constitution. 12
I hope that some one will examine the thioxanthhydrol dyes, and thus
enable the thio-factor also to be stated to three places of decimals.
(3) The imino-linkage.—The acridine dyes are NH-derivatives of the
benzhydrol dyes: similarly the phenazines are related to the indophenols.
(a) Flavine (diaminoacridine) has X about 427, whilst diaminobenzhydrol
has 1548. The NH ratio is here 0-780.
(6) Benzoflavine A (substance No. 27 without the methyl-groups) has
X about 450 (broad). The corresponding substance without the NH group
is Doebner’s violet, which has 1562. The NH ratio is here 0°80. This,
being exceptional, is probably wrong. ‘The acridine dyes analogous to
the phthaleins and rhodamines appear to be little known, and are inaccessible
here (‘ Beilstein,’ iv, supplement, p. 879. Calculated A 438.)
(c) Bisdimethylamino-phenazine has A about 565, and Bindschedler’s
green A730. The NH ratio is here 0°774.
(7) Dioxyphenazine has »X about 510 (broad), and indophenol 2 650.
The NH ratio is here 0°785.
(e) Safranine (without methyl groups) has A 525 compared with indamine
d 660, giving 0°796 for the NPh ratio, whence NH is 0°796 + 1:026 = 0-778.
(4) Whe methylene-linkaye.-—This is inferred from the acridine dyes, this
time compared with the indophenols, on the principle that when two linkages
are present it does not matter which is the oxidised one, 7.e. (CH, + NOH)
= (CHOH + NH).
(a) Flavine / indamine = 427/660. Value of “CH, \ = 0°647.
(6) Dioxyacridine / ndophenol = 421/650. Value of /“CH,\ = 0°649.
The oxidised factors.—Here the supposition 1s made that since a factorial
scheme has been shown to be generally successful, and since Watson and
Meek have established the factor 2 for the change of linkage from CHOH
to CO, this factor 2 will also hold for any other oxidation of the lnkage.
Hence the value of /~CHOH\ is 2 of /CH,\ = 0°39, and /~NOH\ =
3 of 0°78, and so on. That this supposition is correct is proved by the
consistency of the results given in this paper.
The next step is to calculate the colour of the unobservable singly-
linked dicyclic substances :
(1) Dioxy-phenyl ether.—This is (1) dioxyxanthhydrol less CHOH:
calculated X 479 + 0°39 = A 1229, (2) resorufine less NOH: calculated A 576
+ 0°468 = 12380.
(2) The calculation for the sulphur-ether diovyphenylsulphide gives
d 1258.
(3) Dioary-diphenylamine.—This from dioxyphenazine is calculated to
have A510. + 0-468 = X 1090: from dioxyacridine the calculated value is
X 421 + 0°39 =X 1080: from resorufin 4 576 + (2 of 0°891) = 1077 and
so on.
(4) Dioxy-diphenylmethane.—This from dioxyacridine is calculated to
74, Transactions of the Royal Society of South Africa.
have 421 + 0°468 =’900: from dioxyxanthhydrol and the fluoresceins
rather less: the average is A 898.
The final triumph of the scheme comes when it is found that the wave-
lengths of the seven singly-linked substances are themselves related to a
fundamental constant (about 1380) by means of the factors contained in
themselves :
Dioxy-phenyl ether, 1230, divided by the ,/O\ factor 0°891, gives 1382.
Dioxy-diphenylamine, 1083, divided by the “~NHQ\, factor 0°78, gives.
1388.
Dioxy-diphenylmethane, \ 898, divided by the /CH,\_ factor 0°65, gives
1382.
Dioxy-benzhydrol, 1539, divided by the “CHOH\ factor 0°39, gives
1381.
Indophenol, A 650, divided by the “NOH\. factor 0°47, gives 1383.
Dioxy-phenylsulphide, A 1258, divided by the ,/S\_ factor 0°92, gives 1370.
Dioxy-benzophenone, A about 323, divided by the “CO\_ factor 0234,
gives 1380.
This makes it almost certain that the unobserved ones have meaty the
absorption-bands in the infra-red calculated in this paper.
Cy)
CRYSTALLINE STRUCTURE OF ANTIMONY AND BISMUTH.
By A. Oaa.
(With two Text-figures. )
ANTIMONY.
Antimony crystallises in the dihexagonal alternating (calcite) class’ of
hexagonal crystals. The crystalline symmetry is that of a rhombohedron,
the three edges which meet in the trigonal axis being the axes of the crystal.
The angle between any two of these edges is 86°58’. From the geometry of
the rhomb it is easy to find that the angle between the planes (111)
and (110) is 37°23’ and that between the planes (100) and (111) is 56°48’.
If we take the sides of the rhomb to be a and suppose that an atom is
placed at each of the corners of the rhomb, then the distances between the
planes of atoms are—
digg = ‘9978a (1)
dy, = 607la (2)
dip = °7236a (8)
dyin = 688la (4)
By means of an X-ray spectrometer, the bulb having a palladium anti-
cathode, the glancing angles of the first order spectra were experimentally
found to be—
(100) (411) (110)
5°30" : 4°°30' G30;
Applying the formula nA = 2d sin 6, where nis the order of the spectrum,
dX = 0°584 x 10-8 cm., and 6 the observed glancing angle, we find—
a ook Use a(S)
Cig 0 00s AO (6)
Vp oo x Omees, (7)
From (2) and (5) we find a = 6:12 x 10-5 cm.
ere) el SCO ar ok Oo UO alms as
MHS ey els) ens, 5 hse too el OO ree
It is evident from the calculated values of a that the spacings between
the (100) planes and also those between the (110) planes are half what we
76 Transactions of the Royal Society of South Africa.
have supposed them to be by placing atoms merely at: the corners of the
rhomb. If we take a face-centred lattice —
then d,,, = °607la as before (8)
but dy) = *499a (9)
and d= 3670 (10)
hence from (5) and (8). a =612 x 10-8 cm.
(6) 1 CO) Gata 618 ae
COP (RO) tiga x h0ae
mean value of a = 6:14 x 10-8
99
29
Taking the density of antimony as 6°70 grms./em.® the mass of the
rhomb is equal to—
6°70 xX 614? x 099732 104
= 1545. x 10—-4erm,
Since the atomic weight of antimony = 120-2, and the mass of the
hydrogen atom = 1:64 x 10-24 gprm., then » x 120-2 x 164 x pee =
1545 x 10-24 grm., where » is the number of atoms per unit rhomb. This
gives n = 7°85.
It is clear, then, that there are 8 atoms per unit rhomb.
Let us take 8 atoms per unit rhomb, find the mass of the rhomb, hence
its volume and then find a. We find a = 620 x 10-8 em.
Assuming a face-centred lattice we find then—-
dino = 3 00) xX Om? ecm.
dn) = ooex mae.
dip = cet 10
dig, = els eee,
We can now calculate the glancing angles for the spectra from the
different faces.
Planes.
(100) ai (110)
Observed angle 5°30! 4°°30! (3
Calculated ea Ae FOR
Arrangement and Spacings of the Atoms.
(a) (111) planes :
We conclude that the underlying structure is the face-centred lattice,
but a face-centred lattice gives only 4 atoms per unit rhomb. ‘To determine
the positions of the other 4 atoms we must investigate the relative intensi-
ties of the spectra from different faces. The observed intensities of the
spectra of five orders from the (111) planes were—30 : 100 : 33 : 4: 12.
The first order being weaker than the second shows that there is a plane |
of atoms dividing the distance (3°76 x 10-8 cm.) between the (111) planes.
Let £ be the phase difference between the two sets of planes, and taking the
Crystalline Structure of Antimony and Bismuth. 77
intensities* of a normal set of spectra to be 100 : 34: 14: 7 : 4, we have
380 — 1001 4 cos
100 =. 84: 1- + cos28’
which gives 6 = 148° approximately.
The distance between one set of (111) planes would be divided by
another set of (111) planes in the ratio of 0°412 : 0°588.
The calculated ratios of the intensities for this spacing are 30 : 100 :
31 : 5 : 15, which is in close agreement with the cbserved ratios.
tim = SIOG rare -2
( 200) fr ei oo.
— R24 a
( 110) eae ae
oO
Fig. 1.—Spacines of planes in Angstrom units (10-8 cm.).
The spectra could be explained by a structure similar to that of the
diamond but distorted along the trigonal axis,t one set of (111) planes
dividing the distance between those belonging to the other in the ratio of
about 3 : 2 instead of 3 : 1 as in the diamond.
(b) (100) planes :
To settle whether a structure similar to that of the diamond will fit the
observed facts we must examine what this ratio would give for the spectra
* James and Tunstall, ‘ Phil. Mag.,’ 8S. 6, vol. 40, No. 236.
y+ Sir W.H. Bragg and Prof. W. L. Bragg arrived at this conclusion in 1914:
‘“X ray and Crystal Structure,’ p. 227.
78 Transactions of the Royal Society of South A frica.
from the (100) planes. We find at once that it will not explain the (100)
spectra.
The observed (100) spectra are nearly normal, and there can be only a
small difference of phase between the two sets of planes. James and
Tunstall* have shown that this can be accomplished in the following way.
Divide the unit face-centred rhomb into eight equal rhombohedral cells.
Place an atom at each of the unoccupied corners of the small cells and then
/
ie 2
push each atom along the diagonal by an amount equal to 0:074 of the
length of a diagonal of one of the small cells.
The spacings we have determined for the (111) planes would make this
amount to be 0:059 instead of 0-074.
The spacing for the (100) planes would be 0:059 : 0:'941. The two sets
of planes differ in phase by about 21°, and would give a nearly normal set
of spectra in agreement with observation.
(c) (110) planes :
We must now examine whether such an arrangement would suit the
spectra from the (110) planes. The planes would contain an equal number
of atoms, and the spacings are 0118 : 0°862.
* James and Tunstall, loc. cit.
Crystalline Structure of Antimony and Bismuth. WY
The intensities of the first three orders were observed to be 100: 17 :
0, while calculation gives 100 : 20 : 3.
(d) (110) and (111) planes :
James and Tunstall examined the spectra from these planes and found
that they agreed with this arrangement.
The spacings of the planes have been determined by the relative inten-
sities of the spectra from the (111) planes ; hence the determinations of these
intensities are of importance. The ratios were found to be 30 : 100 : 33 :
4: 12, while James and Tunstall found 60 : 100 : 48: 0: 15, giving a
phase difference of 140° between the two sets cf planes. We found a phase
difference of 148°.
The spacings of the planes are given in Fig. 1. Fig. 2 shows the
arrangement of the atoms on one of the eight small cells into which the
rhombohedron can be divided.
The shortest distance between atomic centres is 2°92 x 10-8 cm.
BISMUTH.
Bismuth, like antimony, crystallises in the dihexagonal alternating system
(calcite class). The three edges of the rhombohedron meet in the trigonal
axis and the angle between any two of the edges is 87°34’. The angle
between the faces (100) and (111) is 56°°24".
Again using an X-ray bulb with a palladium anticathode the glancing
angles for the first order spectra were—
(111) (100)
4°18" 5°83!
Taking the density of bismuth 9°80 grm./em.? we find that the unit
rhomb contains 8 atoms and that the length of the side of unit rhomb is
G02 10—* cm.
Assuming the structure of crystalline bismuth to be similar to that of
antimony we find the spacings of one set of planes to be—
Planes.
LAS
ia a a
(00) (110), 9 Ci) Cine yes Gad)
Spacings 3:25 2°35 3°92 2°25 3°69
The spacings are given in Angstrom units (10-8 cm.) ‘The relative
positions of the two sets of planes have not been accurately determined.
The intensities of the spectra from the (111) face showed much the same
order as those from the corresponding face of antimony but were much
fainter.
Experiments are in progress for the measurement of these intensities
whereby the second set of planes may be fixed.
UNIVERSITY OF CAPETOWN,
February, 1921.
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(81).
ON THE REPTILIAN GENERA FUPARKERIA BROOM, AND
MESOSUCHUS WATSON.
By 8. H. Haveuton, B.A.
(Published by permission of the Rt. Hon. the Minister for Mines and Industries.)
(With Plates II and IIT.)
The genus Huparkeria was founded by Dr. Broom in 1913 for the
reception of a small Thecodont reptile from Aliwal North. The type of
the genus is an almost complete skeleton including a beautiful skull and
lower jaw. In his description of the form Broom differentiated Huparkeria
from a genus Mesosuchus, thetwo having been grouped previously under the
latter name by Watson, who was unaware of the existence of the skull of
Euparkeria.
The recent acquisition of the famous ‘“ Alfred Brown” Collection of
fossils by the South African Museum has enabled more development work
to be done on the masses of sandstone containing the Huparkeria and
Mesosuchus remains, and the consequent discovery of more skulls and
partial skeletons which throw a little more ight upon the structure of these
two different forms has resulted.
The remains occur in a mass of greenish-grey sandstone which in the
main is highly calcareous. The matrix, however, is comparatively easily
freed from the bones, which are left clean, and the various sutures of the
skulls are clearly displayed. Most of the material recently uncovered
belongs to the genus Huparkeria, but a fairly complete new skeleton jun-
doubtedly belongs to the interesting Acrodont form Mesosuchus. The dis-
covery of a skull of Hwparkeria lying in close proximity to the pelvis described
by Broom as Browniella africana has led to the re-examination of the bones of
Browniella, and the conclusion has been reached that the pelvis and shoulder-
girdle said to belong to that form are in reality specimens of Huparkeria
capensis.
EUPARKERIA CAPENSIS Broom.
In addition to a number of isolated bones the following are the chief
specimens of this species, all of which come from a locality known as the
Krietfontein Spruit on the Aliwal North Commonage, and are of Upper
Beaufort age. (The numbers in brackets refer to the South African
Museum catalogue. )
7
82 Transactions of the Royal Society of South Africa.
Specimen 1 (5867).—The type. Consists of skull, lower jaw, shoulder-
cirdle, cervical and dorsal vertebrae, right fore-lmb without hand, abdominal
ribs, pelvis, and most of the left hmd limb. Associated with this by Broom
is another dentary, and a complete pes.
Specimen 2 (6047).—A complete skull and lower jaw, humerus, pelvis
and shoulder-girdle (described previously as Browniella africana), femur
in articulation with pelvis, vertebrae, ribs, abdominal ribs.
Specimen 3 (6048).—Skull, of which the bones are disarticulated and
scattered but not broken, pelvis, of which the iliuam was figured by Broom
(‘ Proc. Zool. Soc.,’ 1913, pl. Ixxvin, fig. 18), hind limb, some vertebrae and
ribs.
Specimen 4.—Dorsal vertebrae, pelvis (ilium figured by Broom, ‘ Proce.
Zool. Soc.,’ 1918, pl. Ixxvii, fig. 17), hind limb, coracoids and interclavicle.
Specimen 5.—Disarticulated skull, scapula, fore limb, hind hmb, and
some vertebrae.
Specimen 6.—A portion of the antorbital part of the skull and lower jaw
showing a little of the palate.
Skull.—Specimen No. 2 contains an almost complete skull and lower
jaw, of which some of the bones are slightly displaced. Pressure has
cleaved the skull along the middle line except at the snout, and has crushed
the left side so that the left frontal is vertical and the left nasal is horizontal,
but at a lower level than its neighbour. ‘The left ramus of the mandible
is in its correct position but the right ramus has been displaced. All the
sutures are beautifully shown, but accurate measurements are not possible,
save those of each separate bone.
The skull is undoubtedly of the same species as the type and almost of
the same size; but it displays one or two features which are not shown by
the type-specimen.
The front part of the left premaxilla is as conjectured by Broom, being a
slender pillar of bone bounding the front of the large nostril and meeting
the nasal above it. The postnarial portion of the bone is seen to overlap
the descending part of the nasal on the outside, and the suture between the
two is somewhat different from that in Broom’s figure. The premaxillary
teeth are not seen.
The nasal-maxillary articulation is longer than in the type; the fronto-
parietal and fronto-postfrontal sutures are jagged; the maxilla and jugal -
are supphed with a number of small foramina, mostly passing forwards into
the bones; and the distal end of the quadrate is broader than in the figures
given. Apart from these facts nothing can be added to Broom’s description,
with which this skull closely agrees. No sclerotic plates, however, are
preserved. \
The lower jaw shows several features which are not displayed by the
type. The dentary forms the anterior half of the ramus and forms the
On the Reptilian Genera Euparkeria Broom and Mesosuchus Watson. 83
symphysis, articulating posteriorly with the surangular and angular. The
lower view of the ramus shows a splenial lying within the dentary forming
part of the lower border of the jaw, passing back almost to the level of the
front of the external mandibular foramen but not reaching the symphysis
anteriorly. Lying wedged between the lower half of the dentary and the
posterior half of the splenial is the anterior prolongation of the angular.
The angular forms the lower border of the external mandibular foramen,
and the lower border of most of the hinder half of the ramus. The suran-
cular lies dorsal to the angular, and extends from the dentary to the
articular. Just anterior to the articular it has a fairly large nutritive
foramen ; above the external mandibular foramen and just below the dorsal
edge of the bone is a well-defined groove leading backwards to a small
foramen ; and the upper edge of the bone is thickened to form a small
flange. Lying below the posterior half of the surangular and apparently
passing between it and the angular is a slight splint-bone which I take to be
a prearticular. It is small. The limits of the articular are not well defined.
‘At the actual articular surface it is bounded outwardly by the surangular,
which passes back to lie outside the postarticular process. The articular
surface is shallowly concave and broad for the reception of the broadened
quadrate. I can see no evidence of a coronoid.
The teeth, at least in the lower jaw, are serrated on their anterior and
posterior borders, the posterior serrations being much the coarser of the
two sets. I can see no definite trace of anterior serrations in the upper jaw,
and some teeth are certainly without.
Specimen No. 6 is of interest as showing a small fragment of the
palate in association with part of the side of a skull. The specimen is
somewhat unsatisfactory, but it is possible to identify the elements
present. Most of the left orbital and preorbital region is preserved, and the
front of the left ramus of the lower jaw ; the stone containing the specimen
has split almost longitudinally, so that the inner face of the maxilla is seen on
one piece and the outer side of the jugal on the counterpiece. Comparison
of the lower jaw with the type shows that the specimen is not quite full
grown. The lower jaw shows definitely that the splenial plays no part in the
symphysis. The only real point of interest in the specimen is the evidence
of small teeth on the palate. These are short, blunt, smallish teeth exposed
ina row of four or five on a bone, which apparently articulates latero-
anteriorly with the hinder part of the maxilla. The teeth begin behind the
level of the orbit and are placed in the long axis of the skull. It is
probable that they are borne by the pterygoid, as in the somewhat earlier
form Proterosuchus fergust.
It should be noted that there is in the collection another fragment
showing a portion of the maxilla with two thecodont teeth, serrated pos-
teriorly ; and associated with it a bone of the palate carrying a row of eight
84, Transactions of the Royal Society of South Africa.
short pointed slightly backwardly-curved small teeth, apparently unserrate.
It seems indisputable, therefore, that Huparkeria had palatal teeth and was
thus more primitive than Ornithosuchus—a conclusion strengthened by the
discovery of the prearticular in the lower jaw of the South African form.
Humerus.—The humerus of the type is only displayed in its outer
aspect ; but a complete left humerus has been isolated from skeleton No. 2.
The greatest length of the bone is 45 mm., its maximum proximal width
20 mm., and its distal width 16 mm. The shaft is very slender, but the
bone expands considerably at the ends, especially proximally. The lower
end of the deltoid crest is only 14 mm. from the proximal edge of the bone,
and the crest thus lies high up as in Ornithosuchus. The inner face of the
proximal end is mostly flat, but 1s slightly concave laterally. The distal
condyles are separated by a basin-like depression. ‘There is no entepicondylar
foramen.
Ilium.—The specimen from which Broom figured the imperfect ilium
(1913, fig. 18) also contains a perfect right ilium which has been fully
developed. The whole bone is 20 mm. high and the ilac crest 28 mm. long.
The acetabulum is large and closed and there is a very pronounced supra-
acetabular crest. The height of the acetabular portion of the bone is 13 mm.
The anterior spine of the crest is short and bluntly rounded, the posterior
spine long. The acetabular crest is continued on to the pre-acetabular
process, but not to the post-acetabular process. The lateral face of the
upper portion of the bone is strongly supphed with ridges running at right
angles to the upper border, especially in the anterior portion.
Femur.— Directly associated with the pelvis figured by Broom as that of
Browniella africana I have discovered a left femur. The head of the bone
lies just under the ischium and the bone is at right angles to the plane of
the ischium. It 1s closely similar to that of Huparkeria capensis, having
a length of 59 mm., the femur of Browniella being 74 mm. long; and this
coupled with the fact that a skull and lower jaw of Huparkeria was found in
the sand-stone lying directly under the same pelvis opens up the question
of the identity of the pelvis figured by Broom.
Broom has referred to the differences which exist between this pelvis and
that of the type of Huparkeria. ‘The ischium differs in shape in being
constricted near its middle, and the pubis, besides being much broader and
stouter, differs in having only a single pubic foramen.” Further develop-
ment and a careful examination of the ‘“ Browniella” pelvis seem to show
that these differences were rather too strongly accentuated. Ilium, pubis
and ischium are certainly larger than those of the other specimens of
Buparkeria examined; but their relative sizes are apparently the same.
The constriction of the ischial plate does not exist—the appearance of con-
striction is due to a broken edge of the bone, the true shape being evident
on the matrix beneath. The ilium is of exactly the same type as in
On the Reptilian Genera Euparkeria Broom and Mesosuchus Watson. 85
Euparkeria. There is some doubt as to the value of the other point of
difference mentioned. None of the pubes of Euparkeria are quite entire,
but in at least one there is no evidence of a second pubic foramen.
In view, therefore, of the general similarity of the pelvis to that of
Huparkeria, its articulation with a typical Huparkeria femur and its close
association with an undoubted Huparkeria skull and lower jaw, it must be
concluded that this pelvis is a large girdle of Huparkeria capensis, its
difference in size from the type being probably due to differences of sex.
It is worthy of note that Watson found similar differences in two skeletons
of Procolophon, of which he writes—‘ The male and female skeletons are of
exactly the same length, as are their skulls. The dentition is quite similar
in the two specimens, and I think there 1s not the slightest doubt that they
belong to the same species, The female skeleton is, however, much more
lightly built, having narrower and more slender lmb-girdles, and limbs
which are less than three-quarters as long as those of the male skeleton and
much less robust. Mr. Boulenger tells me that differences of this kind
often distinguish the sexes in living lizards, and I think it is extremely
probable that the same reason accounts for the difference between my two
skeletons.” In Huparkeria the limb bones are more closely similar in the
two sexes than in Procolophon, but there is a marked difference in the size
and strength of the pelvis and shoulder-girdle.
BROWNIELLA AFRICANA Br.
In view of the foregoing discussion, I take as the type of Browniella an
isolated femur, longer than that of Euparkeria and somewhat less curved.
The resemblances between the two are great, but from the figures it will be
seen that in Browniella the trochanter is considerably straighter. The distal
end is less broadened than in Huparkeria.
MeEsosucHus BROWNI Watson.
The fortunate discovery of a second specimen of this interesting form in
the Brown collection enables certain new facts to be added to our knowledge
of it. The specimen consists of a fairly complete articulated skeleton
lacking the skull—of which only a portion of a maxilla is preserved—of
which the pelvis, hind limb and fore limb have been developed. The type
of the genus is a somewhat unsatisfactory specimen; but this second piece
agrees very closely with the type in size and specific characters. It is unfor-
tunate that the skull is not preserved.
The fragment of maxilla is about 380 mm. long and carries nine blunt
round acrodont teeth, which are not arranged in a single row but somewhat
irregularly.
Of the fore limb the parts preserved are the distal end of the humerus,
86 Transactions of the Royal Society of South Africa.
the radius and ulna and the hand. The distal end of the humerus is
17 mm. broad. It has two distinct condyles with a broad shallow depression
between them. There are no epicondylar foramina.
The radius has a length of 35 mm., and consists of a thin shaft expanding
at each end to a width of about 7 mm. ‘The distal end is cup-shaped for
the reception of the radiale, which is closely applied to the bone.
The ulna is not fully exposed. Its length may have been slightly less
than that of the radius. The bones of the forearm are crossed, so that the
distal end of the ulna lies on the opposite side of the radius to the proximal
end.
The bony carpus consists of six elements—two fairly large proximal ones
and four smaller distal ones. The largest bone articulates closely with the
radius and is irregularly quadrangular in section. The other proximal element
hes at the distal lateral corner of the radiale, well separated from the ulna.
Its lateral face is excavate as if for the reception of another, possibly
cartilaginous, element. Of the distal elements the second is small, the first
and third are equal to one another in size, and the fourth shghtly larger. The
first, second and third articulate wth metacarpals I, II and III; the fourth
with metacarpal IV and partly with metacarpal V, which also articulates
apparently with the outer proximal element. The metacarpals have the
following lengths: I,5 mm.; I, 10 mm.; III, 12 mm.; IV, 135 mm.;
V,6mm. The digital formula is 2, 3, 4,5, 3 (the claw of the fifth finger
is displaced), and the fingers (excluding the metacarpals) have the following
lengths: 1,10 mm.; 2,13 mm.; 3,16 mm.; 4,18 mm.; 5,8 mmso ite
claws are comparatively long and pointed. All the fingers are slender.
The fore foot is smaller than the hind foot.
The whole pelvis is preserved, practically undistorted. The left ilium is
entire. As Broom has pointed out, it differs considerably from that of
Huparkeria and resembles much more closely that of Howesia. The maximum
vertical height of the bone is 28 mm., the height above the supra-acetabular
ridge 16 mm. The length of the dorsal edge is 25 mm. The posterior
extension of the crest is much less than in Huparkeria. The acetabulum is
closed. |
The ischium is a plate-like bone meeting its neighbour in a nearly com-
plete symphysis, and sloping downwards and inwards from the acetabulum
to form a very obtuse-angled depression. It is considerably shorter than
that of Huparkeria. Its greatest length cannot have been more than 27 mm.,
its anterior width 13 mm. The pubis is 21 mm. broad posteriorly. It is
plate-like, with the anterior half of the plate bent nearly at right angles to
the posterior half, so that the two pubes together form a nearly vertical
wall of bone. The anterior edges are considerably thickened laterally.
The posterior edges of the bone are not quite straight, so that the articula-
tion with the ischia was not complete, a lozenge-shaped area being left
On the Reptilian Genera Euparkeria Broom and Mesosuchus Watson. 87
uncovered by bone in the centre of the pelvic basin. The pubis has one
pubic foramen. The bending of the bone is greater than in Howesia and
equal to that of Huparkeria.
The femur is a short stout bone 50 mm. long, very different from that of
Euparkeria. It is bent in a slight double curve. The head must have been
cartilaginous, as the proximal end of the bone is excavate and finely rugose,
approximating in shape to that of Howesia. There was a strong high
trochanter at the proximal fifth of. the bone. The bone differs from the
femur of Howesia in being less expanded at the distal end and in having a
stronger curve.
The tibia is 49 mm. long, the fibula 45 mm. The proximal surface of
the tibia has a maximum width of 12 mm., the shaft being 5 mm. thick.
The fibula is a much more slender bone with a somewhat expanded distal
end.
The tarsus of the right side is seen from the plantar side, and is slightly
displaced. In the proximal series there are three bones. A large bone,
obviously formed by the fusion of two, hes directly opposite the distal ends
of the lower bones of the leg. It is 14 mm. at its widest and has a maximum
leneth of 7 mm. Lateral to it and apparently lying partly above it is a
bone which has only a small triangular plantar face. Hxternal to this and
partly articulating with the first is another large bone whose inner edge is
raised and whose under-surface is saddle-shaped. The dimensions of this
bone are 11 mm. by 8 mm. In the distal row four tarsalia are seen, of
which the fourth is the largest. ‘The metatarsals are not fully displayed,
with the exception of the first and fifth. The first is a short stout bone ;
the fifth 1s remarkably similar to that of Howesia. The digital formula is
not obtainable.
There can be little doubt that in the structure of the girdles and limbs
Mesosuchus and Howesia are closely allied forms. ‘The pelvis is very
similar in the two genera and so is the general form of the limb-bones,
while the similarity in the fifth metatarsal is very striking. The manus is
not known in Howesia, and its interpretation in Mesosuchus is not beyond
doubt. Much of the carpus was undoubtedly cartilaginous. There were
certainly four distal carpalia. I have interpreted the other two bones as
radiaie and ulnare; the absence of a bony intermedium is strange, but there
is room for it between the two proximal bones of the carpus. There were
almost certainly cartilaginous centralia, and the lateral surface of the
supposed ulnare is excavate as if for articulation with a cartilaginous pisi-
forme. The digital formula of the hand is primitive.
The structure of the tarsus is fairly clear, although the whole foot has
been turned at an angle to the leg. ‘The large bone which articulates with
the tibia and fibula is taken to bea fused centrale and tibiale; lateral to this
and just possibly fused with it (although a suture seems undoubted on the
88 Transactions of the Royal Society of South Africa.
plantar side) is the intermedium. Lateral to this is the large fibiale with a
pronounced heel. This structure differs from that of Howesia in the fusion
of the tibiale and centrale. Such a fusion is paralleled in stage Q of
Sphenodon ; and a further fusion with the intermedium is seen in Procolo-
phon. In the rest of the foot Howesia and Mesosuchus agree. A specimen
from Burghersdorp (8.A. Mus. Cat. No. 5861) collected by Dr. du Toit
shows the fingers and toes ofa species of either Howesia or Mesosuchus. The
fifth metatarsal is of exactly the same type as in those genera, and the
digital formula of the pes is 2,8, 4,5, 3. The third metatarsal is the longest.
The femur in the Howesidae differs from that of the Pseudosuchia such
as Huparkeria and approximates more closely to that of Hrythrosuchus, but
in the latter the large medial trochanter is not terminal. The forms agree
with Mrythrosuchus, too, in the sudden bending down of the anterior portion
of the pubis; but the ilium lacks the posterior prolongation seen in both
Erythrosuchus and Huparkeria.
EXPLANATION OF PLATES IT ann ITI.
FIG. PLATE II.
1. Euparkeria capensis. Outer view of left dentary of Specimen No. 2.
2. Euparkeria capensis. Lower view of same.
3. Euparkeria capensis. Lateral view of humerus of Specimen No. 2.
4. Euparkeria capensis. Medio-anterior view of same.
5. Euparkeria capensis. Femur.
6. Browniella africana. Femur.
7. Euparkeria capensis. Pelvis of Specimen No. 2.
8. Euparkeria capensis. Right ilium of Specimen No. 3.
PLATE III.
Mesosuchus browni, Watson.
. Left ilium.
. Pelvis from below.
. Proximal end of femur.
. Femur.
. Fore limb, showing carpus.
. Hind hmb, showing tarsus.
WTO orf WN
. True view of metatarsal V.
Trans. Roy. Soc. 8. Afr. Vol. X. Plate IT.
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( 89 )
NOTE ON THE PRODUCT OF ANY DETERMINANT AND
ITS BORDERED DERIVATIVE.
By Srr Tuomas Murr, LL.D.
(1) The fundamental result here obtained is the theorem that the product
of two determinants, the second of which is got by bordering the first, is
expressible as a bilinear function of which the quasi variables are the cofactors
of the bordering elements in the second determinant, and the discriminant is
the unbordered determinant. For example, when the order of the initial
determinant is the 3rd we have
i | Ay My Ms. | Oy (Np Oe A, | = a Da Ds A ‘
| @ A oor 4
| by by Bs | by by bs b, eli) aR heey
| ©] Cg Cg} | Cy Cg C3 Cy | 6) C5 C3 | C,
dy dy dy . |
By way of proof we note that on the right-hand side the cofactor of A,
= 4D, + agD, + a3D;
= — Ay | Ay G3 Ay | + Ay | Ay My Ay | — My | Ay Ay Ay |
by bs by by bs b, by by dy
COE SA ang WG Ce, Ca
= — | A a Ae . = — |G & ag .
ly Ay Ay Hy oop ee ae
by by bs by by by bs by |
ily Woy aa O7Ge C3 Cas)
= — a,| a bg c|,
= cofactor of A, on the left ;
and the outcome is similar when the cofactors of B, and C, are considered.
(2) As every bordered determinant is already known to be expressible as
a bilinear, for example,
se dy d, d.,
Gi ho Garay = ;
b b b b — |bycs| |b, c3| —|bye4| 4
inte: aes] — ayes] Iayeal |,
“1 3 & C4 b b ba
Fa a — |agb5| |a93] —|ay9gl | Cy,
1 % G3 -
90 Transactions of the Royal Society of South Africa.
an alternative form for the preceding theorem is
Erie dy dy ds _ D, D, D3
by Oy ie ce ae on ‘ i i - —
batts — lage GCs] —la4Cg! | bg 1 9% 3) 4
|ab,| — |aybz| |adgl | Cg Cy Cy 31 Cy
where it is curious to note that the two bilinear functions are such that the
elements in the square array of the first are the cofactors, in | a,b,c; |, of
the elements in the square array of the second, and the elements in the
laterals of the second are the cofactors, in the bordered determinant, of the
elements in the laterals of the first.
(3) The theorem thus reached recalls another* in which occur two of the
same component parts, namely,
2 3 ie
| rT ie —Iay cz] | ¢g| —lay ey] By
b b, b., by by bs By, i b | b CG
ec, G: ty ty | O, | Vg] —|@, Og |@ Og) Uy
D, De
and, being thus able to combine the two, we deduce the still more interesting
equality,
by c3l —|b, cs} | d, co] Ay]
9
a“
Gags ian | = OATS On|
i a ema? oe. at ae —|a cz) |a, ¢3| —la, co] By,
rn, by bs b, b, bs by nA , |
| C) €2 © | Cy Cy Cg Cy | ty 3| — |a, 3 iy bo| C,
ld, dod, . | D, 1D; D, 2
oO
where the two bordered determinants are related in the matter of their
elements quite similarly to the two bilinears in §2.
As, however, the first factor on the left-hand side of the theorem which
we have quoted is, when general, raised to the power n—2, our deduced
result in its general form is that the product of any bordered determinant
of the (n + 1) order by the (n—1) power of the unbordered determinant is
expressible as a bordered determinant of the (n + 1) order also.
(4) We have next to note that the extent of the border in the foregoing
need not be restricted to one line: for example,
jdzb,C~| —|agb4c;| la, dye;
| ay bg Cg |. | Ay Ag Ag Ay As | SG aCsl la aesl Ia 105
|b, labs} argbs|| Cres!
by by bs by b;
ewe deck [ay C2] aes] [ges] | —|O142¢3|
d, dy dz . | | bye9| Iby¢3| [dacs |a,dye5|
Cues senders |
* «Trans. R. Soc. Edinburgh.,’ xxxii (1885), § 35.
The Product of Any Determinant and its Bordered Derivative. 91
and a quite similar mode of proof suffices. Thus, taking the aggregate
of the first three of the nine terms on the right, namely,
[gb yes] |aybyi — |agbye;| [aybs| + laybye;| od | » |e dyes
we see that it
== | Ay Ay Ae O, Az | . [C)does
= Se oie ° 4 a; ° |c, dye
b, bs
C€) Cy Cg Cy Ce
@ Ay As .
b, by by .
= | ayb; |. | ages | . | eydges | ,
which is also one of the three terms got on the left by using Laplace’s ~
expansion on the five-line determinant.
The order of the square array of the bilinear depends on the order of
the initiating determinant and the breadth of the border; for example,
By Uy My OM M1) A, Ag My As Ue
b, by by by) dy by bs by bs be
Cy Cy Cz Cy) | Cy Cg C3 Cy C5 Ce
d, d,d,d,| |d, d,d, d, d; d,
NM OR
Att ts -
is equal to the bilinear whose square array is
| abe | | aybg | eee | gb, |
| cq | fmyeg] wee | age, |
| ody | Jers | wees | ed, |
and whose laterals are
|asb4e5dq|,—|agbyesdg|, \agbsesdg|, laqbyesdg|, —|abgesdg], laybocsde|,
an
des fy|,—[Didges fy], [Dyeres als ladees yl, —layeresSals [aid2es Sf, |:
The general theorem may with useful enough fulness be enunciated as
follows: The product of an n-line determinant by the determinant got from
it by bordering with r rows and r columns is expressible as a bilinear function
whose quasi variables are the n-line minors of the multiplier that contain
the bordering rows and the n-line minors that contain the bordering
92 Transactions of the Royal Society of South Africa.
columns, and whose square array is that of the r** compound of the original
determinant.
(5) When we pass on to the case where the initiating determinant is
axisymmetric and is axisymmetrically bordered, another previously known
theorem can be brought into service with advantage, namely, the theorem
regarding the determinant whose matrix is the sum of an axisymmetric and
a zero-axial skew matrix. The 35rd order being taken this theorem is *
, 3 A y
| a@ f+y e-—p| = afe| + mae
f-y b ite fbd
e+pd—a ec edc| Wo)
edec| »
Returning then to our first result above, and taking g, h, k for the border
of the basic determinant here, we have
—|afe afek[= =e4Q,
god f.b adh F
if 1b a) Et
ede € 0.40 Gg
ed. |G
iin
and consequently, by the theorem just recalled,
a f+Ge—-H\=|afe|—|afe| afek
Fa) ee We Kl ieee fod| fbdh
e+H d—-K cc ede ede| jedcg
kWg a;
an unexpected result in pure determinants.
(6) When the determinant with the peculiar matrix referred to at the
beginning of §5 is of the 4th order, there is in the expansion an additional
term of a quite different type, namely, we have—
a gtw f—-v ltul=lag fl| +P4+|[w-—v u|?
g—w +b e+z k—y Giro er ke Z—Yy
ftv ez c ht -@ J feast al,
t—u ktyh—a ad ee Om!
where P is the bilinear whose square array is that of the 2nd compound of
the 4-line axisvmmetric determinant, and whose laterals are
DY Be Wh, Canres
On account of the existence of the said additional term
(aw — yv + zu)?
it might fairly he expected that the deduced theorem would no longer hold,
or, at least, not in the same form as before. Considerable importance
therefore attaches to the value which this additional term assumes when
* «Trans. R. Soc. Edinburgh,’ xxxix (1897), p. 222.
The Product of Any Determinant and its Bordered Derivative. 93
@, Y, 2, U, Vv, w are given the values which they hold in our theorem above.
Now, even where the basic determinant is not axisymmetric but is
| a,byce,d, | , as at the close of §4 above, we have
“ew — yu + eu
= | abesdg | | aybesdg | — | aabyesdg | | aybsesdg | + | aabjesdg | | aybscsdg |
and this being an extensional of
| ab, | | abo | — | aby | | abs | + | agbs | | a2, |
must vanish identically as the latter is known to do.
RonpeEsoscu, 8.A.;
27 April, 1921.
(95 )
COMMENTS ON DR. PERINGUEY’S NOTE ON WHALES.
By Artuur F. BEARPARK.
(With one Text-figure. )
With reference to the note on whales by Dr. Péringuey, published in
the ‘Transactions of the Royal Society of South Africa,’ vol. ix, pt. 1, I feel
that certain statements contained therein cannot be allowed to pass without
challenge.
To deal seriatim with the points in question, I will first draw attention
to the fact that the size of a whale cannot be safely estimated from the size
of its jawbones. The most notable factors affecting the size of the jaw-
bones are the age and sex of the animal, therefore more evidence is required
before a length can be assumed for the particular whale which Dr. Péringuey
describes (p. 73) as about 100 ft. long.
Balenoptera borealis (p. 74).—It is not easy to understand why Dr.
Péringuey should say this whale was considered as rare in the Northern
Hemisphere. The very name suggests the North. Those who have studied
whales look upon B. borealis as a very common species occurring in both
hemispheres. For instance, Lydekker, in his ‘Guide to Whales,’ etc.,
exhibited in the British Museum, p. 20, records that 771 of these individuals
were captured in Finmark in 1885. Further records of B. borealis in the
waters of the Northern Hemisphere will be found in the publications of—
Salvesen, T. E., ‘Journ. Roy. Soc. Arts,’ March 29th, 1912, p. 522.
Andrews, R. C., Assistant Curator of Mammals, American Museum
of Natural History, “Shore Whaling,” ‘ National Geographic
Magazine,’ vol. xxu, No. 5, May, 1911, p. 430.
D’Arcy, Wentworth Thompson, ‘The Scottish Naturalist,’ No. 81,
September, 1918, p. 198.
Southwell, T., F.Z.S8., ‘Seals and Whales of the British Seas, 1881,
(te Of
Beddard, F. H., M.A., F.R.S., ‘ Book of Whales,’ 1900, p. 155.
Harmer, 8. F., Sc.D., F.R.S., ‘ Cetacea Stranded on British Coasts,’
1915, pp. 5, 6, and 11; 1918, pp: 5, 12, and 13.
Bower, T. W., ‘ Alaska Fisheries, etc.,’ Report of the United States
Bureau of Fisheries for 1918. Bureau of Fisheries Document
No. 872, p. 64.
96 Transactions of the Royal Society of South Africa.
Cocks, A. H., M.A., F.Z.S., ‘The Zoologist,’ London, vol. ix, 1885,
No. 100, pp. 135, 141, 148; vol. x, 1886, No. 112, pp. 121, 122,
128, 129, 186; vol. xi, 1887, No. 126, pp. 207, 208, 211, 219, 222 ;
vol. xii, 1888, No. 138, pp. 201, 202, 204, 206, 208.
Therefore, in my opinion, the statement that B. borealis was considered
rare in northern waters is somewhat misleading.
Feeding habits.—Investigation of the food of whales is a matter of
vital importance in studying the great question of the migrations of these
mammals, and it is therefore to be regretted that the statements in the
note under review about the feeding habits of whales—as to whether they
are ichthyophagous or “ planctonophagous ”—are difficult to reconcile with
widely observed facts. Dr. Péringuey states (p. 74) that B. borealis, B.
physalus and B. musculus are not planctonophagous, except perhaps occa-
sionally, whereas, as a matter of fact, all these whales regularly feed on the
Plankton whilst in South African waters. From an examination of the
stomachs of some hundreds of specimens of B. musculus, B. borealis and
B. physalus in South Africa, Iam in a position to state definitely that it is
only on some occasions that fish, or the remains of fish, have been found
therein, whereas the small crustaceans are invariably present. I have not,
so far, observed any indication of a fish diet in the stomachs of B. musculus,
and all available literature refers to this whale as feeding solely on the
Plankton. In the case of the two other species I am inclined to agree with
Andrews (‘ National Geographic Magazine, Washington, vol. xxii, No. 5,
p. 427), who states that ‘The whale-bone whales probably never eat fish
of any kind if other food is to be had, although there is some evidence that
B. physalus is in some degree ichthyophagous at certain periods in the
Northern Hemisphere, this state of affairs being probably due to partial
failure of the Plankton supply. Ichthyophagous whales are easily identified
by the presence of fish-scales in the stomach, which scales apparently take a
much longer time to digest than either the flesh or bones of the fish. On
the other hand, a Plankton-eating whale is, as a rule, very easily identified
by the exudation of terra cotta-coloured feces, but if such be not present, an
examination of the stomach will instantly determine the question. Dr.
Péringuey probably knows that I hold no brief for the Norwegian whalers
to whom he refers (p. 75) as being unreliable, yet if he would go for a trip
on a whaler, any experienced Norwegian gunner would probably be able to
show him Fin, Blue and Sie Whales actually feeding on the Plankton.
If the investigator requires further information about the food of the
whalebone whales, I would refer him to the following publications :
D'Arcy, Wentworth Thompson, ‘The Scottish Naturalist,’ No. 82,
October, 1918, pp. 232-233.
Clark, R. 8., M.A., B.Sc., ‘South Atlantic Whales and Whaling,’
1919. (Shackleton’s 1914-1917 Expedition.)
Comments on Dr. Péringuey’s Note on Whales. Si
Jardine, W., F.R.S.E., F.L.S., ‘ Cetacea,’ 1837, pp. 128, 186.
Burn Murdoch W. G., F.R.8.G.8., ‘Modern Whaling and Bear-
Hunting,’ 1917, p. 254.
Lydekker, ‘ Guide to Whales in the British Museum,’ pp. 18, 20.
Beddard, F. E., M.A., F.R.S., ‘Book of Whales,’ 1900, pp. 154, 156,
158.
Southwell, T., F.Z.S., ‘Seals and Whales of the British Seas,’ 1881,
Dae.
Andrews, R. C., Assistant Curator of Mammals, American Museum
of Natural History, “Shore Whaling,” ‘National Geographic
Magazine,’ vol. xxii, No. 5, May, 1911 (Washington), pp. 427
and 431.
Olsen, O., “The Bryde Whale,” ‘ Proc. Zool. Soc. Lond.,’ pt. iy,
December, 1915, pp. 1073-1090.
‘Report of the Interdepartmental Committee on Research and
Development in the Falkland Islands,’ pp. 9, 10, 11, 40, 46, 47,
55, 73, 74, 75, 81, 90, 95, 101, 102, 117, 118, 158, 184.
A study of these, along with the information I give, will, I submit,
demonstrate that Dr. Péringuey is in error about the feeding habits of the
whales.
Dr. Péringuey’s conclusion (p. 74), that B. brydei is ichthyophagous, is
correct, but his premises are wrong. One cannot arrive at a conclusion as
to whether a whale feeds on the Plankton or fish from the evidence of the
baleen fringe. For instance, this fringe is at least as coarse, or probably
coarser in B. musculus than in B. brydei, but there is no evidence that B.
musculus is ichthyophagous. All observers agree that the food of the
B. musculus consists of small crustaceans, therefore Dr. Pcéringuey’s reason-
ing breaks down in the face of established facts. Olsen states (‘ Proc. Zool.
Soc. Lond.,’ pt. iv, December, 1913, pp. 1073-1090) that the food of B.
brydei consists chiefly of fish, but that it occasionally takes crustaceans, with
which observations I agree.
Dr. Péringuey’s remarks about the asymmetry of skeletons of Cetacea are
by no means clear to me, but the difference in shape of the sterna of
B. borealis and B. brydei, as shown by the photographs, is of considerable
interest. Not, however, from the standpoint of demonstrating the specific
distinctness of the two mammals, which has already been established, but
because the sternum of B. borealis, as shown in the photograph, is quite
different in shape from the sternum of a normal specimen of this whale.
The abnormal form of the breast-bone in the photograph is remarkable, but
it is not safe to use this specimen as an example for comparison with the
sternum from another species of whale. I submit a photograph showing
normal specimens of the sterna of B. borealis.
Humpback whales.—I have measured a number of these in South Africa
9
98 Transactions of the Royal Society of South Africa.
which have exceeded 50 ft. in length. Such whales are not rare, therefore
Dr. Péringuey’s statement on p. 75 should be modified.
The statement that this species possesses a penis bone requires confirma-
tion. Eleven humpback whales recently examined by me gave negative
results. The alleged penis bone shown in position in the skeleton of the
Humpback whale in the South African Museum, Capetown, resembles very
closely one of the pelvic bones of a sperm whale.
It is not my intention to speak here on the great question of the possible
extermination of whales. I submitted some suggestions for the control of
the whaling industry to the Dominions Royal Commission, March, 1914
(Minutes of Evidence, pp. 822-329), and, inter alia, the question of Antarctic
A is that of a male 46 ft. 4 in. long.
B is that of a female 51 ft. 9 in. long.
whales migrating northwards in order that their young should be born in
comparatively warm seas was discussed.
I would like, however, to remark that there is no evidence which points
to any possibility of whales being exterminated. ‘There is, however, reason to
believe that the whaling industry will suffer through whales becoming
scarcer.
IT am dealing elsewhere with the whole question of the preservation of
whales and the whaling industry; my object in submitting the present
comment is the purely scientific one of endeavouring to apply the test of
careful observation to assertions claiming to be fact.
(99 )
NOTE ON THE PECTORAL FIN OF THE SOLE, ACHIRUS
CAPENSIS: ITS ORIGIN AND SIGNIFICANCE.
By J. D., F. Giucureist.
(With Plate IV.)
The changes which take place in the body of flat fish in correlation with
alteration of the position of the body are of particular interest from an
evolutionary point of view. These changes are not so far back in the
genealogy of the animal that we cannot hope for some light on the question
of their origin, and the more recent such changes are the more instructive
they will be. The following case may therefore be of special interest in this
respect. It is also suggestive of some wider problems, such as the origin
of paired limbs, on which some remarks are made.
The pectoral fins of the soles have lost their original functions of balancing
the body, and we find that their subsequent fate has been very diverse. In
some species they become very much enlarged; in others they disappear
altogether. This is well illustrated in the South African soles, of which
there is a great variety. Among these there is a small sole, Achirus capensis,
which has been described as devoid of pectoral fins. It is common on the
sandy shores of False Bay, and as it can readily be kept in confinement, it
is a very suitable subject for the study of colour-adaptation to various
surroundings. It was while some experiments of this kind were being
made that a pecuhar action of the branchial aperture was observed, for, in
inspiration, not only did the opercular membrane close down against the
body, but a small flap of transparent skin was seen to rise from the body to
meet it, thus forming a double valve for the closing of the aperture. Other
living specimens were examined, and in all this little valvular membrane
was found, apparently functioning as an accessory organ in respiration.
On one occasion, however, a number of very small soles of this species was
procured, and it was observed that the small flap, though present and well
developed, lay applied to the body, and did not function in any way in the
process of breathing.
This structure may be best examined in an adult specimen. In one,
100 Transactions of the Royal Society of South Africa.
128 mm. in length, it is readily seen by raismg the membranous border of
the operculum, which conceals it. It consists of a transparent fold of skin,
which at its upper end is continuous with the opercular membrane. It
attains its greatest breadth (1°3 mm.) about 2 mm. from the upper angle
of the opercular opening, and from this broader part gradually tapers away,
the total length being 7 mm. The whole structure is situated over the
clavicle, to which, however, it is not attached, and that it represents a
reduced pectoral fin is indicated by the presence of six dermal rays, though
no traces of other elements (radials) of the limb are to be seen. These
dermal rays are widely set apart, and are so arranged as to form a support
for the elongate fold. The rays taper off towards the border, near which
they end. This border is slightly emarginate between the distal ends of the
rays, the emargination being much more marked in movement in the
living condition (P]. IV, fig.b). The two halves of each ray are fused together
throughout their length, except at the proximal end, where they appear as
short stout forks, the function of which in normal fins is to secure the rays
to the radials. As these radials are absent in this case, the meaning of the
marked development of these forks here may be interpreted as a device for
the more effective control of the valvular pectoral in its movements.
The upper and lower ends of this fin may be particularly noticed (Pl. IV,
fig.a). At its upper end it is distinctly continuous with the margin
of the opercular membrane, and but for the presence of the rays the whole
structure might be interpreted as an extension of this margin on to the body.
At its lower end it is continued as a rayless membrane, gradually becoming
narrower. This rayless part is about half its total length, which is
considerable, being about a third of the whole opercular opening.
This structure differs very considerably from the normal fin of fishes.
The mesodermal supporting elements of the fin have been lost, the dermal
rays reduced in length, though comparatively stout; their proximal ends are
well developed and spread out. Though the fin is shorter in length, its base
is relatively large, so as to form a thin vertical membrane, which is supported
by the altered position of the rays. Not only has the structure of the fin
been very much altered, but the function is totally different.
We have therefore here an illustration of the appearance of a new organ,
which apparently does not occur in any other fish,* and is of comparatively
recent origin.
It can be accounted for, as any modification of the body can be accounted
for, by a process of natural seléction of favourable germinal variations.
Such an explanation cannot, however, with our present knowledge of cell
structure and function, either be proved or disproved, and we may meantime
look round for some other possible suggestions.
* Since this paper was in print, I have noticed a similar fold, but without rays, in
some of the Chimaeridae, and that, in the living Callhorhynchus, it functions as an
accessory valve in respiration.
Note on the Pectoral Fin of the Sole, Achirus capensis. 101
We may assume that the altered position of the body of the flat fish
was not assumed suddenly and by all its members at once, and that it came
about probably in correlation with a changing external environment. <A
change ip environment, brought about primarily by a simple change in
physical conditions, may be followed by much more complex changes in the
actual surroundings of the organism. By way of illustration we may
suppose that the first change was a purely physical one, as, for instance, the
gradual deposition of mud. Secondary changes would follow, the mud
becoming populated by a variety of its characteristic forms of life. These
forms in their turn would attract other predatory fish, etc., which feed on
them, and so a great many new factors—these and others—would arise
which could possibly lead to changes in the ancestral sole. One possibility is
suggested by an observation on a particular species of fish, the blenny, Clinus,
of which there is a great variety at the Cape. It is active in its habits,
and preys on other small fish, crabs, etc. Its eyes are well developed, and
resemble the motile eyes of the chameleon. In captivity it was seen
frequently to seize and devour small mullet swimming above it at the
surface, and it was observed that, in order to watch its prey more readily, it
frequently lay over on one side, thus directing an eye upwards. It can be
readily concluded that if this device is adopted for detecting prey, it will
serve equally well for detecting enemies, and may have been adopted by flat
fishes; another possible factor is the changed source and changed kind of
food, consisting of small animals in the muddy ground. The reason for the
change of position need not, however, be considered here, and these possible
factors are merely mentioned to indicate how complex the external environ-
ment may become. The suggested change in environment by a gradual]
deposition of mud is also purely illustrative. A change in environment
may be brought about in other simple ways, as, for instance, by gradual
migration of the animal itself to other localities.
With the change from a vertical to a horizontal position the function of
the pectoral fin of maintaining the body in a vertical position would be lost,
and, owing to disuse, it would tend to diminish in size. This diminution in
size, accompanied probably by more complex physiological changes, would
of course appear in the succeeding generation if the identical factors which
brought it about in the parent were present, and it follows that if these were
more pronounced, as they would-be in a changing environment, the effects
would be more marked. This process of diminution would be arrested at
any stage, in which it began to function in another capacity, as has occurred
in the Achirus.
It might be suggested that the reduction of the pectoral fin is in co-ordi-
nation with the elongation of the body as happens in the case of many fish.
This, however, is not the case in Achirus, where the body is not elongate, but
rather short in proportion to its depth.
102 Transactions of the Royal Society of South Africa.
If reduction of the fin in length to its present dimensions can thus be
accounted for, how can we explain the relative elongation of its base of
attachment to the body? Apparently this cannot be explained as an effect
of use of this portion of the fin, for, as noted above, in the early stages this
feature is well marked, though the organ is functionless. It is then merely
a dermal membrane in which rays are not developed. It may be borne in
mind, however, that the paired as well as the unpaired fins arise in this
way, the membrane being replaced by the permanent fin, whose base of
attachment to the body is, as a rule, relatively shorter. The presence of a
well-developed though functionless pectoral fin of this nature at an early
stage may therefore be accounted for by the facts, or rather the mechanism
of development.
A modification of the body acquired in this way by functional and
environmental changes would be hard to distinguish from a germinal or
“innate ” character.
The question as to whether such acquired characters may become fixed
in another way, viz. by so affecting the germ-cells that they would be repro-
duced in succeeding generations, if their original cause were removed, is
another though quite legitimate inquiry, namely that of the inheritance
of acquired characters.
There are so many pitfalls to be avoided in the discussion of the question
of heredity, largely due to the use of terms not strictly defined, that one
must walk warily. Thus the word “environment” has been here used in
two senses, first as the external surroundings (physical and biological) of the
body as a whole, and secondly as the external surroundings of an organ of
the body, including the other organs. It is necessary to use the term in the
wider sense, for the change in position may not have been in direct co-ordi-
nation with a changing external environment, but with some gradual
internal change in the organs of the animal itself. Wath this explanation
it will probably be admitted that the acquired characters referred to are
transmitted, or rather repeated in succeeding generations, for the same
cause acting under the same conditions will produce the same effect. It
may even be admitted that the changed environment so defined may become
so complex and fixed that these characters will be indistinguishable from
hereditary characters, acquired by a process of natural selection of favour-
able variations which have not been brought about by use and disuse. In
this particular case it is not at all obvious that the new organ is of any great
advantage to the animal. On account of its accidental position near the ebb
and flow of the current of water at the opercular opening it is kept in
activity, and, so long as it does not seriously interfere with the normal
functions of the body, its continued existence will be tolerated. Other
cases apparently of this nature are known in the animal kingdom, and
perhaps this may be added to the list.
Note on the Pectoral Fin of the Sole, Achirus capensis. 103
Whatever the explanation of the process may be, however, there is little
doubt but that we have here the reduction of the pectoral fin to a simple
fold of the epidermis of the body, lying parallel with and resembling the
membranous epidermal border of the operculum. So striking is the resem-
blance that the idea of their possible homology is suggested, though such
resemblance is no proof of homology. The reduction of a pectoral fin to a
lateral fold of the body also recalls one of the oldest theories of the origin
of paired limbs, and is an inducement to inquire a little further into the
matter.
The simplest mechanism of respiration by gill slits would appear to be
the establishment of a current of water by ciliated epithelium, as in some of
the lowest vertebrates (Balanoglossus, Ascidians, Amphioxus). In the higher
forms, the Cyclostomes, there are gill pouches connected by tubular passages
to the exterior, where they open by small rounded apertures, guarded by
more or less circular rings of cartilage. In the Elasmobranchs, which
are provided with a more expansible pharyngeal cavity for inspiration,
there is a simple mechanism consisting of a series of vertical folds which
overlap the slit-hke external openings of the gills, and act as effective
valves, being mechanically closed in inspiration and opened in expiration.
These are very well developed in some Elasmobranchs (Chalamydo-
selachidae). In others (Chimaeridae) there is a great development of the
anterior of these flaps, which grows backwards over the gill openings, and
acts as a valve for the whole of the branchial apparatus forming the
operculum proper of the fishes.
In bony fishes there is a further elaboration: this enlarged operculum
strengthened by dermal bones takes over the functions of inspiration, while
the membranous posterior border retains the original valvular function.
Further, the bony operculum may come to subserve other functions, chiefly
defensive, and, in exceptional cases (Anabas scandens), locomotory. We
have therefore an illustration of how the simple epidermal flap which covers
the external opening of a gill slit may be transformed into a somewhat
elaborate organ, modified to perform other than its original functions.
There is some reason to suppose that in the primitive vertebrates the
number of gill slits were more numerous, and extended further along the
body than in later forms. The reduction in the gill slits began probably in
the more posterior ; they became closed up, and their gill arches and opercula
disappeared, but it is quite possible that some of these opercula may, like the
anterior operculum, have become developed into more elaborate organs
taking on some other function, and should the evolution of the vertebrate
body have been in the direction of a decrease in length and an increase in
depth, the function now performed by the paired fins is a likely enough one.
How the simple opercular flap could have tided over the critical period when
the gill opening which it covered became closed up may be indicated by the
104. Transactions of the Royal Society of South Africa.
case of the Achirus, for it could still perform a valvular function in con-
junction with the operculum in front of it, or it may well have been the
case that in the primitive vertebrate with its increased number of gill slits,
not only the opercular flap of the hyoid arch became enlarged to cover some
of the gill clefts, but that this happened also in two of the branchial arches.
It need not, however, necessarily find any form of useful activity to justify
its continued existence.
It may be significant in connection with this suggested opercular origin
of the paired limbs that in the lower vertebrates, such as Cephalochordata
and Cyclostomata, in which no opercular flaps are developed, though there
are branchial skeletal elements, there is no indication of the existence of
paired limbs at any time in their development or life-history.
The origin of paired limbs from such opercula is not a new suggestion, as
itis part at least of the hypothesis associated with the name of Gegenbaur,
who suggests that the limb-girdle and pterygia are derived from a visceral
arch and its rays, and (2) the pinna or distal part from the flaps or external
free part of the gill septum. He, however, did not apparently entertain
the idea that one of these parts of his theory might be correct while the
other might be wrong. Another theory (Graham Kerr) is that external gills
were in primitive vertebrates developed on each visceral arch and that these
gave rise to the paired limbs. This theory is not inconsistent with
Gegenbaur’s suggestion that the girdles are derived from visceral arches.
A third theory (Balfour) is that the paired limb is derived, not from any
external modification of the gill septum, but from an epidernal fold which
in primitive vertebrates is supposed to have occurred along each side of the
body. It seems to have been generally taken for granted throughout the
history of the discussion that this is inconsistent with the derivation of the
supporting structures of the limbs from visceral arches, but it is not quite
evident that it is necessarily so. It is true that in the elaboration of the lateral
fold theory it has been suggested that the differentiation into hmb rudiments
began at the anterior and posterior regions of the folds, but this is not a
necessary corollary of the hypothesis, for the differentiation may have
commenced at the anterior extremity in association with a disappearing gill
arch, and, on the movement backwards of this combined structure (epidermal
fold and supporting visceral arch), the process may have been repeated to
give rise to the anterior pair of limbs.
On the other hand, while all three theories look for an external epidermal
organ as the origin (in part at least) of paired fins, the derivation of the
supporting structures (girdles and radials) from gill arches is not bound up
with this origin. Any of the three views in this limited sense are consistent
with the origin of these elements de novo. Thus even Gegenbaur’s or Graham
Kerr’s hypotheses may be correct so far as the limb proper is concerned, and
yet open to question as regards the origin of its supporting structures.
Note on the Pectoral Fin of the Sole, Achirus capensis. 105
There are many illustrations in the body of how ready the mesoderm is
to collaborate with any process initiated by the ectoderm or endoderm, by con-
tributions of muscular or supporting tissues. Examples of this are abundant,
for instance, in the elaboration of the nervous system and the sense organs,
or, to take a more kindred example, in the provision of the supporting radials
for the vertical or unpaired fins. In fact this collaboration and assistance
is sometimes carried so far that the mesodermal elements usurp the place
and function of the original organs entirely, as, for instance, in the case of
the notochord initiated by the endoderm, and perhaps in the apparently
primitive pectoral of the Dipnoi initiated by the ectoderm.
The question, therefore, whether or not the skeletal elements of paired fins
are derived from the visceral arches or arise as in the case of unpaired fins
may be looked upon as supplementary to the question of the origin of fins as
such, the origin of radials and girdles being another though interesting and
important enough question in itself, particularly in connection with the
possible derivation of the skeleton of the pentadactyle limb from that of the
fish limb.
If, therefore, we must look to some epidermal structures or superficial
modifications of the body as the precursors of paired limbs, we have—(1)
external gills whose primary function is respiration, (2) a problematic longi-
tudinal fold of the body with the function, also problematic, of balancing the
body, (3) vertical folds of the body, whose function is primarily connected
with the mechanism of respiration by gill slits.
With regard to the external cill theory, it may be helpful to keep in view
the probable meaning of the external gills or other additional organs of
respiration, which appear to have displaced the simple external respiratory
organs so characteristic of animals devoid of gill slits. The most obvious
significance of the appearance of these simple organs is that the supply of
water to the gill slits has in some way or other become partially cut off.
This may occur under various altered conditions of environment, as, for
instance, in a prolonged embryonic stage, rendered possible by a more
abundant supply of food in the form of yolk. A protecting shell is present
in such cases, and this, of course, seriously interferes with the free access of
water. In some such cases the yolk and body may become highly vascularised
as in the large egg of Galeichthys feliceps, which is carried about in the mouth-
cavity of the parent and thus has the benefit of its respiratory apparatus,
or the egg of the Cape Bellostoma in the shell of which there are numerous
slit-like apertures. In the pelagic egg of a Cape fish the pectoral fin was
observed to be in constant motion—yet another device connected with
respiration. Compare the branchial character of the paired limbs in
Ceratodus.
In other cases, more particularly the embryonic Elasmobranchs, more
specialised supplementary respiratory organs are developed in the form of
106 Transactions of the Royal Society of South Africa.
external gills which project from the gill septum into the surrounding liquid
medium. The egg case is constructed so that the water can pass in at one
end and out at the other, and, in the somewhat elaborate egg of Callorhynchus,
there is a very perfect device for providing a current of water, as the long
flat tail of the embryo projects into a narrow part of the egg case, and keeps
up a constant undulatory movement.
This same diminution of the supply of water may occur in post-embryonic
life, and here we find similar devices adopted, both the simple increase of
vascular surface and the development of external gills.
T'wo simple cases of the first which do not seem to have been previously
recorded may be mentioned. One is that of a species of Goby (Trypauchen)
found in masses of mud brought up by dredge off Delagoa Bay on one or two
occasions. When alive it was of a bright red colour, which microscopical
examination showed to be due to blood contained in a network of fine vessels
spread over the whole body under the thin transparent skin. The other is
that of the blind deep-sea fish (Barathronus), a specimen of which was
brought up from a bottom of Globegerina ooze off the Cape. Another
example is the highly vascularised skin of the clawed toad, Xenopus, which
spends most of its life under water.
In post-embryonic stages we may also have the development of external
cills, under conditions unfavourable for an adequate supply of oxygen, as, for
instance, in the muddy or semi-aquatic environment of the Dipnoi, in which
external gills are developed.
It is obvious that a semi-aquatic environment must have been passed
through by the numerous vertebrates which now lead a terrestrial life, and
these, in all probability, passed through an external gill stage. Further, it
is not improbable that migration to the land conditions began as early as
migration to the pelagic conditions or at the Amphioxus—Myzine stage.
(In this connection it may be mentioned that the Cape Bellostoma is said to
make occasional excursions from its aquatic element.) What more probable
than that the primitive limbless vertebrates, equipped with external gills,
should, as has been suggested, use these as organs for the support and locomo-
tion of the body under its new conditions of life, and from these the
pentadactyle limb be developed ?
If the tetrapod limb has been evolved in this way from an external gill,
developed in connection with the transition of the primitive aquatic
vertebrate to terrestrial life, it can hardly be supposed at the same time
that from this organ arose the paired limbs of fishes, developed as organs
necessary in the transition to pelagic life, probably in association with the
shortening and flattening of the body.
There have been belated attempts among fishes to acquire the terrestrial
habits. The most successful, however, such as that of the Periophthalmus or
Anabas, have shown no approach to the acquisition of an organ of terrestrial
Note on the Pectoral Fin of the Sole, Achirus capensis. 107
locomotion at all similar to the pentadactyle hmb. Attempts to trace any
structural resemblance between a paired limb of any fish and that of a
terrestrial animal cannot be said to have been much more successful.
The theory of the origin of paired limbs from a lateral fold of the body
has at least this in its favour—that its suggested function was essentially in
the direction of an adaptation for an active pelagic life. The various
arguments for and against this theory need not be repeated, and only some
points noted which seem to indicate the origin of paired fins from vertical
folds rather than lateral folds.
(1) Vertical opereular folds of the epidermis exist in fishes at the
present day, and there can be no doubt but that they existed in the primitive
vertebrates, in which an expansible pharynx and its associated valvular
opercula first appeared, not unlikely before balancing paired limbs suitable
for a more pelagic life appeared, whereas, not only does no such longitudinal
fold as the hypothesis requires occur in any living fish, but there is no
satisfactory embryological or palaeontological evidence that it ever existed.
(2) The function of the vertical folds is an obvious one at the present
day, and even a necessity in the first primitive vertebrates, in which the
respiratory current of water was drawn in by the mouth by means of an
expansible pharynx provided with a framework of jointed branchial arches,
whereas the suggested function of the lateral fold, a keel for steadying the
body or a parachute-like expansion, are at the best conjectures, the first
perhaps not mechanically justifiable.
(3) That such opercular folds may attain greater dimensions is shown in
some primitive sharks, and more especially in higher fish, in which they may
have assumed the form of large organs taking on other functions, such as
assisting in inspiration, acting as defensive and offensive organs or organs
of locomotion.
(4) It accounts for the absence of paired limbs in the Cephalochordata
and the Cyclostomata in which no opercular folds are developed, and their
presence, except when secondarily absent, in all vertebrates in which such
folds appear. Gull septa provided with skeletal elements are present in the
Cyclostomata as in other fishes, but there are no opercula and therefore (?)
no limbs.
A great many objections have been raised to Gegenbaur’s theory as to
the origin of paired limbs from branchial septa from two points of view,
which may be distinguished from each other: first the suggested origin of
girdles and radials from the skeletal parts of a gill septum, and second, the
implied origin of the distal part of the limb from the gill flap or
operculum.
The objections to the first, which appear to be weighty, need not be gone
into, but the objections to the second apply equally to the opercular origin
and may be considered.
108 Transactions of the Royal Society of South Africa.
(1) The gill septa with their skeletal elements of the apparently primitive
Cyclostomes and Elasmobranchs are fixed, not projecting beyond the surface,
and therefore not likely to give rise to external organs like paired limbs.
This may be admitted in the case of the Cyclostomata, but it is just these which
have no paired limbs. In the case of the Elasmobranchs the opercula are
external organs and are well developed in Chladoselachidae and Holo-
cephah.
(2) Another objection to the origin of pectoral fins from vertical folds is
that it has been observed in most cases examined that when they arise as
folds of the body these are more or less longitudinal in direction, and it is
only subsequently that they assume a vertical position, and that by rotation
sometimes in one direction, sometimes in another. This certainly does not
corroborate this vertical fold origin, but it may be noted that the position of
the developing paired limb has not been investigated in many kinds of fish,
and that the paired limbs of Elasmobranchs are not necessarily primitive in
this respect. In this group and that of the Dipnoi, indeed, the endo-
skeletal elements seem rather to be taking the place of the primitive fin. In
some Achirus and in the larval form of a very large South African sole
(Synapturs microlepis) the rudiments of the pectoral have been observed to
be vertical in position.
In this connection, however, another theory of the origin of paired limbs
which has been much discussed may be noted, namely, that they arise from
a longitudinal lateral fold, not such as Balfour supposed to have occurred in
primitive fishes, but as it arises in the formation of the metapleural folds in
Amphioxus (Thatcher, ete.) These folds now function as a_ protective
covering for the external opening of the gill slits. Evolution of organs
is often in the direction of a reduction in numbers in co-ordination with the
elaboration of a few, and it is not impossible that with muscular pharyngeal
in place of the more primitive ciliary respiration the fold may have become
differentiated into valvular opercular flaps for the remaining gill opening.
This suggestion is very speculative as is also this theory itself, but it is
mentioned as a possibility which might indicate that the origin of paired hmbs
in ontogeny as longitudinal folds which only later become vertical folds is
not so inconsistent with their derivation from opercula as appears.
(3) It is objected that transverse folds across the body, such as the
opercula, would rather tend to arrest than to assist in the forward movement
of a swimming fish, but it is just for this purpose that paired fins are frequently
used, as may.be observed in the commonest types of fish in life. With
the shortened and deep body assumed by the vertebrates in taking to
a free-swimming life there is no such provision for the checking of the
forward movement as is found in the more elongate bodies, in which advance
or retreat can be effected by a reversal of the undulatory movement of the
body and paired fins are necessary for this purpose. This is confirmed by
No'e on the Pectoral Fin of the Sole, Achirus capensis. 109
the tendency in elongate forms for the pectorals to disappear. High speed
is usually associated with some means of controlling it.
The checking of forward progress in the primitive limbless vertebrates
could be effected by the only available structures, the transverse opercular
folds, and it may again be noted that the Cyclostomes which have no such
transverse folds or opercula have apparently never advanced to the paired
limb stage.
(4) It may be objected that there is no evidence of the free opercular
part of the gill septa becoming separated from the more internal part with
its supporting skeleton. The evidence against that part of Gegenbaur’s
theory dealing with the origin of girdles and radials from gill arches
and rays is in part a reply to this, and need not be repeated. It has
carried so much weight that his whole hypothesis of the origin of
paired limbs (from the vertical opercular folds as well as the gill arches) is
generally viewed with less favour than the lateral fold hypothesis. His
theory might still be partly true, even though the girdles be derived
from some one of the other sources which have been suggested—ribs and
other organs. (The extra-branchial cartilages, which are well developed in
some present-day sharks (e.g. Odontaspis) and form the main supports for
the gill flaps, are possible sources of the girdles, more especially as they are
situated in the somatopleure like the girdles, and not in the splanchnopleure
like the visceral arches.) ‘To the evidence already adduced of the possible
persistence of the opercula, when gills, gill-clefts and the rest of the gill
septum have disappeared, may be added that the operculum of living Teleostei
may find other or additional supports, as, for instance, in the Scleroparsi, and
that in the many cases of reduction of gill clefts the gill arch may be as
greatly reduced as the gill cleft.
It might be suggested that the fact that the pectoral fin can become an
epidermal fold supports the suggestion that an opercular fold may have
become a pectoral fin, as this is merely a reversal of the process. This
argument might have had more weight at one time, but recent investigations
seem to indicate that such reversals do not occur in the evolution of animal
life. This has been inferred from the fact that no clear cases are known,
but this inference does not, however, imply the existence of any law, and
cannot be advanced as an argument against the above suggestion, if indeed
the particular case of the Achirus may not point to the possibility of such
reversal, at least in measure. The argument, however, is not a good one.
It might be used also to indicate that the opercular fold disconnected from
the gill septum, for in the case of the Achirus the fold has lost its attachment
by radials to the shoulder girdle. It might be used also in favour of
another suggestion, that paired fins have arisen from median fins, for in
several soles the pelvic fin of one side may assume a median position and
become continuous with the anal.
110 Transactions of the Royal Society of South Africa.
Perhaps it may be necessary to note another point of view which has
been taken up with regard to this question of the origin of paired limbs. It
is pointed out that in development it is the mesodermal elements which
seem to take the initiative, that in the origin of the skeletal elements the
successive appearance of parts is from within outwards, and that a mere
epidermal fold cannot be looked on as the actual origin of limbs, for the
real structure of a fin begins only when the supporting mesodermal cartila-
ginous elements appear. With regard to the first two points, it is true that
ontogeny is often a valuable guide in tracing the rise of an organ, but not an
infallible one, and with regard to the last, the finished organ is often
totally different from its initial phases, as for instance in the case of the
axial skeleton of vertebrates. Besides, it is difficult to imagine that an
external organ of locomotion originated entirely from an internal structure.
It seems, therefore, quite legitimate to discuss the origin of paired fins apart
from that of the radials or girdles.
Reviewing the position as a whole with regard to the origin of unpaired
fins, apart from the question of the source of their secondary supporting
structures, the following hypothesis has been proposed: (1) The external
valvular flaps of the gill septum and (2) external gills attached to the gill
septum ; (3) longitudinal lateral folds of the body conjectures from certain
evidences to have been present in some primitive fishes; (4) longitudinal
lateral folds such as occur in Amphioxus; (5) unpaired fins.
The only one of these which seems entirely exclusive of the others is the
last, for which there is little or no evidence. What is suggested is that the
paired limbs of fishes arose as modifications of the external gill flaps, called
here for convenience valvular opercula, which appeared as separate active
external organs in co-ordination with muscular pharyngeal inspiration and
expiration, and that these opercula in their turn may have originated as
segmental differentiations of a supra-branchial lateral fold, originally used
in raising and sustaining the body in the water, and which now appears in
ontogeny as more or less disconnected longitudinal folds. This is not
opposed to the hypothesis that external gills which are vascularised gill
septa may have given rise to the paired limbs of Amphibia and terrestrial
animals.
Much more could probably be said in elaboration of the supposed oper-
cular origin of paired limbs. Thus a specially pleasing feature, and there-
fore perhaps to be regarded with caution, is that it appears to account for
the absence of paired limbs in the Cyclostomes which have a branchial arch
skeleton but no opercula, and shows that this negative feature is not secondary
but essentially connected with one of the three fundamental characters of
the Chordata, namely pharyngeal respiration by gill slits: in fact the phylum
could be divided into two sub-phyla on the basis of its pharyngeal
respiratory system and its derivatives (paired limbs) thus :
Note on the Pectoral Fin of the Sole, Achirus capensis. 111
(a) CYMNOPLEURA, with simple respiratory mechanism and no opercula
nor limbs, including Hemichorda, Cephalochorda, Urochorda and Cyclo-
stomata, each of which might also be rearranged on the same basis.
(b) CLADOPLEURA, with complex respiratory mechanism and paired limbs,
including fishes with paired limbs derived from opercula (Tetraptera), and
Amphibia, etc., with paired limbs derived from opercula modified as external
cills (Tetrapoda).
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ON THE SENONIAN AMMONITE FAUNA OF PONDOLAND.
By Ly EF; Spare, D.Se., F.G.8.
(With five Plates.)
CONTENTS.
A INTRODUCTION . ‘ ’ . 114 |. Sup-FamMiILty PACHYDISCINAE:
B. DEscRIPTION OF SPECIES . erage, SNF; Genus ParapacHypiscus, Hyatt:
Famity PHYLLOCERATIDAE : 17. P. aff. —_ ootacodensis,
Genus PHYLLOCERAS, Suess : Stoliczka sp. . 132
1. Ph. woodsi, v. Hoepen . 117 18. P. simplex, v. Hoepen
2. Ph. umzambiense, v. Spe : : . 133
Hoepen . : pth 19. P. umtafunensis (Crick
Famity LYTOCERATIDAE : MS.), Spath . - 3s
Genus GAUDRYCERAS, Grossouvre : 20. P. antecursor, v. Hoepen
3. G. tenuilineatum, v. sp. : a ise
Hoepen . : eas ie ey . P.? sp. nov.? é 134
4 G. varicostatum, v. ne FAMILY KOSSMATICERA-
Hoepen . : a a Aa TINAE:
5. G. cinctum (Crick MS.), Genus Maprasitss, Kilian and
Spath . : . 118 Reboul :
6. G. amapondense, v. 22. M. natalensis (Crick MS.),
Heepetie a) ok 118% Spat ity cc lane 134
7. G. sigcau, v. Hoepen . 118 23. M. acuticostatus ( Crick
Genus TETRAGONITES, Kossmat : MS.), Spath . a bad:
8. 7’. superstes,v. Hoepen. 119 24. M. faku, v. Hoepen
9. T. nuperus, v. Hoepen. 119 sp. s . 135
Genus PSEUDOPHYLLITES, Koss- 25. M. africanus, v. Hosnen
mat : Sp. : , . 135
10. P. andra (Forbes). Sea Ke) 26. M. aff. africanus, v.
Kamity DESMOCERATIDAE: Hoepen sp... Sy ess)
SuB-FAMILY PUZOSINAE: 27. M. sp. ind. oS
Genus Parapuzosis, Nowak : 28. M. similis, Saath . 135
ll. P. haughtoni, nov. + -k28 Incertae sedis—
Genus SCHLUTERIA, Grossouvre : Genus HopLoscapuHItEs, Nowak:
12. S. simplex, v. Hoepen sp. 129 29. H. sp. (cf. -semilaris,
13. S. crassa, v. Hoepen sp. 129 Stoliczka ?). . 136
Genus HAUERICERAS, Grossouvre : 30: Hf. ‘sp. (el. pavana,
14. H. gardeni, Baily sp. . 129 Forbes ?) A . 136
15. H. rembda, Forbes sp. . 131 31. H. sp. nov.? J 4 136
16. H.? sugata, Forbes sp. . 131 32. H.sp. ind. . : a, LoG,
10
114 Transactions of the Royal Society,of South Africa.
Famity ‘PRIONOTROPIDAE?’: 44, S. tenue (Crick MS.),
Genus Mortoniceras, Meek : Spath . ; . 144
33. M. soutoni, Baily sp. . 136 45. S. minor (Crick MS.),
34. M. stangeri, Baily sp. . 137 Spath . : . 144
35. M. stangeri (Baily), var. 46. S. amapondense, _ v.
sparsicosta, NOV. +. 138 Hoepen sp... . 144
36. M. stangeri (Baily), var. 47. S. wmzambiense, v.
densicosta, nov. . 188 Hoepen sp... . 145
Genus PSEUDOSCHLOENBACHIA, Famity NOSTOCERATIDAE:
Spath : Genus BostrycHocERAsS, Hyatt :
37. P.umbulazi, Bailysp. . 139 48. B. ? amapondense, v.
38. P. pseudofourniert, nov. 140 Hoepen sp. . 145.
39. P. papillata (Crick MS.), Genus DipLomoceras, Hyatt:
Spath . : . 141 49. D.? indicum (Forbes) . 145
40. P. griesbachi (Crick MS.), Genus OXYBELOCERAS, Hyatt :
Spath . : «14h 50. O. amapondense, v.
Incertae sedis— Hoepen sp... . 145
41. Gen. nov. (Muniericeras ?) Genus NEocrRIOcERAS, Spath:
cricki, Spath . = 41 51. NV. cf. spinigerum, Jimbo
Genus EULOPHOCERAS, Hyatt : sp. : . 146
42. E. natalense, Hyatt . 142 Famity BACULITIDAE:
Genus SPHENISCOCERAS (Crick Genus Bacuuirses, Lamarck :
MS.), Spath . 5 ibe 52. B. capensis, Woods . 146
43. S. africanum (Crick MS.), 53. B. sulcatus, Baily. . 146
Spath . : . 143 54. B. bailyi, Woods . . 146
A. INTRODUCTION.
In the description of the Upper Cretaceous fauna of Umkwelane Hill
in Zululand,* and in a paper-on “ Upper Cretaceous Ammonoidea from
Pondoland,” + reference was made by the writer to a collection of Pondo-
land Cephalopoda in the British Museum (Natural History). This collection
was presented by the Natal Government, through Sir F. Abel, in 1894,f
and thefossils were referred to by Kossmat,§ who examined them in the same
year. The late Mr. G. C. Crick described part of this collection, and the
writer originally had the intention of revising and completing the MS.
The description, apparently, was begun some time after Kossmat’s visit,.
but before the publication of Mr. Woods’s ‘“‘ Cretaceous Fauna of Pondo-
land ’’; || for, e.g., a specimen of Baculites had been given a new name by
Crick, which was altered to B. capensis after the appearance of Woods’s
* “Cretaceous Cephalopoda from Zululand,’ Annals S.A. Mus., xii, pt. vii, No. 16,
OZ, p. 223.
t+ Annals Durban Mus., iii, pt. 2, 1921, p. 39.
t See Catal. Natal Contrib. Colon. and Indian Exhib., 1886, p. 44.
§ “Die Bedeut. d. Siidind. Kreideform,” Jb.K.K.R.A., xliv (1894), Hefte 3 und 4,
(1895), pp. 463-4 ; also Rec. Geol. Survey India, xxviii (1895), pt. 2, p. 42.
|| Annals S.A. Mus., iv, pt. vii, No. 12, 1906.
On the Senonan Ammonite Fauna of Pondoland. 115
monograph in 1906. In this paper * Woods also mentioned that Crick
hoped to give an account shortly of other Pondoland species of Hulophoceras
in the British Museum. The omission of any reference to this genus, pro-
posed by Hyatt in 1903, in the (meaningless) description of the new genus
Spheniscoceras as “intermediate between Placenticeras and Sphenodiscus,”
may indicate that Crick’s account was written before 1903; a copy of
Hyatt’s description of the genus Hulophoceras, however, was added at the
end of the MS. by Mr. Crick with the remark, “ Not represented in this
collection.” This the writer is at a loss to understand, considering the
close affinity, if not identity, with Hulophoceras, of Crick’s genus Sphenisco-
ceras. There are, also, discrepancies, e.g. in the naming of some of the
forty-three examples of Mortoniceras, none of which is described in the MS.;
but a revision of Crick’s MS. has now become unnecessary, for recently
Dr. van Hoepen 7 dealt with many of the forms described by Crick. Accord-
ingly, only a few short extracts from Crick’s MS. will here be given, and it
is hoped that a general account of the collection and a revision of the generic
nomenclature of all the Pondoland Ammonoidea will prove of general
interest.
It has been considered advisable to include in the present paper a review
of the very important genera Pachydiscus and Parapachydiscus. When
describing some of the forms of this group in the “ Cretaceous Cephalopoda
from Zululand,” the writer recognised that Nowak’s f treatment of the
genus Pachydiscus, including in that one polyphyletic genus a host of
Upper Cretaceous Ammonites of many horizons, was unsatisfactory.
Much additional work, however, remained to be done, and even now, in the
absence of original material, it is feared that the interpretation of a number
of species here referred to must be based merely on the published descrip-
tions and figures. The genotype species, however, or at least representative
forms of the various lineages here recognised, are preserved in the British
Museum collections. This revision was necessary to enable the writer to
place some undescribed new forms, simply labelled ‘“ Pachydiscus’”’ by
Mr. Crick, but is provisional. The whole family Desmoceratidae, with its
trachyostracous descendants, including the ‘‘ Prionotropidae,”’ will be
dealt with in the writer’s forthcoming “‘ Monograph of the Gault Ammo-
nites.”’ _
After the writer’s observations on the Pondoland fauna § were penned,
an account of the stratigraphy of the Umzamba Beds by Mr. W. J. Plows
* Annals §.A. Mus., iv, pt. vii, No. 12, 1906, p. 337.
7 “ Cretaceous Cephalopoda from Pondoland,” Ann. Transv. Mus., viii, 1921, pt. 1.
t “ Untersuch. wi. d. Cephalop. d. Ob. Kreide in Polen,”’ pt. iii, Bull. Acad. Sc. Cracovie,
Cl. Sc. Math. et Natur., sér. B, Sc. Nat., Juin 1913, pp. 337 and ff. ;
§ Loc. cit. (Pondoland), pp. 53-56.
116 Transactions of the Royal Society of South Africa.
was published.* His paper is of interest, since he records Mortoniceras
soutont from bed 14, which is considerably higher than the basement bed
from which the Survey had recorded Pseudophyllites indra. Mr. Plows
also points out that this basement bed “is visible for the whole length of
exposure at low tide, and is also probably the basement on the right bank
oftheriver.” f There should then have been no difficulty in identifying this
basement bed. The writer also agrees with Mr. Plows in considering that the
fossil contents of these beds could and should be zoned.
Dr. van Hoepent is ‘‘inclined to regard the Pondoland Beds as of Upper
Santonian age.” Butit should be pointed out that the type of Grossouvre’s
Gaudryceras rouviller,§ which species was cited by Dr. van Hoepen, is from an
unknown horizon, whereas the very immature specimen of fig. 10, of Upper
Santonian age, is more depressed than G. sigcau, v. Hoepen. According to
Dr. van Hoepen, there are also differences in the suture-line. Again, the
South African species of Schliterva do not resemble the type of Grossouvre’s
Desmoceras pyrenaicum. ‘‘ Lenticeras”’ jullient, Pervinquiere, is not con-
sidered to be related to the Pondoland genus Spheniscoceras, which has
priority before Dr. van Hoepen’s “‘ Pelecodiscus’”’; and “‘ Schloenbachia ”’
fournert, Grossouvre,|| was on a previous occasion characterised by the
writer as being more nearly allied to Gauthiericeras § of the Coniacian. The
Pondoland deposits, or rather the great majority of the Ammonites, may
thus still be considered to be of Uppermost Senonian, or more precisely of
“Campanian plus Maestrichtian’”’ age, as stated in the writer’s previous
accounts, where it was pointed out that the five known zones of this Upper
Senonian and Maestrichtian probably represent only part of the true
succession of horizons. The chances, then, are that a fauna like that of
Pondoland, which does not quite agree with, e.g., either the Upper Campanian
of Galicia or the Valudayur Group of Southern India, if homogeneous at all,
may belong to a hitherto unrecognised intermediate horizon or even horizons.
Perhaps, since non-sequences are possible, these horizons are not even
consecutive.
The writer’s thanks are due to Dr. A. Smith Woodward and to Dr. F. A.
Bather of the British Museum, to Mr. Henry Woods of the Sedgwick
Museum, Cambridge, to Dr. Rogers, to Dr. Peringuey, and to Mr. EH. C.
Chubb for assistance in various ways.
* “The Cretaceous Rocks of Pondoland,” Annals Durban Mus., iii, pt. 2, pp. 58-66,
pl. viii.
+ dbid., p. 62. PoC. cit. ps4.
§ “Les Ammon. d. ]. Craie Supér.,”> Mém. Carte Géol. Dét. France, I], Pal., 1893 (4),
p228, pl. XXxXvil, fis. 7.
|| Lb1d., p. 112, pl. xxxv, fig. 1.
q{ Loc. cit. (Zululand), p. 240.
On the Senonian Ammomte Fauna of Pondoland. iG
B. DESCRIPTION OF SPECIES.
Famity PHYLLOCERATIDAE.
Genus PHyYLLOCERAS, Suess.
1. Phylloceras woodsi, v. Hoepen.
1921. Phylloceras woodsi, v. Hoepen, loc. cit., p. 3, pl. i, figs. 1-6,
text-fig. 1.
1921. Phylloceras nera (Forbes), Spath, Pondoland, p. 40.
The writer examined only one immature example of this species, but
Dr. van Hoepen has since shown that the Pondoland form differs in suture-
line from the Indian type.
2. Phylloceras umzambiense, v. Hoepen.
1920. v. Hoepen, ‘‘ Descr. of some Cret. Amm. from Pondoland,”’
Ann. Transvaal Mus., vii, pt. ii, p. 142, pl. xxiv, figs. 1-3.
1921. Spath, Pondoland, table, p. 50.
This is a more inflated form than the last, and was compared by the
writer to the Patagonian form of Ph. ‘ nera,’ described by Paulcke, and to
Ph. pergenst, Grossouvre sp.=Ph. velledae, Sharpe non Michelin sp.
Famity LYTOCERATIDAE.
Genus GAUDRYCERAS, Grossouvre.
3. Gaudryceras tenuilineatum, v. Hoepen.
1921. Gaudryceras tenuilineatum, v. Hoepen, loc. cit., p. 5, pl. u, figs.
7-9, text-fig. 2.
¢ 1921. Gaudryceras, sp. juv., Spath, loc. cit., p. 41.
The immature example described by the writer may belong to the
finely striated species established by Dr. van Hoepen, but it differs in
dimensions. The latter are nearer those of G. varicostatum, v. Hoepen,
but in this species and in G. conctum (Crick MS.) the striation is coarser.
4. Gaudryceras varicostatum, v. Hoepen.
1921. Gaudryceras varicostatum, v. Hoepen, loc. cit., p. 7, pl. ii, figs.
10-12, text-figs. 3-4.
1921. Gaudryceras kayei (Forbes), Spath, loc. cit., table to p. 50.
Dr. van Hoepen may be right in including in G. varicostatum the A. kayet
recorded by Griesbach and figured by Woods. The writer compared this
118 Transactions of the Royal Society of South Africa.
species to G. pulchrum, Crick, which has the dimensions of van Hoepen’s
species, but six labial ridges on the outer whorl.
5. Gaudryceras cinctum (Crick MS.), Spath.
(Plate LX, figs. 3a, 6.)
1921. Spath, Pondoland, loc. cit., pp. 41-2, and table to p. 50.
This species is represented by a single, well-preserved example (No.
C 19415), the dimensions of which are 64:5—-40—-37—-34. Mr. Crick’s
long description of this form is not given, since the species is extremely close
to G. varicostatum, v. Hoepen, and may only be the adult of the latter form.
G. varicostatum, however, at a smaller diameter, has H>Th, and has a
larger umbilicus. Crick also had compared his species to Griesbach’s
A. kayei, Forbes. The whorls of G. cinctum, however, are fewer, and the
striation is coarser than in the typical Pondicherry examples of G. kayea in
the British Museum.
The writer compared this species to G. mite (Hauer), which is very close,
but according to de Grossouvre * belongs to the Santonian.
G. varagurense, Kossmat, which had been thought by Crick to be very
close to G. cinctum, has much finer ornamentation, and a different rib-
curve. G. varagurense, var. patagonica, Paulcke,t is comparatively in-
volute.
6. Gaudryceras amapondense, v. Hoepen.
1920. v. Hoepen, loc. cit., p. 143, pl. xxiv, figs. 4 and 5.
1921. Spath, loc. cat., table to p. 50.
The holotype of this species, compared by the writer to G. colloti,
Grossouvre, and G. glaneygense (Redtenbacher), has its inner whorls too
badly preserved for exact determination, but is more involute than G.
cinctum. Its ornament is also far less regular.
T. Gaudryceras sigcau, v. Hoepen.
1921. Loc. cit., p 9, pl. ii, figs. 13-16, text-fig. 5.
This species is based on immature specimens, compressed already at a
very early stage, and thus different from any of the known Pondoland
forms of Gaudryceras.
* Craie Super., 1894, pp. 227-8.
+ “Ceph. d. Ob. Kreide 8. Patagon.,’’ Ber. Naturf. Ges. Freiburg i. B., xv, 1907.
p. 6, pl. xvii (viii), figs. 1-2.
On the Senonian Ammonite Fauna of Pondoland. ia)
Genus TETRAGONITES, Kossmat.
8. Tetragonites superstes, v. Hoepen.
(Plate VI, fig. 6.)
1921. Tetragonites superstes, v. Hoepen, loc. cit., p. 10, pl. u, figs. 17-20,
text-fig. 6.
1921. Tetragonites cf. epigonum (Kossmat), Crick, in Spath, loc. cit.,
p. 42.
The figured example, clearly belonging to v. Hoepen’s species, is the
specimen (No. C 19416) referred to by the writer, but the Tetragonites (¢) sp.
ind. described in the same place may be a young (and crushed) Pseudo-
phyllites.
9. Tetragonites nuperus, v. Hoepen.
1921. Tetragonites nuperus, v. Hoepen, loc. cit., p. 18, pl. i, figs. 3
and 4, text-fig. 8.
71921. Tetragonites aff. cala (Forbes), Spath, loc. cit., p. 43.
This species differs from Forbes’s type in having greater inclusion and
constrictions that describe a slightly different curve. The example
compared by the writer to the Valudayur specimens in the British Museum
may be still closer to Forbes’s species, but is more evolute than the example
figured by Woods.* The suture-lines of all these forms of Tetragonites are
very similar.
Genus PSEUDOPHYLLITES, Kossmat.
10. Pseudophyllites indra, Forbes sp.
1906. Pseudophyllites indra (Forbes), Woods, loc. cit., p. 334, pl. xh,
fig. 6.
? 1920. Tetragonites teres, van Hoepen, “‘ Descr. of some Cret. Amm.
from Pondoland,”’ Ann. Transvaal Mus., vii, pt. 1, p. 144, pl. xxv, figs.
land 2.
1921. Tetragonites (%) sp. ind., Spath, loc. cot., p. 42, pl. vu, fig. 3.
1921. Pseudophyllites indra (Forbes), 7b., table to p. 50.
¢1921. Tetragonites virgulatus, v. Hoepen, loc. cit., p. 11, pl. i, figs.
1 and 2, text-fig. 7.
This well-known and widely distributed species is represented in the
present collection by three specimens (No. C 19417-9), the largest of which
Crick described as being, ‘‘ as Kossmat has already stated, by far the largest
* “Cret. Fauna of Pondoland,” Ann. S. Afr. Mus., iv, No. 12, 1906, p. 335, pl. 41,
fig. 7.
120 Transactions of the Royal Society of South Africa.
example of the species yet known. It is 290 mm. in diameter (nearly 114
inches), and its greatest thickness 148 mm.; there are indications of the
former existence of at least another half whorl, so that the specimen when
complete must have attained a very large size. The specimen is entirely
septate, and the portion of the whor! that is missing was probably occupied
by the body-chamber.”
‘* Another example (No. C 19418) is only 70 mm. in diameter, but is
without doubt referable to the same species.”
‘“‘ A third example (No. C 19419), in a somewhat harder and darker sand-
stone, appears to be referable to this species. At first sight it seems to be
more compressed, and to have less rapidly increasing whorls and a steeper
umbilical wall than Pseud. ondra, but these differences may in part be due
to the imperfection of the anterior part of the outer whorl. Portions of
the mouth-border are preserved on each side, and these show that the
aperture was very oblique, the ventral (peripheral) portion being projected
considerably forward. The body-chamber is seen to occupy rather more
than one half of the outer whorl.”
Famity DESMOCERATIDAE, Zittel.
The genera Pachydiscus, Parapachydiscus, Parapuzosia, Kossmaticeras,
and the many allied developments discussed in the following pages can all
be derived from Desmoceratid stocks that persisted during the Upper
Cretaceous, and the unmodified types of which, owing to lack of exact
information, are at present included in the genus Puzosia, Bayle, though
the latter really should be restricted to the Upper Albian-Cenomanian
planulata-group. Some of these Puzosid developments will be referred to
below, under Puzosinae, a convenient but polyphyletic assemblage of
successive waves of modified Phylloceratids; but with regard to the
Pachydiscinae, a sub-family that includes a number of developments of
Puzosinae, the genus
PacHypiscus, Zittel,
is here taken to include only the group of P. peramplus, Mantell sp., as
suggested by Kilian and Reboul * after Pervinquiére and other writers.
Other species of this genus are: P. prosperianus, d’Orbigny sp., P. lewesiensis,
Mantell non Sharpe sp., P. catinus, Mantell sp., P. rhodanicus, Roman
and Mazeran, P. juvencus, Laube and Bruder sp., P. vaju, Stoliczka sp.,
and probably P. seppenradensis, Landois. A. undatus, J. de C. Sowerby,t
* “Les Céphalop. Néocrét. des es Seymour et Snow-Hill,” Wiss. Ergebn. Schwed.
Stdpol. Exp., iii, pt. vi, 1909, p. 41.
t+ Min. Conch., vi, 1827, p. 134, pl. dlxix, fig. 2, B.M., No. 43940a.
On the Senonian Ammomte Fauna of Pondoland. 121
which Sharpe * long since had considered to be “‘ better omitted from our
lists,” may be a crushed Pachydiscus of this group. ‘‘ Pachydiscus ”’
vectensis, Sharpe sp., like Sowerby’s A. cinctus, apparently is based on a
malformation. Whether the supposed Turonian Pachydiscus laevicaniculatus
(Roemer), Lasswitz,t belongs to this restricted Pachydiscus seems doubtful.
The simple suture-line, with its small external saddle, and the special-
isation in size mark this genus as a very distinct assemblage of closely
allied forms. Nowak derived it from UAligella, but strong costation appears
repeatedly in Desmoceratidae during the Cretaceous, not only in the
Aptian, to which formation the true Uhligella is restricted, though probably
the Pachydiscus ancestor is nearly related to such a Puzosid as that figured
by Pervinquiere as Pachydiscus sp.t and quoted by Nowak.§ This stock
continued into the Senonian, and gave rise, in addition to Kossmaticeras, to
at least two more developments, resembling the young Pachydiscus peram-
plus, namely, Nowakites, mentioned below, which attaches itself to such
forms as ° Puzosia’ le marchandi, Grossouvre,|| of the Lower Senonian,
and the later Canadoceras, which is a close ally of the Puzosid group here
separated as Kitchinites.
PSEUDOJACOBITES, gen. nov.,
is here proposed for those Turonian forms that, like P. farmeryi, Crick sp.
(genotype: “ Two Cephalopods from the Chalk of Lincolnshire,’’ Geol. Mag.,
N.S., decade v, vol. vii, 1910, p. 345, pl. xxvii, figs. 1, 2, B.M., No. C 12220),
develop multituberculation, and attach themselves to the true Pachy-
discus, possibly via such forms as ‘ Pachydiscus’ anapadensis, Kossmat sp.
The exact relations, however, of this latter form, as of ‘ P.’” rotalinus,
Stoliczka sp., which Kossmat thought might come from the same beds, and
which is classed as Cenomanian by Nowak, are not clear. Boule, Lemoine
and Thévenin,** and Kilian and Reboulft record it from the Upper Senonian,
and the last-named authors put it in the genus Jacobites, but this is a late
Campanian development of the sparsicosta-group of Kossmaticeras; and
the inner whorls of ‘ Pachydiscus’ rotalinus do not suggest affinity with
* “ The Fossil Moll. Chalk of England,” II, Cephalopoda, iii, 1857, p. 46.
+ “ Kreide Amm. v. Texas,” Geol. und Pal. Abh., N.F., vi, 1904, p. 16, pl. iii, fig. 2.
t “ Amm. du Crét. Algér.,’? Mém. Soc. Géol. France, Pal. xvii, No. 42, 1910, p. 37,
pl. iii, figs. 1-3.
Seo. Cit... Dp. 347.
| ““Les Ammon. d. 1. Craie Supér.,” Mém. Carte Géol. Détail. d. 1. France: II,
Paléont., 1894, p. 173, pl. xxii, fig. 5.
a woexcit.., 1913; p. 34:7.
ee “ Pal, de Madagascar: III, Céph. Crét. d. Envir. de Diego-Suarez,”’ Ann. de Pal., i,
fasc. 4 (1906), pl. vi, fig. 5, and pl. vii, fig. 1.
tT Loc. cit., 1909, pp. 37 and 52.
122 Transactions of the Royal Society of South Africa.
this stock. On the other hand, ‘ Pachydiscus’ rotalinoides, Yabe,* from
the Senonian Toyayo series, which may include any Senonian horizons
below the Parapachydiscus-beds of the Hokkaido, and also, apparently,
not a Jacobites, may possibly belong to the genus Menwites, described
below. |
PARAPACHYDIScuUS, Hyatt,
was proposed for A. gollevillensis, d’Orbigny, which is very close to Parap.
crishna and to Parap. egertoni, Forbes sp., the latter again a very near ally
of the Kuropean Parap. neubergicus, Hauer sp., Parap. perfidus, Grossouvre
sp., and Parap. jacquoti, Seunes sp. Kossmat’s Indian “‘ P. gollevillensis,” +
here renamed Parap. compressus, n.n., is different from the European type
as figured by d’Orbigny and Seunes.t A specimen in the British Museum
from the Baculite Limestone of Valognes, Manche (No. 50135), represents
a new form (Parap. valognensis, nov.), with a whorl-thickness of 30 per cent.,
more closely costate and flatter even than Grossouvre’s fig. 9 of plate xxxi.
Sharpe’s A. gollevillensis § represents a more coarsely ornamented type, and
the new name Parap. sharpei is here proposed for it. A series of Irish
specimens in the British Museum includes a further new form, Parap.
hibernicus, nov. (type No. C 15126), which differs from Parap. oldham,
Sharpe sp., in the coarseness of its ornament and from Parap. sharped in
having the whorl-section more rounded and the inner whorls continuously
costate. Other varieties of Parap. oldhami are more finely costate than
the type or Grossouvre’s form,|| e.g. specimen No. C 14830.
A. soma, Forbes, which is not identical with the more inflated A. ganesa,
Forbes, as Stoliczka and Kossmat had thought, belongs to this group of the
genus Parapachydiscus, also P. haldemensis, Schliter sp. and, probably, the
compressed forms figured by Nowak on plate xl, but the group is not equiva-
lent to Nowak’s egertonianus-series.
Apart from these compressed forms, which may tend to differentiate the
ornamentation on the ventro-lateral edges, Parapachydiscus also includes
many forms that develop inflated whorls and that are closely connected with
the previous group by such forms as Parapachydiscus ganesa (Forbes), P.
cricki (Kossmat), and P. complezus (Meek). The numerous species grouping
themselves round Parap. colligatus (Binkhorst) and P. deccanensis (Stoliczka)
belong to this division of Parapachydiscus, including the three forms
* “Notes on some Cret. Foss. from Anaga, ete.,” Science Rep., Tohoku Imper.
Univers., 2nd ser. (Geol.), iv, No. 1, 1915, p. 21, pl. ii, figs. 5 and 6.
t ‘* Unters. 8S. Ind. Kreideform.,” iii, 1898, p. 97 (162), pl. xxi (xv), fig. 1.
t “Contrib. Céph. Crét. Supér. France,’ Mém. Soc. Géol. France, Pal., No. 2, 1891,
pl. 5, fig. 1 only.
§ Loe. cit. (111, 1857), p. 48, pl. xxii, fig. 2.
|| Loc, cit., 1894, pl. xxii, fig. 1.
On the Senonman Ammomite Fauna of Pondoland. 123
described by the writer from Zululand, and the Parapachydiscus aff. ootaco-
densis (Stoliczka) described below. P. neubergicus, var. stallauensis,
Imkeller,* better renamed Parapachydiscus stallauensis, apparently belongs
to this second division rather than the first.
Within this comprehensive genus Parapachydiscus several lineages, no
doubt, could be separated, perhaps by means of the suture-line, but with
the data available at present such subdivisions appear to the writer to be
artificial, and he would class as Parapachydiscus, e.g., both Pachydiscus kolu-
turensis (Stoliczka), Yabe and Shimizu,f and Parapachydiscus fascicostatum,
Yabe sp.f
A bituberculate development, derived from the more globose forms of
the genus Parapachydiscus, leads to A. menu, Forbes, with scaphitoid
coiling ; this development is here separated as |
MENUITES, gen. nov.
(genotype: A. menu, Forbes, “ Report on the Fossil Invertebrata from
Southern India, collected by Mr. Kaye and Mr. Cunliffe,” Trans. Geol.
Soe. ser. ii, vol. vu, 1845, p. 111, pl. x, fig. 1, B.M., Geol. Soc. Coll., No.
10482). M. stwri, Redtenbacher sp., and M. portlocki, Sharpe sp., well
figured by Griepenkerl § and ama and M. selbiensis, Pervinquiere
sp., belong to this stock.
The original figure of Schliiter’s A. proteus suggests that A. auritocostatus,
Schliiter, and ‘ Pachydiscus ’ ambiguus, de Grossouvre, represent compressed
forms of this genus Menuwites.
The possibility of certain Scaphitids of the Campanian (Hoploscaphites,
Acanthoscaphites *) having been derived from the present group of Desmo-
ceratids has already been pointed out by the writer. Other late Senonian
Scaphitids, however, at present included in these genera, are derived from
Hoplitoplacenticeras (=group of * Hoplites’ vari).
The Campanian and Maestrichtian genus Parapachydiscus, as here
understood, is not a direct descendant of the Turonian Pachydiscus. It has
already been mentioned that in the Lower Senonian there is a group of
forms, namely,
* “Die Kreidebild. u. ihre Fauna a. Stallauer Eck u. Enzenauer Kopf bei Tolz,”’
Palaeontographica, 48 (1901), p. 57, pl. iii, fig. 5.
+ ““ Notes on some Cret. Amm. from Japan and California,’ Science Reports, Tohoku
Imper. Univ., 2nd ser. (Geol.), v, No. 3, 1921, p. 55, pl. viii, fig. 3
t “Notes on some Cret. Amm. from Japan and California,’’ Science Reports, Téhoku
Imper. Univ., 2nd ser. (Geol.), v, No. 3, 1921, p. 57, pl. viii, fig. 5, and pl. ix, figs. 2-5.
§ “‘ Die Verstein. d. Senon. Kreide v. KG6nigslutter, etc.,”’ Pal. Abh., iv, 1889, p. 99,
(401), pl. xii, fig. 2.
|| ““ Fauna d. Lineburger Kreide,” Abh. Preuss. Geol. Landesanst., N.F., Heft 37.
1902, p. 104, pl. vi, figs. 2-3.
124 Transactions of the Royal Society of South Africa.
NOWAKITES, gen. nov.
(genotype: NV. carezi, de Grossouvre sp., loc. cit., 1894, p. 190, pl. xxv, fig. 3),
attaching itself to such contemporaneous (Coniacian) forms as ‘ Puzosia’
le marchandi, Grossouvre. Nowakites draschei, Redtenbacher sp., N. yoko-
yamar, Jimbo sp., NV. canali, N. linderi, Grossouvre sp., and N. pazllettianus,
d’Orbigny sp., probably all belong to this genus, the last apparently entirely
different from the Texas A. flaccidicosta, Romer, with which Thomas and
Peron and Nowak (loc. cit., p. 354) had identified it.
Nowakites savini, Grossouvre sp., represents a lateral offshoot. The
forms also bear some resemblance to certain tuberculate Madrasites, Pseudo-
kossmaticeras, and Canadoceras, all of which, however, are of uppermost
Senonian age. On the other hand, the Lower Utatur
HOLCODISCOIDES, gen. nov.
(genotype: A. cliveanus, Stoliczka, ‘‘ Foss. Ceph. Cret. Rocks S. India,”
1865, p. 157, pl. Ixxvu, fig. 3), which also includes H. moraviatoorensis,
papillatus, and paravati, Stoliczka sp., and which has wrongly been in-
cluded in Kossmaticeras, is not directly related either to Nowakites or to
Madrasites.
For * Sonneratia’ janeti, Grossouvre, which forms a separate develop-
ment, allied to Nowakites,
PACHYDISCOIDES, gen. nov.
(loc. cit., 1894, p. 145, pl. xxii, fig. 4) is proposed. % Pachydiscoides
pervinquieret, n.n.=P.n. sp. ef. P. janeti, Pervinquiere non Grossouvre,*
probably belongs to the same stock.
From the Coniacian stock, here separated as Nowakites, forms with
increasingly strong ornamentation, like Redtenbacher’s A. zsculensis, result,
for which the genus
HUPACHYDISCUS, gen. nov.
(genotype: ‘ P.’ rsculensis (Redtenbacher), Grossouvre, pl. xxvi, fig. la, b
only), is proposed. Nowakites lindert (Grossouvre, pl. xxiv, fig. 4), one of
the largest species of that genus, already foreshadows the present develop-
ment ; on the other hand, Eup. levy: (Grossouvre) and Hup. grossouvrer
(Kossmat) are transitional to the later and much more abundant Parapachy-
discus, in which tuberculation seems to begin already in the smooth
initial stage. Hup. jeant (Grossouvre) is another species, and a Japanese
example, studied by the writer, apparently allied to ‘ Pachydiscus’ haradaa
and P. teshionensis, Jimbo, may belong to the same stock.
* « Bt. de Pal. Tunis., I, Céph. d. Ter. Second.” (1907), p. 174, pl. vii, fig. 11.
On the Senonian Ammomite Fauna of Pondoland. 125
The genus
CANADOCERAS, gen. nov.
(genotype: A. newberryanus, Meek [Trans. Albany Inst., iv, 1857, p. 47,
“‘ Deser. and Illustr. Foss. Vancouver’s and Sucia Islands, etc.’’; Bull. U.S.
Geol. and Geogr. Survey Terr., ii, 1876, p. 367, pl. iv, fig. 3], B.M. Geol. Soc.
Coll., specimen figured Plate VII, fig.5, Plate VIII, fig. 4), of the Upper Chico
Formation, is allied to the Puzosid development Kitchinites, referred to
below under Puzosinae, 2.e. to the group of “‘ Holcodiscus”’ pondicherryanus,
Kossmat, which persists in the Campanian, together with the true Kossmati-
ceras and its allies. A. newberryanus, Gabb non Meek * =“ Pachydiscus ”’
henleyensis, Anderson, is probably a Parapachydiscus of the colligatus-group,
though Anderson,{ perhaps wrongly, puts it as of Lower Chico age, and
confuses with “ Pachydiscus’”’ even Horsetown species of Pleuropachydiscus
(‘‘ Pachydiscus ”’ sacramenticus) and of Desmoceras s.s.= Latidorsella, Jacob
(“ P.? merriamt). <i" Bal
A. fraternus, Meek, and A. suciensis, Gabb non Meek,t belong to Canado-
ceras, aS does a closely allied species in the British Museum. The true
A. suciensis, Meek, or, at least, smooth Vancouver examples in the British
Museum, are more doubtfully included here; ‘‘ Pachydiscus’’ neevest,
Whiteaves, is probably a Grossouvrites.
Some of the more inflated forms, like C. multisulcatum, binodatum,
and perplicatum, Whiteaves sp., though having inner whorls of the new-
berryanus-type, resemble some forms of the other genera here separated ;
e.g. there may be a tendency to acquire the oblique costation of Koss-
maticeras, like K. pachystoma, Kossmat sp., or of Kitchinites japonicus,
n.n.=Desmoceras gaudama, Yokoyama non Forbes,§ testifying to the
Puzosid ancestry of the pachydiscoid ‘° Kossmaticeras’ here named
Canadoceras.
The smoother Canadoceras of the suciensis-group above mentioned
resemble a somewhat parallel development, provisionally left in the genus
Parapachydiscus, and including slowly or quickly coiled (globose) forms,
retaining the varices or constrictions of the Desmoceratid ancestors; e.g.
the Southern Patagonian P. hauthali, P. steenmanni, and P. patagonicus,
Paulcke sp., probably also the Japanese ‘ Pachydiscus’ naumanni, Yokoyama.
P. antecursor, v. Hoepen sp., and P. umtafunensis (Crick MS8.), Spath, con-
nect this group with the true Parapachydiscus, and after a short hauthali-
stage develop ornamentation foreshadowing that of the cricki-group of
* Pal. Calif., 1 (1864), p. 61, pl. xxvii, figs. 199, 1996, c; pl. xxviii, fig. 199a.
tT “Cret. Depos. Pacif. Coast,” Proc. Calif. Acad. Sc. (3) Geol., ii, (1902), p. 104.
(Wrongly quoted as A. suciaensis.)
i coe. cit an (1869), po 132, pl. xxi5.figs. 11, Ila; 6:
§ “ Verstein. a. d. Japan. Kreide,” Pal., xxxvi (1890), p. 184, pl. xix, fig. 5.
126 Transactions of the Royal Society of South Africa.
Parapachydiscus. P. amarus, Paulcke sp., is a transitional species from
Kitchinites or Kossmaticeras.
The young of ‘ Pachydiscus ’ conduciensis, Choffat,* somewhat resembles
P. umtafunensis, and Choffat’s gigantic species may have to be considered
as a special offshoot of this group of Parapachydiscus. The new genus
LYTODISCOIDES, gen. nov.
(genotype: Pachydiscus conduciensis, Choffat, loc. cit., p. 18, pls. u—v and
frontispiece), is proposed for this development. The ornamentation here,
though also increasing in strength, is of quite a special type.
As regards the sub-family Kossmaticeratine, the genus Madrasites, Kilian
and Reboul, is a descendant of the true Kossmaticeras (theobaldianus-group),
and Gunnarites and Grossouvrites are closely allied genera, but Seymourites
and Grahamites are still doubtful, and at any rate cannot be regarded as
subgenera of Kossmaticeras. A late Campanian type that, unlike Madrasites,
does not tend to involution, but leads, via forms of the group of A. brandti
and A. direri, Redtenbacher, to Brahmaites, requires a new name, and
PSEUDOKOSSMATICERAS, gen. nov.
(genotype: A. pacificus, Stoliczka, loc. cit., 1865, p. 160, pl. lxxvii, fig. 9),
is here proposed. Most of the European “ Kossmaticeras’”’ belong to this
stock.
SuB-FAMILY PUZOSINAE.
Nowak introduced his genus Parapuzosiafor Ammonites of the ‘denisoni’-
type, and he erroneously thought that they could be traced from the
Cenomanian to the uppermost Cretaceous. It might, of course, be held
that A. denisonianus, Stoliczka, is thus the genotype; but this species is
not cogeneric with Parapuzosia daubreer (Grossouvre), the only species
described and figured by Nowak, so that the writer selected this Santonian
form as the type of the genus Parapuzosia, which also should include
Parap. leptophylla and P. (%) wenica, Sharpe sp., P. corbarica, Grossouvre
sp., P. gaudama, Forbes sp., P. indopacifica, Kossmat sp. and P. stobaei,
Nilsson sp., the latter possibly transitional to Parapachydiscus.t The
unornamented A. griffith, Sharpe, may be close to the last species, as
de Grossouvre thought; for Sharpe’s drawing of its suture-line is
obviously quite sketchy.
* “ Contrib. Connaiss. Géol. Col. Portug. d’Afr.: 1, Le Crét. de Conducia,’? Comm.
Serv. Géol. Portug., 1903, p. 18, pls. ii-v and frontispiece.
+ This form was considered to be a Parapachydiscus in Cret. Ceph. fr. Zululand, 1921,
p. 225.
On the Senonian Ammomte Fauna of Pondoland. 127
Parapuzosia has the course of the radial line and the constrictions of
the strongly projected subplanulata-type different from the straight orna-
mentation of the Campanian
KITCHINITES, gen. nov.
(genotype: Holcodiscus pondicherryanus, Kossmat, loc. cit., 1897, p. 40
(147), pl. vi (xvii), fig. 6; British Museum, Geol. Soc. Coll.). This also
includes K. japonicus, n.n. (= Desmoceras gaudama, Yokoyama non Forbes),
and a South American form before the writer, near to K. darwin (Philippi
in Steinmann). The genus is very close to the stock from which also
originated Kossmaticeratinae (namely, Kossmaticeras, Madrasites, Gun-
narites, Jacobites, Pseudokossmaticeras, Brahmaites), but not the isolated
Holcodiscoides, which is a much earlier offshoot from Puzosinae. ‘ Puzosia’
ishakawai, Jimbo, is probably a young Kitchimites, hardly distinguishable
from the ancestral Parapuzosia.
That the genera Parapuzosia and Kitchinites, which continue the true
Puzosid type into the Senonian, are not derivatives of the Cenomanian
A. denisonianus, is clear; for the latter already develops a pachydiscoid
stage, and
PACHYDESMOCERAS, gen. nov.
(type: A. denisonianus (Stoliczka), Kossmat, loc. cit., 1898, p. 121 (186),
pl. xv, figs. 5a, 6), is here proposed for this form. It is probably closely
related to the contemporary Puzosia of the typical planulata-group rather
than to the Aptian Uhligella, and the Cenomanian Holcodiscordes is a close
ally. But in the Turonian, true Puzosids still persist and give rise to at
least two developments, one specialising in size (Austiniceras) and the
other in crenulation of the periphery (Tragodesmoceras). The genus
AUSTINICERAS, gen. nov.
(genotype: A. austen, Sharpe, loc. cit., 11, 1855, p. 28, pl. xii, figs. la and 0,
non 2, B.M., No. C 3382), has inner whorls of the type of A. dibley2, n.n.(=
A. austeni, Sharpe, p.p. pl. xii, fig. 2 only, holotype, B.M., No. C 13912,
Dibley Coll. from Holaster subglobosus zone), which is more involute than the
type species, and on the outer whorl develops costation of a different type,
foreshadowing that of the later Parapuzosia. The gigantic A. mobergi,
A, poctai, Grossouvre, and A. montis-albi, Laube and Bruder, belong to this
genus Austiniceras.
TRAGODESMOCERAS, gen. nov.
(genotype: Desmoceras clypealoides, Leonhard, ‘“‘ D. Fauna d. Kreidef. i.
Oberschles.,”’ Pal., vol. 44 (1897), p. 57, pl. vi, fig. 2), 1s proposed for a group
128 Transactions of the Royal Society of South Africa.
of desmoceratid forms sometimes included in Mumniericeras. This was
referred to by the writer on a previous occasion,* and forms the lineage
leading from the Turonian Tr. clypealoides (Leonhard), Tr. miilleri (Gross-
ouvre), and Z'r. hernensis (Schliiter) to the Lower Senonian Tr. clypeale
(Schliiter). This lineage is of interest on account of the probable derivation
therefrom of keeled forms.
Neither of the genera Austiniceras and Tragodesmoceras is the direct
ancestor of Parapuzosia; for in the Turonian forms occur (cf. Puzosia
curvatisulcata, Chatwin and Withers, and P. marlowense, Noble) which are
still of the regular Puzosia type. The latter was compared to Desmoceras
pyrenaicum, Grossouvre, which, however, is a Schliiteria; the former
(P. curvatisulcata), with a very complex suture-line (B.M., No. C 12229),
may be an Austiniceras, and resembles A.? gaudemarist, Roman and
Mazeran sp., but is too fragmentary for exact determination.
Genus Parapuzosia, Nowak.
11. Parapuzosia haughtont, sp. nov.
(Plate VIII, fig. 1.)
21921. Parapuzosia, sp. nov.? ind., Spath, loc. cit. (Zululand), p. 224,
plimix, fie 2. plvnex, fos. la. plascmne Ome.
The large fragment (No. C 19439) on which this description is based,
labelled ‘‘ Pachydiscus sp.”’ by Crick, was referred to fT as probably a Para-
pachydiscus of the colligatus-supremus type on account of its resemblance
in smoothness and section of the outer whorl and suture-line to those of
the example here described as Parapachydiscus aff. ootacodensis (Stoliczka).
On preparation of the umbilical and dorsal portions of the fragment, how-
ever, the ornamentation of the inner whorl was revealed, showing the primary
and secondary ribs characteristic of Parapuzosia daubreet.
At a whorl-height of 260 mm., the thickness was probably not more than
180 mm. (not 200 mm., as stated in the footnote on p. 229, loc. cit.). The
whorl-section thus agrees with that given by the writer for the Zululand
form (loc. cit., pl. xx, fig. la), and is not so compressed as that of P. daubreer
as figured by Nowak. { The umbilical wall is higher, and the whorl.is
thicker at the umbilical border in the South African form; on the other hand,
there is no definite umbilical edge, as noted of P. daubreer by Grossouvre,
but the sides are gradually rounded off into the umbilicus, as illustrated by
* Loc. cit. (Zululand, 1921), p. 237.
t+ Loc. cit. (Zululand, 1921), p. 229.
t * Untersuch. i. d. Ceph. d. Ob. Kreide i. Polen.”’: III, Bull. Acad. Sc. Cracovie,
Classe Sc. Math. and Nat., ser. B, June 1913, fig. 32, pl. xliii.
On the Senonan Ammomte Fauna of Pondoland. 129
Nowak and by the writer for the Zululand example. Since the description
of the latter was based only on the photographs and plaster cast of the
dorsal area, it is uncertain, of course, whether it belongs to the present
species. A comparison of the casts of the dorsal areas of the two forms,
however, shows that at any rate the inner whorls, with the typical strongly
projected Parapuzosia ribbing, are closely similar.
Genus ScHLUTERIA, Grossouvre em.
12. Schliiterra sumplex, v. Hoepen sp.
1921. Desmoceras simplex, v. Hoepen, loc. cit., p. 19, pl. 11, figs. 11-16,
text-fig. 10.
1921. Schltiterta woodsi, Spath, Pondoland, p. 45, pl. vu, fig. 1.
The writer does not lay stress on minute differences in the convexity of
the sides, proved to be due to sexual dimorphism in the recent Nautilus.
The dimensions of the two forms, quoted in the synonymy, agree, but
whereas Dr. van Hoepen had comparatively small examples, the specimen
figured by the writer measured 85 mm. in diameter.
13. Schliterva crassa, v. Hoepen sp.
1906. Phylloceras sp., Woods, loc. cit., p. 332, pl. xh, fig. 5.
1921. Desmoceras crassum, v. Hoepen, loc. cit., p. 20, pl. iv, figs. 3-4,
text-fig. 11.
This form shows more inflation than the last species. It is interesting
to note the complexity of its suture-line at a small diameter as compared
with that of the Santonian form figured by Grossouvre.*
Genus HAvERICERAS, Grossouvre.
14. Hauericeras garden, Baily sp.
1921. Spath, Zululand, loc. cit., p. 238, text-fig. A.
1921. v. Hoepen, loc. cit., p. 27, text-fig. 15.
1921. Spath, Pondoland, loc. cit., p. 50 (table).
To the writer’s notes on this form in the above-mentioned paper it is
only necessary to add that the late G. C. Crick took measurements of 20
out of the 37 examples in the present collection, and the mean of these
measurements Is :
whorl-height |=35 per cent. of the diameter.
whorl-thickness =19 se 63 as
umbilicus =39 a, s3 0
* Loc. cit., 1894, pl. xxxvii, fig. 9 (as Desmoceras pyrenaicum).
il
130 Transactions of the Royal Society of South Africa.
The percentages given by v. Hoepen are -34—-17—-40 for an example of
120 mm. diameter. The specimens range from 60 mm. (B.M., No. C 18521)
to 145 mm. (C 18516) in diameter, and the relatively thickest form (C 18518)
has a narrow umbilicus (dimensions : 130 —-34—-215 —-40), whereas the
thinnest form (C 18517) has a wide umbilicus (dimensions: 138 —-30—-16
---46). This, as Mr. Crick observes, is contrary to the general rule
given by Grossouvre,* namely, that in the same specific type individuals
with a narrow umbilicus show a corresponding compression of the
whorls.
The transverse section of an example 106-5 mm. in diameter (B.M., No.
C 18520), very close to the type-specimen, allowed of measurements being
taken at different stages, and Crick’s figures are here appended :
Diameter of |106-5 (1 )j78-0 (1 55-0 (1 )}38-0 (1 )|26-0 (1 yj18-5 (1 )J12-75 (1 )
shell
Height = of [37-5 (0-352)|29-5 (0-378)|20-75 (0-377)|14-75 (0-388)|10-0 (0-384)| 6-5 (0-351)| 4:5 (0-352)
outer whorl
Thickness of |21-5 (0-201)/16-5 (0-211)|11-5 (0-20 )| 8-25 (0-218)| 5-75 (0:221)| 3-5 (0-189)| 3-5 (0-274)
outer whorl
Width of |41-5 (0-389)|28-25 (0-362)|19-5 (0-354))/13-5 (0-355)|10-0 (0-384) 6-75 (0-364)| 4:5 (0-352)
umbilicus
The same transverse section shows that at a diameter of 13 mm. there is
no keel, whereas at 18 mm. it first begins to be indicated very feebly. Up
to a diameter of 50 mm., the constrictions, also, are biconvex, like those of
H. fayoli, Grossouvre, or of Forbes’s small types of H. rembda (=A. durga).
The course of the constrictions, if it remains biconvex in larger examples of
H. rembda, as drawn by Stoliczka,} may serve as a distinction between the
two species, as, é.g., Pervinquiere { had thought ; for the larger whorls of
H. garden have semicircular constrictions, sloping forwards towards the
umbilical suture as much as the outer half is projected forwards towards
the keel. In several examples the apex of this semicircular curve is
slightly flattened ; in others there may be a small indentation in the semi-
circle, either near the umbilical edge (No. C 18540) or nearer the periphery
(C 18537), suggestive of the original biconvexity, and in. the transitional
stage the course of the constrictions would be as described for H. sulcatus
by Kner and Nowak.§
The last example has the periphery of the cast (but not that of the test)
still rounded at a diameter of 110 mm., as in Baily’s larger specimen (No.
113871, Geol. Soc. Coll.). Another example (C 18522) has lateral grooves
* Loc. cit. (1894), p. 8.
t Loc. cit. (1864), pl. xxxiui, fig. 5.
{ Loc. cit. (1907), p. 169.
§ Kner, “ Verstein. d. Kreidemerg. v. Lemberg.,”’ Naturw. Abh., iii (1850), 2, p. 8,
pl. i, fig. 3: and Nowak, loc. cit., p. 371, pl. xli, fig. 12, pl. xlv, figs. 44-45.
On the Senonian Ammomte Fauna of Pondoland. 131
accompanying the keel (of test only) at 90 mm. diameter, and specimen
No. C 18535 has a very strong keel, just beginning to become undercut, as in
H. rembda, at about 110 mm., set on the far less acute periphery of the cast.
Forbes’s type of H. rembda, on the other hand, shows the acute periphery
of the cast, and the peculiar polygonal keel already at 30 mm. diameter.
This keel is a more distinct characteristic of H. rembda than the biconvex
constrictions.
The suture-line of H. gardeni (Baily’s paratype 11371) differs from that
of H. sulcatum, as figured by Nowak, in the very regular decrease of
the three auxiliary saddles down to the umbilical lobe, a character shown
also in some Japanese examples before the writer (Geol. Soc. Coll.). In
the latter, however, the periphery is more compressed than in H. gardeni
typus, or in H. fayoli, which latter species they resemble in their con-
strictions.
H. garden, in the present collection, is associated, in the same hand-
specimens, with Mortoniceras, Pseudoschloenbachia, and Spheniscoceras.
15. Hauericeras rembda (Forbes).
1921. v. Hoepen, loc. cit., p. 28.
1921. Spath, loc. cit., table to p. 50.
This species is not represented in the present collection or apparently in
Dr. van Hoepen’s material, but was recorded by Griesbach and Woods.*
16. Hauericeras ? sugata (Forbes).
1921. Desmoceras compactum, v. Hoepen, loc. cit., p. 21, pl. iv, figs. 5-7,
text-fig. 12.
1921. Hauericeras * sugata (Forbes), Spath, Pondoland, p. 46, pl. vi,
figs. 3a, b.
Dr. van Hoepen’s single immature example hardly justifies the creation
of a new species, though it is possible that the Pondoland form may not be
of the same date as Forbes’s type. The writer assumed (loc. cit., p. 54) that
Desmoceratids of the group of A. sugata have a fairly wide vertical range.
Probably derivatives of several desmoceratid stocks, in the present case
Schliiterva, successively became carinate and consequently have all been
included in this well-known species. There is, however, no satisfactory
character to warrant the separation of the immature Pondoland examples
from Forbes’s species.
* Loc. cit. (1906), p. 333.
132 Transactions of the Royal Society of South Africa.
SuB-FAMILY PACHYDISCINAE.
Genus ParapacHypiscus, Hyatt.
17. Parapachydiscus aff. ootacodensis, Stoliczka sp.
(Plate VII, fig. 6.)
1921. Parapachydiscus sp. ind., Spath, loc. cit. (Pondoland), table to
p. 50.
In the description of Parapachydiscus sp. n. aff. colligatus, Binkhorst sp.,
from Zululand, the writer referred to two large specimens in the present
collection, not described by Crick, that “‘ may belong to Parapachydiscus of
the colligatus-supremus type.’ One was described as having “‘ an umbilicus
of 19 per cent. and a thickness of 45 per cent. of the diameter,” the other as
showing “at a whorl-height of 260 mm. a thickness of 200 mm.” Both
were stated to be more compressed than the Zululand example described,
and to be intermediate in sectional outline between fig. 30 (P. collegatus)
and fig. 31 (P. oldham) in Nowak.*
The larger fragment (C 19439) is described above as Parapuzosia haugh-
toni, sp. n., since development of its dorsal area has shown that the resem-
blance of its smooth outer whorl to the Parapachydiscus here described was
only superficial. The smaller example (No. C 19438), also labelled ‘* Pachy-
discus sp.” by Crick and not described, has the following dimensions :
at diameter =150 mm., -50, -45, -19.
At the diameter of 59 mm. the whorl-height = 29 mm., and the thickness
= 31 mm.; on the other hand, where the height of the outer whorl = 150
mm., the thickness is only 125 mm., so that whereas, at first, the whorl-
section is round or slightly depressed, with increase of size it becomes com-
pressed. The specimen is still septate at this stage.
P. ootacodensis shows a similar change of proportions, with age, 7.e.
from the more inflated inner whorls (Stoliczka’s fig. 4 of pl. liv) to the more
compressed adult figured by Stoliczka on pl. lvi. The ribbing also is
similar, though the inner whorls seem more distinctly costate in the Pondo-
land form, almost as strongly ornamented as those of the much more inflated
P. sp. n. aff. colligatus, figured by the writer. The present form is thus
transitional to the more coarsely ornamented P. grossouvrer (Kossmat) =
A. ootacodensis, Stoliczka, pl. lvu, non cet., but none of the ribs reaches
to the umbilical border. The umbilical edge is perhaps not quite so marked
as it is in Kossmat’s smooth Indian example.t
The suture-line differs from that of P. ootacodensis in having the four
* Loc. cit. (1913), pl. xliii. + Loc. cit. (1898), fig. 16, pl. xvi (xxii).
On the Senonian Ammonite Fauna of Pondoland. 133
bifid, outer saddles gradually and regularly decreasing to the umbilical
border. The four or five inclined simple saddles of the umbilical wall
also slope down to the umbilical suture more regularly even than those of
P. tweenranus (Stoliczka).*
18. Parapachydiscus simplex, v. Hoepen sp.
1921. Pachydiscus simplex, v. Hoepen, loc. cit., p. 25, pl. v, figs. 3 and 4,
text-fig. 14.
This species, belonging to the ganesa-cricki group of Parapachydiscus
(see ante, p. 132), is not represented in the present collection.
19. Parapachydiscus umtafunensis (Crick MS.), Spath.
(Plate IX, figs. 4a, b.)
1921. Spath, loc. cit., Zululand, p. 229, footnote 5.
So? » Pondoland, p. 49.
This species, represented by the two examples (Nos. C 19434—-5) of
dimensions 75 —-35 —:36 —-40 and 72-5 —-34 —-35 —-39, already referred to
by the writer, at first sight seems identical with P. antecursor, v. Hoepen
sp., but has Th>H, whereas v. Hoepen’s single example is described as
having the whorl-thickness equal to or less than the height. The real
difference, however, appears to be in the suture-line, that of the present
species showing very good agreement with the suture-lines of the South
American forms of the group of P. hauthalr, Paulcke sp., and especially
with that of P. amarus, Paulcke sp.t
P. tweenianus (Stoliczka), to which the species had been compared by
Crick, has a different type of ornamentation.
20. Parapachydiscus antecursor, v. Hoepen sp.
1921. Pachydiscus antecursor, v. Hoepen, loc. cit., p. 24, pl. v, figs.
1 and 2, text-fig. 13.
This species was compared by its author to a number of pachydiscoid
developments, of which only P. tweenianus (Stoliczka) is distantly related.
In this species, however, the ribbing increases in strength, relatively, on the
outer whorls, and the inner whorls are smooth.
* Loc. cit. (1865), pl. lv, fig. 1b.
t “Ceph. d. Ob. Kreide Siidpatagon.,’ Ber. Nat. Ges. Freiburg., xv, 1907, p. 62,
text-fig. 29.
134 Transactions of the Royal Society of South Africa.
21. Parapachydiscus ? sp. nov.?
1921. Spath, Pondoland, p. 48, pl. vii, figs. 5a, 6.
This form, possibly a young Kztchinites, is not an immature example of
one of the species of Parapachydiscus described above, but is too small for
accurate identification.
SuB-FAMILY KOSSMATICERATINAE.
Genus MaprasITEs, Kilian and Reboul.
22. Madrasites natalensis (Crick MS8.), Spath.
(Plate V, fig. 3.)
1921. ‘ Cret. Ceph. fr. Zululand,’ Ann. §. Afr. Mus., xii, pt. vii, No. 16,
p. 300.
1921. “ Up. Cret. Amm. fr. Pondoland,’ Ann. Durban Mus., iii, pt. 2,
table to p. 50.
This species is based on a well-preserved specimen (No. C 19432) of dimen-
sions 81—-34—-40—-30. It shows the aperture well, like the form: of
Madrasites described below, and was characterised, in the writer’s previous
papers, as being closely allied to M. faku, van Hoepen sp. The latter has
a smaller umbilicus and coarser costation therein, and the suture-line is
drawn with comparatively simple elements, whereas in the present species
it is as ramified as that of Grossouvrites gemmatus (Huppé in Steinmann).
Mr. Crick’s long original description is not now given, since fig. 3 of Plate V
will show how close the present species is to M. faku, v. Hoepen sp.
23. Madrasites acuticostatus (Crick MS.), Spath.
(Plate VIII, fig. 2.)
1921. Spath, Zululand, p. 300.
1921. ,, Pondoland, table to p. 50.
This species is represented by a single well-preserved example (No.
C 19433) of dimensions 100—-37-—-36—-28, and was described at length
by Mr. Crick as Holcodiscus acuticostatus. It may suffice to quote the
following sentence :—
‘“ Compared with the previous species (H. natalensis), the present species
has a relatively coarser sculpture on the inner, and a finer sculpture on the
outer, whorls; itis also a more compressed shell, with a wider umbilicus, and
a larger number of tubercles on the umbilical margin.”
On the Senonian Ammonite Fauna of Pondoland. 135
24. Madrasites faku, v. Hoepen sp.
1920. Holcodiscus faku, v. Hoepen, loc. cit., p. 144, pl. xxv, figs. 3-4,
pl. xxvi, figs. 1-2.
1921. Madrasites faku (v. Hoepen), Spath, loc. cot., p. 47, and table
to p. 50.
This species is close to the last two, as pointed out by the writer on a
previous occasion.
25. Madrasites africanus, v. Hoepen sp.
1920. Holcodiscus africanus, v. Hoepen, loc. cit., p. 146, pl. xxvi, figs. 3-5.
1921. ” ae os loc. ctt., p. 23.
1921. Madrasites africanus (v. Hoepen), Spath, loc. cit., p. 48, and table
to p. 50.
This was compared by the writer to M. buddhaicus (Kossmat), from
which it differs merely in proportions.
26. Madrasites aff. africanus, v. Hoepen sp.
1921. Holcodiscus africanus ?, v. Hoepen, loc. cit., p. 23.
A specimen of Madrasites was described by Dr. vau Hoepen as being
relatively thicker than M. africanus and showing other differences.
27. Madrasites sp. ind.
1906. Holcodiscus sp., Woods, loc. cit., p. 336, pl. xl, fig. 2.
1921. Madrasites sp. ind. (Woods), Spath, Pondoland, p. 48, and table
to p. 50.
This form has a smaller umbilicus and finer ornament than M. africanus.
It was compared to Jacobites anderssont by Kilian and Reboul,* and shows,
indeed, close resemblance in the inner whorls to the form -figured by
these authors on pl. vil, fig. 2, if one may judge by Professor Kilian’s casts
of his Antarctic specimens in the British Museum. Nothing like the adult
Jacobites, however, seems to have been found in Pondoland, so that Mr.
Woods’s form is here considered to be nearer to the typical Madrasites
madrasinus (Stol.).
28. Madrasites similis, Spath.
1921. Spath, Pondoland, p. 48, pl. vi, fig. 1.
This form was described as standing in the same relationship to (the
tuberculate) M. faku as Kossmaticeras sparsicostatum (Kossmat) does to
(the costate) K. theobaldianum (Stol.).
* “ Céph. Néocrét., etc.,” Wiss. Ergeb. Schwed. S. Pol. Exp., iii, 6, 1909, pp. 62-3.
136 Transactions of the Royal Society of South Africa.
INCERTAE SEDIS.
Genus Hoploscaphites, Nowak.
At least four species of this genus have been described, but the examples
are mostly too fragmentary or too immature to allow of definite specific
identifications.
29. Hoploscaphites sp. (cf. semilaris, Stoliczka °).
1906. Scaphites sp., Woods, loc. cit., p. 348, pl. xliv, figs. 8a-c.
30. Hoploscaphites sp. (cf. pavana, Forbes ?).
1921. Scaphites cunliffer. v. Hoepen (non Forbes), loc. cit., p. 28, pl. v,
figs. 5-7, text-fig. 16.
21921. Hoploscaphites sp. juv., Spath, Pondoland, p. 49.
31. Hoploscaphites sp. nov. ¢
1921. Scaphites sp., v. Hoepen, loc. cit., p. 29, pl. v, figs. 8, 9.
32. Hoploscaphites sp. ind.
1921. Scaphites sp., v. Hoepen, loc. cit., p. 30.
Famity PRIONOTROPIDAE.
Genus MortonicEras, Meek.
33. Mortomceras soutom, Baily sp.
(Plate VII, fig. 4.)
1921. Spath, loc. cot. (Zululand), p. 234, pl. xx, fig. 4.
1921. v. Hoepen, loc. cit., p. 38, pls. x, xi, text-figs. 19-22.
1921. Spath, loc. cat. (Pondoland), table to p. 50.
Of the twelve specimens referable to the group of M. souton, five are
too fragmentary to be definitely identified with any of the types mentioned
below, but the remaining examples show great variability, as do the specimens
described by Dr. van Hoepen. The specimen (No. C 19441) referred to as
close to Baily’s type, the suture-line of which was figured by the writer,*
seems to be the most typical, though it has a considerable number of bifurcat-
ing ribs. A fragment (C 19469) is probably also close to the type, whereas
* Loc. cit. (Zululand, 1921), pl. xx, fig. 4.
On the Senonian Ammonite Fauna of Pondoland. 137
specimen C 19461 is nearer to the (at a corresponding size) slightly less
closely costate and more evolute example figured by Woods. In this last
specimen (C 19461) the costation is regular, single, and relatively closer on
the inner whorls; in the first (C 19441) the beginning is more distantly
costate, and closeness of the ribbing and bifurcation set in only at a later
stage.
A still more closely costate type than C 19461 is C 19442, but here the
sharp umbilical edge of the holotype is lost on the outer whorls. The suture-
line of this example is figured on Plate VII, fig. 4. C 19454 has a more
distantly costate beginning even than C 19441, but otherwise is a character-
istic M. soutona. Specimen C 19451, on the other hand, in which the distant
costation of the inner whorls persists to a comparatively late stage, and
which also has a more inflated outer whorl, is transitional to M. aff. umkwel-
anense, Crick, described by the writer.* Still another and more evolute
form (C 19460) has the costation of the outer whorl more distinctly reclined
than the holotype or other examples. Finally, No. C 19447, similar in the
inner whorls to the example figured by Woods, has costation simple on the
outer whorl and more distant than the ribbing of any other specimen.
The Zululand form described by the writer as M. aff. soutoni is not
represented in the present collection.
34. Mortonceras stangeri, Baily sp.
(Plate IX, fig. 2.)
1921. Spath, loc. cit. (Zululand), p. 297.
1921. a », (Pondoland), table to p. 50.
Thirty-one examples, including a few impressions and _plaster-casts
thereof, are referable to this species and its varieties, the great majority
of the examples being close to the specimen figured by Woods and to Baily’s
holotype (Geol. Soc. Coll., 11366) and two paratypes (11367 and 68). They
include specimens C 19440 and C 19443, the suture-lines of which were
figured in text-figs. lc, 1b of the writer’s Zululand paper (p. 297), and
C 19459, of which a sectional outline of the inner whorls and the internal
suture-line were given (loc. cit., pl. xxiii, figs. 3a, b).
The dimensions of Baily’s types are :-—
Holotype (11366). Paratype (11367). Paratype (11368).
Diameter . ; . o20 mm. 101 mm. tS mmr
Height of outer whorl . 26 percent. 31 per cent. 27 per cent.
Thickness of outer whorl 19 _,, Si iee es Dae
Umbilicus . : eG. «oe 47, Se. tes
* Loc. cit. (Zululand, 1921), p. 234, text-fig. D 2, p. 297.
138 Transactions of the Royal Society of South Africa.
The outer whorl flattens out in all the large examples, but the two quadrate-
whorled paratypes do not quite agree in ornamentation either with the
holotype or with each other. No. 11367 is transitional to the var. sparsicosta,
described below, and has a very indistinct median lateral tubercle ; in 11368
this tubercle is as pronounced as in the example figured by Woods.
35. Mortoniceras stangeri (Baily), var. sparsicosta, nov.
(Plate V, fig. 1.)
In some of the examples of M. stangeri, described above, the inner
whorls are less closely costate than in the type. These are transitional
to the present variety, which is characterised by the comparatively coarse
and distant tuberculation of the inner whorls, and the depression until a fairly
late stage of the whorl-section, Baily’s paratype 113684, the internal
suture-line of which was figured on pl. xxiii, fig. 3c, of the writer’s Zululand
paper, also specimen No. C 19444 (text-fig. la, p. 297, loc. cit.), belong to
this variety. The lateral view of the latter specimen is represented in fig. 1
of Plate V, and has dimensions : 295—-27—-25—-53.
Crick included specimen C 19444 in what he listed as an ‘‘ intermediate
group between WM. stangert and M. soutoni,” but none of the examples in his
list has any resemblance to M. soutont. Moreover, there is no description of
any of these Mortoniceras, and even the most typical example of M. soutona
was erroneously listed as M. stangert.
36. Mortoniceras stangeri (Baily), var. densicosta, nov.
(Plate V, fig. 2.)
This is a more finely costate variety, typically represented by specimen
C 19456, here figured, of the following dimensions: 125—-28—-25—-52. The
inner whorls of this specimen also seem more compressed, though slightly
crushed. Some transitional examples (C 19450, C 19450a, C 19462)
connect this variety with the typical M. stangeri, just as this again is
connected by a series of gradations with the var. sparsicosta. The outer
whorls of the three forms here described are very similar, and tend to
weaken the tuberculation of the costae.
The occurrence of distantly and closely ribbed varieties is of interest in
connection with a recent paper by Professor Salfeld.* In the varieties of
M. stangeri, however, there is no correlated change in whorl-section, and
the var. densicosta is not so common as the other forms, being typically
* “Bemerkungen zu v. Bubnoff, etc,’ Zeitschr. f. indukt. Abstamm. und Vererb.
Lehre, 1921.
On the Senonian Ammonite Fauna of Pondoland. 139
represented by only one example. The writer is of opinion that the common
occurrence of these types of variation in Ammonites (not mutations, since,
apparently, they are contemporaneous) may be taken to indicate a form of
“dimorphism,” or rather of “ polymorphism.” The “ polymorphism,”
however, necessitates agreement in the other characters, which would have
to be ascertained to exist in a larger number of specimens than are available
in the present instance. Moreover, it is difficult to stipulate what would
be agreement in, e.g., the suture-lines ; and since the change of a species-
group into another and later one occurs by means of adaptive radiation,
graphically represented by fans or a multitude of more or less radiating
lines rather than by a single line in the customary genealogical tree,
opinions would differ greatly as to the “‘agreement’”’ in a given species-group.
The fact that the adult whorls of the various forms here discussed become
similar, but not “ catagenetic,’ shows that this “‘ polymorphism” is a
phenomenon quite different from the simplification of the ornament on the
outer whorls of many Ammonites, which may tend to perfection of adapta-
tion in, e.g., Oxynoticeras, and which often, like the approximation of the last
few septa, marks only the slackening of growth on reaching maturity, but
which, also, may be simply correlated with the overlap of the mantle in
the mature stage.
Genus PSEUDOSCHLOENBACHIA, Spath.
37. Pseudoschloenbachia umbulazt, Baily sp.
(Plate VI, fig. 5.)
1921. Pseudoschloenbachia umbulazi (Baily), Spath, Zululand, p. 240,
pl. xx, fig. 2, text-fig. B, p. 241.
1921. Schloenbachia umbulazi (Baily), v. Hoepen, loc. cit., p. 35, pl. viii,
figs. 6-9, pl. ix, figs. 1 and 2.
1°21. Pseudoschloenbachia umbulazi (Baily), Spath, Pondoland, table
to p. 50.
The measurements of typical examples agree with those given by
Dr. van Hoepen, e.g. No. C 19427 has dimensions 50—-52—-28—-14._ The
inner whorls of this example were figured by the writer (loc. cit., text-fig.
Bl, p. 241). They show that at a diameter of 832 mm. the thickness = 30
per cent., and the umbilicus = 18 per cent., so that the whorls become
relatively thinner with age, whereas the umbilicus narrows. At a diameter
of 21 mm. the thickness has increased to 33 per cent. In the var. acuta, at
the same diameter, the whorl-thickness is only 26 per cent.
The writer has pointed out that these Pondoland examples represent
more strongly ornamented varieties, with increasing tuberculation round
140 Transactions of the Royal Society of South Africa.
the umbilicus. This is partly due to the larger size of the Pondoland
examples ; for at a size corresponding with that of the geno-holotype,
figured on pl. xx, fig. 2, of the Zululand paper, the umbilical tuberculation
of the present specimens is pushed back to only a very slightly earlier stage.
The term variety, thus, is hardly applicable to these forms. In example
No. C 19426, however, here figured (Plate VI, fig. 5), of dimensions 46—
‘52—-29-—-13, the umbilical tubercle is decidedly more distinct than in
the neotype, which seems to agree in this respect with Baily’s lost holo-
type. Dr. van Hoepen’s figs. 1 and 2 of pl. ix represent a similar form.
This was again selected as neotype by Dr. van Hoepen, but the writer had
fixed the geno-neotype just previously. This form is transitional from
the true P. umbulazi to the species described below as P. pseudofourniert,
and thence to P. papillata.
The new Egyptian species of Pseudoschloenbachia, mentioned by the
writer on previous occasions and figured on Plate V, fig. 4, as P. humez,
nov.,* is distinguished from P. umbulazi by more distinct costation and
tuberculation, especially at the ventro-lateral edges. This Hgyptian
specimen is preserved as a chalcedonic cast, resembling the mode of
preservation of a cast in flint of a new Sphenodiscus, allied to S. acuto-
dorsatus, Noetling, from beds of the same age near Jerusalem
(B.M. C 22136).
Specimens No. C 19436-7 are associated with Hawericeras gardenia and
Mortoniceras stangeri, in addition to other mollusca, in the same blocks.
38. Pseudoschloenbachia pseudofourmerr, nov.
(Plate VI, fig. 3.)
This species is based on example C 19425 of dimensions 57—-48
—-27—-19, referred to previously (Zululand, p. 242) as being slightly
constricted and transitional in ornamentation, from P. umbulazi to P. papil-
lata. It differs from the former in the largeness of its umbilicus and the
presence of the constrictions, in addition to the greater strength of its orna-
mentation, with fewer, but prominent, umbilical tubercles. P. papillata
represents a still more coarsely ornamented type, with a yet wider umbilicus.
Ornamentation and constrictions of the present species recall Grossouvre’s
‘ Schloenbachia’ fourniert,t probably belonging to the genus Gauthiert-
ceras; but the acute periphery of Pseudoschloenbachia and the numerous
* Blanckenhorn’s label, in German, attached to this specimen, reads as follows:
“ Schloenbachia n. sp. ind. aff. varians, yet not two, but three or four auxiliary lobes,
i.e. three auxiliary lateral lobes and one umbilical lobe. The first lateral lobe unusually
broad-stemmed, deeply bipartite. Cenoman ?”’
f Loc. cit. (1894), p. 112, pl. xxxv, fig. 1.
On the Senonian Ammonte Fauna of Pondoland. 141
elements of its suture-line are important distinctions from the earlier
stock.
P. humei, nov. (Plate V, fig. 4), has more numerous but less prominent
inner tubercles, and more distinct and fewer outer nodes.
39. Pseudoschloenbachia papillata (Crick MS.), Spath.
(Plate IX, figs. la, 6.)
1921. Pseudoschloenbachia papillata (Crick MS.), Spath, Zululand,
pp. 240 and 242.
1921. Schloenbachia umbulazi, var. spinifera, v. Hoepen, loc. cit., p. 37,
pl. ix, figs. 3-7.
1921. Pseudoschloenbachia papillata (Crick MS8.), Spath, Pondoland,
p. 43, pl. vi, figs. 2a, b.
This form is sufficiently distinct from P. umbulaz to be accorded specific
rank.
40. Pseudoschloenbachia griesbachi (Crick MS.), Spath.
(Plate VII, fig. 1; Plate VI, fig. 4.)
1921. Pseudoschloenbachia griesbacht (Crick MS.), Spath, Zululand,
p. 240, text-fig. B8 on p. 241, p. 312. |
1921. Schloenbachia umbulaz, var. griesbachi, v. Hoepen, loc. cit., p. 35,
pl. vu, figs. 3 and 4, pl. viii, figs. 1-5, text-figs. 17-18.
1921. Pseudoschloenbachia griesbachi (Crick MS.), Spath, Pondoland,
table to p. 50.
Crick’s long description of this species is omitted. The dimensions of
the holotype are: 103—-51—-25—-14; those of the paratype: 33—-52
—-26—-13. The suture-line of the holotype was figured by the writer
on a previous occasion.
INCERTAE SEDIS.
41. Gen. nov. (Muniericeras ¢) crickr, Spath.
1921. Loc. cit., Pondoland, p. 44, pl. vu, figs. 4a, b.
It is interesting to note that Dr. van Hoepen remarked the resemblance
of young examples of Pseudoschloenbachia papillata (his Schloenbachia um-
bulazi, var. spinifera) to Grossouvre’s Muniericeras lapparenti. The tuber-
culate periphery of the present form, however, separates it from the genus
Pseudoschloenbachia, and it may be only a heterochronous homcomorph of
the Lower Senonian Mumniericeras.
142 Transactions of the Royal Society of South Africa.
Genus HULoPHOcERAS, Hyatt.
42. Hulophoceras natalense, Hyatt.
1921. Spath, Pondoland, table to p. 50.
As will be mentioned in the description of Spheniscoceras, Hyatt’s form
may be only one and an uncommon type of the development separated
from Eulophoceras by Crick and again by Dr. van Hoepen on account of
minute differences in what appears to be a very variable suture-line.
Genus SPHENISCOCERAS (Crick MS.), Spath.
1921. Spath, loc. cit. (Zululand), p. 242, text-fig. C1, p. 248.
Genotype: S. africanum (Crick MS.), Spath, ebid., p. 248, fig. C la
(Plate VI, fig. 1).
In the original diagnosis this group is described as “intermediate
between Placenticeras and Sphenodiscus, Meek,” and after giving the char-
acteristics of these two genera Crick continues :
‘In the present genus the shell is discoidal, lenticular, with a very
small or a closed umbilicus, the sides with obscure radiating ridges; the
periphery is acute and usually pinched in a little on either side. The lobes
and saddles of the suture-line are rather numerous, but not so many as in
Placenticeras and Sphenodiscus. In the suture-line the saddles are feebly
incised and have not entire margins as in Sphenodiscus ; the external lobe is
_ wide and very deep; the external saddle is deeply divided by a secondary
lobe.
“This genus is represented in the present collection by three [four]
examples and a fragment. The examples, which range from 102 to more
than 157 mm. in diameter, appear to belong to four different species.”
The suture-lines of Crick’s three species were figured by the writer on
a previous occasion ; that of Sphenrscoceras cf. amapondense, here repro-
duced, is of interest in that it has the external saddle less indented than
the other “ species,’ and the ventral lobe is very shallow.
The absence of any reference to Hulophoceras, Hyatt, to which genus
Crick refers the specimens in 1906 (see Woods, loc. cit., p. 837) is the more
remarkable as the four “ species’ of Crick are very close to Hulophoceras
natalense, Hyatt, and it might be held that the development of what may
be a thinner keel, and the shght differences in suture-line and ornamenta-
tion, are not sufficient to justify the creation of even different species.
The ornamentation of the earlier whorls of Sph. minor has great resem-
blance to that of Diaziceras ; but since the early whorls of D. tissotiaeforme
are compressed (loc. cit., Zululand, pl. xix, fig. 1c), Spheniscoceras cannot be
On the Senonian Ammonite Fauna of Pondoland. 143
a direct descendant of Diaziceras, but must be looked upon as a more or
less parallel development therewith of the normally lobed Pseudoschloen-
bachia and the form described as gen. nov. (Mumericeras ?) crickt, Spath.
Eulophoceras, if distinct at all, is a closely similar development. From
Hyatt’s description, however, it appears that the ornamentation of his
somewhat imperfect holotype of Hulophoceras is not strongest at the um-
bilical and the ventro-lateral shoulders, as in Diaziceras and in the early
whorls of Spheniscoceras.
The fact that Dr. van Hoepen, who again created a new genus for the
group here discussed, did not record the true Huluphoceras natalense, con-
firms the writer’s opinion that the separation of Spheniscoceras from Eulo-
phoceras is not well founded, and Hyatt’s type, which is not well preserved,
may represent only an extreme variation.
Lenticeras * gullient, Pervinquiere, which was included in the present
stock by Dr. van Hoepen, is here considered to belong to the Coniacian
family Tissotedae. It was selected as type of Prelybicoceras, Douvillé,*
already in 1912.
The internal suture-line of Sph. tenue is here figured for comparison
with that of Sphenodiscus, in which genus more pronounced reduction has
resulted in monophylloid saddles. This character induced Salfeld + to
consider the genus Sphenodiscus and the families Placenticeratidae, Coilo-
poceratidae, and KHngonoceratidae to belong to the _ super-family
Phylloceratida.
43. Spheniscoceras africanum (Crick MS.), Spath.
(Plate VI, fig. 1.)
1921. Spheniscoceras africanum (Crick MS.), Spath, loc. cit. (Zululand),
p. 242, fig. Cla, p. 243.
1921. ? Pelecodiscus capensis, v. Hoepen, loc. cit., p. 32, pl. v, fig. 11.
1921. Spheniscoceras africanum (Crick MS.), Spath, Pondoland, table
to p. 50.
The holotype of this species (C 19421) of dimensions 157—-59—-28
—-03, was described in detail by Mr. Crick, but there is now no need to
give the original diagnosis. Dr. van Hoepen’s small form is doubtfully
included here. Its suture-line, however, shows inclination towards the
umbilicus, like the suture-lines of the other species mentioned below.
* « Hvolut. and Classif. d. Pulchell.,’’ Bull. Soc. Géol. France (4), xi, 1911, fase. 4—5.
(1912), p. 315. ;
Ub. d. Ausgestalt. d. Lob. Lin. b. Jura und Kreide Amm.,”’ Nachr. K. Ges. Wiss.
Gottingen, Math.-Phys. K1., 1919, pp. 8 and 9 (of separate copy).
144 Transactions of the Royal Society of South Africa.
44, Spheniscoceras tenue (Crick MS.), Spath.
(Plate VIII, fig. 3; Plate VII, fig. 3.)
1921. Spath, loc. cot. (Zululand), p. 242, text-fig. C le, p. 243.
1921. 3 ,, (Pondoland), table to p. 50.
This species is represented in the collection by one example, almost
completely septate, of the following dimensions: 122-5—-63—-23—-00.
Crick described this species in detail, but it is characterised in the one
sentence :-—-
‘ Besides the differences in the suture-line, this species is easily dis-
tinguished both from Spheniscoceras africanum and Sph. minor by its
greater compression and much more feeble ornaments.”’
45. Spheniscoceras minor (Crick MS.), Spath.
(Plate VI, fig. 2.)
1921. Spath, loc. cit. (Zululand), p. 242, text-fig. C 1b, p. 243.
SPA dle - (Pondoland), table to p. 50.
This species was based on specimen C 19422, the dimensions of which
are: 102—-64—-27—-00. These dimensions were taken at the begin-
ning of the body-chamber. A portion of this is preserved, but is very im-
perfect. Of Crick’s description only the following need be quoted: “ At
first sight this appears to be an immature specimen of Sph. africanum, but
the form of the suture-line does not justify that conclusion. The character
of the umbilicus [perfectly closed], as well as the fewness and promin-
ence of the undulations on the inner half of the lateral area, serves also to
separate this species from Sph. africanum. The ornaments of the test
and the form of the suture-line also distinguish this species from Sph.
tenue.”
A second example, not noticed by Crick, is enclosed in a block of matrix
attached to one of the Mortoniceras, referred to under M. soutonz, and con-
sidered to be transitional to M. aff. umkwelanense (C 19451). The strong
ornamentation 1s even more pronounced in this second specimen.
46. Spheniscoceras amapondense, v. Hoepen sp.
(Plate VII, fig. 2.)
1921. Pelecodiscus amapondensis, v. Hoepen, loc. cit., p. 33, pl. vii,
figs. 1 and 2.
A fragment (No. C 19424) in the collection, described by Mr. Crick as
* Spheniscoceras, sp. ind.,’ agrees with the smooth outer whorl! of the type-
On the Senonian Ammonite Fauna of Pondoland. 145
species, S. africanum, but its suture-line differs from those of the known
“species ” of this genus, notably in the external saddle. The suture-line of
S. amapondense is probably nearer than any other to that of the present
fragment, but the half of the suture-line of S. africanum, opposite to that
figured, is also similar, so that by splitting one of these specimens in two
still further “ species ”’ could be established.
47. Spheniscoceras umzambiense, v. Hoepen sp.
1921. Pelecodiscus umzambiensis, v. Hoepen, loc. cit., p. 30, pl. v, fig. 10,
pl. va, fig. 1.
The suture-line of this form differs slightly from those of the other
““ species ’’ here recorded.
Famity NOSTOCERATIDAE.
Genus BostrycHocerRas, Hyatt.
48. Bostrychocerus ? amapondense, v. Hoepen sp.
1921. Bostrychoceras * sp. nov., Spath, Zululand, p. 252.
1921. Heteroceras amapondense, v. Hoepen, loc. cit., p. 17, pl. iv, figs.
1 and 2.
1921. Bostrychoceras ? sp. (Woods), Spath, Pondoland, loc. cit., table to
p. 50 (Nos. 37, 38).
1921. Diplomoceras ? sp. (Woods), Spath, ibed., No. 40.
Genus DreLtomoceras, Hyatt.
49. Diplomoceras ? indicum (Forbes).
1906. Hamaites (Anisoceras) indicus, Forbes; Woods, loc. cit., p. 340,
pl. xliv, figs. 2a, b.
1921. Diplomoceras ? indicum (Forbes), Spath, loc. cit., table to p. 50
Hilo. 39).
Genus OXYBELOCERAS, Hyatt.
50. Oxybeloceras amapondense, v. Hoepen sp.
1921. * Hamites”’ cf. quadrinodosus, Jimbo; Spath, Zululand, p. 255.
1921. Oxybeloceras ? sp., cf. onterruptum, Schliiter sp., Spath, zbed.
1921. Hamites amapondensis, v. Hoepen, loc. cit., p. 15, pl. iii, figs, 5, 6,
text-fig. 9.
1921. Oxybeloceras ? cf. quadrinodosum, Jimbo sp.; Spath, Pondoland,
p. 50, pl. vii, figs. 2a, b.
12
146 Transactions of the Royal Society of South Africa.
Genus NEOCRIOCERAS, Spath.
51. Neocrioceras cf. spinigerum, Jimbo sp.
1921. Spath, Pondoland, pp. 51-2, pl. vii, figs. 6a-c.
Famity BACULITIDAE.
Genus BacuitrEs, Lamarck.
52. Baculites capensis, Woods.
1921. Spath, Pondoland, table to p. 50.
This species is represented by one example (No. C 19420) of a length of
45 mm.
53. Baculites sulcatus, Baily.
1921.-v;> Hoepen, loc. cvt., p. 18, ploimi,) figs (28:
1921. Spath, Pondoland, table to p. 50.
54. Baculites bailyx, Woods.
1921. v. Hoepen, loc. cit., p. 18, pl. ii, figs. 9, 10.
1921. Spath, Pondoland, table to p. 50.
EXPLANATION OF PLATES.
PLATE V.
FIG.
1. Mortoniceras stangeri (Baily), var. sparsicosta, nov. Umzamba Beds, Pondoland.
B.M., No. C 19444. (Suture-line figured in Spath, loc. cit. (Zululand), p. 297,
text-fig. D la.) Reduced?. (P. 138.)
2. Mortoniceras stangeri (Baily), var. densicosta, nov. Umzamba Beds, Pondoland.
B.M., No. C 19456. Reduced 3. (P. 138.)
3. Madrasites natalensis (Crick MS.), Spath. Umzamba Beds, Pondoland. B.M.,
No. C 19432. (P. 134.)
4. Pseudoschloenbachiahumei,nov. Near Bir Mellaha, Egypt. No. I, 2288. 4a =sectional
outline. (Pp. 140, 141.)
PLATE VI.
1. Spheniscoceras africanum (Crick MS.), Spath. Geno-holotype. B.M., No. C 19421.
Reduced 3. (P. 143.)
2a, b. Spheniscoceras minor (Crick MS.), Spath. B.M., No. C 19422. Lateral and
sectional views. (P. 144.)
3a, b. Pseudoschloenbachia pseudofournierit, nov. B.M., No. C 19425. Lateral view
and sectional outline. (P. 140.)
4. Pseudoschloenbachia griesbachi (Crick MS.), Spath. Paratype, B.M., No. C 19429.
(P. 141.)
On the Senonian Ammonte Fauna of Pondoland. 147
FIG.
5. Pseudoschloenbachia umbulazi (Baily), var., with more pronounced umbilical tubercles.
B.M., No. C 19426. (P. 139.)
6. Tetragonites superstes, v Hoepen. With mouth-border. B.M., No. C 19416. (P. 119.)
(All the specimens on this plate are from the Umzamba Beds of Pondoland. )
PLATE VII.
la, b. Pseudoschloenbachia griesbachi (Crick MS.), Spath. Holotype, B.M., No. C 19428.
Side view and sectional outline. (P. 141.) !
2. Spheniscoceras cf. amapondense, v. Hoepen sp. Portion of suture-line, with external
and first lateral saddles. S=siphonal line. B.M., No. C 19424. (P. 144.)
3. Spheniscoceras tenue (Crick MS.),Spath. Internal suture-line, enlarged and diagram-
matic, at diameter=115 mm. A-=antisiphonal line. B.M., No. C 19423.
(P. 144.) See Plate VIII, fig. 3.
4, Mortoniceras aff. soutoni (Baily). Suture-line of, B.M., No. C 19442, last (?) septum,
at about 250 mm. diameter. S-=siphonal line. (P. 136.)
(Specimens 1-4 from the Umzamba Beds of Pondoland.)
5. Canadoceras newberryanum (Meek). Geno-holotype, British Museum (Geol. Soe. Coll.),
Upper Senonian of Vancouver Island. Reduced 3. (P. 125.) See Plate VIII,
fig. 4.
6. Parapachydiscus aff. ootacodensis, Stoliczka sp. Outline-section of inner whorls of
specimen C 19438 from the Umzamba Beds of Pondoland. (P. 132.)
PLATE VIII.
1. Parapuzosia haughtoni, sp.nov. Lateral view of portion of cast of dorsal impression
of specimen, B.M., No. C 19439, to show ornamentation of(missing) inner whorl.
Reduced 3. (P. 128.)
2. Madrasites acuticostatus (Crick MS.), Spath. B.M., No. C 19433. (P. 134.)
3. Spheniscoceras tenue (Crick MS.), Spath. B.M., No. C 19423. (P. 144.) See Plate VII,
fig. 3.
(Specimens 1-3 from the Umzamba Beds of Pondoland.)
4. Canadoceras newberryanum (Meek). Reduced 2. Peripheral view of specimen figured
on Plate VII, fig. 5. (PR. 125.)
PLATE IX,
la, b. Pseudoschloenbachia papillata (Crick MS.), Spath. Holotype, B.M., No. C 19430.
Lateral and peripheral views. (P. 141.)
2. Mortoniceras stangert (Baily), typus. B.M., No. C 19440. Reduced about 2.
Penultimate septum, figured in Spath, loc. cit. (Zululand), p. 297, fig. D le.
(Ps 137.)
3a, b. Gaudryceras cinctum (Crick MS.), Spath. B.M., No. C 19415. Lateral view and
sectional outline. (P. 118.)
4a, b. Parapachydiscus umtafunensis (Crick MS.), Spath. Holotype, B.M., No. C 19434,
Side-end peripheral views. (P. 133.)
(All the examples are from the Umzamba Beds of Pondoland.)
Trans. Roy. Soc. 8. Afr. Vol. X.
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( 149 )
A FUNGUS—-GIBELLULA HAYGARTHII, SP.N.—ON A
SPIDER OF THE FAMILY LYCOSIDAE.
By Paut A. VAN DER BuJL.
(With four Text-figures.)
In March 1921 Mr. W. Haygarth sent us from Krantz Kloof, Natal, a
specimen (fig. 1) of a spider with an interesting fungus growing on it.
The fungus belongs to the genus Gibellula,* Cavara, and this collection
is the first record of the genus from South Africa. It is evidently an un-
described species, and we suggest for it the name Gibellula Haygarthii, sp. n.,
and give the following brief diagnosis :—-
Gibellula Haygarthi, sp. n., P. v. d. Bijl.
Coremia white, many from back of spider (fig. 1), terete, erect, un-
branched, 1-3 mm. long x 80-133 p diam., base bulbous, 266-500 p, apex
more or less sterile for 133-173 w; fertile hyphae unbranched, covering
sides of the coremia (fig. 2), entire height 35-266 p, of two distinct segments,
* In the genus Giibellula the mycelium and conidia are colourless, and the conidia
are aggregated together at the apex of conidiophores borne laterally on the coremia.
These characters distinguish the genus from related genera.
150 Transactions of the Royal Society of South Africa.
lower segment 28-239 w long x 6-7 x diam., densely incrusted, rough, upper
segment 18-26 uw long x 3-6 w diam., faintly rough ; head globular, basidia
short and thick, 4-5-5x3-5 w; sterigmata radiating from the basidia
.
”
a yd Ed od
(As ae
5A =e tot
Fre: 3.
(fig. 4), 6-7-5 pw long x1-5-2 pp diam.; spores hyaline, abstricted singly
from apex of sterigmata, 3-6—-4-4 x 1-5 p.
Habitat.—On a spider of the family Lycosidae; collected by W. Haygarth
at Krantz Kloof, Natal. The specimen was preserved in formalin, and it was
hence not possible to grow the fungus and try inoculation experiments.
The pathogenicity of these fungi requires investigation.
( 151 )
SOME SOUTH AFRICAN STEREUMS.
By Pau A. VAN DER Bust.
(With nine Text-figures.)
Very little attention has thus far been given to the South African fungi
belonging to the family Thelephoraceae,* and of which the largest genus is
undoubtedly Stereum.
Several species of this genus have so far been collected, and Mr. C. G.
Lloyd kindly rendered assistance in identifying them.
They occur on trees and old logs principally, and a few grow on the ground.
The fructifications of some species are stipitate, whereas others form shelf-
like outgrowths on the wood or trees on which they grow or occur flat on
these substrata. The hymenium or spore-bearing part is on the under
surface of the fructification (except, of course, when these are flat on the
substratum), and between the basidia there may be variously formed colour-
less outgrowths called cystidia.
While not at this stage attempting to give a key to the species, we will
for convenience divide them into (1) Stipitate species, (2) Sessile species.
1. STIPITATE SPECIES.
Stereum glabrescens, Berk. and Curt. (Fig. 1.)
Plants stipitate, single, or a few rising from a common mycelial pad ;
Imigee tl
* A smooth hymenium is characteristic of the Thelephoraceae, and the genus Stereum
is distinguished from other genera of the family by its context tissue of several layers, of
which the intermediate is normally composed of longitudinally arranged hyphae.
152 Transactions of the Royal Society of South Africa.
pileus flabelliform, 1-5-5 cm. long, 1-5-6 cm. broad, when young evidently
at times infundibuliform and splitting on one side, zoned, finely velvety,
chestnut-coloured ; margin thin, lobed, lighter than pileus ; stalk -5-1 cm.
long x 1-3 mm. thick, velvety, brown; hymenium even, pinky-buff,
spores hyaline, subglobose, 4 w diam.
Not an unfrequent fungus in the native bush of Zululand. W. Haygarth.
Stereum prozimum, Lloyd. (Fig. 2.)
Plants usually growing densely clustered; pileus thin spathulate to
suborbiculate, attached by a reduced base, upper surface finely velvety,
light cinnamon to light ochre, partly glabrous with age, concentrically
Fic. 2.
zoned, -5-5 cm. broad, 1:5-3-5 cm. high; hymenium reddish bay, with a
waxy appearance; encrusted cystidia scanty, 49 uw x 14 pw, projecting
beyond the hymenium ; spores globose, 4 wu diam.
Gillitts, Natal. W. Haygarth.
The finely velvety surface, combined with habit of growth and waxy,
reddish-bay hymenium, should aid in recognising this fungus.
2. SESSILE SPECIES.
Stereum australe, Lloyd. (Fig. 3.)
Pileus coriaceous, attached by the reflexed side or broadly centrally,
densely tomentose, becoming concentrically furrowed, brown to ashy ;
about -5-1 mm. thick; margin entire; hymenium even, smooth, grey to
light fawnish, with coloured conducting ducts (lacticiferous ducts) which curve
unto the hymenium between the basidia ; ducts, 4 diam. ; spores hyaline,
flattened on one side, 3 x 4 p.
Occurs in Natal and the Transvaal.
Some South African Stereums. 153
Few Stereums have conducting ducts in the hymenium. Their presence
should aid in recognising the above.
Fie, 3.
Stereum bicolor, Fr.
Plants sessile, largely resupinate; pileus soft, 1-5-3-5 cm. broad x
1-1-5 cm. long x -5—2 mm. thick, surface zonate or azonate, faintly villous,
becoming glabrous, light umber brown; hymeniun even, glabrous, white ;
intermediate tissue umber brown, darker than surface ; cystidia encrusted,
spores hyaline, 4-5 x 3-4 wp.
Decaying wood at Durban, by P. van der Bij].
This species is recognisable by its soft consistency, the contrast in colour
between the upper and the under surface, and the colour of the intermediate
layer.
Fia. 4.
Stereum cinereum, Lev. (Fig. 4.)
Plants sessile, efitused—reflexed, pileus coriaceous, cuneate, thin, some-
154 Transactions of the Royal Society of South Africa.
what flexible to firm, laterally connate, 1-3-2-5 cm. broad x 1-4cm. long x
‘5-1 mm. thick, surface concentrically zoned, sulcate, tomentose, brown to
pale and ashy, margin undulate to lobed; hymenium even, pinky-buff,
cystidia absent ; spores hyaline, flattened on one side 4-5 x 2 p.
Gillitts, by W. Haygarth.
This appears close to Stereum lobatum, with surface more sulcate and
more densely pubescent and general appearance more firm.
Stereum hirsutum (Wild.), Fr. (Fig. 5.)
Plants sessile, effused—reflexed, coriaceous, drying firm; pileus imbricate,
surface strigose—hirsute, zoned, concentrically furrowed, ochraceous buff
Fig. 5.
becoming grey in age, connate, 1-5-4 cm. broad x 1-5-6 cm. long x 1 mm.
thick ; hymenium even, buff-coloured ; spores hyaline, flattened on one
side, 6-5-7:5 x 2-2-5 w; cystidia absent.
A very common fungus on old logs all over South Africa. Also a wound
parasite on fruit and other trees.
Fic. 6.
Stereum Kalchbrennert, Mass. (Fig. 6.)
Plants sessile, pileus coriaceous, dimidiate to reniform, attenuated at
Some South African Stereums. 155
attachment, usually clustered, laterally connate, 3-4-5 cm. across, ‘5-1 mm.
thick ; surface concentrically zoned and furrowed, densely strigose, with
bright cinnamon-brown hairs ; hymenium smooth, ochraceous to lilaceous,
spores (teste Massee) ellipsoid, hyaline, 7 x 4:5 pw.
Locality.—Eastern Cape Province, on logs of Rhus laevigata, by J. D.
Keet.
Like St. vellerewm, this species is also close to St. hirsutum. The surface
is brighter coloured, and the plants as a whole usually larger and more
robust. Appears to be rare. |
Stereum lobatum (Kunze), Fr. (Fig. 7.)
Plants sessile, imbricate; pileus coriaceous, thin, somewhat flexible,
cuneate, laterally connate, narrowly attached ; 5-10 cm. broad x 3-7 cm.
long x -5 mm. thick, surface with multi-coloured zones, finely pubescent
Gate
to subglabrous; margin thin, undulate to lobed; hymenium even,
pinky-buff, cystidia absent; spores hyaline, flattened on one side,
4x 1-5 p.
A common South African species ; Cape, Natal, and Transvaal.
Stereum durbanense, sp. n., P.v.d. B. (Fig. 8.)
Plants perennial, sessile, effused—reflexed ; pileus thick, corky, drying
firm and brittle, 2-6 cm. broad x 1-3 mm. thick, imbricate ; surface con-
centrically sulcate, tomentose, cinnamon, changing to grey in old specimens ;
margin entire ; hymenium even, light fawn, spores hyaline, 3-4 p.
At Durban on old logs.
St. subpileatum has encrusted cystidia. These we do not find in the
Durban specimens. Apart from this, the specimen appears close to Sé.
156 Transactions of the Royal Society of South Africa.
subpileatum, and is so referred by Mr. Lloyd, who notes the absence of
cystidia.
Fic. 8.
Stereum tomentosum, n. sp., P. v.d. B. (Fig. 9.)
Pileus coriaceous—corky, largely resupinate, with free margin, -5-1 cm.;
surface tomentose with pad up to 1 mm. thick, tan to old gold and becoming
chocolate with age, concentrically sulcate in better-developed specimens ;
Fig. 9.
margin thick, entire; hymenium pale hlac; in structure 1-2 mm. thick
(including pad), with a narrow, horny, reddish-brown zone beneath the pad,
beneath which the tissue is a ight fawn ; spores not found.
Locality.—Collected at Durban on dead log by P. van der Bij.
Stereum vellereum, Berk.
Plants thin, almost membranous, sessile or effused-—reflexed ; pileus
coriaceous, tough, imbricate, laterally connate, often attenuated at attach-
Some South African Stereums. 157
ment, 1-5-2-5 cm. long x 1-2 cm. broad xX -3 mm. thick; surface densely
strigose with greyish-white hairs, concentrically furrowed ; margin thin,
entire to lobed, incurved in drying ; hymenium pale ochraceous, smooth ;
spores (/este Massee) hyaline, subglobose, 4—5 wp.
Locality.—On old logs of Quercus sp. in the Paarl district, by P. van
der Bil.
Close to the common St. hirsutum, but thinner and paler in colour.
UNIVERSITY OF STELLENBOSCH,
STELLENBOSCH,
UnIon oF SoutH AFRICA.
( 159 )
COLOUR AND CHEMICAL CONSTITUTION.
Part XV.—A Systematic Stupy oF FLUORESCEIN AND
RESORCIN-BENZEINE.
By James Morr.
In Part XI of this work (These T., Vol. IX, part 2, p. 129) a systematic
study of phenolphthalein was undertaken, as a result of which the colour-
value of substitution (by bromine taken as a standard substituent) of every
one of the twelve hydrogen atoms in this compound was approximately
determined. .
As the nature of this investigation does not seem to have _ been
fully understood overseas, e.g. by the abstractor of the London Chemical
Society, I think it desirable to reprint the results in another form, viz.
that of the “ dicyclic colour-factors ’’ explained in Parts X and XIV.
Table of Colour-factors for Bromine in Phenolphthalein.
Colour-factor.
Bromine substituted for hydrogen in the a position =1-0180
9 2 7 ‘ b oe — 0020
» 53 ; c ee = Ol4S
9» a ; d » ~=1-0180
9 ; a : e 10270
; ~ : = i ONS
. » 45 3 g ee OleES
53 : - - h Om)
> 7 5 a 10235
; 9 $5 j ee OLS
y 55 ig as k OS
» is ap as l 3 =1-0285
Note.—The numeration of the positions is that given on p. 130 of Part
XI, and is also repeated here under fluorescein.
Thus to calculate the colour of phenol-tetrabromophthalein (which is
160 Transactions of the Royal Society of South Africa.
phenolphthalein with bromine for hydrogen in the a, b, c, and d positions),
we have A,=A,)(554) x 1-018 x 1-002 x 1-0145 x 1-:0180 =583-6.
Again, it was shown in Part IX, p. 225, that the factors for chlorine
and iodine are almost the same as those for bromine. Putting in the
numerical values into the algebraical formula given on that page, we find
that the colour-factor for chlorine differs from that of bromine by
18 x 0-000037, or 1 part in 1500, and that the colour-factor for iodine is
greater than that of bromine by alike amount. Hence, for example, the
calculated colour of phenol-tetrachlorophthalein is 583-6 (that of phenol-
/
“Ale ae 1499\ 4
tetrabromophthalein just calculated) multiplied by al , chlorine having
replaced bromine four times. Thisis 582-1. This agrees with observation,
viz. A581 and A582 from two specimens. I am greatly indebted to
Professors Partington of London and Mackenzie of Dundee for specimens
of tetrachlorophthalic acid which they sent me after I had vainly tried to
purchase it all over the world for three years.
1500 \*
Similarly, phenol-tetraiodophthalein should have A=583-6 x Ge )
=585-2, but I have not been able to make (or obtain from its discoverer)
this compound.
In the same way, the calculated colour of ordinary (fqjk) tetrabromo-
phenolphthalein is 554 x (1-0145)? x (10125)? =A 584-0, and the calculated
1499 \4
colours of tetrachloro- and tetriodophenolphthalein are 581-0 x( 72 |
1500\4
and A 584-0 x Cay respectively. These agree with observation.
A.—-Derivatives of Fluorescein.
geen.
fon
cL Je
b
Employing the same numeration of the positions as was used for phenol-
phthalein in Part XI, 7.e. assuming that fluorescein is e—l-oxophenol-
phthalein, it is to be expected that the same colour-factors will hold good.
Unfortunately, the experimental difficulties have in some cases proved
Colour and Chemical Constitution. 161
too much for me, particularly as regards substitution in the h and 2
The following table contains my observations (note:
X fluorescein
Substitution-products of Fluorescein.
positions.
= 4934) :—
A
a-monobromo- : : : . 501
a—d-dibromo- : : . 609
c-monobromo- 3 3 : . 499
a-—c-dibromo- 3 ; : . 506
a—c—d-tribromo- : Oke
a—b—c—d-tetrabromo- : : . O14
a-methoxy- . ‘ : . 497
b- or c-methoxy- . : ‘ . 492
a-b-—c-trichloro- . : : . 505
a-b—c-d-tetrachloro- : 2-513
g—j-dibromo- : : . 509
f-g-j-tribromo- . : ; > old
Beast verre DOM o- an 521
(cosine) {
g-j-dimethyl- . 500
(cresorcin)
f-k-dibromo-g-j-dimethyl- . 516
h-i-dimethyl- 4894
(“* y-orcinphthalein ”’)
f-9 j-k-tetrabromo-, der. of above . 521
h-1-dioxy- (phloroglucin- af 489 neutral
phthalein) | 494 alkali
h-i-dimethyl-abed-tetrachloro- 5114
a-c-dichloro-fgjk-tetrabromo- anette)
b-c-
” 29 29 9
(Phloxin iy poss
epenucuracnlore-tpjk tetra bromo- ne 54d
(Phloxin proper) {
abcd -tetrachloro-fgjk-tetraiodo- 548
(Rose bengale) uf
[‘ a-orcinphthalein ” . 520 vague]
Oxyquinolphthalein 550-+527-+ 509
A
a-nitro- é : : ’ . 500
c-nitro- : : : 3 . 498
Or i 489 neutral
| 494 alkali
b- or c-oxy- : : : . 492
a—d-dioxy- . : : . 493
ad-dimethoxy- . : ‘ . 499
b-c-dimethoxy- . : : . 496
abc-trimethoxy- . . 499
a—f-g—j—k-pentabromo- : 529
c—f-g—j-k- 5 : : . 526
a-—c—f-g—j—k-hexabromo- : . 535
a—d—f—g—-j-k- S; , . 939
a-nitro-f—-g—j—k-tetrabromo- . . 530
CEI am ash ; O20)
f—k-dinitro-g—j-dibromo- at
22
(‘‘ methyleosine ’’) of :
f-g—j-k-tetrachloro-. : . 518
abedfgjk-octochloro- . : = 007
ss -octobromo- . : . 544
f-g—j—k-tetriodo- | 595
(erythrosine)
acdfgjk-heptabromo- . ; . 540
abe -trichloro-fgjk-tetrabromo- . 536
abed -tetrachloro-g-iodo- ‘ . 520
g-iodo-fkj-tribromo- 543
29 29
[‘‘ B-orcinphthalein ” 440]+
[‘‘ Tribromo-f-orcinphthalein ” 463]
if 494 neutral]
|. 499 alkali]
Oxyquinoltetrachlorphthalein 577-530
[‘‘ Quinolphthalein ”
* This may be abd: there is no means of deciding.
+ The substances in brackets are related to fluorescein but are not direct derivatives.
13
162 Transactions of the Royal Society of South Africa.
It is probably unnecessary to describe the method of preparation of
any of these substances, the information in Part XI about derivatives of
phenolphthalein sufficing by analogy. It may be noted also that positions
f and g are radically different in fluorescein, although they are almost the
same in phenolphthalein. I have assumed, in assigning constitutions to
these substances, that a substituent will enter the g and j positions first,
since the other two ortho-positions, f and k, are “ protected ”’ by the oxo-
linkage.
Deductions from the Observations.
A. The observations become consistent for the four “ phthalic ”’
positions, a, b, c, and d, if we assume a-monobromo- = d-monobromo-
— +8, and also assume b-monobromo- to have no effect, and c-monobromo-
= +4}.
Hence a-monobrome colour-factor in fluorescein = 1-0162
b- ” 9 ” = 1-0000
C- re) ”? ”? =1-0091
d- ” om) 9 =1-0162
These four factors therefore are smaller than those obtained for phenol-
phthalein, but their mutual variations resemble those of the phenolphthalein
series, and the actual discrepancy is not much more than 1 part in 500,
or than the error of observation. It follows therefore that the phthalic
part of the fluorescein molecule does not differ much spatially from the
phthalic part of the phenolphthalein molecule.
B. It has been impossible to make monobromo-compounds of the
fgjk class, but by difference I estimate that {-bromo=k-bromo= +6, and
that g-bromo=]-bromo=7#.
Hence {-monobromo colour-factor in fluorescein = 1-0122
eS as oe) 29 =e ONi
If 7 : My =1-0157
k- 29 ” ” =1-0122
These four factors are not smaller than those obtained for phenol-
phthalein, and vary irregularly like an. experimental error: nevertheless
I think the small differences are real, and are due to the presence of the
oxo-linkage.
As already stated, the h and 7 bromo colour-factors could not be
ascertained by direct experiment. By inference from y-orcinphthalein
and phloroglucin-phthalein, however, I find them to be less than unity,
viz. both the h-monobromo and the 7-monobromo colour-factors are
Colour and Chemical Constitution. 163
about 0-990. This is completely discrepant with the phenolphthalein
figures.
C. The chloro-derivatives follow as before from the bromo-derivatives
1499
by multiplying by 1500
as often as the change from bromine to chlorine
is made.
D. The nitro-derivatives are not materially different from the bromo-
derivatives. The a-,d-, g-, and j-nitro colour-factors are all about 1-016,
and the c-, f{-, and k-nitro colour-factors are all about 1-012.
EK. The methyl colour-factors are considerably smaller than the bromo
ones, the methoxyl] colour-factors are smaller still, say 1-006 for the biggest
of them, viz. the a-methoxyl, and the hydroxy colour-factors are smallest
of all, practically unity. Indeed, when caustic alkali is employed all the
oxy-derivatives of fluorescein have their absorption band in the same place
as that of fluorescein. Gallein and oxyquinolphthalein constitute excep-
tions to this rule.
Two examples of the method of calculation may be given :—
1. Octobromofluorescein : A, = 493-5 x 1-0162 x 1-0091 x 1-0162 x 1-0122
x 1-0157 x 1-:0122 x 1-:0157=543-0. The observed value was 544.
1500
This is 540-0. The observed value was 537. Further experiment may
show that the ratio Cl/Br is not so near to unity, the error only showing
in this extreme case.
1499\8
2. Octochlorofluorescein: A, = 2 for octobromofluorescein x ) :
B.—Derwatives of Resorcin-benzerne.
HO 2 -— OH
J ee yy
i | g
afm
de
b
Resorcin-benzeine is phenyl-2—7-dihydroxy-xanthhydrol and is fluores-
cein minus the CO,H group. The same numeration is therefore used, with
m for the position occupied by the new hydrogen. Resorcin-benzeine in
alkaline solution appears identical with fluorescein in colour and intensity
of flucrescence. The absorption-band is at A 492.
164 Transactions of the Royal Society of South Africa.
Substitution- products of Resorcin-benzerne.
A A
pee nr Atiss neutral b-bromo- : : : . 493
492 alkaline a-orc-bromo- . : : . 497
484 neutral 490 neutral
b-oxy- 492 just alkaline Fe Ocoee tee, alkaline
544 alkaline d- or m-bromo- . ; : . 505
b-methoxy- . : : : . 493 b-nitro- : é : : . 500
d- or m-oxy- : : . 430+492 fgjk-tetrabromo- . : ; . org
d- or m-sulphonic acid \ 498 fgjk-tetrabromo-b-nitro- ; mys 47
‘‘resorcin sulphophthalein ”’ b-nitro-d- or m-sulphonic acid . 6506
Sojk-tetrabromo-m-sulphonic acid . 527
It is to be noted that the list is limited from the fact that sulphuric
acid cannot be used as condensing agent in making derivatives of resorcin-
benzeine, because at an elevated temperature resorcin itself condenses to
a coloured fluorescent compound “ resorcin-ether,’ the nature of which
is at present unknown. It has a dirty rose colour in alkali with AA 490 and
550 and a bluish-green fluorescence. Phosphorus oxychloride, if not over-
heated, forms a suitable condensing agent.
The conclusions to be drawn in the case of resorcin-benzeine are :—
A. The f, g, 9, and k colour-factors are the same as those of phenol-
phthalein and fluorescein, viz. about 10135 on the average for bromine
taken as standard. Further experiment is necessary to decide whether
the g-factor is greater than the f-factor, and whether the g- and j-factors
are identical, as they should be by symmetry.
B. In the phenyl ring the effect diminishes in the order d, c, b, but in
this case the effect is still positive at b, although it appears to be nil in
fluorescein and is sometimes negative in benzaurine. Much further work
is necessary to clear up these small anomalies.
Addendum, 12th June 1922.—Professor G. T. Morgan kindly sent me a
sufficient specimen of 1-3-dioxynaphthalene to enable naphthofluorescein
to be made and examined. It has AA 535 and 496. The ortho-meta
naphthol/phenol factor is thus 1-042 in the dicyclic series. This could not
be ascertained from naphtholphthalein, because the latter is a diortho-
compound and therefore cannot be compared with phenolphthalein. This
naphthalene factor can be employed to calculate the naphthosafranines
and naphthacridines.
The effect of the naphthalene ring in the dicyclic series is practically the
same as that of two bromine atoms substituted in the gh or 1] positions :
v.e. the outer carbons of the naphthalene ring have almost no effect. This
is of great theoretical importance.
( 165 )
THE “ ACCOUNT-BOOK” OF JAN HASZING.
By C. Pisper, M.D. (Leiden), Johannesburg, and
H. Zwarenstein, M.B., Capetown.
Jan Haszing practised as a surgeon at Capetown in the middle of the
eighteenth century. His ‘‘ Account-book,”’ which in some way or other
has found its way to the archives at Capetown, starts at 1736 and runs on
continuously till 1767. The following pages contain a verbal transcript
of some of its entries, together with an English translation. They give some
idea of what medical practice was like then in an “outpost” like Capetown.
No explanations or footnotes are necessary ; we only beg to draw attention
to the fact that the Rd. (“ rix-dollar ”’) was divided into eight “schellings”’
(sixpences). It is a pity that our colleague occasionally has forgotten to
note down for himself what he charged his patients; may we infer that
medical men were but indifferent bookkeepers even then ?
[ AANTEEKENINGH.
166 Transactions of the Royal Society of South Africa.
AANTEEKENINGH OFF SCHULT BOEK VAN JAN HASZINGH
CHIRURGIIJN. Beginnende Ao. 1736.
1763. Christiaan Persoon Deb.
Den 28 Febr: uw huysvrouw met den chirurgijn Uijting gevisit:d.
lab: aan een febris Biliosa dewelke in een febris Hectica nader-
hand overging. Wijdersdaagts een en tweemaal ge¥: tot den
8 Maij. dat overleeden is komt voor practijk en visitens
samen - - - - - - - - Rds. 25.
Den Inhoud deses Voldaan.
1765. Mons: Pieter Soermans Deb:
5 N:r Uw beminde seer geluckig van een dood kind ’twelk een
waterbuijkje had verlost en waarvan de placenta op een extra
ordinaire wijse vast zat, sodat deselve met veel moeijte moest
afgehaald worden. Verder ’t nodige in *t kraambed besorgt
en wederom volkome herstelt.
Ten dank voldaan.
1765. Juff:w D: Wed:we Hazewinkel Deb.
18 Junij Ued: met de Heeren practizijns Van der Riet, Nelson, d:
Vrij en van Nierop, ge¥: lab: aan Colica Biliosa beneevens een
consid:ble Hernia umb: komt voor gedaane consultatien en
visitens tot den 1 Julij samen - - - - Rds. 10.
Den Inhoud deses Voldaan.
1766. Den Adsist Alleman Deb.
15 Augts: Voor uw huijsvrouw 1 bott: Garg. sac A
20---: ,, ,, Hen bott: vin: medicat: >» =~6
21---: ,,,, 1 flesje Elir: stom c: tct: absinte a as
27 - - - : Voor uw kind 1 flesje sp: nitr: dulé: 55 RE
29- -- 2 |,, » een dos: sirp: cich: c: Rhas 1) » 4
1 Sbr: Voor uw huysvrouw 1 haust: Corral 2) spo ine
3 Sbr: Voor ad jdem - - - - - ae a
5--: 59, 1 g: flesje sp: carm: sil¥: c: tet: Castoril yp ile
22-- : Voor de meyd Lea | Flesje sp: nitr: dulce » 4
PASTE 59) ad jdem - : - - - Se yon) et
The ‘*‘ Account-Book”’ of Jan Haszing. 167
REGISTER OR ACCOUNT-Book oF JAN HASsZzINGH, SURGEON.
Beginning Ao. 1736.
1763. Christiaan Persoon, Deb.
28 Febr. Went to your house, together with surgeon Uijting, to
see your wife, suffering from febris biliosa, which later on
changed to a febris hectica. Thereafter every day one or
two visits up to 8th May, the date she died; comes for
treatment and visits together ; . Rds. 25
Paid.
1765. Pieter Scermans, Esq., Deb.
5 Nov. Your beloved very successfully delivered of a dead child
which had a water-swollen abdomen, while the placenta was
fixed in an extraordinary way, so that the removal was
attended with great difficulty. Thereafter attended to the
lying-in, resulting in complete recovery.
Paid with thanks.
1765. The Widow Hazewinkel, Deb.
18 June. Came to your house, together with the doctors Van der
Riet, Nelson, d. Vrij and van Nierop; found you suffering from
colica biliosa complicated by a considerable hernia umbilicalis ;
comes for consultations and visits up to Ist July . Rds. 10
Paid.
1766. The Assistant Alleman, Deb.
15 Aug. For your wife, 1 bottle gargarism . A eet ee!
20'<; , 1 bottle vinum medicat. . ; ; : PG
Ai, », 1 vial elix. stomach. c. tinct. absinthi Pople
27. ~,, For your child, 1 vial spirit. nitr. dulce. . Ae. ee |
Oe 5 3 92 dose syrup. -cich. c. thas. 1). ean wee
1 Sept. For your wife, 1 haust. corral 2) . : oy. eee te?
Die ss ,, the same ; : : : : Me ee
Dia ad ,, Ll vial spirit. carmin. sylv. c. tinct. castori oy ed:
22 ,, Fortheservant-girl Lea, 1 vial spirit. nitr. dulc. » 4
Se iss ,, the same ; : : cece
168 Transactions of the Royal Society of South Africa.
11 Obr: Voor een meyd een Vomet: - - - =) | | agg eres
HD ae: » 5, een haust: carmin: - - - =) gg ey
26 Xbr: Voor uw huijsvrouw 1 haust: carm: Annod: yn ae
Overgedaane Visitens van tijd tot tijd same 5, 20-
Somma Rds. 27.2
Den Inhoud deses Voldaan.
1) 2 2) 2
1766. Anthony Barrange de Jonge Deb:
1 Maijy Uw meyd Gev¥: met een Consid:ble Antrax
beneevens Verscheyde farunculae aan de Rug
en geneese Rds: 4.4
26 -- : Jonge gev: met een gecond: solutie aan de groote
toon en geneesen | Rds. 1.4
1 Julij Voor UE en huysvrouw ieder een pur®: er ele
9-- : ,,,, UE een bott: garg: eer 4
10 - - : Voor Uw huysvrouw ad jdem . -4
Somma Rds. 8-
Den Inhoud deses Voldaan.
1766. Jan Victor Kelders Deb:
2 Maij Uw huysvrouw met de Heere d: Vrij & V: Nierop
gev: Lab: aan een febris ardentis sijnde ontrent
—-— maande swanger vervolgens geconsul: &
gepract: tot den 12-: sijnde same 20 visitens __Rds: 6-
21 - : Wederom alleen ob nieuws gev: ’snagts ge-
aborteerd hebbende sijnde flux: Loch: seer
wijnig waar voor ’t nodige besorgt tot volkome
herstelling gelijk meede Lab: aan verscheyde
farunculus in de nek daags verb: & geneesen 6
Somma Rds. 12-
Den Inhoud deses Voldaan.
The ‘‘ Account-Book”’ of Jan Haszing. 169
11 Oct. For a servant-girl, a dose of emetic ; : RE 5
Ae ;) sarhalst. carmin, 9. : p : Oy
26 Dec. For your wife, 1 haust. carmin. anodyn. : Re ech
For visits . : : ; : : els
Together Rds. 27-2
1766. Anthony Barrange de Jonge, Deb.
1 May. Visit to your servant-girl, suffering from a con-
siderable anthrax and several furuncles on
the back; cured same. . Rds. 4-4
26 May. Visit to your house-boy, ee an an
inflamed contusion in his ae toe ; cured
same . Rds. 1-4
1 July. A purgat. for S ocelt a one on your ai Bc aoe eS
9 ,, <A bottle of gargarism for yourself . ae 4
10 ,, The same for your wife : ; wii 4
Together . Rds. 8-
Paid.
1766. Jan Victor Kelders, Deb.
2 May. Went to see your wife, together with the doctors
de Vrij and van Nierop; found her suffering
from a febris ardentis while being about ——
months pregnant; for consultations and
treatment up to 12th May, 20 visits altogether Rds. 6
21 May. Came to see her again, alone, having passed an
abortion during the night, the flux. loch. being
very scanty; done what was necessary to
remedy this, with perfect success; treated her
at the same time for furuncles in the neck,
dressed those daily, cured same . 6
Together . Rds. 12
Paid:
170
Transactions of the Royal Socrety of South Africa.
1766. Mons: Michiel Smuts Deb:
4 Maij
18 Julij
26 > -
27 - -
18 Augt:s
22
ede se pd)
6 Sept:r
ieee
12 Octob:
1766. Jufi:w D: Wed:we Abel Deb:
De Jonge Jacob gevisit: met een solutie inde
5 Maij
18 Junij
10 Ob:r
Voor UE een purg:
Uw klhjn dogtertje gev: Lab: aan een febris
ardentes waar voor de noodige med: besorgt
tot med: Augt:s & geneese komt voor
practijk visitens & gel: med: same
: Voor uw dogter Maria 1 bott: Garg:
: Voor uw dogters Barbara & Maria jd: & 1
purg:
Voor de oude Jonge 1 pl: Resol¥: op Leer
: de klijne jonge gev: met een gecontund: solutie
in de pink, waarvan de nagel weggenome
Verders daags verbonde & geneese komt
voor de geneesing
De meijd Gilida gevisit: Lab: aan een Con-
sid:ble geinphlameerde keel overgaande
skielijk tot gangr:
ons een singulier en sorglijk geval *twelk
verder door de Heer van Nierop behandelt
en door mij de medicam: besorgt sijn komt
voor practijk visitens en gel: medicam:
samen
& sphacelus sijnde
: De madagascarse jonge gevisit: met een
consid:ble solutie aan de hiel beneevens
contusie aan de tendo Achilles ’t selve daags
verb: den tijd van ontrent 2 maande &
geneese
Uw dogter Maria gevisit: met een geobstru-
eerde knie wijt een sinking stoffe veroor-
saakt waar voor *t noodige besorgt en
geneesen
Somma
Den Inhoud deses Voldaan.
Vinger & geneesen
Ken Vomet:
Voor Een flesje oogwater
Den Inhoud deses Voldaan.
Rds. -4
Ge
-4
ie
-4
9-
10
6-
pe
Rds. 29.4
Rds: 1.4
29 4
99 4
Rds: 2.4
The <‘ Account-Book”’ of Jan Haszng.
1766. Michiel Smuts, Esq., Deb.
4 May. A purgative for yourself
18 July. Came to see your little daughter, suffering from
a febris ardentes, wherefore I supplied the
necessary remedies till medio Aug. and cured
her; comes for treatment, visits, and medicine
For vour daughter Maria, 1 bottle gargarism .
For your daughters Barbara and Maria, | bottle
of gargarism each and | dose of purgative
18 Aug. For the old ee 1 resolv. plaster on
leather
22 Aug. Came to see the young ree Hoe piece
from an inflamed contusion in the pink,
wherefore I took away the nail. Thereafter
daily dressings and cured; comes for the cure
6 Sept. Came to see the servant-girl Gilida, suffering
from a considerably inflamed throat promptly
developing into gangraena and sphacelus ;
this being a singular case which has caused us
much anxiety ; further treated by doctor van
Nierop, the medicines being supplied by me ;
comes for treatment, visits, and Medicines
together .
Came to see the Medaaseeas ee ce ere from
a considerable inflammation in his heel, com-
plicated by a contusion in the tendo Achilles ;
dressed the same daily for a period of about
two months and cured same
Came to see your daughter Maria Mice rane
an obstructed knee arising out of a fluxion ;
performed what was necessary and cured
26
27
99
99
17
99
12 Oct.
Together
Paid.
1766. The widow Abel, Deb.
5 May. Came to see the house-boy Jacob, suffering from
an inflammation in his finger; cured same .
18 June. An emetic
10 Oct. Hye-wash
Paid.
Rds
171
10-
. Rds.29-4
172 Transactions of the Royal Society of South Africa.
1766. Den Herw: Heer Serrurier Deb:
16 Maij Snagts of smorgens vroeg uw oudste dogtertje
ge¥: met convulsio & St: Vitus Dans waar
voor ettelijke Rijsen geclisteerd. Verder
de noodige med: besorgt tot den 24- &
geneesen Rds. 4
28 -- : Uw Eerw: gev: Lab: aan Angina die seer hard-
neckig was wederstreevende veele middele
en waar voor de nodige med: sijn besorgt
tot pr:mo Julij & geneese $e
25 Junij UE beminde Gev¥: met inphlamatie in de keel
waar voor 't nodige besorgt en geneesen ae
6 Augt:s Voor uw oudste dogtertje Een purg: a -4
15 - - :-,, ,, ad jdem I -4
LOM bei ,,45 1 Hes] e" Collin: ss -4
23 == 2 3.) Henionsimann? Calabr: a -4
1 Sb: wo) ad jdem a -4
18 X:br Voor UE beminde 1 purg: & 1 bott: garg: ae
Somma Rds: 15.4
Hiervoor Ten Dank Voldaan.
1766. Matthijs Van Wieling d: Jonge Deb:
24 Maij. Uw vrouw Gev: klagende over swaare naween alsoo
daags bevoorens hadde geaborteerd waar voor ’t , nodige
besorgt en geneesen.
Ten dank Voldaan.
1766. De Wed:we N:s Mulder Debet.
15 Junij UE Gev¥: met een sware coorts waar voor de
nodige med: besorgt & geneesen komt voor
practijk Visitens en Gel: medicam: same Rds. 6:
1 Juhj Voor de Jonge Thomas Een purg: . a)
Sulla y yes. ema cadena ; oO
Somma Rds. 6.6
Den Jnhoud deses Voldaan.
The ‘* Account-Book”’ of Jan Haszing. 173
1766. The Rev. Serrurier, Deb.
16 May. Came to see, during the night or early morning,
your eldest daughter, suffering from con-
vulsions and St. Vitus’s dance; wherefore I
administered several clysters. Thereafter I
supplied those medicines as were necessary
(till the 24th), and cured her : . Rds. 4
28 ,, Came to see your Reverence, suffering from a
very obstinate angina, which several medi-
cines had been unable to conquer ; wherefore
I supplied medicines up to Ist July, and
cured same . : : 5
25 June. Came to see your beloved, suffering from in-
flammation in her throat ; wherefore supplied
what was necessary, and cured . : ' Syke
6 Aug. A dose of purgative for your eldest daughter . ,, -4
bb, Lhesame ~. oe: é » A
10 Sept. An eye-wash for same . tn, (gues
Zo 4, . One ounce of mann. calabr.. . ot
Oct.) Lhe same. : : ‘ : : . oe wee
18 Dec. One dose of purgative and 1 bottle of garg. for
your beloved ; ; : : ; ee
Sum total . 15.4
Paid with thanks.
1766. Matthijs van Wieling, Junior, Deb.
24 May. Visited your wife, complaining about severe after-pains.
having suffered an abortion the day before; wherefore done
what was necessary, and cured.
Paid with thanks.
1766. The widew N. Mulder, Deb.
15 June. Came to see yourself, suffering from a serious
fever; wherefore supplied what was neces-
sary, and cured ; comes for treatment, visits,
and medicines . : ZR Gs a6
1 July. A dose of purgative for your house-boy Thomas ,, -3
Dut) he samernr ‘ : . ; : , ees:
Together . Rds. 6.6
Paid.
174 Transactions of the Royal Society of South Africa.
1766. Mons: Thobias Rogiers Deb:
4 Julij Uw huysvrouw verlost van een vrugtje van ontrent 3
maande ’twelk bijna gebore was, en waarop Verder ontlasting
Van een mola, en een verrotte placenta volgde. Verder
‘t nodige besorgt & herstelt.
Ten dank Voldaan.
1766. Christiaan Pieter Brand Deb:
8 Juli) Uw vrouw geaborteerd hebbende en geen
placenta gevolgd sijnde waar voor late
clisteeren en verder de benodigde med:
besorgt tot volkome herstelling Rds: 6.-
21 Augt:s Voor uw Jonge Ken Vomet: ee EG
>> >» Wenspurg: F =ory
Somma Rds: 6.6
Den Inhoud deses Voldaan.
1766. Rijnier Keet Debet.
14 Julij Uw vrouw gev: Lab: aan een sware bloedstorting
waarop door *t gebruik van gepaste med: en ’t
sette van eenige clisteeren ontlasting van een mola
gevoled is.
12 Augt:s ,, op nieuws wederom gev: hebbende reets 10 dagen
lang sterk gevloeit hier voor wederom de nodige
medicam: besorgt tot pr:mo 7b:r & geneese: komt voor
practijk, visitens en Gel: medicam: same Rds: 10.
Den Inhoud deses Voldaan.
1766. Pieter Malet Debet.
16 Julij Uw huysvrouw seer geluckig & voorspoedig verlost van
een kwalijck gekeert kind *twelk een levendige soon was—
verder ’t nodige int kraambed besorgt tot volkome herstelling—
Ten dank Voldaan.
The ‘*‘ Account-Book”’ of Jan Haszing. 175
1766. Thobias Rogiers, Esq., Deb.
4 July. Delivered your wife of a foetus of about Tet) months
which was very nearly born, and whereafter a mola and a
putrid placenta came to light. Thereafter done what was
necessary, and cured.
Paid with thanks.
1766. Christiaan Pieter Brand, Deb.
8 July. Your wife having suffered an abortion where no
placenta had come to light, clysters admin-
istered to her and supplied what was
necessary, resulting in complete recovery . Rds. 6
21 Aug. An emetic for your house-boy Oe
A purgative for same . oe Co
Together . Rds. 6.6
1766. Rijnier Keet, Deb.
14 July. Came to see your wife, suffering from a severe hemor-
rhage, whereupon appropriate medicines and the
administering of some clysters brought to light a mola.
12 Aug. Came to see her again, having suffered already for the
last ten days from severe loss of blood; supplied what
was necessary up till lst Sept.; comes for treatment,
visits, and medicines together. . Rds. 10
Paid.
1766. Pieter Malet, Debet.
16 July. Delivered your wife very successfully and prosperously
of an awkwardly bent-up child, which turned out to be a
living son; supplied what was necessary for the lying-in,
resulting in complete recovery.
Paid with thanks.
176 Transactions of the Royal Society of South Africa.
1766. Juff:w Elsje Colijn Deb: ;
26 Julij Voor UE na de Verlossing een ord: drank tot
voortsetting van de kraam en vandenaween Rds: 1-
15 Tb:r Voor Een purg: ni -4
28--: ,, ,, ad jdem 3 -4
28: Obie 17,4, adadem me -4
1 Xb:r_ Voor ’tkind 1 ons mann: calabr: & -4
Somma Rds: 3-
Den Inhoud deses Voldaan.
1766. D. Wed:we Huijsman Debet
28 Augt:s Haar EH: met de praktizijns V: Nierop & Nijting gev: Lab:
aan een duyster & singulier geval in de buijk denkelijk door
geobstrueerde ingewande voortgebragt & waarvoor een grote
pL: op Leer vande EmpL: Rustingii 1) geappld: hebbe ben-
evens verscheijde onse sp: sal: vol: ol: om te ruijke komt
voo gel. medica: beneevens eenige gedaane consultatien,
& Visit:s same Rds: 5-
Segge Voldaan.
1)?
1766. Joh:s Guil:m V. Helsdinge Deb:
1 Ob:r Uw vrouw int kraambed gev: klagende over swaare pijn
in de buijk Etx: Waar voor eenige rijse doen clisteeren verder
‘t nodige besorgt en geneesen.
Ten dank Voldaan.
1766. D: Paruijkemaker Joni Debet.
4 8b:r Uw vrouw gev: Lab: aan flux: menss: waar voor ’t nodige
besorgt & geneese. Rds: 1.4
Hier Van Voldaan.
1766. Dirk D: Jongh Debet.
16 9b:r Uw meyd seer geluckig van een dood kind verlost verder
‘tnodige besorgt & hersteld.
Ten dank Voldaan.
The “* Account-Book’”’ of Jan Haszing. Eek
1766. Mrs. Elsje Coliajn, Deb.
26 July. For yourself, after the confinement, the ordi-
nary mixture to succour the lying-in and the
after-pains et ; » Rds: -1-
15 Sept. A dose of purgative
28 ,, #The same :
28 Oct. The same
1 Dec. 1 ounce of mann. calabrin. for the child
Sumtotal’ .> Rds: 32
Pare
1766. The widow Huijsman, Debet.
28 Aug. Visited your Honour (together with the doctors van
Nierop and Nijting), suffering from a perplexing and singular
affection in the abdomen, probably caused by obstructed
intestines, wherefore we applied a large plaster of empl.
rusting 1), on leather, while at the same time we made you
snuff up several ounces of spir. sal. vol. ol.; comes for
medicines, together with consultations and visits . Rds. 5
Paid.
1)?
1766. Joh. Guil. V. Helsdinge, Deb.
1 Oct. Visited your wife during her lying-in, complaining about
severe pains in her abdomen, etc.; caused her to be
clystered a couple of times, and supplied what was
necessary ; cured.
Paid.
1766. The wig-maker Joni, Deb.
4 Oct. Visited your wife, suffering from flux. mens.; where-
fore supplied what was necessary, and cured . Rds. 1.4
Paid.
1766. Dirk de Jongh, Debet.
16 Nov. Your servant-girl very successfully delivered of a dead
child ; supplied what was necessary, and cured.
Paid.
14
178
Transactions of the Royal Society of South Africa.
1766. Mons:r Poul:s Hen:s Eksteen Deb.
1766.
1766.
1766.
1766.
21 9b:r UE beminde seer geluckig van een levendige soon met een
geklemt hooft op sijn Rhoonhuysiaans verlost sonder eenige
beletselen dog *tkind ontrent 18 uuren hierna overleden.
Verder de moeder int kraambed van ’tnodige besorgt en
volkome hersteld.
Ten dank Voldaan.
. De Boode Ab:m Paling Deb:
13 Xb:r Uw meyd geluckig van een kwalijck gekeert dood kind
verlost na lang in arbeijd geseten te hebben verder ’tnodige
besorgt tot volkome gesontheijd.
Ten dank Voldaan.
24 Xb:r Uw meyd seer voorspoedig & geluckig van een levendige
jonge *twelk de nek en linker schouder aanbood verlost.
Verder *tnodige besorgt en hersteld.
Ten dank Voldaan.
De Heer Johannes Breeser Deb:
15 Xb:r Uw beminde met den onderchirurgijn Ramol gev: met
abces onder den arm ’tselve ’sanderendaags geopent verders
eenige rijsen geconsul: en gevisit:d—
Komt voor practijk en voor geconsuleerde visitenssamen Rds: 3-
Den Inhoud deses Voldaan.
Ernst Philip Sparenberg Deb.
17 Jann. Uw huysvrouw gev: Lab: aan een ongemak uijt een
kraam gehouden waar voor ‘tnodige besorgt tot ult:mo
Maart komt voor practijk visitens en gel: medicam: samen
Rds: 5-
Den Inhoud deses Voldaan.
Joh: Jacob Le Roé Deb.
6 Jann. Uw vrouw met den chirurgijn V: Nierop gev: lab: aande
geelsugt—komt voor gedane consultatien tot 12-:
Voldaan met een Vragt Houdt.
The ‘‘ Account-Book’”’ of Jan Haszng. 179
1766. P. H. Eksteen, Esq., Deb.
1766.
1766.
1766.
1766.
1766.
21 Nov. Your beloved very successfully delivered, by means of
the method of van Rhoonhuysen, of a living son, whose
head had caught fast, without any adversity ; the child,
however, having died about 18 hours later. Thereafter
supplied what was necessary for the lying-in, resulting in
complete recovery.
Paid with thanks.
The messenger A. Paling, Deb.
13 Dec. Your servant-girl, who had been a long time in labour,
successfully delivered of an awkwardly turned dead child ;
thereafter supplied what was necessary till complete health
was restored.
Paid with thanks.
The widow Michiel de Kok, Deb.
24 Dec. Your servant-girl very successfully and _ propitiously
delivered of a living son, which had proffered its neck and
left shoulder. Thereafter supplied what was necessary, and
cured.
Paid with thanks.
Mr. Johannes Breeser, Deb.
15 Dec. Visited, together with the under-surgeon Ramol, your
beloved, suffering from an abscess under the arm, which we
opened the next day; thereafter several visits and consulta-
tions :
Comes for treatment, together with visits in consultation Rds. 3
Paid.
Ernst Philip Sparenberg, Deb.
17 Jan. Visited your wife, suffering from an ailment originated out
of a child-birth ; supplied what was necessary till the end of
March; comes for treatment, visits, and medicines Rds. 5
Joh. Jac. Le Roé, Deb.
6 Jan. Visited, together with the surgeon van Nierop, your wife,
suffering from jaundice ; comes for consultations . 12
Paid by a cart-load of firewood.
( 181 )
I. ON SOUTH AFRICAN PARAMPHISTOMIDAE (Fiscu.).
I]. SOME TREMATODES IN SOUTH AFRICAN ANURA, AND THE
RELATIONSHIPS AND DISTRIBUTION OF THEIR HOSTS.
By C. S. GropBetaaR, M.A.,
Lecturer and Demonstrator in the Department of Zoology,
University of Stellenbosch, Stellenbosch, S.A.
(With nine Text-figures.)
I. On SoutH AFRICAN PARAMPHISTOMIDAE (Fisch.).
The occurrence of Paramphistomids (=Amphistomes) in sheep and
cattle is fairly general in South Africa wherever suitable conditions prevail
to ensure its life-history. The conditions required are identical with those
required for the life-cycle of the common liver-fluke, viz. swampy vleis
where there is a fairly permanent water-supply to ensure both the develop-
ment of the eggs and the existence of the intermediate host, which in the
case of P. calicophorum is the ubiquitous Lsrdora tropica Krauss.
These light, flesh-coloured parasites, from 8 to 10 mm. long, occur singly
or in groups covering a few square inches in the rumen of sheep and cattle. .
They attach themselves by the well-developed posterior sucker to the
mucous membrane of the rumen between the numerous short villi. The
eggs are shed in the stomach, and gradually find their way down the intestine,
and are deposited with the feces. When the feces with the eggs happen
to be dropped in swampy vleis, or in such places where they are immersed
under water for some time, development of the eggs proceeds at the advent
of the rainy season, and infection of the intermediate host, if present,
takes place. The sheep and cattle infect themselves by feeding on the grass
or pasture along the margins of pools containing infected snails, and on
which the cerceriae have encysted themselves.
In the south-western districts of the Cape Province infection of stock
takes place during October, November, and December. In the eastern
districts, viz. Molteno, Dordrecht, Barkley Hast, and wherever Param-
phistomids (not necessarily P. calicophorum), occur in Natal, Orange Free
State, and the Transvaal, infection takes place from April to June.
182 Transactions of the Royal Society of South Africa.
I had the opportunity of examining a large number of sheep infected
with Paramphistomum calicophorum and P. cotylophorum. In one case I
counted as many as 150 individuals in the rumen of an ox, and yet the
host seemed none the worse for it, both as regards vitality and condition.
Fortunately, therefore, no loss of stock infected with these parasites need
be feared.
Paramphistomum calicophorum (Fisch.) (11) (fig. 9) is the species
prevailing in the south-western districts of the Cape Province. I also have
specimens from Dordrecht, Burghersdorp, and Molteno. It probably
occurs all along the lower reaches of the Zumbergen and Drakensbergen
into Natal.
Fischoeder records it from Queensland, China, and the Cape Province
(Kapland).
Paramphistomum cotylophorum (Fisch.) I have found in sheep from
Dordrecht, Burghersdorp, and a few specimens were sent me from
Onderstepoort.
The Life-history of Paramphistomum calicophorum Fisch.
(See figs. 1-8.)
The commonest fresh-water snail in the Cape Peninsula, and in the dis-
tricts of Stellenbosch, Paarl, Tulbagh, and Ceres is the ubiquitous [sidora
(Physa) tropica (Krauss). The eggs of the previous season, deposited
Fig. 2.
under the leaves of the common water-lilies (A ponogeton) and round the
bases of stems of a dwarf variety of rush, hatch in July, in the middle of
the rainy season.
During the months of August to November I undertook the examina-
On South African Paramphistomidae (Fisch.). 183
tion of Isidora tropica in a restricted locality in what is locally known as
the Stellenbosch Flats, situated in the town commonage. From previous
observations carried out on the occurrence of rediae and cerceriae in fresh-
Fic. 3. Fia. 4.
water snails from that particular locality, I knew that the snails harboured
small black cerceriae in abundance. These cerceriae readily encyst on
the grass stems growing round and in the pools of water, and thus easily
reach the stomach of sheep and cattle that graze on the commonage.
Animals that had grazed on the commonage were found to be heavily infected
184 Transactions of the Royal Society of South Africa.
with Paramphistomids about three months after they had grazed on the
infected grass. It needed no particular ingenuity to regard the common
Isidora tropica as the intermediate host of the locally known Param-
phistomid. After a careful examination of the rediae and cerceriae in the
<0. ™ > A
oO oo
Ce me) \
AGP YOrg,
Tors ry i g~ \
LOG OS Ng SIONS one
intermediate host Isidora tropica, I carried out a feeding experiment.
Three young sheep and a young lamb were used for the purpose. The sheep
were obtained from a dry locality in the Karoo from a small flock I knew
to be free of Paramphistomids ; the lamb I had reared myself, to make sure
that it was free of infection from any source whatsoever. For three weeks,
at the rate of two to three feeds per week, they were fed on grass heavily
infected with encysted cerceriae. As a result of continued and careful
On South African Paramphistonidae (Fisch.). 185
observations, it was quite certain that the snails from which the cerceriae
had escaped to encyst on the grass used in my feeding experiment harboured
only these particular dark-coloured cerceriae. In the locality selected
there were no other species of fresh-water snails to be found; animal life
in the pools, as far as vertebrates are concerned, was confined to the develop-
ment of tadpoles that died off as soon as the pools began to dry up. The
cerceriae encyst very readily, and in the laboratory I caused a large number
to encyst on grass stems placed in a wide tube of water into which the
cerceriae of infected snails were removed. Such grass was also used as a
feed. I was therefore perfectly sure that my experimental animals were fed
only on the cysts of the cerceriae under observation.
A few months after the first feed I found on examination that my
experimental animals were heavily infected with Paramphistomids. In
some cases these flesh-coloured and cream-coloured parasites were found in
all four divisions of the stomach, and a large number had found their way
down to the small intestine. The death of one of the young sheep towards
the end of the second month was evidently due to a confined life and an
abnormal heavy infection which was revealed on a post-mortem examination.
The adult parasite I identified as Paramphistomum calicophorum
Transactions of the Royal Society of South Africa.
186
Fia. 7.
Fic. 8.
On South African Paramphistomidae (Fisch.). 187
(Fisch.) (11), and the intermediate host is the small, dark-brown fresh-water
snail Zsedora (Physa) tropica, Krauss.
Observations on the Early Development of the Eggs of
P. calicophorum Fisch.
I collected a number of eggs in a flat dish, filled with water about half
an inch deep. The eggs are light yellow, elliptical, tapering a little more
towards the opercular pole than to the opposite pole. Length -17 mm.,
greatest breadth -09 mm. The embryonic cells lie towards the opercular
pole, and are surrounded on all sides by yolk-cells.. The operculum is small.
Development proceeded under water kept at room temperature (16°-17° C.).
By segmentation the embryonic cells had on the eighth day extended to
the opposite pole; the yolk-cells began to break up and diminish in size.
A membrane of thin flattened cells had grown round the entire cell-mass,
and one or two large covering cells lay over the anterior extremity of the
188 Transactions of the Royal Society of South Africa.
embryonic cell-mass. By the fourteenth day the embryo had become
quite distinct : the segmentation cells had lost their outline, and the remains
of yolk-cells were scattered round the embryo, which now almost filled half |
of the egg-case and extended from pole to pole.
On the twentieth day the embryo, the future miracidium, began to show
sight movements of contraction and subsequent expansion. Evagination
and invagination of the future alimentary canal were distinctly seen, accom-
panied by the ingestion of food-products into the embryonic digestive
tract. The digestive tract could be clearly seen, on account of its lumen
being filled with coarse granules.
On the twenty-third day the embryo had so much increased in size
as to lie with its mouth right up against the operculum. The wriggling
movements, the flickering vibration of the cilia, indicated that it had now
merely to lift the operculum to emerge from the egg-case as a free-swimming
miracidium. As soon as the eggs were now exposed to sunlight, the mira-
cidium burst open the operculum and soon escaped to swim about in
a smooth, swift-gliding fashion. It is, of course, well known, from experi-
ments carried out by Leuckart, Thomas, Looss, and others, that sunlight is
necessary for the miracidium to finally break through the operculum, and
thence swim about freely in search of the intermediate host.
I now introduced a snail into the dish containing the miracidia. At
first they swam about aimlessly, but as soon as they happened to come to
about half an inch or so from the snail, they suddenly changed their course,
swam straight up to the snail, and began to attach themselves to the exposed
parts. Some soon let go their hold, swam away, again approached the snail,
and attached themselves to it. Others, meantime, crawled over it with the
definite object it appeared of entering somewhere. It was apparent that
sooner or later some would find their way through the pulmonary aperture
into the mantle cavity, encyst themselves round or in the kidney, or work
their way up the visceral mass and encyst themselves in the liver.
As Looss (15) had as early as 1896 completely worked out the life-
history of Paramphistomum cervi in Egypt, where Physa alexandrina
(Bourg.) and Physa micropleura (Bourg.) are the intermediate hosts, I
did not consider it worth while to actually infect snails for the purpose
of my investigation. He found (loc. cit., p. 186) that after a period of
fifteen days the sac-like sporocysts contain immature rediae, and about
fifteen days afterwards the first generation of rediae appear.
The Rediae and Cerceriae.
The rediae are colourless transparent organisms containing daughter-
rediae in various stages of development. The more developed daughter-
rediae are confined to the anterior region of the parent redia. The birth-
On South African Paramphistomidae (Fisch.). 189
aperture is situated a short distance behind the sucker in the adult rediae.
I counted as many as seventeen daughter-rediae in various stages of develop-
ment in a parent redia. In many there are less. In some rediae one
could determine immature from two to three cerceriae with their incipient
but clearly defined eye-spots and well-defined tails, together with daughter-
rediae; and, as one might expect, the immature cerceriae were confined
to the anterior region and the daughter-rediae to the posterior extremity.
It seems safe to conclude that the second generation of rediae give rise to
rediae only, and the third generation give rise to rediae and cerceriae, and
that the fourth and subsequent generations will behave as the third.
With regard to the development and generations of rediae and cerceriae,
Looss (loc. cit., p. 189) states in connection with the life-history of Param-
phistomum cervr :—
‘* Les produits de ces rédies commencent 4 se différencier de tres bonne
heure; ils prennent naissance et se développent de la méme maniére
que les germes des rédies de l’Amphistomum subclavatum. ... Mais,
tandis que chez celui-ci les germes des premicres rédies se transforment
de suite en cercaires, chez l’Amphistome conique ( x Paramphistomum
cervt) la seconde génération donne encore des rédies. Ces derniéres
quittent leur mére par un orifice d’accouchement, situé a une courte distance
en arriére de la ventouse ; elles s’établissent a cdté des rédies plus Agées,
s’accroissent et produisent une troisiéme génération qui peut étre encore
une fois des rédies. De cette manieére, il résulte finalement un assez grand
nombre de rédies offrant 4 peu prés le méme aspect et qui commencent dés
lors & produire des cercaires. Chez le ver qui nous occupe je n’ai pas des
germes de cercaires en compagnie avec des germes de nouvelles rédies dans
une rédie mére, comme cela se produit chez le Gastrodisque. Mais cela
peut étre purement accidentel, car je n’ai pu suivre, aisni je l’ai dit, que
pendant dense mois le cycle évolutief de ce ver.”
The mature rediae are -6 mm. to 1 mm. in length. The small anterior
sucker is circular, with a diameter of -05--0O7 mm. A daughter-redia just
before birth is -25 mm. long, and the fairly well-developed sucker -05 mm.
in diameter. The sac-like alimentary canal is well defined.
The cerceriae are born in a very immature state. They live for some
time in the liver of the intermediate host, and when mature and capable of
greater activity they leave the snail. In adult infected snails, as well as
uninfected specimens, the shells are worn off at the apex, and a part of the
liver is exposed. It is, therefore, certain that the adult cerceriae emerge
from the intermediate host through the decaying apex of the shell. They
may, of course, also leave the snail by the pulmonary aperture.
As soon as they escape they swim about actively in the water by the
rapid wriggling movements of the tail. The body is almost entirely opaque
190 Transactions of the Royal Society of South Africa.
or black, owing to pronounced superficial pigmentation. This pigment is
more pronounced in and round the eye-spots ; it then extends forwards and
backwards in heavy radiating streaks. In the young immature cerceriae
the pigment is confined to the eye-spots a short distance behind the anterior
sucker. The body of a mature cerceria is oval, -42 mm. long, and greatest
breadth -25mm. The tailis-5--6mm.long. The anterior sucker is small and
circular, with a diameter of -03 mm., and the larger posterior ventral sucker
has a diameter of -09 mm. The mouth leads into a slightly elongated
cesophagus which bifurcates at a point just behind the eye-spots to form
the two short limbs of the forked intestine. The tail springs from the
dorsal posterior margin of the body just round the exterior margin of the
posterior sucker.
The excretory system consists of two lateral trunks. Hach main trunk
starts anteriorly in the region of the eye-spots, runs downwards and out-
wards, and suddenly turns inwards. Here each trunk gives off a branch ;
the branches from each side meet to form a transverse trunk. The main
trunks on each side now run outwards and backwards and join just in front
of the posterior sucker. At the junction is a well-defined excretory pore,
that opens on the dorsal surface. After the two trunks have joined they
continue as a single tube down the tail bifurcating just before it reaches the
distal extremity to open on each side through two minute pores.
Cystogenous cells are scattered throughout the organism. These cells
aggregate to form chains around and along the course of the vascular trunks
when the cerceria becomes less active and begins to encyst itself.
The cerceriae readily encyst themselves on the sides of the glass tube in
which they may be collected, and on grass stems introduced into the tube.
The tail is now cast off, the anterior and posterior extremities are drawn in
under the body, and a small round black speck about -2 mm. in diameter is
formed. Encystation is accompanied by the expulsion of a granular mass
containing small dark rhabdite-like bodies. This granular mass hardens
when exposed to the action of sun and air. Before encystation takes place
the cerceriae crawl or wriggle up the grass stems for a short distance, and
encyst just above the water-level.
I have no hesitation in regarding the cerceriae described above as
identical with Cawston’s Cerceria frondosa (7), which he removed from
Isidora schakoi Jickeli, from Potchefstroom (Tol.). It is also the same
as Gilchrist (13) recorded.
Some Trematodes in South African Anura. UTS)
II. Some TREMATODES IN SouTH AFRICAN ANURA, AND THE
RELATIONSHIPS AND DISTRIBUTION OF THEIR Hosts.
It is a well-established fact that trematodes, as well as most parasitic
worms, occur in fauna groups. If we investigate the trematodes in a par-
ticular class of host in a defined zoo-geographical region, we will find that
closely allied or similar trematodes probably live in the same class of host
in a different zoo-geographical region.
During the last four years I devoted a good deal of time to the examina-
tion of frogs in search of trematodes. In view of the cosmopolitan occur-
rence of the Anura throughout the Union, the present paper deals only with
such trematodes as I have from time to time removed from frogs in the
neighbourhood of the Cape Peninsula. A search for trematodes in frogs
from the more inland districts, viz. Worcester, Middelburg (C.P.), Molteno,
and Dordrecht thus far yielded no results. The material, however, which I
have collected clearly bear out the truth of the above-mentioned statement.
Methods employed.—In looking for trematodes in frogs, I more or less
followed the methods suggested by Johnston (14) and Looss (19). After
inspecting the buccal cavity, the ventral body-wall was slit from vent to
chin, the digestive tract laid open, and the whole length of the alimentary
canal slit open, and the internal surface was examined with an eye-lens.
The trematodes found in the body-cavity, gall-bladder, urinary bladder,
lungs, etc., were immediately removed to glass dishes containing normal
saline. Worms intended for whole mounts were placed in a drop of normal
saline on a slide, and covered with a cover-glass or another slide according
to the size of the worm. To prevent the cover-glass or the slide being
washed away when the fixing fluid is added, it is necessary to put a small
weight on the cover-glass or covering slide. The addition of a weight at
the same time flattens out the object intended for a whole mount without
appreciably changing its length. In the case of larger and stronger worms
a good deal of pressure is often necessary to cause the worm to flatten out.
In the case of small worms a small glass tube about 8 mm. in diameter and
20 mm. long was used ; the pressure can then be regulated by the addition
of a little mercury poured into the bottle. For larger and stronger worms
small wooden cubes were placed on the upper slide and weights were then
placed on these wooden blocks. In view of the action of fixatives on the
metal weights, it is necessary to avoid such reactions by first using small
wooden or glass cubes, and increase the pressure by adding small metal
weights. The whole is now flooded with the fixing solution.
I obtained the best results by using boiling or a hot saturated solution
of corrosive sublimate in water with about 5 c.c. of 2 per cent. glacial acetic
acid to every 100 c.c. After a few seconds the weights are lifted off for a
192 Transactions of the Royal Society of South Africa.
moment, and one notices that the fluid has already taken effect by the
opacity of the tissues round the edges. The weights are replaced, and
finally removed after a period varying from five minutes to half an hour,
according to the bulk of the worm.
The worms are further left in the fixing fluid for fifteen minutes and
longer. They are subsequently washed in 40-50 per cent. alcohol, left for
a day or so in iodised 70 per cent. alcohol, and they are then ready for the
after treatment of staining and mounting.
I also obtained very good results by using 10 per cent. formalin as a
fixative for whole mounts.
The following trematodes, referred to their systematic position, have
been found :—
Order HETEROCOTYLEA Mont.
Family POLYSTOMIDAE Taschbg.
Sub-family PotystominaE Van Ben.
Genus Polystomum Zeder.
1. Polystomum integerromum (Rud.) Frol.?
In 1758 Roesel von Rosenhof (21) first figured and described this trema-
tode from the urinary bladder of the frog without apparently naming it.
In 1792 M. Braun (6) found it in the bladder of the green frog Rana esculenta
(L.), and described it as Planaria uniculata, n. sp.
Zeder (25) in the year 1800 first attempted a classification of the then
known trematodes. The various helminthologists had previously often
described and figured the same animal under different names. The genus
name Planaria, to cite one instance, was in turn used to denote entirely
different worms. Zeder founded the genus Polystoma, and referred the
above-mentioned Polystomum integerrimum to this genus as Polystoma ranae.
Rudolphi (22) was the first to describe and figure it under the name by
which we now know it, viz. Polystomum integerromum (he, however, wrote
it Polystoma integerrimum).
Von Baer and Van Beneden (1827), Stieda and Willemoes-Suhm all sub-
sequently contributed to its anatomy and development. It was, however,
left to Zeller (26), who first in 1872 and afterwards in 1876 gave a complete
description and fully worked out the life-history of this parasite.
Polystomum wintegerrimum is found in the urinary bladder of Xenopus
laevis from Stellenbosch. About 5 per cent. of these frogs, popularly
known as “ platanas,” harbour this parasite. I observed from one to
six in the urinary bladder. Curiously enough, I never came across it in a
large number of Rana fuscogula from Stellenbosch, Paarl, and Worcester.
Some Trematodes in South African Anura. 193
Order MALACOTYLEA Mont.
Family FASCIOLIDAE Rail.
f PuaciorcHInaE Lihe.
‘\. =LEPopERMATINAE Looss.
Genus Opisthioglyphe Looss (18).
Sub-family
2. Oprsthiogly phe endoloba Dujardin.
Frélich (10) included this parasite with Diplodiscus subclavatus under
the name Fasciola ranae. Rudolphi referred Frélich’s Fasciola ranae to his
own Distomum clavigerum, and in this was followed by a number of later
writers, e.g. Dujardin and Diesing. Dujardin found the real O. endoloba,
however, and, recognising it as distinct from D. clavigerum, described it
in 1845 as Distomum endolobum. It was subsequently also described by
Looss (17).
These small trematodes were found in the posterior region of the in-
testine, and in the rectum of Xenopus laevis from Stellenbosch. They
are small oval worms with a body-length of 2-255 mm. Anteriorly the
body tapers slightly more than posteriorly. The posterior extremity is
shightly emarginate. I only came across it once. In about a half-inch
length of the intestine, just before it opens into the rectum, I found as
many as fifteen individuals.
O. endoloba is the common European frog parasite in Rana temporaria,
occasionally in Bufo variabilis, calamitas, and vulgaris, and in Triton cristatus.
The genus renamed by Looss (loc. cat., Zool. Jahrb., p. 588) is closely related
to the genera Dolichosaccus (Johnston) and Brachysaccus (Johnston) found
in Australian frogs (14).
Sub-family SyNCOELINAE Looss.
Genus Halipegus Looss.
3. Halipegus ovocaudatus Vulpian.
Vulpian first records this trematode in 1860. It was subsequently
described by Creutzburg (8) and Looss (18). Sonsino (1893) also found it
in the stomach and in the anterior region of the intestine, whereas it is
usually found in the buccal cavity under the tongue.
From the buccal cavity of Rana fuscigula, Stellenbosch.
Family PARAMPHISTOMIDAE Fisch.
Sub-family CLADORCHINAE Fisch.
Genus Diplodiscus Diesing.
4. Diplodiscus subclavatus Goeze.
Frélich (10) included Opisthioglyphe endoloba with Dviplodiscus sub-
clavatus under the name Fasciola ranae. Previously, in 1787, Goeze (12)
15
194 Transactions of the Royal Society of South Africa.
described it under the name Planaria subclavata. Rudolphi (22) placed it
in his own genus Amphistoma. Diesing (9) established the genus Dviplo-
discus, and referred Amphistoma subclavata to Diplodiscus, calling it Diplo-
discus subclavatus. Later authors—Filippi, 1855; La Valette de St.
George, 1855: and Pagenstecker, 1857—devoted themselves to observa-
tions on its anatomy and life-history. Von Linstow described the repro-
ductive system fully, and observed the independent movements of the
vitellogenous granules. It was Looss (16), however, who, in 1892, published
a complete account of this well-known parasite and fully established its
life-history.
Judging from the size of the suckers, length of intestinal limbs and shape
of the testes, I found the South African specimens from the rectum of
Aenopus laevis (Stellenbosch, Paarl, and Kuilsrivier) identical with the
Kuropean representatives. The Australian representatives constitute the
species D. megalochrus Johnston and D. microchrus Johnston (14).
I wish here to record the discovery of two other trematodes found in
Aenopus laevis, which I have thus far been unable to refer to their natural
position among the Malacotylea. The following is a preliminary account.
Preliminary Account of a Distomid found in the Gall-bladder
of Xenopus laevis.
The presence of these worms in the gall-bladder of Xenopus laevis
imparts a pronounced yellowish-green colour to that organ. On opening up
the gall-bladder the worms were found shrouded in a dense mass of a partly
coagulated substance. They occur singly or in pairs, more often in pairs.
The body, about 7 mm. long, is divided into two distinct regions of more or
less equal length: an anterior narrow region ‘5 mm. wide, and a posterior,
flattened, oval region about 2mm. wide. Posteriorly it ends in a moderately
smoothcurve. Theanterior halfis mobile, carrying the anterior and posterior
suckers, of which the former has a diameter of -17 mm. and the latter of -1 mm.
It has a well-developed bulbous pharynx, short cesophagus, and intestinal
limbs which do not extend to the posterior extremity. The testes are
rounded, and placed symmetrically at the same level on each side of the
body just within the intestinal limbs in the posterior region of the body.
The compact rounded ovary is almost symmetrically placed in front of the
testes, slightly to the left of the middle line. The coils of the uterus, filled
with yellowish-brown eggs, are confined to the region between and behind
the testes, and do not extend over the intestinal limbs on each side. The
vitellaria are lateral, extending from the level of the posterior sucker to a
point a short distance in front of the termination of the intestinal limbs.
They form dendritic clusters arranged on each side external to the intestinal
Some Trematodes in South African Anura. 195
limbs, and only overlapping the limbs anteriorly. A vesicula seminalis is
present. The genital aperture is situated between the suckers, nearer to
the posterior sucker.
With respect to such external characters as the division of the body
into a narrow mobile anterior and a broad flattened posterior region, it
certainly belongs to the Gorgoderinae (cf. Spathidium folium v. Olfers),
but it differs from them in the presence of a well-developed pharynx,
short cesophagus, vitellaria confined to the sides, and a poorly developed
posterior sucker.
It differs from the Brachycoelinae in respect to pharynx, length of
intestinal limbs, and position of the vitellaria, but agrees with them in
respect to the position of ovary and testes and the coils of the uterus con-
fined to the region behind the testes. It certainly shows affinities with
both the Gorgoderinae and Brachycoelinae. I am of opinion that it is
more closely allied to the latter than to the former.
A Mstomd from the Body-cavity of Xenopus laevis belonging to
the genus Heterolope Looss (sp. inq.).
This trematode I found in the body-cavity of Xenopus laevis. On one
occasion I found it under the peritoneum on the outer surface of the
lungs. The worm measures 9 mm. from end to end, greatest breadth 2 mm.
It tapers more posteriorly than anteriorly. The suckers are well developed,
and approach each other. The ovary lies between the testes, which are
situated one behind the other in the middle line in the posterior region of
the body. Cirrus-sac, ductus ejaculatorius, and cirrus present; a vesicula
seminalis lies outside the cirrus-sac. The genital aperture lies just in
front of the anterior testes in the posterior region of the body, far
behind the posterior sucker. The coils of the uterus lie in front of
the testes.
So far I have not been able to find a frog-trematode recorded in which
the genital aperture lies far behind the posterior sucker in the posterior
region of the body. It undoubtedly belongs to the genus Heterolope
Looss, or a closely allied genus.
General Conclusions.—In considering the relationships and distribution
of the hosts of South African frog-trematodes, one can, it appears to me,
only arrive at such conclusions as Johnston (14) did with respect to the
Australian frog-trematodes. The following is a reproduction of his con-
clusions so modified as to include the South African representatives.
The occurrence of the trematodes recorded above from South African
frogs gives a striking illustration of the tendency of helminths to occur in
faunal groups. That is to say, that helminths found parasitic in any
196 Transactions of the Royal Society of South Africa.
particular class of host, in a defined zoo-geographical region, find their
nearest relatives not in that region in which they themselves occur, but in
the same class of host living in other zoo-geographical regions.
Five such faunal groups from Amphibia are now more or less well known,
viz. European, North American, Asiatic, Australian, and South African.
In each region we find, in hosts of this class, trematodes corresponding
or closely corresponding with representatives in the other regions. The
group in the European region, as the longest known and most extensively
investigated, shows the greatest number of genera; in point of numbers,
the American group, to which a good deal of attention has been given of
late years by Stafford and others, follows closely on the European. The
Australian, Asiatic, and South African groups show smaller numbers, partly,
perhaps, because less completely worked up. The facts at present seem to
indicate that in Asia, Australia, and South Africa the number of representa-
tives may be further, more or less extensively, added to by subsequent inves-
tigations. Besides the frogs, other classes of hosts, as mammals, birds, and
reptiles, show faunal groups of parasites with relations analogous to those
exhibited by the group from frogs, as the small collection of trematodes
I have from mammals thus far go to show, viz. Paramphistomidae and
Fasciolidae from sheep and cattle, for example. Leaving the latter in the
meantime out of account, and confining ourselves to the groups from frogs,
we find: Opisthioglyphe endoloba, occurring in the intestine of Huropean
and South African frogs, is represented in North America by Glypthelmius
quieta and in Australia by Dolichosaccus trypherus and Dolichosaccus
ischyrus, which also live in the proximal part of the intestine of their hosts.
The three European species of Pneumonoeces, found in the lungs, are
represented in America by no less than six species (P. longiplexus
Stafford, breviplerus Stafford, P. varioplexus Stafford, P. similiplexus
Stafford, P. medioplecus Stafford, and P. complexus Seeley), while they
are represented in Asia by Pneumonoeces capyristes Klein, in Australia
by P. australis. The only representative of Pneumonoeces I have thus
far come across was a specimen given me by my chief, Dr. EH. J, Goddard ;
it was removed from the lungs of Rana fuscigula. Unfortunately it was
damaged, and did not lend itself to specific determination.
The Gorgoderinae, represented in European frogs by the two genera
Gorgodera and Gorgoderina, comprising between them, according to Ssinitzin,
five separate species, all living in the bladder of frogs, are represented in
America by four species of Gorgoderina and one of Gorgodera. No species
of Gorgodera has yet been described from Asia, and thus far I have not
yet found any in South Africa, but the genus is represented by one species—
G. australiensis (Johnston)—from Australia. The European Brachycoelium
crassicolle R., found in the intestine, is represented in America by B. hospi-
Some Trematodes in South African Anura. Loe
tale; in Australia by three species of Mesocoelium, viz. Mesocoelawm mesem-
brinum (Johnston), Mesocoelium megaloon (Johnston), and Mesocoeliwm
oligoon (Johnston) ; in Asia by M. sociale; and the form I have prelimi-
narily described from the gall-bladder of Xenopus laevis may turn out to
be the South African representative of the Brachycoelinae.
The Pleurogenetinae, occurring in the anterior part of the intestine,
are represented in Europe by three genera (Prosotocus, Pleurogenes, and
Brandesia), comprising between them seven species; in America by
Loxogenes arcanum, in Asia by Pleurogenes gastroporus and P. sphaericus,
and in Australia by P. freycineti and P. solus. Halipegus ovocaudatus,
occurring in the buccal cavity of European frogs, is also represented in
South Africa; in North America represented by H. occidalis ; in South
America by H. dubius; and in Asia by H. longispina—all living in similar
situations ; but, up to the present, no representatives of this fluke have
been found in Australia.
Diplodiscus subclavatus in the rectum of European and South African
frogs is represented in America by D. temperatus, and in Australia by two
species of Diplodiscus. The European Polystomun integerrimum in the
bladder of frogs does not seem to be represented in America in frogs, but
three species (P. coronatum Leidy, P. hassall1 Goto, and P. oblongum
R. Wright) occur there in the bladders of Chelonians; in Australia this
heterocotylean genus is represented by Polystomum bulliensi Johnston, and
in South Africa by the Kuropean species. No representatives of Diplodiscus
nor Polystomum have yet been described from Asia. The American
Cephalogonimus americanus in the intestine of frogs may be represented
in the Old World by C. lenowr Poir. The genus Ganeo, described by Klein
from the Indian Rana hexadactyla, seems to stand alone, unrepresented in
any other region. Likewise the “species inquirenda,” belonging to the
genus Heterolope, from the body-cavity of Xenopus laevis, referred to above,
seems to be unrepresented in the frogs of other zoo-geographical regions.
It is a remarkable fact that, of the six species of flukes described from
frogs inhabiting Southern Asia, four of them appear to find their nearest
relatives in flukes from Australian frogs. Mesocoeliwm sociale Lithe is
certainly more closely related to the Australian species of Mesocoeloum than
to Brachycoelium crassicolle R., its Kuropean, or B. hospitale Stafford, its
American representative. Pnewmonoeces caprystis Klein has been shown
(Johnston, loc. cit., p. 325) to be more nearly related to P. australis than
to any other Huropean or American species of this genus, and the Asiatic
Pleurogenes gastroporus and P. sphaericus and the Australian P. freycinete
and P. solus have likewise been shown to be more nearly related to one
another than any of them are to the American or European Pleurogenetines.
The South African group, on the other hand, is certainly directly related to
198 Transactions of the Royal Society of South Africa.
the European frog-flukes, the Asiatic, as it were, standing midway between
the Australian and European. The American frog-flukes, many of which
have evolved into distinct genera, are not so nearly related in their structure
to the Kuropean as are the Asiatic. And in addition to this, the American
genera, generally speaking, contain more species than the same genera in
Asia, Australia, and South Africa, and this may be taken to indicate that
the American frogs, with their flukes, have been longer separated from the
parent-stock.
The great similarity of the five groups of flukes from frogs found in the
five regions mentioned, points to the fact that the flukes are a very old
group of animals, and existed in the ancestors of present-day frogs a very
long time ago, when their distribution was much less extensive than it is
to-day. The mutual relationships of these groups of trematodes support
the view that the Anura originated somewhere about the centre of the
Palearctic region, and migrated westwards, southwards, and south-west-
wards. They may have reached the western portion of the Boreal land-
mass, existing right across from Asia to North America, in early Tertiary
times ; or they may have made their way westwards in Pliocene times,
when a considerable migration of vertebrates westwards is known to have
taken place. The Australian forms must have found their way there before
the separation of the Australian continent from South-eastern Asia, a
separation which is generally supposed to have taken place somewhere
about late Cretaceous or Eocene times. The South African forms must
have found their way down here during late Pliocene times. The greater
diversity of the North American frog-trematodes would seem to indicate
that they have been longer separated from the parent-stock than the
Asiatic, Australian, and South African forms, so that the America-wards
migration probably took place in the earlier of the two periods mentioned.
In view of the probable land connection between Australia and South
America through the Antarctic, a connection which is supported by a good.
deal of biological evidence, it is unfortunate that practically nothing seems
to be known about the frog-trematodes of South America. There are only
two indirect references to such trematodes, viz. in Braun (2, p. 906) and
Klein. <A pretty close similarity has, however, been shown by Zschokke
to exist between some cestode-parasites of South America and the Australian
Marsupials.
The close similarity existing between the respective representatives in
the five groups of frog-trematodes in question here reminds us that the
trematodes, owing to the conditions under which their lives are passed,
have probably evolved much more slowly than their hosts, for the
Amphibian ancestors of our own present-day groups, at the time of
Some Trematodes in South African Anura. 199
their dispersal, must have been considerably different from their repre-
sentatives now living.
LITERATURE REFERRED TO.
Those papers marked with an asterisk (*) I have not been able to see, but have had to
depend for any knowledge of their contents on such works as (1), (2), (17), (18).
(1) Bennam.—‘ The Platyhelmia, Mesozoa, Nemertini,’”’ Lankester’s Treatise of
Zoology, pt. iv.
(2) Braun.—‘‘ Vermes,” in Bronn’s Classen u. Ordnungen des Thierreichs (1892-3).
(3) Braun.—“‘ Zur Verstaindigung tiber die Gultigkeit einiger Namen von Fascioliden-
Gattungen,” Zool. Anz., xxiv (1901).
4) Braun.—‘‘ Trematoden der Chelonier,” Mitt. a. d. Zool. Mus. in Berlin (1901).
) Braun and Litun.—Handbook of Practical Parasitology, English edition, 1910.
*(6) Braun, M.—Fortsetzung der Beytrage zur Kenntniss der Eingeweidewirmer (1792).
) Cawston, F. G.—‘‘ The Cerceriae of the Transvaal,’’ Parasitology, vol. xi, No. 1,
1918.
*(8) CREUTZBURG.—Untersuchungen iib. d. Bau u. d. Entwickelung d. Dist. Ovocau-
datum Vulpian, etc.
*(9) Drestnc.—‘‘ Monographie der Gattungen Amphistoma und Diplodiscus,’ Ann. d.
Wiener Mus. der Naturgeschichte, Wien, 1835.
*(10) FroLicH.—Beytrage zur Naturgeschichte der Eingeweidewiirmer (1792).
(11) FiscHorpER.—“‘ Die Paramphistomiden der Saugethiere,’? Zool. Anz., xxiv, Bd.
No. 646, 1901.
*(12) GornzE.—Versuch. einer Naturgeschichte der Eingeweidewiirmer thierischen Korper,
Leipzig, 1787.
(
(5
(6
(7
(13) GitcHrRist.—*“ The Life-history of Distoma lutewm, n. sp., with Notes on some
Cerceriae and Rediae found in South Africa,”’ Parasitology, vol. x, No. 3, 1918.
(14) Jounston, S. J.—‘‘ On some Trematode Parasites of Australian Frogs,’’ Proc. of
the Linn. Soc., N.S.W., vol. xxxvii, No. 146.
(15) Looss, A.—‘‘ Recherches sur la Faune Parasitaire de l’ Egypte,’’ Mémoires |’ Institut
Egyptien, tome iii, fascicule premier, Le Caire, 1896.
(16) Looss, A.—‘‘ Uber Amphistomum subclavatum Rud.,” Festschrift, Leuckart, 1892.
(17) Looss, A.—‘‘ Die Distomen unserer Fische u. Frésche,’”? Bibliotheca Zoologica,
Heft 16, 1894.
(18) Looss, A.—‘‘ Weitere Beitrige zur Kenntniss der Trematoden-Fauna Agyptens,
u.s.w.,” Zool. Jahrb. Syst., xii, pp. 521-784 (1899).
(19) Looss, A.—“ Zur Sammel- und Conservierungstechnik von Helminthen,” Zool. Anz.,
xxiv (1901), p. 302.
*(20) PAGENSTECKER.—Trematodenlarven u. Trematoden, Heidelberg, 1857..
*(21) RoESEL von RosENHoF.—Historia naturalis ranarum, etc. (1758).
*(22) RupDoLpHI.—Entozoorum sive vermium intestinalium historia naturalis (1809).
(23) StmEs and Hassatyt.—‘* An Inventory of ... the Fasciolidae,’? Archiv de
Parasitologie, i, 1898, p. 81.
*(24) Vutpr1an.—‘‘ Note sur un nouveau Distome de la grenouille,’’> Compte rendu, 1859.
*(25) ZEDER.—Erster Nachtrag zur Naturgeschichte der Eingeweidewirmer, etc., Leipzig
(1800).
(26 ) ZELLER.—“‘ Untersuchungen tiber die Entwicklung und den Bau des Polystomum
integerrimum,” Zeit. Wiss. Zool., xxii, 1872.
200 Transactions of the Royal Society of South Africa.
EXPLANATION OF FIGURES.
Figs. 1-4 illustrate the development of the egg up to the hatching of the free-swimming
miracidium emerging from the egg-shell.
J. =Intestine.
E.C.T. =External cellular envelope.
Gg. =Germ-cells and yolk-cells.
Ex. T. =Excretory tube.
Op. =Operculum.
N.S. =Cells of nervous system.
C. =Ciliated coat.
S. =Sucker.
G., =Developing mass of germ-cells to give rise to redia.
Fig. 5. A redia of the first generation.
Fig. 6. An immature redia of the second generation.
Fig. 7. An immature cerceria just after it has escaped from the redia.
Fig. 8. A mature free-swimming cerceria, Cerceria frondosa.
Fig. 9. Mature Paramphistomum calicophorum Fisch.
S., =anterior sucker.
S.. =posterior sucker.
Oe., =cesophagus.
Vs. =Terminal region of vas deferens
Ut. =uterus.
Todo Lestes.
Ov. =Ovary.
Vit. D. =Right vitelline duct from lateral vitelline gland.
Oot. =Ootype.
L.C. =Laurer’s canal.
G.P. =Genital aperture.
The figure represents a ventral view with posterior extremity slightly turned up to
show posterior sucker.
{ 201 3
STUDIES IN THE MORPHOLOGY OF SELAGINELLA
PUMILA, SPRING.
Part I.—THE VEGETATIVE ORGANS OF THE SPOROPHYTE.
By A. V. DuTHIE.
(With twenty-six Text-figures.)
With the exception of a doubtful record from Natal, this small annual
species of Selaginella is known only from the Cape Province. It has been
collected in most of the coastal districts from the Cape Peninsula to
Alexandria, and has recently been found as far inland as Tulbagh. As the
species is inconspicuous and might easily escape the notice of collectors, it
is probable that its distribution is more general than is usually supposed.
In the Stellenbosch district it occurs abundantly from July to October,
and is generally found on clayey soil associated with liverworts and mosses
and forming part of the annual flora of flats and hill slopes. It has also
been found growing luxuriantly in water-logged turf by the side of
trickling water.
So far as I am aware, the only recent work dealing with the species is
contained in a paper by Sykes and Stiles (15) which appeared in the Annals
of Botany in 1910.
The observations embodied in the present paper have extended over
several years, and have brought to light certain interesting features, some
of which do not appear to have been recorded for other species. The paper
deals chiefly with the external morphology and anatomy of the vegetative
organs of the sporophyte. A general discussion of the results will be
postponed until after the completion of the work.
Sclechtendal (10) figured two varieties of S. pumila, namely, pygmaeum
and bryoides, differing in size, the latter being the larger and more pro-
cumbent. Hieronymus (6) recognises two distinct species under the names
S. pumila, Sclecht., and S. bryoides (Kaulf.), Hieron., the former character-
ised by lanceolate, the latter by ovate, leaves. The characters upon which
the two species or varieties have been based are found, on the examination
of material from different parts of the Stellenbosch district, to overlap to
202 Transactions of the Royal Society of South Africa.
some extent. Specimens gathered in shaded situations on hill slopes are
delicate and often more or less procumbent, with ovate leaves ; while those
gathered on exposed flats are wiry and erect, with lanceolate leaves. Varia-
tions in the leaf form may occur in a single specimen, especially if the
environment be altered during growth (fig. 1). Two instances of lobed
leaves were noted (fig. 2); but these are obviously to be regarded as abnor-
malities. Individual plants vary greatly in size. Fruiting specimens of
1 cm. are not uncommon, while the tallest plants noted reached the height
of 19cm. By far the greater number are branched ; but small unbranched
specimens are occasionally met with.
An excellent figure of the leafy stem and cones of the flats type occurs
in Marloth’s Flora of South Africa, vol. i, fig. 60. 1.; while the more
Fig. 2.
delicate hillside type is figured by Sim (14). Neither of these drawings,
however, gives a correct impression of the root-system of the plant. As in
the majority of described species of Selaginella (6), three roots first arise at
the base of the hypocotyl in the neighbourhood of the spore, the middle
one being the main root of the sporeling (fig. 3). Hach of these roots may
branch laterally. The erect and slender hypocotyl, which may reach the
height of about 4 mm., bears the two oppositely placed cotyledons and the
plumule which is later continued into the leafy stem (fig. 4). Additional
erect or sub-erect stems with independent root-systems may arise laterally
from the upper part of the hypocotyl.* The first of these secondary stems
originates midway between the cotyledons either before or after the appear-
ance of its root-supply (fig. 5, a-d). A second stem with its root may
arise at the same level in the neighbourhood of the first, and this process
may be repeated until, in hardy specimens, a tufted habit results (fig. 6,
* It is probable that this apparent lateral shoot should be regarded as a delayed
branch of the first stem dichotomy, the erect axis representing the produced branch.
Studies in the Morphology of Selaginella pumula, Spring. 203
a,b). All such secondary axes arise on one side only of the hypocotyl, and
their roots grow down through the air for about 2 mm. before striking the
soil (fig. 5). Attempts to induce the formation of adventitious roots at
other points by means of cuttings or layerings have been unsuccessful.
With a little care the secondary stems with their roots can be removed,
leaving the primary stem with the cotyledons and trident root-system
intact (fig. 7). Both primary and secondary stems may remain unbranched
above or produce in one plane a varying number of short lateral branches
Fic. 4. Fia. 6.
each ending in a cone. One or more of the secondary stems may eventually
overtop the main stem. The secondary roots may also branch laterally.
The detailed anatomy of the tissues from which these secondary axes
and their roots originate, also the minute structure of the growing points of
root and stem, will be dealt with in a later paper.
The symmetry of erect and evenly illuminated shoots is radial. The
leaves are all of the same size, and are arranged in four vertical rows along
the surfaces of the four-sided stem. Though at first sight the phyllotaxy
appears to be somewhat irregular, it is found on examination to be a modi-
fication of the usual decussate arrangement. The two leaves belonging to
each pair do not arise at exactly the same level, and later may become more
or less widely separated as the result of the elongation of the intervening
204 Transactions of the Royal Society of South Africa.
stem tissue (fig. 8). This difference in level may sometimes be observed
in the cotyledons themselves. Fig. 9 shows the cotyledons and the suc-
ceeding leaf pair cleared in caustic potash. It will be noticed that not
only do the leaves of each pair differ in size, but also that the leaf-traces
of the two cotyledons are inserted on the stele of the axis at slightly different
levels. Decussate leaves are often present at the base of the plant, and may
appear at other points as well. In the specimen, a part of which is illus-
trated in fig. 10, the two opposite cotyledons were followed by four evenly
spaced leaves. After these came five approximately decussate leaf pairs,
then one whorl of three leaves, and finally eight leaves evenly spaced. Tall-
g
=
IG. 7% Fic. 8. Fic. 9. Fia. 10.
stemmed plants growing in shaded situations often show a considerable
interval between the two leaves of a “ pair,” combined with a certain
amount of stem torsion which serves to obscure the phyllotaxy.
While specimens from the flats are usually wiry and erect-stemmed and
show typical radial symmetry, the dorsiventral condition is occasionally
approximated to, especially in hardy plants where several basal stems form
a close, erect tuft. On such stems the leaves appear to be shifted slightly
so as to face towards the periphery of the fascicle (fig. 11). Small plants
collected in 1918 on sloping ground which had been ploughed over some
time before and from which all larger plant growth was absent, showed for
the most part erect stems with radial symmetry. Specimens were, however,
found in shaded hollows of the uneven soil which exhibited prostrate
secondary axes and a marked dorsiventrality. In one of these specimens
the leaves on the shaded side were distinctly larger than those on the
illuminated side of the stem (figs. 12, 13). This transition from the iso-
Studies in the Morphology of Selaginella pumila, Spring. 205
phyllous to the anisophyllous condition does not appear to have been
observed before in this species.
The root of S. pumila is entirely devoid of root hairs. The mycelium of
an endophytic fungus was found to be present in all the material examined,
ig
Fie. 13.
Ere. Li.
and is especially abundant in the large cells of the inner cortex (figs. 14,
15). It is probable that infection of the root cells takes place through the
epidermis, as hyphae are often observed in contact with the outer surface
of the root and occasionally in the cavities of the epidermal cells. The
hyphae appear to be restricted to the subterranean parts of the root system.
The nature of the fungus, its mode of entry into the root and the relation-
ship existing between the two plants, will be dealt with in a later paper.
Bruchmann has recorded the presence of an endophytic fungus together
with the absence of root hairs for S. sponulosa (1) and S. prewssiana (3),
while Janse (6) has reported fungus mycelium in the roots of certain Javanese
species of the genus. More recently Uphof (16) has described and figured
traces of fungus mycelium in S. rubella.
206 Transactions of the Royal Society of South Africa.
The epidermal cells of the root are somewhat papillate and, as age
advances, the outer walls may become markedly thickened. This thicken-
aw: Aa
nx XS
e» vO
Fic. 15.
ing of the walls may extend to the outer cortex, or (fig. 15) the epidermis
and cell layer immediately next to it may remain comparatively thin while
a zone of the middle cortex is strongly thickened. Chlorophyll is present
in the cortex of the aerial region of the young root (fig. 16).
It is worthy
of note that in all essential features this aerial part of the root is similar
in structure to the subterranean part.
An interesting feature of the root of S. pumila which does not appear
to have been recorded for any other species of the genus is the presence of
Studies wn the Morphology of Selaginella pumila, Spring. 207
a well-marked air cavity in the inner cortex lying on the phloem side of the
stele (figs. 14, 15, 16). This intercellular space arises by the separation
and breaking down of some of the inner cortical cells, the remains of which
may often be seen still adhering to the endodermis. The resemblance of
a transverse section through an old root of S. pumila, with its enlarged air
cavity and eccentric stele, to that of Lsoetes (12) or Stigmarza (11) is very
striking (fig. 15). The resemblance to the latter was found to be still more
marked in hardy, tufted plants preserved in formalin alcohol. Here the
cavity in the basal region of old roots appeared to have extended right
round the endodermis, isolating the central stele completely.
The endodermis of the root is well defined. The cells when young show
the usual radial cuticularisation (figs. 14, 16), but later the entire walls
Hre: 7. Fig. 18.
become strongly thickened (fig. 15). As in other investigated species of
Selaginella (5, 16), the root is monarch. In small roots the vascular
tissue 1s poorly developed, the xylem consisting of a small group of from
twe to five tracheides with the phloem in contact with the metaxylem. In
the largest roots examined as many as twelve tracheides were observed,
most of them of small size. These small tracheides are sometimes arranged
in two groups separated by the larger tracheides. A single layer of peri-
cycle cells appears to be present which may be interrupted by one or more
of the tracheides abutting directly on the endodermis.
The stem of S. pumila is typically four-sided, with the leaves inserted
along the sides (figs. 17, 18). The cells of the epidermal layer are covered
by a cuticle and, in young material, the walls are either uniformly lignified
or the lignification is confined to the surface walls. In old stems the ligni-
fication may extend through the outer cortex, resulting in the formation of
a distinct hypodermis. Chlorophyll is specially abundant in the thin-walled
cells of the inner cortical layers. Intercellular spaces are present between
the cells. The trabeculae which bridge the air space between the inner
208 Transactions of the Royal Society of South Africa.
cortex and pericycle are of a simple type, each consisting of a much elongated
endodermal cell showing the usual cuticular band (figs. 17, 18). The single
stele is placed somewhat obliquely in the middle of the air space. The one-
layered pericycle is usually separated from the xylem by one or more layers
of thin-walled phloem cells. Occasionally a xylem tracheide is found to
abut directly upon the endodermis. The stele is usually somewhat elliptical
in form, with two protoxylem groups.
Stomata are confined to the margin (fig. 19, a) and the aligular surface
of the leaf. They occur along the vein, often extending beyond its termina-
tion, and may appear also on the two wings. With the exception of the
Fig. 19.
cotyledons, a conspicuous leaf-base is present below the insertion of the
ligule, and here a definite group of stomata is always found (fig. 19, 6). Up
to twenty stomata have been counted in this region.
The guard cells are placed parallel to the leaf-margin (fig. 19, c). Only
one instance of a transverse position was noted (fig. 19, d). As many as
seven epidermal cells may abut upon a single stoma. ‘The number of
stomata present may vary within wide limits. When very numerous,
adjacent stomata such as those shown in fig. 19, e, may occur.
As mentioned above, the leaf-base, except in the cotyledons, is well
developed. It contains a mass of aerenchyma roofed over by an epidermis
which is perforated by numerous stomata (figs. 19, b, 20, 21). Sykes and
Stiles (15) describe and figure a similar air cavity in the projecting base of
the sporophyll of certain species of Selaginella, and compare it with the
mucilage cavity of Lycopodium and the parichnos of fossil genera. It is
Studies in the Morphology of Selaginella pumila, Spring. 209
also somewhat suggestive of the aerenchyma in the leaf-base of Miadesmia
(11). This feature does not appear to have been noted before in the
vegetative leaf.
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2x0
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Fig. 20.
The epidermal cells of both surfaces are usually elongated, with sinuous
The mesophyll of the leaf-blade shows no differentia-
lateral walls (fig. 19).
It extends right to
tion into palisade and spongy parenchyma (fig. 22).
the leaf-margin.
The vascular bundle in the cotyledons and lower leaves of the axis is
simple, and consists of from four to five rows of delicate spiral and annular
tracheides. It does not extend to the apex of the leaf, and, before dying
16
210 Transactions of the Royal Society of South Africa.
out, it expands slightly owing to the development of several short, flanking
tracheides. The next leaves in order of development are found to possess
a more complex vascular tissue. The protoxylem of the vein is accompanied
by two more or less interrupted groups of spiral and reticulate transfusion *
wisersrs7s)
yy TT yet
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ave.
By
van
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TAH IN]
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Fig. 23: Fic. 24.
tracheides, which may appear at any point between base and apex. All the
upper leaves of the plant, including the sporophylls, show two well-developed
bands of transfusion tracheides to right and left of the protoxylem (figs. 22,
23, 24). These bands of tracheides do not accompany the leaf-trace into
the stem. There is evidently a connection between the amount of trans-
fusion tissue and the number of stomata on the leaf-blade. The only other
Fia. 25.
species of Selaginella in which a similar triple mid-rib has been noted is
S. laevigata, Bak., var. Lyallii, Spr. (5). Bruchmann in 1909 published an
account of the vegetative organs of this species (2), but I have not had the
opportunity of consulting his paper, nor yet of obtaining material of the
species for comparison with S. pumilu.
The shape of the ligule was found to vary considerably, as shown in
* The use of this term implies no expression of opinion as to the phylogeny of the
tissue in question.
Studies in the Morphology of Selaginella pumila, Spring. 211
fig. 25. The base is surrounded by a well-developed group of reticulate
tracheides which is usually continuous with the transfusion elements already
described (figs. 24, 26).
BIBLIOGRAPHY.
*(1) BrucumMann, H.—Untersuchungen iiber Selaginella spinulosa, A. Br., Gotha, 1897.
*(2) BrucuMANN, “ Von den Vegetationsorganen der Selaginella Lyallit,’? Flora, 1909.
*(3) BrucHMann, “‘ Selaginella preissiana,’’ Flora, 1910.
(Review of this paper in Bot. Gaz., 49, 399.)
(4) GorpeL, K.—Organography of Plants.
(5) Harvey-Greson, R. J.—“ Contributions towards a Knowledge of the Anatomy of
the Genus Selaginella, Spr.”
Part 1. ‘‘ The Stem,” Ann. Bot., 8, 1894.
Part 2. “‘ The Ligule,’’ Ann. Bot., 10, 1896.
Part 3. ‘“‘ The Leaf,’ Ann. Bot., 11, 1897.
Part 4. ‘‘ The Root,’ Ann. Bot., 16, 1902.
(6) Hizronymus.—Naturliche Pflanzen Familien., 1902.
(7) Hitt, T. G.—“‘ On the Presence of Parichnos in Recent Plants,’? Ann. Bot., 20.
(8) Lotsy, J. P.—Vortrage tiber Botanische Stammesgeschichte.
(9) Martotu.—Flora of South Africa, Vol. I.
10) ScLtEcHTENDAL.—Adumbriationes Plantarum, 1825.
11) Scort, D. H.—Studies in Fossil Botany, Part 1.
12) Scorr, D. H., and T. G. Hiri.—*‘ The Structure of Jsoetes Hystrix,’’ Ann. Bot., 14.
13) S—ewarp, A. C.—Fossil Plants, Vol. II.
14) Sim, T. R.—Ferns of South Africa, 1915.
15) Syxzs, M. G., and W. Srites.—“‘ The Cones of the Genus Selaginella,’ Ann. Bot.,
1910.
(16) Urnor, J. C. Tu.—“‘ Contributions towards a Knowledge of the Anatomy of the
Genus Selaginella : The Root,’’ Ann. Bot., 34, 1920.
EXPLANATION OF FIGURES.
Fig. 1. Variations in leaf form ; from one plant.
Fig. 2. Abnormal leaf forms.
Fig. 3. Development of first roots at base of hypocotyl of sporeling.
Fig. 4. Plant with unbranched stem and three roots at base of hypocotyl.
Fig. 5, a-d. Development of first secondary axis with its root, midway between bases
of cotyledons.
Fig. 6. Plant with five secondary axes developed at base. a. Hypocotyl shown.
b. Reverse of fig. a.
Fig. 7, 6. Group of secondary axes removed from primary axis a.
Fig. 8. Part of radially symmetric stem with four leaf rows.
Fig. 9. Cotyledons and succeeding leaf-pair cleared in caustic potash.
Fig. 10. Part of stem with three leaf-pairs, one whorl of three leaves, and two alternate
leaves.
* T have had no opportunity of consulting papers (1) to (3).
212 Transactions of the Royal Society of South Africa.
Fig. 11. Part of stem of unequally illuminated plant showing approach to dorsiventral
condition.
Fig. 12. Unequally illuminated plant with two prostrate secondary axes showing
dorsiventrality and anisophylly.
Fig. 13. Secondary axis of 12, on a larger scale, showing difference in size of dorsal
and ventral leaves.
Fig. 14. Transverse section of young root showing endophytic fungus and cortical
cavity.
Fig. 15. Transverse section of old root.
Fig. 16. Transverse section through aerial part of young root of secondary axis.
Fig. 17. Part of transverse section of young stem.
Fig. 18. Section of older stem with base of leaf and ligule.
Fig. 19. a. Part of leaf-margin with stomata. 6. Part of epidermis of leaf-base with
stomata abutting upon thickened epidermis of stem destitute of stomata. c. Stoma with
neighbouring epidermal cells elongated in direction of leaf-margin. d. Normally placed
stoma and stoma with guard cells transverse to long axis of leaf.
Fig. 20. Vertical section through leaf-base showing aerenchyma, stomata, and
tracheides at base of ligule. The leaf-trace is cut obliquely.
Fig. 21. Horizontal section through superficial aerenchyma of leaf-base.
Fig. 22. Transverse section across leaf-blade.
Fig. 23. Outline of leaf showing triple vein, position of ligule, and of stomatal area
of leaf-base.
Fig. 24. Part of leaf cleared in caustic potash showing the protoxylem and two parallel
bands of transfusion tracheides continuous with reticulate tracheides at base of ligule.
Fig. 25. Variations in shape of ligule. The glossopodial cells are not shown.
Fig. 26. Transverse section through base of ligule showing tracheal sheath.
oom
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(2130)
SOME OBSERVATIONS ON THE EFFECTS OF A BUSH
FIRE ON THE VEGETATION OF SIGNAL HILL.
By Marecaret R. MicHeE11,
Lecturer in Botany, University of Cape Town.
(With Plates X to XII and one Text-figure.)
The practice of burning the veld in South Africa is probably an ancient
one. According to Dr. Marloth (3), the early Portuguese navigators saw
bush fires while sailing along the coast of the Cape Province, thus indicating
that the custom was not one introduced by the early white settlers.
At the present time there is no exact information as to the effects that
this periodic burning has on the vegetation, though certain experiments
in connection with this problem are being conducted at Pretoria by Dr. EH. P.
Phillips of the Division of Botany (5, 6). In South Africa, however, the
problem is not a uniform one, as the types of vegetation to be found in
different parts of the Union show fundamental differences. Thus the area
which Dr. Phillips has under observation is typical grassland, such as covers
large tracts of country in the Transvaal and Orange Free State, while the
area with which the present paper is concerned is normally covered with
sclerophyllous bush, the characteristic vegetation of the south-western
region. In regions of grassland, as a rule, burning is carried out system-
atically, year by year, whereas in the south-western districts this is not the
case, fires occurring at irregular intervals. These latter fires may be due to
natural causes, but more often they are started deliberately either by the
farmer who wants young shoots on which to graze his cattle, or by the poorer
members of the population who, after some time has elapsed, collect the
dead branches for firewood. It follows from the nature of the vegetation
that in grassland the fires are more readily controlled than in the bush-
covered country of the south-west.
Signal Hill, or Lion Mountain, on a portion of whose slopes the following
investigation was carried out, separates Cape Town from the sea on its
western side. As will be seen from the accompanying map (fig. 1), the hill
runs in a north-north-easterly direction, the highest point (about 2200 feet),
Ai
Zs
214
Transactions of the Royal Society of South Africa.
TABLE MOUNTAIN SANDSTONE .
x
Ww
a
<x
—
]2
<
2
aQ
UJ
TGel
= MALMESBURY SLATES ,
= RA'IGED BEACH,
ras a
0 6OO U)
04° 9,0
52 O0°0° 0
Adapted from the
Fic. 1.—Geological map of the northern part of the Cape Peninsula, showing Signal Hill and the burnt area.
Geological map of the Cape Division, 1899. Scale 144 inch to a mile.
Effects of a Bush Fire on the Vegetation of Signal Hull. 215
known as Lion’s Head, being situated at the southern end of the hill.
The rest of the hill consists of a ridge about a mile and a half long and of
considerably lower altitude than the head, the average height being about
1100 feet.
The soil of that part of Signal Hill with which this paper is directly
concerned is clay, derived from the Malmesbury slates, of which the northern
end of the hill is composed. The burnt area is by no means uniform in
character. Several valleys (fig. 2) of more or less equal depth cut
up the hillside, and these have a vegetation which in its constituents
shows certain marked differences from that of the ridges in between
the valleys.
The bush fire, the effects of which are recorded in this paper, broke out
on the 5th February 1919, and owing to a strong south-east wind it was
not extinguished till late on the following day. The approximate extent
of the burnt area is shown in fig. 1. Owing to the somewhat sheltered
position of the water-course in each valley, the vegetation there was not
badly burnt, while elsewhere the fire killed the aerial parts of every plant
with the exception of a few isolated pines and one eucalyptus tree. These
trees were the only ones on the burnt slope, and they were odd specimens
which had invaded the area from neighbouring plantations.
At one spot at the southern end of the area the ground is fenced off,
and this protected area was valuable in affording a means of judging the
effects of cattle and man on the regenerative processes after the fire.
In our * investigations valley A (fig. 2) and its immediate slopes
provided us with a suitably restricted field for more detailed observations.
The other parts of the burnt area were visited from time to time, but an
intensive study was made of valley A alone.
Detailed observations on the vegetation of this part of Signal Hill were
not made prior to the bush fire, and consequently it was necessary to rely
upon the evidence of charred bushes, and the character of a few small
patches which had escaped the fire, in estimating the original composition of
the flora. The plant which dominated these slopes as seen from the town
was Rhus lucida, bushes of which were dotted at frequent intervals over this
part of the hill. The green colour of these bushes was in marked contrast
with the prevailing grey-green colour of the smaller shrubs growing in
between the Rhus bushes.
The first visit to the burnt area was made eighteen days after the fire,
and during that interval there had been some rainy days interspersed with
fine, hot days.
* During the year 1919 this work was carried out in conjunction with three students—
Miss 8. Garabedian, Miss E. G. Torrance, and Miss W. Wilson. The investigation formed
part of the work of the M.A. course in botany for that year.
216
Transactions of the Royal Society of South Africa.
The following shrubs were noted on a small unburnt area at the foot of
valley A :—
Rhus lucida (a).
Hlytropappus Rhinocerotis (a).
Cliffortia ruscifolia (f).
Passerina sp. (f).
Athanasia trifurcata (f).
Leyssera gnaphaloides (f).
Asparagus capensis (f).
Psoralea hirta (f).
Noltea africana (l, f).
Rhus glauca (0).
Rhus angustifolia (0).
Passerina filiformis (0).
Anthospermum ciliare (0).
Borbonia cordata (o).
Solanum sodomaeum (0).
Berkheya rigida (0).
Aster fruticosa (0).
Cliffortia polygonifolia (0).
Kuclea racemosa (0).
Gymnosporia buxifolia (0). In water-course only.
Leonotis Leonurus (0).
Conyza ivifolia (r).
99 99
29
In addition the following undershrubs or herbs were found :—
Cynedon Dactylon (I, a).
* Lobelia Erinus (I, a).
Selago adpressa (I, f).
Chironia baccata, (o).
Pelargonium grossularioides (c).
Rumex acetosella (0).
* Krodium malachoides (0).
Andropogon hirtus (0).
Briza maxima (0).
Helichrysum crispum (0).
Helichrysum cymosum (0).
Scabiosa columbaria (o).
Oenanthe filiformis (0).
+ The usual abbreviations have been used. a=abundant, f =frequent, o =occasional,
r=rare, | =locally.
Effects of a Bush Fire on the Vegetation of Signal Hull. 217
Gorteria personata (0).
Pellaea auriculata (0).
* Lobelia lutea (zr).
(* indicates that the plants were in flower.)
In one spot an outcrop of slate was seen, and this was covered with
crustaceous lichens. Associated with this rock were Mesembryanthemum
asperum and Crassula muricata, both succulents, and it may be noted here
that apart from these rock exposures succulents were rare.
The outstanding feature of this first visit was the vigorous growth of
aerial shoots in Asparagus capensis and Andropogon hirtus. Both these
species had been burnt down to the ground; but in spite of the fact that less
than three weeks had elapsed since the fire, the whole hillside was dotted
with these plants, their green colour being in vivid contrast with the pre-
vailing blackness of the scene. In the case of Asparagus capensis many
bushes were a foot and a half high. Both these species were more abundant
on the southern slopes of each valley than elsewhere, and it was noted later
that these slopes which are more exposed to sunlight than the northern
slopes bear a more open type of vegetation. Another plant which was
growing actively on the burnt area was Hibiscus aethiopicus, but owing to
its small size it did not strike a conspicuous note, as was the case with the
two other species.
The next visit was paid about a month later, on the 20th March. The
condition of the area was little changed. One plant of Asparagus capensis
was in full flower, and it is of interest to note that the usual flowering period
of this species (May to June) had been anticipated. Hibiscus aethiopicus
was in flower all over the area. Besides these, Gladiolus brevifolius and
Haemanthus cocconeus were flowering, the latter being very conspicuous.
In the following years (1920 and 1921) flowers of this plant were rarely found,
and it seems safe to conclude that the fire must have induced the burst of
activity in 1919.
All over the hillside young shoots were being given off in profusion
from the bases of the charred stems in the various species of Rhus which
characterise these slopes. The following species were noted :—
Rhus lucida (a).
R. angustifolia (I, f).
R. glauca (0).
R. tomentosa (0).
R. mucronata (0).
R. rosmarinifolia (r).
Other plants here and there were seen to be behaving in a similar fashion,
218 Transactions of the Royal Socrety of South Africa.
but the phenomenon was not as general in these as amongst the species of
Rhus. The following list of such plants was made :—
Montinia acris (I, f).
Zantedeschia aethiopica (0).
Sonchus Kcklonianus (0).
Cliffortia polygonifolia (0).
Salvia africana (0).
Anthospermum ciliare (0).
Athanasia trifurcata (0).
Mohria caffrorum (0).
Leonotis Leonurus (r).
Bobartia spathacea (r).
Peucedanum galbanum (2).
Gymnosporia buaifolia was giving off new shoots, but as the plants were
confined to the water-courses they had been somewhat protected and the
aerial shoots had not been killed. A common plant in the water-course
at the foot of valley B was Clutyia pulchella, and although the old stems
appeared badly charred, they were giving off new shoots over the whole
surface. This was an unusual method of regeneration, the only other
plant behaving in a similar way being a young Kucalyptus tree which was
growing on the slopes above valley C.
A week later another visit was paid, and the only change noted was that
certain plants were now in flower, viz. Curculigo plicata, Oenanthe filr-
formis, Andropogon hirtus, and Ozalis lida. The two first-mentioned
species belong to the same class of plant as Haemanthus coccineus, the flowers
in all cases appearing some considerable time before the leaves.
In the Cape Peninsula there are two marked periods of flowering activity.
The first begins about the end of March and attains its maximum with the
first winter rains. Numerous species of Oxalis are characteristic plants of
this period. The second and major period is in the spring, towards the end
of the rainy season. These two flowering periods are well marked on the
area under discussion, and it is noteworthy that in general there was no
obvious forestalling of the periods in 1919. However, though the times of
flowering remained as in normal years, the same cannot be said of the quan-
tity and quality of the flowers produced. In both these respects the flowers
on the burnt area were far above the average, and this in a year when
elsewhere the flowers were not particularly fine.
During April the following plants came into flower :—
Oxalis tomentosa (a). On the lower slopes only.
Oxalis variabilis (a). Slightly later than O. tomentosa.
Effects of a Bush Fire on the Vegetation of Signal Hull. 219
Oxalis hirta (f).
Oxalis sericea (0).
Melica racemosa (0).
Polygonum atraphaxoides (0).
All the species of Oxalis which were found on the burnt area have
deeply situated bulbs, and several new bulbs are formed each year, vegetative
reproduction seeming to be the rule. As a result, the ground is covered
with Oxalis plants and the whole hillside presents an extremely gay appear-
ance. In 1919 the brilliant display of Oxalis was noteworthy.
In April the following additions were made to the March list of plants
which were producing new shoots from the uninjured parts below :—
Adiantopsis capensis (a). Under bushes and on northern
slopes of valleys.
Pellaea auriculata (f). 2 my af
Cliffortia ruscifolia (0).
Royena glabra (0).
Bulbine favosa (0). Valley only.
Kiggelaria africana (r). ,, -
Lycium afrum (r).
It was noted at this time that certain plants showed no signs of recovery
after the fire, notably EHlytropappus Rhinocerotis, and Protea mellifera,
which was represented by a single specimen.
The winter rains began to fall in May, and on the next visit on the 13th
June many changes were noted.
In the winter the northern side of each valley gets very little sun, and the
differences in vegetation between the two sides of each valley were very
marked. Valley A bore evidences of a great rush of water during the rains
which had fallen shortly before this visit, and the stream bed was character-
ised by a large number of young shoots of Ranunculus. Later in the year
two species were identified, R. pubescens and R. muricatus. On the banks
at the side of the water-course and also on the northern slopes of the valley
an abundance of small ferns, mosses, liverworts, and lichens were noted.
The most abundant fern was Adiantopsis capensis, which here and there
formed close sward-like patches. Pellaea auriculata and Mohria caffrorum
were plentiful. The following genera of liverworts were noted: Fossom-
bronia, Fimbriaria, Lunularia, and Riccia. The mosses and lichens were not
identified. Thelatter were confined to the exposed rock surfaces and branches
of various shrubs. The following herbaceous plants, all of which possess
underground organs of perennation, were noted on the northern slope :—
* Crassula septas (a).
* Rumex cordatus (f).
220 Transactions of the Royal Society of South Africa.
Scabiosa columbaria (f).
Cyphia Phyteuma (f).
* Oxalis variabilis (f).
* Oxalis glabra (f).
* Oxalis lanata (f).
Oxalis tomentosa (f). Flowers dead.
* Oxalis cernua (0).
* Oxalis compressa (0).
* Oxalis sericea (0).
* Curculigo plicata (0).
Rumex sp. (0).
* Crassula saxifraga (0).
These plants covered the ground, and gave an air of luxuriant growth
which was entirely lacking in the case of the southern slope. On the latter
the cryptogamic element was poorly represented, and Crassula septas,
which owing to its beautiful clusters of white flowers was most conspicuous
on the northern side, was entirely absent. Rumex cordata and Scabiosa
columbaria were also lacking on this side. The various species of Oxalis,
however, were well represented, and in addition Curculigo plicata, which was
at this time in leaf, though in many cases the flowers were still persisting.
Asparagus capensis was flowering freely.
On both slopes of the valley, and also on the ridge between valleys A and
B, the leaves of a large number of Monocotyledons were seen.
On the ridge seedlings of Borbonia cordata were common, and were
localised in a number of “ schools.”” This was the first example of a plant
coming up from seed after the fire. The normal fruiting period of this
species coincided with the time of the fire, and it seems safe to conclude
that the seeds, which are fairly large and heavy and thus not wind distri-
_ buted, are able to withstand a high degree of heat.
Several plants of Antholyza revoluta were seen in flower on the ridge.
Plants of Arctopus echinatus were also fairly common.
Valley B was visited on this occasion, and was found to agree fairly
closely with A, both in the composition of the flora and the distribution of
species.
The whole area, with the exception of the portion enclosed by the fence
already mentioned, was used as a grazing-ground for cattle. The result of
this was twofold. Certain plants, notably Andropogon hirtus, were almost
entirely eaten down. Inside the fence, on the other hand, the tall inflor-
escences of this grass were quite a feature of the vegetation (fig. 3). A
second result of the cattle grazing was the formation of a number of tracks
all over the hillside and the consequent loosening of soil. This was very
Effects of a Bush Fire on the Vegetation of Signal Hull. 221
conspicuous on the ridge at the northern boundary of the burnt area.
Here several new water-courses were formed asa result of the washing
away of the loosened soil.
Man was also instrumental in loosening the soil. Wood-gatherers were
constantly seen, as is always the case after a bush fire, and the tracks caused
by dragging large bundles of burnt brushwood down the hill were very
noticeable.
The outstanding feature of the next period of activity, which began
towards the end of August, was the predominance of Monocotyledons
amongst the herbaceous plants in flower. The following were noted :—
MoNOCOTYLEDONS.
Water-course of valley A.
Moraea tristis (f).
Moraea papilionacea (0).
Valley slopes and ridge.
Sparaxis grandiflora (a).
Geissorhiza secunda (a).
Geissorhiza sp. (a).
Pterygodium catholicum (a).
Homeria collina (f).
Moraea tripetala (f).
Lachenalia orchioides (f). Not in full flower.
Pterygodium alatum (f).
Disperis villosa (1, f).
Hesperantha sp. (0).
Babiana plicata (o). End of flowering period.
Satyrium bicorne (0).
Hypoxis serrata (0).
Romulea chloroleuca (0).
Lachenalia unifolia (r).
Gladiolus gracilis (r).
Disperis circumflexa (r).
Rudge only.
Romulea hirsuta (a).
Baeometra collumellaris (f).
Galaxia ovata (0).
Ornithogalum tenellum (0).
Aristea cyanea (0).
Galaxia graminea (r).
222 Transactions of the Royal Society of South Africa.
Northern slopes of valley A.
Hypoxis stellata (f).
Southern slopes of valley A.
Babiana stricta (f).
DICOTYLEDONS.
Valley slopes and ridge.
Dimorphotheca pluvialis (f).
Drosera cistiflora (f).
Diascia sp. (f).
Cenia turbinata (0).
Oxalis cernua (0).
Tripteris clandestina (0).
Zantedeschia aethiopica (0).
Oxalis variabilis (0).
Anagallis arvensis (0).
‘Nemesia barbata (o).
Nemesia parviflora (0).
Zaluzianskya divaricata (0).
Oxalis sericea (0).
Oxalis compressa (0).
Silene Burchellii (0).
Cerastium capense (0).
Heliophila diffusa (0).
Adenogramma galioides (0).
Cyphia volubilis (r).
Microloma lineare (r).
Northern slopes of valley A.
Scabiosa columbaria (f).
Southern slopes of valley A.
Ursinia anthemoides (f). This plant was also found in-small
numbers on other parts of the area,
Gazania pinnata (0).
With regard to the vertical range of these plants, the majority gradually
diminished in numbers towards the top of the area. There were a few
exceptions to this general rule. Hypowxis serrata was present on the higher
slopes only. Ozalis cernua, which was present on the lower slopes
of the area, was replaced towards the top by the closely related species
Oxalis sericea. The orchids, with the exception of Pterygodium catholicum,
were all confined to the lower slopes.
Effects of a Bush Fire on the Vegetation of Signal Hill. 223
Towards the end of October a further visit was paid, and most of the
plants that had been noted in September were found to have finished
flowering. Other plants had taken their place, the most conspicuous of
these being Moraea pavoma var. lutea. This plant was abundant on the
lower slopes of the burnt area, and the tall stems tipped with comparatively
small yellow flowers provided the dominant note in the vegetation. On a
more detailed survey of the area it was found that a large number of grasses
were in flower, and these together with Moraea pavonia may be said
to characterise the late spring period. The following list of grasses
was made :— :
Vulpia bromoides (a).
Lasiochloa ciliaris (a)
Aira caryophyllea (0)
Briza major (a).
Melica racemosa (f).
Khrharta calycina (f).
Festuca scabra (f).
Andropogon hirtus (f).
Bromus molliformis (I, f).
Briza minor (0).
Khrharta erecta (0).
Lolium rigidum (0).
Avena barbata (0).
Brizopyrum capense (0).
Bromus unioloides (0).
Hordeum murinum (0).
Aristida angustata (rz).
These grasses were widely distributed over the lower slopes of the whole
area.
In addition to the grasses, the following plants were noted as being in
flower :—
Water-course of valley A.
Lobelia Erinus (f).
Ranunculus pubescens (f).
Ranunculus muricatus (f).
Caucalis africana (f).
Valley slopes and ridge.
Moraea pavonia var. lutea (a).
Trifolium angustifolium (a).
Rhus lucida (a).
224 Transactions of the Royal Society of South Africa.
Passerina sp. (f).
Psoralea decumbens (f).
Leyssera gnaphaloides (f).
Felicia tenella (f).
Sebaea aurea (white variety) (f).
Pelargonium tabulare (f).
Medicago denticulata (I, f).
Trifohum agrarium (I, f).
Trifolium procumbens (I,
Trifolium glomeratum (I,
Geranium dissectum (0).
Chrysocoma coma-aurea (0).
Sebaea exacoides (0).
Pelargonium grossularoides (0).
Pelargonium myrrhifolium (0).
Wahlenbergia capensis (0).
Prismatocarpus sessilis (0).
Lessertia tomentosa (0).
Hibiscus aethiopicus (0).
Gorteria personata (0).
Kuphorbia genistoides (0).
Fumaria muralis (r).
Pelargonium lobatum (r).
Disa micrantha, (r).
)
)
f
f
Northern slopes of valley A.
Cyphia Phyteuma (f).
Southern slopes of valley A.
Aster fruticosa (1, a).
Solanum sodomaeum (0).
The following were seen coming into flower, but were not fully out :—
Micranthus plantagineus (tf).
Cyanella capensis (f).
Psoralea hirta (1, f).
Micranthus fistulosus (0).
As the above list indicates, there were a large number of plants in flower
at this time, but the general effect was not as showy as that of the previous
month. The differences between the northern and southern slopes of the
valleys were less marked than before, and it is noteworthy that at this time
Effects of a Bush Fire on the Vegetation of Signal Hull. 225
of year the sun’s rays fall much more uniformly on the two slopes than
during the winter months.
_ The whole area showed a deficiency in annuals, and certain of these had
clearly been introduced by cattle since the fire. Amongst these were noted
Trifolium agrarium, Trifolium procumbens, Trifolium glomeratum, and
Medicago denticulata, which were found associated with animal manure.
The foregoing concludes the observations made during the year
immediately following the fire. In 1920 the first visit took place on the
4th April. The earlier months of this year had been very dry, and the
hillside presented a very barren appearance. On this occasion the charac-
teristic feature was the bright green of the bushes of Rhus lucida, this shrub
having grown very actively during the winter months of the previous year.
The average height of the bushes was about three feet. The ground in
between the bushes was occupied by a large number of smaller plants of a
grey-green colour. A careful examination was made, and a large number of
them were found to be seedlings. The following list of seedlings was made :—
Hlytropappus Rhinocerotis (a).
7 Athanasia trifurcata (1, a).
Psoralea hirta (1, a).
Borbonia cordata (I, a).
Arthrosolen laxus (f).
y+ Anthospermum ciliare (f).
Psoralea uncinata (I, f).
7 Peucedanum galbanum (I, f).
Erica viridi-purpurea (I, f).
Mesembryanthemum scabrum (0).
Cliffortia ruscifolia (0).
7 Cliffortia polygonifolia (0).
Conyza ambigua (0). Chiefly growing in bushes
of Rhus lucida.
7 Aster fruticosa (0).
Aspalathus spinosa (0).
Aspalathus sp. (possibly A. thymifolia), (0).
Senecio rigida (0). Mostly confined to the damper
parts of the area.
Senecio pubigerus (0). Chiefly growing in bushes
of Rhus lucida.
Senecio Burehellii (0). ar re
Kuryops abrotanifolius (0).
Passerina sp. (0).
Osteospermum moniliferum (0).
226 Transactions of the Royal Society of South Africa.
Stoebe alopecuroides (0).
7 Salvia africana (rz).
Tt Olea verrucosa (r).
Hakea suavolens (0). Very local.
The outstanding feature is the large number of representatives of the
families Compositae and Leguminosae (fig. 4). It is illuminating to compare
this list with the following, which contains the names of all the shrubs which
had rejuvenated from underground stocks :—
Rhus lucida (a).
Asparagus capensis (f).
Rhus angustifolia (1, f).
Myrsine africana (Il, f).
Montinia acris (I, f).
Rhus glauca (0).
Rhus mucronata (0).
Rhus tomentosa (0)
EKuclea racemosa (0)
Royena glabra (0).
7 Olea verrucosa (0).
Hermannia cuneifolia (0).
7 Salvia africana (0).
Polygonum atraphaxoides (0).
7 Athanasia trifurcata (0).
Leucadendron adscendens (0
+ Cliffortia poly gonifolia (0).
7 Anthospermum ciliare (0).
Leonotis Leonurus (0).
Rhus rosmarinifolia (r).
EKuclea tomentosa (r).
+ Aster fruticosa (r).
7 Peucedanum galbanum (r).
Cluytia pulchella (r).
(+ Common to both lists.)
It will be noted that only seven plants are common to both lists, and, with
the exception of a few specimens of Athanasia trifurcata, the two families
so conspicuous in the seedling list areabsent. On the other hand, two families
which did not appear in the seedling list are an outstanding feature of this
list, viz., Anacardiaceae and Ebenaceae.
The distribution of the seedlings is of interest. Certain of the Legu-
minosae, viz. Borbonia cordata, Psoralea hirta, and Psoralea uncinata, were
Effects of a Bush Fire on the Vegetation of Signal Hill. 227
very local in their distribution and formed more or less isolated patches
all over the area. It was obvious that these seedlings were all growing
close to the spot that had borne the parent plant (or plants), (fig. 5).
Another example of this localised distribution of seedlings was found in
Hakea suavolens. In this case a single, large shrub near the top of the area
had been burnt, but owing to its very woody nature it could be recognised.
All round the burnt plant was a host of seedlings, and it called to mind
a note made by Dr. Harvey in Australia (2) that certain of the Proteaceae
native to that country only shed their seeds after prolonged basking in
the sun, or after bush fires. Hakea is one of the gencra mentioned in this
connection. The seeds of members of the Compositae on the area (with
the exception of Osteospermum moniliferum) are all wind scattered, and
consequently their distribution was much more general. In point of numbers
the plants of Elytropappus Rhinocerotis were far in advance of any others,
and, with the exception of the northern slopes of the valleys, they were evenly
distributed all over the area. This suggests that, as in the case of the
Leguminosae, the seeds had withstood the fire and had not been introduced
from plants growing in the adjoining unburnt part. Had this been the case,
one would have expected to find the numbers of plants at the margin in
excess of those near the centre.
The northern and southern slopes of the valleys showed a somewhat
striking difference in the bushes which grew on them. The southern slopes
were the more sparsely populated, and the chief bushes were Rhus lucida,
Elytropappus Rhinocerotis, Athanasia trifurcata, and Borbonia cordata.
The northern slopes differed from the southern in the diminution in numbers
of Elytropappus Rhinocerotis and Athanasia trifurcata, and the presence of
Myrsine africana, Anthospermum ciliare, Raus angustifolia, Stoebe alopecu-
roides, and Royena glabra.
Asparagus capensis, the young shoots of which in the previous year
had been such a feature of the landscape, was most inconspicuous. After
a careful survey of the area it was clear that the number of plants was about
the same as in 1919, but this year no young shoots had appeared before the
winter rains, and the old dull-green shoots were easily overlooked. Another
noteworthy feature was the absence of flowers in Haemanthus coccineus.
On the other hand, Andropogon hirtus and Hibiscus aethiopicus were in full
flower, as in the previous year. A plant that had either been absent or
overlooked in 1919 was Linum thesioides, which was rather sparsely distri-
buted on the southern slopes of valley A.
The area was visited once more in June, but there was little to record.
The early months of 1920 were remarkably dry, and the whole vegetative
period was somewhat later than usual over the whole Peninsula. The
same plants were found as in 1919 (with the exception of the
228 Transactions of the Royal Society of South Africa.
seedlings already mentioned), and the only additions to the previous lists
were :—-
Kriospermum lanceifolium (0).
Lobostemon fruticosus (0).
* Hragrostis brizoides (zr).
* Disa tenuis (r).
* Chenopodium ambrosioides (r).
Watsonia rosea, (r).
Cotyledon grandiflora (r).
The winter rains this year were exceptionally heavy, and at one point
on the southern slope of valley A there was a small landslide (fig. 6) which
caused a large amount of debris to be washed down the water-course,
altering this considerably (fig. 7). In several places a deep channel was
cut by the rush of mud and stones. |
The spring flora, though in composition the same, was by no means as
rich as in the previous year, and as elsewhere on the Peninsula the season
was considered a good one, one is forced to conclude that the previous year’s
display was an effect of the fire.
In October records were made of a few plants that had not been noted
the previous year, viz. :—
Indigofera incana (f).
Gnaphahum parvulum (f).
Lessertia pulchella (0).
Silene anglica, (0).
Lessertia excisa (0).
Hypochoeris glabra (0).
Carduus pycnocephalus (0).
Avenastrum sp. (0).
Aizoon sarmentosum (0).
Hermannia prismatocarpus (0).
Polygala bracteolata (r).
Urospermum picroides (zr).
Pelargonium hirsutum (r).
During the winter months several water-holes for cattle were dug on
the ridge between valleys A and B. In October these holes were nearly
dry, and the plants growing in them were interesting in that they were
totally distinct from any growing elsewhere on the area. The follewing
were seen :—
Cyperus tenellus.
Crassula brevifolia.
Effects of a Bush Fire on the Vegetation of Signal Hull. 229
Juncus bufonius.
Small grass. Not in flower; possibly
Agrostis lachnantha.
This year a visit was paid in November. The only record of note was
the abundance of Ornithogalum pilosum, a plant which until recently had
been considered rare on the Peninsula.
In March 1921, a little over two years after the fire, a few plants of
Antholyza lucidor were in flower at the top of the patch, but as the particular
region in which the plants were found had not been visited in any previous
March, there were no records of the behaviour of this species. Senecio
pubigerus, which in 1920 had been confined to the shelter of the bushes,
had spread widely over the slope of valley A and elsewhere was quite a
common plant apart from the larger bushes. One young silver tree
(Leucadendron argenteum) was seen near the top of the patch.
During the late winter and spring months rapid growth took place over
the whole area, and on the northern slopes of the valleys a number of
seedlings of Hrica viridi-purpurea were noted coming into flower for the
first time. On the slopes between valleys B and C a single plant of Protea
grandiflora was noted. The shoots of this had obviously come up from the
underground stock. On the ridge between valleys A and B a group of young
plants of Relhania ericoides and several young plants of Selago ramosissima
were seen.
The spring flowers of this year showed a still further reduction in number
and quality, with the exception of Babiana stricta. This species was
exceptional in that no obvious effects of the fire manifested themselves in
this or in previous years. This may be due to the fact that the corms are
unusually deeply seated.
GENERAL CONCLUSIONS.
It is clear that in certain plants rapid growth was induced after the fire,
while in others the vegetative parts were wholly killed and the plants had
to rely upon their seeds for the continuance of life.
The exact way in which fire influences the plant is not clear. Several
factors may be at work. Possibly the heat during the fire acts directly on
the underground organs, mobilising all the forces of the plant. It is a well-
known fact that by employing high temperatures the normal time of flower-
ing in certain plants may be anticipated (4). In the case of those plants,
notably Asparagus capensis, in which the flowering period was obviously
induced at a somewhat earlier time than usual, it is probable that the heat
acted directly on the underground organs. It is also likely that the direct
action of heat was responsible for the production of the large number of
18
230 Transactions of the Royal Society of South Africa.
flowers of Haemanthus coccineus, though in this case the actual time of
flowering was the normal one.
In those cases, however, in which the time of flowering was normal, but
the number and size of the flowering shoots were greater than usual, it is
probable that additional factors were involved. As an example of this
type of behaviour we may take the spring Monocotyledons, which have
underground storage organs, and whose leaves appear soon after the early
winter rains. Owing to the removal of all the bushes their leaves were well
supplied with light, and hence we may suppose that their photosynthetic
activity was greater than would have been the case had they been over-
shadowed by bushes. Another factor which may have operated to some
extent was the increased warmth of the soil, due to the exposure of the
ground to the direct rays of the sun. The winter, however, in the Cape
Peninsula is very mild, and it seems likely that light rather than heat is the
predominating factor.
The following suggestions are put forward as to the ways in which fire
may influence the soil and thus benefit the plants :—
1. The physical effects of fire on the upper layers of soil may be such
that increased aeration is brought about.
2. Soil protozoa, which are known to have a retarding influence on the
erowth of plants, are very sensitive to high temperatures (1), and
the number of these organisms may be considerably reduced.
3. The ash from the burnt plants may have a certain manurial value,
thus enriching the soil chemically.
No definite example was met with of a species which had been eradicated
by burning, though there is no direct evidence to show that such a species
did not exist, since the area had not been surveyed before the fire. The
evidence that one relies upon in drawing the conclusion is the composition
of the unburnt surroundings. This, however, is far from satisfactory,
owing to various factors such as cultivation, ete.
One fact is clear, and that is that burning favours the spread of the
‘ Rhenoster Bush ” (Elytropappus Rhinocerotis). The type of vegetation
now established on the site of the fire is similar to that termed by Marloth
Rhenosterveld ” (fig. 6), and the foregoing lends support to his view that
this formation is an artificial one (3).
Another result of a bush fire on a slope such as this is obvious. The
soil is laid bare, and cases of erosion—some slight, some marked—were
common all over the area.
While making the observations just recorded, several interesting facts
were noted, which, though they have no direct relation to the subject
under discussion, are nevertheless striking and may be mentioned here.
Effects of a Bush Fire on the Vegetation of Signal Hill. 231
The Peninsula flora is characterised by the presence of certain families
which are typical of the south-western districts, but are either lacking or
scarce in South Africa beyond these limits. The area under discussion is
singularly deficient in a number of these families. Bruniaceae and Pene-
aceae are entirely absent, the family Restiaceae is represented by a single
specimen, and members of Proteaceae, Rutaceae, and Ericaceae are only
occasionally seen.
It has been suggested, especially in the case of the Proteaceae, that bush
fires have been largely instrumental in eradicating large numbers of species
from these slopes. This suggestion, however, does not explain why it is
that most of the families mentioned above are well represented on the
slopes below Lion’s Head, which slopes in their vegetation incline to the
true ““ Macchia ”’ type. It is significant that the soil here is derived from
granite (fig. 1), and as bush fires have occurred in plenty, it seems reasonable
to conclude that the soil may be the determining factor in the case.
The problem needs further investigation, but certain facts such as the
dominance of Blaeria ericoides at certain spots on the granite soil, and the
scarcity of the family Ericaceae on similar positions on the slate, lead one to
the conclusion that the soil in this particular instance has a profound
influence on the vegetation.
My thanks are due to Professor Thoday for his kindness in taking the
photographs which illustrate this paper, and for the many helpful criticisms
which he has offered from time to time during the progress of the work.
SUMMARY.
1. The bush fire, the effects of which are recorded, broke out on the
5th February 1919 and burned for two days, killing all the aerial parts of
plants on the slope.
2. About three weeks later considerable growth had taken place in
Asparagus capensis and Andropogon hirtus. Shortly afterwards Haemanthus
coccineus and a few other less notable plants were in flower. Various species
of Rhus were putting up shoots.
3. The early winter and spring flowering periods were characterised by
an abundance of vigorous flowering shoots. These appeared at the usual
time. In the majority of cases these plants possess underground storage
organs. Progressive decrease in numbers and vigour of plants flowering
during these periods was noted in 1920 and 1921. Suggestions are made to
account for the phenomenon.
4, Numerous seedlings came up during the winter of 1919, but in general
these were not identified till the next year.
232 Transactions of the Royal Society of South Africa.
5. The northern, shaded slopes of the valleys showed a conspicuously
different plant population from the southern exposed slopes during the
winter months. In the summer months the contrast was not as sharply
marked. This is attributed largely to the fact that in winter the sun
shines on the northern slopes for a short period only of the day, while the
southern slopes get most of the available sunlight.
6. During 1920 the shrubs were divided into two classes: (a) those
where the underground parts had survived the fire and from which new
shoots arose, and (b) those which were killed by the fire and which repro-
duced themselves by seed. The Rhenoster bush falls in class (6), and is.
clearly favoured by burning.
7. The removal of the vegetation by the fire helped the process of soil
erosion. This process was also aided by man and cattle. Several small
washaways occurred on the area subsequent to the fire.
8. The area is shown to be deficient in several typical south-western
families, but to what extent this may be attributed to the influence of
repeated fires is not clear, and the view is brought forward that soil may be
the determining factor in this case.
LITERATURE CITED.
(1) Hatt, Sir A. D.—The Soil, third edition, 1920.
(2) Harvey, W. H.—Memoirs of Dr. Harvey, 1869.
(3) MartotH, R.—Das Kapland, Jena, 1908.
(4) Pattapin, W.—Plant Physiology, edited by B. E. Livingston, 1918.
(5) Pures, E. P.—‘‘ A Preliminary Report on the Veld-burning Experiments at Groen-
kloof, Pretoria,’’ South African Journal of Science, xvi, 1920.
(6) Puitures, E. P. — “‘ Veld-burning Experiments at Groenkloof,’ Union of South
Africa, Department of Agriculture, Science Bulletin, No. 17.
Trans. Roy. Soc. 8. Afr. Vol. X. Platexrx.
A B' B ra
Fic. 2.—Photograph of burnt area on Signal Hill, taken from the roof of the Natural
Science Building, University of Cape Town, two and a half years after the fire.
Be
Fic. 3.—Southern part of the burnt area, looking up the hill. The foreground is covered with
bushes of Acacia horrida, and was not touched by the fire. The fenced-in region may be
distinguished from the open hillside by the presence of numerous old inflorescences of
Andropogon hirtus.
Neill & Co., Ltd.
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Trans. Roy. Soc. 8. Afr. Vol. X. Plate XT.
Fic. 4.—Detailed view of some of the seedlings which have grown since the fire. In the fore-
ground is a patch of Borbonia cordata and a few plants of Cliffortia ruscifolia. The plants
in the background are mainly Elytropappus Rhinocerotis, the two larger shrubs being
Rhus lucida.
Fie. 5.—A “ school” of young plants of Psoralea hirta. The darker bushes are Rhus lucida.
Neill & Co., Lid.
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( 233 )
COLOUR AND CHEMICAL CONSTITUTION.
Part XVI.—FuRTHER MISCELLANEOUS OBSERVATIONS.
By James Morr.
5)
The main lines of the theory of the colour of “ cyclic”’ coloured sub-
stances having been established in Parts X, XIII, and XIV, it only remains
to put on record observations of the absorption wave-length of all the
‘““monocyclic’’ and “ dicyclic”’ colours which I have used in making the
discovery, but have not published for fear of overloading these theoretical
papers with unnecessary detail.
A. DERIVATIVES OF THE PROTOTYPE COLOURS.
1. Orthopara- or (2-4’)-dioxybenzhydrol, from salicylaldehyd (see
Beilstein, ii, 1114). This has A 543 (broad) in alkali, A 486 in HCl, and
A about 495 when in suspension in neutral water. The isomeric p—p-com-
pound has A 539 in dilute alkali, but in HCl has the same A as the o—p-
compound.
2. 2-oxy-4’-dimethylaminobenzhydrol, from salicylaldehyd and di-
methylaniline, has A 561 (broad) in alkali, with A500 in HCl. The isomeric
4—-4’-compound has A 572 in alkali and A 504 in acid. This was predicted
(as A 571) in Part XII, top of p. 211.
3. 2-4-4’-trioxybenzhydrol, from p-oxybenzaldehyd and_ resorcin,
appears to have A 494, whereas A 550 was expected : possibly the reaction
is abnormal.
4, 2-4-dioxy-4’-methoxybenzhydrol, from anisaldehyd and resorcin, has
A about 380 in alkali. This substance appears to be monocyclic, since
2—4-dioxybenzhydrol has a similar X.
5. 2-4-dioxy-3’—4’-dimethoxy benzhydrol-2’-carboxylic acid, from opianic
acid and. resorcin, has A 390 in alkali. This is also monocyclic, being a
derivative of monophenolphthalein (Part XIII, p. 38).
6. Mono-a-naphtholphthalein, from phthalaldehydic acid and a-naphthol,
has A 370, and is also monocyclic.
234 Transactions of the Royal Society of South Africa.
7. C-phenyl-derivative of foregoing, from benzoylbenzoic acid and
a-naphthol, has A 401 in alkali. In strong sulphuric acid, however, it has
the much higher colour A 543.
8. 1-4'-dioxybenznaphthydrol, from p-oxybenzaldehyd and a-naphthol,
is violet with A 590 in alkali. I have assumed the constitution to be
OH
i ae Seas
HO¢ >—CHOH 1)
Wa
on the analogy of naphtholphthalein (Berichte, 1920, 1445).
9. The corresponding substance from f-naphthol,
HO< \—CHOH< _.»,
SS OS
or 2-4’-dioxybenznaphthydrol, has A 556, with a pink colour.
10. Phenol-$-naphtholphthalein, the benzoic acid-derivative of the
foregoing,
nt OH
yao a
HO-C>-COH-K_ YD
7
>?
is now found to have A 570 (broad). My statement in Part IT (1918, p. 113),
that it has A 637, is erroneous.
The naphthol/phenol colour-factor from these two substances is quite
different from that obtained when the C,H, group of the naphthalene-ring
is joined metapara to the benzhydrol linkage, viz. 1-026 for orthometa- as
against 1-082 for metapara-attachment.
From these data I can roughly predict the wave-length of the still
unknown p—p—a-naphtholphthalein to be about A 628, that of the commercial
o—o-a-naphtholphthalein being A 662.*
C,H,CO,H
B, FurtHer DERIVATIVES OF PHENOLPHTHALEIN (Part [V—continued).
(a) 3- (or 6)-oxyphenolphthalein, from 3-oxyphthalic acid, has A 556
in NaOH and is higher in bicarbonate (A 562).
* When, however, the hydroxy! is para to the benzhydrol linkage, a third
naphthol/phenol factor comes into play. Its value is 1-042 (see Part XV, Naphtho-
fluorescein). The true calculated 1 of p—p—a-naphtholphthalein is 602. This is also
the factor for safranines like Magdala-red.
Colour and Chemical Constitution. 235
(b) 3-6-dioxyphenolphthalein, from Thiele’s dioxy-acid, gives a large
difference between caustic and bicarbonate colour, viz. A 549 and A 563
respectively.
(c) 4-5-dioxyphenolphthalein, from normetahemipinic acid, gives a
similar difference, viz. A 558 in NaOH, and A 568 in NaHCO, .
(d) 5-methoxyphenolphthalein, from 4-methoxyphthalic acid, has
A565.
(ec) 3-6-dimethoxyphenolphthalein has A 568.
(f) 4-5-dimethoxyphenolphthalein (‘ phenolmetahemipineine”’) has
A 556.
(g) 5-6-dimethoxyphenolphthalein (‘‘ phenolhemipineine ’’) has A 571.
I have to thank the Institute of Chemistry for obtaining for me small
quantities of the necessary acids from unknown donors in the English
universities.
(h) 3-nitrophenolphthalein, from 3-nitrophthalic acid, has A 570. The
value A 559 was given in 1918, and is erroneous.
(7) a-nitro-fgjk-tetrabromophenolphthalein has A 597 (blue-violet).
(See Part XI, p. 130, for the nomenclature.)
(j) 5-nitrophenolphthalein, from 4-nitrophthalic acid, has A 572. Its
tetrabromo-compound has A 599.
(k) 3-4-5-6- (or abed)-tetrachlorophenolphthalein (“‘ phenoltetrachloro-
phthalein ’’) has A 581, scarcely different from that of the common (fgjk)
isomer, viz. A 583.
(1) Phenoldimethylalphanaphthylaminephthalein is green, A 625.
C. DERIVATIVES OF GHOSH’S QUINOLINIC ACID.
These are phthaleins with N for CH in the phthalic ring.
I. Phenolquinolineine has A 533.
II. Orthocresolquinolineine has d 544.
III. Thymolquinolineine has A 593.
IV. Resorcinquinolineine has A 490.
V. Orcinquinolineine has A 496.
D. DERIVATIVES OF TRIPHENYLCARBINOL.
1. Ortho-oxy-malachite green (salicyl-green) has A 627 in neutral solu-
tion, A 572 in alkali, and A 505 in acid.
2. 2'-oxy-4-dimethylaminofuchsone (foregoing with OH for NMe,) has
A 490 neutral, A 533 in alkali, and A 479 in acid.
3. p-methoxy-malachite green, from anisaldehyd, has only one colour,
viz. A 605.
236 Transactions of the Royal Society of South Africa.
4. p-Methoxybenzaurine (=aurine methyl ether) has A 551.
5. Dimethylaminodioxyphenyldinaphthylearbinol, from dimethyl-
aminobenzaldehyd and a-naphthol, has A 730 when neutral and A 492
when acid.
6. 2-2'-4-4’-tetraoxytriphenylcarbinol has A 516 (non-fluorescent). On
heating with zinc chloride this gives resorcin-benzein, A 492 (very fluorescent).
7. Paranitro-malachite green has A 642 in water. Formdnek gives A 637
in alcohol. The band is vague and not easily measured.
8. Nitro-oxy-dimethylaminotriphenylearbinol, from 4-nitro-4'-oxy benzo-
phenone, has apparently A 610, whereas calculation givesA575. This requires
reinvestigation.
9. Para-amino-malachite green (unsymm. tetramethylfuchsine), from
reduction of No. 7, has A 581 in acetic acid, A 610 vague in stronger acid,
and is yellow in HCl (A about 420).
10. 2-2’-4-4’_4”-pentaoxytriphenylearbinol (dioxyaurine) has A 545.
11. Unsymm. dimethy]parafuchsine, from dimethylaminobenzaldehyd
and aniline hydrochloride, has AA 547 and 493 when neutral, but when acidic
shows vague bands at about A 575 and near the red end (over A 700), which
1s surprising.
E. Vartous SIMPLE COLOURED SUBSTANCES.
(a) Quinonoxime. This has two vague bands at both ends of the
spectrum, viz. about AA 680 and 350.
(6) Sodium salt of foregoing (Na-p-nitrosophenol). In water A 398.
Baly gives A 415 in alcohol.
(c) Nitrosodimethylaniline. The green base has A 415+A 725. The
acetate has A 457 alone: the chloride in HCl A 390.
(d) 2-3-dicyano-quinol. This is colourless, but with a very strong
fluorescence: A about 330 neutral, A 410 alkaline. The fluorescence of the
salts of this substance is extraordinary, so that this is probably the simplest
highly fluorescent substance.
(e) 3-6-dioxyphthalimide (from hydrolysis of above). This is yellow
with A 405 when neutral, but A 480 in NaOH : highly fluorescent.
(f) 3-6-dioxyphthalic acid. When neutral this is nearly colourless with
A370. Alkali turns it yellow (A 437) with a marvellous fluorescence. The
fluorescence must depend on the juxtaposition of ONa and COONa, since I
find that sodium salicylate, when examined in presence of NaOH in sunlight,
shows a fluorescence (violet).
F. UNCLASSIFIABLE OBSERVATIONS.
I. Quinizarine: chief band A 598.
II. 2-chloroquinizarine : chief band A 601.
Colour and Chemical Constitution. aes
III. a-Naphthofluorane in conc. H,SO, has AA 500 and 475.
IV. Phenosafranine (‘‘ desensitol’’) has AA 525-+-505 in water, and
AA 532+497 in alkali.
V. Quinhydrone in AmHCO,: A 480 broad.
VI. Tetrabromo-quinhydrone in NaOH: A510. (2-2’-6-6’ variety.)
VII. Quinhydrone of dimethylphenylenediamine: AA 558+513.
WATT. x e tetramethylphenylenediamine: AA 6144568.
IX. — x dihydrophenazine: AA 701+640.
Note.—The quinhydrones obey the dicyclic colour laws, but the
oxidation-factor is =, not 2.
( 239 )
ON HYALITE.
By J. S.'v. p. Lingen and A. R. I. WALKER.
The investigation of hyalite, some of the results of which are recorded
in the following note, first suggested itself to the authors whilst they were
engaged in studying the optical behaviour of liquid spherulites.
An attempt was being made to find a solid spherulitic body analogous
—hboth as regards internal structure and optical behaviour—with the type
of liquid spherulite described by Lehmann, in which the molecules of the
substance are regarded by him as being arranged either along lines radiating
outwards from the centre of the spherulite, or on the surfaces of spheres
possessing a common centre—the centre of the spherulite.
As a result of a preliminary examination of a number of substances
selected for this purpose, hyalite was chosen as being the most promising.
In this way one was led to consider the nature of what is commonly
termed the “‘ anomalous double refraction ” of hyalite. In this connection
Dana * writes as follows :
(Opal) often shows double refraction similar to that observed
in colloidal substances due to tension. The mammillary form Hyalhte
often yrelds the uniaxial interference cross of a negative substance in
parallel polarised light; + this is referred to tension by Schultze, Ber.
nied. Ges., 69, 1861.”
Similarly Rosenbusch f :
“ Hyalite is doubly refracting, and, under certain conditions,
exhibits the interference cross of a uniaxial substance with isochromatic
curves. This appearance is referred to conditions of strain occasioned
by the concentric shell-like structure of the mineral. The character
of the double refraction is negative. The hyalite cross often separates
into hyperbolas during a rotation of the section between crossed nicols,
which would necessarily be the case if the layers were not regular
spherical shells.”
* Dana, A System of Mineralogy, 6th ed., p. 194.
+ The italics are ours.
t Rosenbusch (Iddings), Microscopical Physiography of Rock-making Minerals,
4th ed., p. 194.
240 Transactions of the Royal Society of South Africa.
The material investigated was supplied to the Geological Department
of the University of Cape Town by Foote of Philadelphia, and is labelled
as having been obtained from Guanajuato, Mexico.
The specimen represents a portion of a succession of deposits which pre-
sumably originally lined a rock cavity. Several “‘ forms ”’ of silica—some
chalcedonic, others opaline—are represented, and, in our opinion, judging
from this single specimen, a definite order of deposition is observable.
We infer that, throughout a period of continuous deposition, the chalce-
donic varieties were deposited at the commencement, and the opaline
varieties towards the close, of such a period.
These varieties, enumerated in the order in which we consider them to
have been deposited, are as follows :—
Chalcedomic Silica.
1. Dark chocolate-brown variety resembling jasper.
2. Opaque, white ; in parts bluish-white.
3. Pale blue or bluish-white, semi-translucent ; botryoidal (pale brown
by transmitted light).
Opaline Silica.
4. Clear, colourless, botryoidal hyalite.
5. Extremely pale bluish-white, opalescent or slightly clouded hyalite ;
spheroidal in form, and much less common than the colourless and water-
clear hyalite. (Since this variety generally occurs forming semi-spheroid.
masses on the surface of the clear hyalite, no difficulty is experienced in
distinguishing between the two—particularly when both are immersed in
water,—but, when the slightly clouded variety is detached from the clear,
optical examination proves the surest method for its identification. It is
important to note also that, in certain parts of the specimen, the small
semi-spheroids of this variety are coated with a thin layer of clear hyalite.)
Apparently the forms of silica enumerated above were not the result of
one continuous period of deposition, but of several such periods ; nor was
deposition uniform over the whole surface of the cavity throughout each
period. Several repetitions of the sequence 2, 3, 4, 5 are observable.
In the following we refer principally to two kinds of thin sections,
which, for convenience, we propose to name “ diametral ”’ and “ tangential ”
respectively. By “diametral section’? we mean one cut approximately
through the centre of a spherulite ; by “‘ tangential section ” one prepared
from the surface layer of such a body.
Throughout the preparation of any one section the material was closely
observed in order to see whether the grinding—the process by which all
On Hyalite. 241
the sections were made—had caused to appear within the material any
structures which might be attributable to a release from internal strains
consequent on the destruction of the spheroid. Nothing of this nature was
observed. Occasionally a section would break along irregular cracks,
but, generally speaking, they retained their circular outline unbroken and
did not show, when examined under the microscope in ordinary transmitted
light, any signs of a strained internal concentric structure.
THE COLOURLESS AND WATER-CLEAR HYALITE.
Diametral Sections.
(a) Examination in plane polarised light.
A diametral section of this variety, when placed on the stage of the
microscope so that the axis of the microscope passes through the centre
of the spheroid, exhibits, when examined in plane polarised ight between
crossed nicols, a figure consisting of a black cross with isochromatic rings
resembling that given by a uniaxial mineral in convergent polarised light.
On the insertion of a quarter undulation mica plate the figure behaves in
a similar manner to that of a negative uniaxial substance ; many sections
were tested in this way, and in all cases the sign was negative. A move-
ment of the section, however, causes a movement of the figure, so that
whenever the section is excentrically placed the figure resembles and behaves
like a partial uniaxial figure. On this account we'do not regard this figure
given by hyalite as being analogous with that of a uniaxial mineral in con-
vergent polarised light; sections of radiating chalcedony (described by
Miers as a biaxial mineral) and diametral sections of oolitic grains have
been observed to give, under similar conditions, figures resembling, both
in general appearance and optical behaviour, those given by hyalite.
(b) Examination in convergent polarised light.
The behaviour of diametral sections in convergent polarised light is
essentially similar to their behaviour in plane polarised light.
Tangential Sections.
(a) Examination in plane polarised light.
Tangential sections, between crossed nicols, are either quite isotropic
or show very slight graying marginally.
(b) Examination in convergent polarised light.
In convergent light a tangential section yields a figure composed solely
of a black cross ; owing to the thinness of the section and the weak strength
242 Transactions of the Royal Society of South Africa.
of birefringence of the mineral, no colours are produced. The cross remains
stationary within a comparatively wide range of motion of the section ; on
the rotation of the latter it breaks up into two hyperbole—but this separa-
tion, although apparent, is so slight that we hesitate to proclaim the
biaxial nature of the figure. We do, however, regard it as being a true
axial figure.
The sign of the mineral, determined from these figures by means of a
quarter undulation mica plate, is negative.
THE OPALESCENT OR SLIGHTLY CLOUDED HYALITE.
Thin sections of this variety have a very faint brownish tint when viewed
by transmitted hght. The important difference between this and the
clear hyalite, so far as optical behaviour is concerned, is in the sign of the
double refraction ; whereas the nature of the double refraction of the clear
is negative, that of the slightly clouded is positive. In other respects, what
has been written concerning the clear applies also to this variety.
COMPOUND SPHEROIDS.
Mention has been made of the occurrence of spheriods of the slightly
clouded hyalite having an external layer of colourless hyalite. A section
intermediate between a tangential and a diametral section was prepared
from one of these. When this section was examined in plane polarised light
it exhibited an extinction figure composed of a black cross and a black ring
—the latter being comparatively close to the margin of the section. On
testing this figure with a quarter undulation mica plate it became apparent
that the black ring was of the nature of a compensation band resulting
from the overlapping of the two varieties—positive and negative—conse-
quent on the curvature of the spheroid.
Subsequently a section was made through a cluster of such spheroids,
which, on examination, revealed the fact that in some spheroids there is
a repetition of concentric layers of positive and negative material.
FuRTHER EXPERIMENTS AND OBSERVATIONS.
(a) Examination by means of X-rays.
X-rays were passed through a thick diametral section of the clear
hyalite, and the effect of the mineral on the rays was ascertained by means
of a photographic plate—after the method of von Laue. The “ pattern ”
showed that, for a given radius, the dispersion had been uniform around
the central spot.
On H yalite. 243
(b) The effects of heat.
It was observed that, when a fragment of clear hyalite was heated on
platinum foil in a bunsen flame, thin flakes split off from its surface, and
that these flakes eventually curled and became portions of spherical surfaces ;
in addition they lost their glassy appearance and turned opalescent.
A somewhat thick diametral section, placed on a glass slip and heated
as above, became opalescent and curled into a concavo-convex shape,
whilst at the same time a small spherical kernel split out from its centre.
When the concavo-convex portion was examined under the microscope,
it was found to have a pale-brown colour by transmitted hght; further,
between crossed nicols in plane polarised light it yielded an extinction figure
having a positive sign—that is, having the same sign as the naturally
occurring opalescent hyalite previously described.
Portions of some of the flatter opalescent shells obtained by heating
behaved in a very similar manner to tangential sections of the naturally
occurring form.
(c) Chemical examination.
Material was submitted to Mr P. R. v. d. Riet Copeman for analysis,
with a view to finding out what chemical change had accompanied the change
in sign produced by heating. From a series of silica determinations, and
assuming the general formula of hyalite to be Si0,.nH,0, he arrived at
the following conclusions :—
1. The loss—presumably water—on heating the clear hyalite at bunsen
heat was 2-98 per cent.
2. The clear contains 10-28 per cent. water.
3. The naturally occurring opalescent hyalite contains 7-55 per cent.
water.
From the above it appears that the artificially produced opalescent
hyalite has almost the same water content as the naturally occurring
opalescent form.
REMARKS IN CONCLUSION.
Our chief object in writing this note has been to record observations
pending the arrival of more material for further investigation. At present
we feel we can only write tentatively regarding the reasons why hyalite
is doubly refracting. We consider that the explanation that it is due to
tension is, at the best, an insufficient one; we are inclined to regard the
occurrence of spheroids composed of concentric layers of positive and
negative material as making this view—considering it as being the sole
explanation—untenable.
We would suggest that hyalite is a substance of which the component
244 Transactions of the Royal Society of South Africa.
molecules are anisotropic unities ; that their arrangement in the naturally
occurring forms may be due to strains, but that their crystallographic
orientation, on which the sign of the double refraction depends, is due to
their water content.
One of us * has shown in a previous paper how, by gradually driving off
the water which it contains, the internal molecular structure of brucite is
changed ; this change is tantamount to an alteration of the axial ratio of
the mineral. Keeping this in mind, and taking into consideration also the
fact that whenever the clear negative hyalite is heated to redness it becomes
converted into an opalescent positive material resembling the naturally
occurring opalescent hyalite, we cannot help but feel that the water content
is the most important factor in deciding the nature of the double refraction
of hyalite.
That tangential sections of hyalite behave as uniaxial—or biaxial ?—
mineral substances we regard as evidence in support of our contention that
the molecules are in themselves anisotropic.
In conclusion, we wish to thank the Royal Society of South Africa for
the grant in aid of research.
* J. 8S. v. d. Lingen, Trans. R. Soc. 8. Afric., Vol. VII, Pt. 1, 1918, p. 59.
( 245 )
CIRCUMCISION REGIMENTS AS A NATIVE CHRONOLOGY.
By the Rev. Professor W. A. Norton, M.A., B.Litt.
The regiments of the successive circumcision lodges, holden about every
five years in most tribes, form a rough method of fixing the chronology,
by comparing the mephato or regiments circumcised in different tribes at
the time of the same event or reign. For example, the well-known chief
Sechele, Livingstone’s friend, was remembered in the neighbouring tribes
as having been contemporary with such-and-such a chief, by circumcision,
asa youth. Having fixed the date in last century, therefore, of his circum-
cision, one can deduce the dates of mephato in the other tribes. At the
beginning of my work as a Government research grantee I cast about for
some subject which would be virgin ground, and at the same time have a
wide general bearing on the comparative history and folklore of the tribes.
I lit upon this interesting subject, and found my researches well rewarded ;
seeing that I was able to collect regiment lists in some thirty of the central
South African tribes, some of these lists running well back into the eighteenth
century. That veteran scholar, Fr. Bryant of Natal, has worked out a
valuable list of Zulu regiments: but practically nothing seems to have been
done in the case of the more Western tribes.
The interest of these regiment lists is manifold. They are the natural
scaffolding of native history. For years, as a missionary, I wasted hours,
as had my predecessors, in dating the births of baptismal candidates by
wars or comets, before discovering that the terse query “ O leng ?”’ brought
the terse answer “ Ke le-tlhaselwa”’ or “ Ke le-fitlha-Kgosi,” at once
approximately dated the person’s circumcision (usually about the age of
eighteen). However adequate dating by wars may be in the war-worn
South-East Africa, the wars of the Bechwana or Basuto were mostly short
raids, and infinitely more cumbersome and uncertain for dating purposes
than the regiments. Of course the regiments must first themselves be
dated once for all, to which necessary work this paper is contributory.
I withhold further remarks on the lists in detail till readers, who are
working with the individual tribes, have had the opportunity and goodness
to check them, as I hope they will, by researching into tribal history with
their help. I shall be deeply grateful if those who are not arranging to
publish such research themselves will kindly put it at our disposal, to be
I)
246 Transactions of the Royal Society of South Africa.
correlated and, if possible, published at an early date, with due acknow-
ledgments.
I am convinced that what native history is still to be gathered must be
gathered at once, from the old men with memory of it. Time and again
have I been told, during the last few months, that old people have just
died who could have answered my question (the younger generation are
woetully uninterested). Could not those in touch with such make the future
their debtor by an effort to record their recollections within the next few
months? Even where such record were not available for co-operative
history, as I suggest, I should be grateful to have the names and whereabouts
of such sources of information, in case some doubtful point within their
memory needed to be cleared up. Suggested names of officials, missionaries,
or settlers who have studied the tribes, or have long experience of them,
would also be gratefully received.
One thing I especially ask: that those who co-operate in this urgent
work of salvaging the last remnants of these recollections would take care
to give the regiment (mophato) names contemporary with each incident
as well as the date A.D. where that can be fixed.
In conclusion, I might mention that the significance of the regiment name
itself often throws lght on the history, as will be seen in two examples
given above (v. index under -tlhaselwa and -fitlha-Kgosi), and also on the
feeling of the tribe at the time. The spelling even reflects the variant pro-
nunciation, as (seTlhaping) hachane for fatshwane and (seTlokwa) hori for fire.
N.B.—The following meteorological dates have been kindly checked
by H.M. Astronomer at the Cape, and may be useful in fixing others :—
Comets in 1811-2, Nov. 7°48, ’58-9, ’°82; meteors, Nov. 27, 1885; total
solar eclipses, Nov. 19, 1835, and April 16, 1874.
I regret that data I have been looking for concerning the ancient tribe
of the ba-Hurutse have not yet arrived: I trust workers in that and other
spheres will kindly help me with information and correction. I beg to
thank the reigning chiefs Kgama, Lenewe, Sebele, Lotlamoreng, Seboko,
Gaborone, and their counsellors, the chieftainess Moletshi, the chief Segale
and others of his family, especially the teachers Amos and Seth Pilane,
and masters and scholars at Pietersburg Training College, Tyger Kloof,
etc., for valuable help in preparing this paper: also the Government
officials of the Native Affairs Department and of Bechuanaland, Basutoland,
and the Transvaal, for constant kindness and help in every way.
Note to InpEXx oF REGIMENT NAMES.
N.B.—In all but the two or three cases where I have given the special
prefix, the prefix ma is to be understood before the form given in the index
as collective name of the regiment: the individual member of it uses the
singular prefix Je-; hence the formula for asking one’s regiment, “‘ O le-ng ?”’
z.e. ““ You are le-what?”’ To find the mophato preceding or following,
Circumcision Regiments as a Native Chronology. 247
one asks: “‘ A rupisitswe ke mang?” or “ A rupisitse mang ?”’ (7.e. they
were circumcised by, or circumcised, whom ?), and so builds up or checks
the list of successive mephato.
Lists 1-14 of the Transvaal tribes very rarely have the composite names
which are characteristic of 15-25, those of Bechuanaland, showing a different
strain of cultural borrowing where not of original descent.
INDEX TO REGIMENT NAMES OF TRIBES OF TRANSVAAL AND
BECHWANALAND.
ma-abaN Kwe, dividing the sheep, 22.
abaNtwa, ordering the host, 20.
alola, removers (intrans.), 15; alosa, do. (trans.), 25.
abaKgomo, sharing the cattle, 19, 23-25.
aparaNkwe, wearing leopard (skins), 8, 17.
apaPula, spreading the rain, 25.
bapoga, outstretched, erect (? aba Phoha, distributing feathers), 15.
betlwana, those cut or chipped (¢.e. in circumcision), 17.
bitsi, callers, 1, 4.
butswane, the ripe ones, 11.
busa, turning back (the enemy), 24 f.
betla-Kgang, fashioning strife, 23.
chologa (for thologa, q.v.), 23.
cecele, clever (for tsele-tsele), 8*, 9%.
chedi, gushers, 13.
chama, undaunted, 8, 17.
ea-ka-Kgang, going with strife, 18, 20.
ea-ka-Kgosi, going with the chief, 23.
ea-ka-Thata,, going with strength, 17-19, 21.
ea-ka-Thebe, going with a shield, 19, 24.
emelwa, championed, 15 (or waited for, because kept long uncircumcised)..
ea-ka-Ntwa, going with a host, 18-21.
ea-ka Pula, going with rain, 18 f.
fatlha, defenders, 8, 12.
fatshwa, dappled cattle, 15 (or phatshwa).
fatshwana (dim. of above), 8*.
firi, hyenas, 8, 16, 17 (hiri).
fitlha-Kgosi, buriers of the chief, dying during rites, 18 (of Chief
Ikaneng), 20 (Gopane), 26 (Thapuse).
folosa, those that bring down, 15.
fulaca, turners (of the enemy), 1.
fenya, defenders, 24 ? f.; a Ramphelana (of R.=Bogacwi, chief of
RES.) O:
fenyaKgosi, chief's bodyguard, 21.
fikang, making fast, 26.
ga-e-Kgomo, no cattle, 22 (27); gaenaKgang, having no strife, 22.
lo-gaga, the cave (collective name of regiment), 19, 23.
gata, tramplers, 8, 16 f., 24 f.
gasa, scatterers, 9 f., 13, 24.
gata-Kgomo, treading on cattle, 19, 22 f.
248 Transactions of the Royal Society of South Africa.
gana, refusers, 14; and a ba-Modimosane ba-Matau regiment, 1901.
ganelwa, those put off, refused, or quarrelled with, 8, 14, 16 f., 20, 22
(? ganela, refusers), 23, 25 f.
gana-Kgomo, refusing cattle.
gana-Mokewa, refusing the custom (i.e. of circumcision), shirking
circumcision, 16, 20 f.
gana-Tsatsi, avoiding the daylight (7.e. beasts of prey), 16.
godimo, those above, 16.
godiri (a plant-name), 16.
goiwa, those acclaimed, 17.
gala-tladi, shrieking like the thunderbolt, 23.
godu, thieves, 23.
golopo, 9f., who are asked ?
gora-Poto, lickers of the pot; a baModimosana ba-Matau regiment,,
1900 2
gwasa, roaring like the waters, 17, 19, 23, ? 26.
mo-golodi, liberators or blue crane (ac. to tone), 22.
“ gou-Kom!” (“ Come at once!” prob. a tribute to the emphatic ways:
of the Voortrekkers, or the Bastaards before them), 22.
German, c. 1814, 20.
gapela, seizing for (the chief %), 24.
hirios dani 7,
hachane, v. fatshwane, 22.
lo-haha, the (ostrich) feather (collective name of regiment wearing it ?), 22..
henyaKgosi=f-Kg, 22*, 27.
hlaba (v. tlhaba), stabbers, 13.
hl[wlana (v. thhwana), 1, (6).
hohu, herd of eland (phofu), 3*, 4-7.
ilwa, avoided, 25.
isakKoma, bringing the war-song, 8.
isaKgosi, bringing the chief, 21.
isaNkwa, bringing the host, 19, 21.
ipofhi, binding themselves, 18.
ilaMotse, avoiding the stad, 17.
ilaNkwe, avoiding the leopard, 8.
ilaKgang, avoiding strife, 23.
jaNko[u], those that eat the nose (sheep), 1, 8.
jane, little vessels, 14.
jaKgomo, beef-eaters, 14, 22, 24 f.
jaPoho, bull-eaters (? sacrificed before circumcision), 18, 24 ?.
jaTau, hon-eaters, 22.
jweTshepe ?, stones of iron, 15.
kaka, thorn trees, or the stout ones, 16.
keretla, bellowers, 1.
[k]galwa, those (de)cried, 1*, 2, 9f., 13.
kgari (? kgare, wreath of grass on head of carriers), 14, 25.
kgau, corn dried up in sun @, 2, 6.
kgOba [Motse], troubling (the stad), (10), 15.
kgomoca, comforters, stilling cries (? ironical), 14.
kgwadiba, ox-tripes, 9*.
Circumcision Regiments as a Native Chronology. 249
kewa, spitters, [10], 16.
koba, driving away in contempt, 2, 7f., 10.
Kompon (modern name “ compound ”’ of mine, etc.), 17.
kuka, the lifters, 8.
kwa, discoverers, 10 f.
kete-kete, so-and-so, 8 (-soswe).
lataKgosi, bringing the chief, 19, 23.
letaKgosi, awaiting the chief, 22, 25.
leta-Pene, bringing the vanguard, 22.
llelaKgosi, lamenting the chief, 17.
lat[l]hwa, castaways, (3)*, 6 (lahlwa).
lalaMetlwa, bivouacked among thorns, 8 (also called lallaDira, ambushed
for the enemy ; and lalAbilwe, sleeping in battle-order, in list 16*).
lau (the parts of) the lion (tau), 1, 19, 22 f.
lekana, mates, 4”.
letaNtwa, awaiting war, 16.
lekaNtwa, attempting war, 15, 19.
lobela, ? denying falsely, 11.
lomaKgomo, biting the ox, 15-17.
lwelaKgosi, fighting for the chief, 16, 19.
lwelaNaga, fighting for the land, 19, 23.
lwelaMotse, fighting for the stad, 8, 15, 17f., 20.
lokwana, bead necklace (perhaps orphans, so decorated ; the regiment
following Montsiwe’s death so called among baRolong), 19, 23, 26.
losaNkwe, leopard-slayers, 19 (1.-Tlou, elephant, do.).
loto, warning (or from a plant name), 12.
dila, sour milk, or the smearers, 23.
dima, cattle out at grass, 7, 8*, 9-11, 13.
dingwana, little cannibals, or whirlwind, 3*, 4f., 7f., 10, 13.
disa| Kgomo], herding (oxen), [10], 23*, 26.
dupe, fine rain, 15.
dikwa, surrounded (a forlorn hope), [10].
disaKgosi, tending the chief, 21.
dihaMotse, letting down the stad, 22.
N.B.—The D’s are placed here as the J sound in Suto-Chwana
is really a cerebral L, and connected with that letter etymologically.
metsaPhala, great snakes (lit. rooibok-swallowers), 23.
nala, nail (? naila), [10].
nare, wild-oxen, ? 26.
naila, shins, 6, 9, 11, 13, 2 14.
nanne, rascals, 8, 16.
nkwe, leopards, 3*, 9f., 13*, 17.
nkwenyane (dim. of above), 3.
noko, porcupines, 22.
noga, snakes, 8, 18*, 19, 22 f.
ntwa, wars, troops, 23 f.
nousho, black, 1f., 7, 12.
ntwane, a little troop, 8.
ntshwabisa, those that grieve me, 16.
ngana (from a tree name), 4, 6-8, 10*, 11, 13*, 22, 24 f.
250 Transactions of the Royal Society of South Africa.
ngo[a]naPelo, stout of heart, 23*.
ngOpe, water-courses, 8, 17, 26 ?
ngolo, goat with yellow belly 2%, [10].
notwane, little hammers ?, 22, 27.
OpaThebe, those that drum on their shields, 15.
oketsa, those that add, enlarge, 15.
pula, rain, 26.
phutha, gathering, 24.
petu, necklets (? as part of uniform), 23.
pena-pena, the vanguard, 16.
petlwana, little adzes or skin-scrapers (so Brown, petlo), 9*.
phaswa, dappled (black and white oxen), 23*, 15 (phatshwa).
Portuguese, 17.
pondo, from “* pound ” on appearance of sovereigns, 9.
pota, turning round, 11.
pulana, a little rain, 1 f., 12*, 14.
rataKgosi, loving the chief, 15 f.
rataKgomo, loving the cattle, 16 f.
ratwa, beloved (? latlhwa, q.v.), 3.
rema, the woodsmen, 4, 6, 17.
rema-ka-Cheka, felling with axes, 15.
retela, smooth, or very hard; a regiment of baModimosana baMaTau,
? 1902.
retlwa, the tree Grewia cana, do., ? 1918.
me-rileba, raw meat or ? maiden, 5, 9*, 11 f.
rutla, pull up (tree), 1*, 2, 3*, 14.
robaMetlwa, breaking the thorns, 15.
rooi- Baaitje, red-coats, e.g. of Major Warden, with whom baRolong
allied against Moshesh, v. 24, or after British took over Tvl., v. 26.
setlaoka, red ants, 16.
sKpa, putting things straight, 23.
s[h Jotla-Kgosi, scorning a chief, 17, (18).
sita[ Kgosi], outstripping (the chief), 22, (24), 27.
sitaPheho, outstripping the wind, 15.
swene, monkeys, (10), 14.
sonya, grinners, 7 f., 12 (=soswe in list 8).
soswe (mo-soso, a tree), 8.
sOkola, the passionate, 15.
sole (from soldiers, prob. in first Matebele War), 20.
swele, burnt (and so fearing the fire, shirking; but perhaps swene), 10.
senya, destroyers, ? 25.
teane, meeting, 16.
di-thaga, yellow finks, 3*.
theoga, descenders, 26.
thamaga, red ox with white back, 9*, 10, 13*, 14.
thametsu, making an enterprise too hard for someone, 16.
thiba, those that stop the way, 8, 14.
tekEKta, talkers, 1.
thibaKgosi, stopping the chief, 22.
thibaKgomo, stopping the cattle, 24.
Courcumcision Regiments as a Natwe Chronology. 251
thologa, overflowing, wasters, 19.
tlhware, pythons, 18, 26.
tlogela, forsakers (or tlh., shoots), 15.
tlakane, devourers, 4, 7 f., 9*, 11-18, 17.
tlatsa, helpers, 26.
tladi, lightning, (1), 2, 3*, 4-6, 10, 12, 18.
tlhaba, stabbers, 9,13; tlh.-tau, stabbing the lion, 22.
tlhaselwa, those attacked (? in mophato or circumcision lodge) by
CER, 235 20:
tlhwana, brown cows, 4 and 10 (if without h, snares ?).
tlhola-Kgang, continuing strife, 16.
tlhoma-Thebe, fixing shields, 16*, 23.
thopa-Kgomo, raiders of cattle, 23.
tlotlaKgosi, honouring the chief, 22 f.
thubaNtwa, breaking up the host, 16, 22.
thubaPula, rain, 23*.
thul[w]a, the hammered ones (hammerers), (1*), 8f., 11.
tsaKgosi, taking the chief, 18.
tsa[ea|Kgang, taking up strife, 17.
tshela, leapers, 6f., 9, 14 (¢ tsHla, very warm).
tshelaPhala, leaping the buck, 16f., 19%, 22 f.
tsetse, fleas, 23.
tsie, locusts, 14, 17.
tsuba, burning, 14; ts.-Tlali, do., lightning.
tuba, thumpers or “ guts,”’ 9*, (10*).
tuku, flame, 8, 10.
tswaKgotla, those that come from court, 16.
tshepe, men of iron, 6.
tsosa, rietbok (? tshosa, hartebeest-bulls), 15.
tsapa, hungry ones, 15.
tla-kaMokgwa, coming acc. to custom, 26.
tica, dappled (black and white spots), 16.
tshela-Tau, leaping the lion, 18.
tshola-Tau, lion-hunters, 18.
tlholaLibe, spying out bad things, 19 f.
ut{lwlaKgang, hearing (or tearing away) strife, 15.
N.B.—O and E are pronounced broad, ng as in sing, ¢ as English ch,
ch this aspirated, tsk as aspirate of ts.
The numbers refer to the mephato name-lists adjoined. When letters
of the regiment name are bracketed (as sometimes omitted), the bracketed
numbers and meanings omit them.
In the lists the small letter denoting each successive chief of a tribe is
suffixed to the name of his own regiment, and prefixed to that of the first
regiment indicated in his reign.
I shall be specially grateful for further information and lists from the
following tribes :—baTauana of Ngami, bagaMotlhware, bagaMananwa,
baKwenaba-Kubung, and baFokeng, baPhalane, etc.; baPhiring, baTaung
of Rustenburg, baNtwane, baNareng, baPo, baTlako and baModjadji and
baMamabolo, and from the Kaffir and Rhodesian tribes.
[Z'o face p. 252.
24, 25. 26, 27.
anes : baTAUNG. 7
baSELEKA. baRAPULANE. bagaMATLAKU. baTLHAPING,
er After 1830
800 a. Mosomi, 1 a. Mankurwane
b. Thapuse, d. 93 b. son Molale, d.
1920
c. Gasebone, 93-6, 2-13 ce. son Hendrik
d. Sehume, 96-02
e. Mosome, ii, d. 13
h. Koikoi o. Molekane f. Molatlhegi
j. Moroka p- Makgwe
k. Sefunelo, d.c. 30q. Matlabe
1. Moroke r. Shwedintle
m. Tshepinare s. Ramolekana
n. Samuel
hout baRolong (cf. 16, f. 19) ?
ngana ngana
abaKgomo aba Kgomo
a, 28 ntwa jaKkgomo (hachane)
do. ntwa (‘‘ gou-kom”’), 36
jaKgomo™ abaNkwe
r eakaThebe ? gwasa® (gaeKgomo)
rootBaartje ilwa ngana
gata, thibaKgomo ? senya? ? nare lau
busa® busa ? ngope jaTauP
fenya ? lokwanaP, 75 jaKgomo
jaPoho ? gata: fikang, 79 (notwane)
rootBaaitye
alosa gang, 82 lo-haha
tlakaMokgwat*, 85 d(henyaKgosi)
disaKgomo®, 89 bmogolodi, 85
tlhaselwa® fitlhaKgosi, 93 ganelwa
sita ganelwa, 97 nosi, tlatsat, 00 sita Kgosi, 96
gasa leta Kgosi etheoga, 06 tlotla Kgosi, 02 f.
eaka or apaPula_ tlhware, 09 thibaKk gost
gapela Skgare, 08 fpula, 12 henyaKgosi¢
phutha, 16 ganelwa, 16 letaK gos, 14
eakaThata
(anon.), 18
Trans. Roy. Soc. S. Afr., Vol. X, No. 4.]
[Zo face p. 252.
1. 2. 3. 4
2 . 5. 6. 7. g&. 9. 10. 1l. 12. 13 14 15. 16 17 18. 19. 20. 21. 22. 25. 24, 25. 26
baTLHALOGA. maGANANWA. baZOKW 3 : 3 4 ° : : 2 ; p 27.
OKWA. bagalf ASIBI. MASHISHANE. baKEKANA, bagaMATLALA. EGU bagaMAKAU. buPEDI. bagaMUTSHA ba MOSETLHA. baMPAHLELA. baKWENA, baMAGOPA, bamaNGWATO. baK WEN A. baTLOKWA, bagaMALEYE. baNGWAKETSE, AUR ONE do. bagaMOLLWA. bagaMOTLHUWARE. vaROLONG. baSELER A. voRAPULANE. eee AK. baTLUAPING,
a. Moshebudi Lebogo <exratt “Ls : After Molefe After Mathibi After Legwafana After Bogacwi After Powe After Moletsek, 90 * After TSIDI ; After 183
b. Ramanyoba Bae nen neh skontaemDs Mapela a: ee d a. Pheto, it Moshe Tulare, d. 24 Nehaupe Matsobane Sekwati a. Kgame, i Mocwasele Kgosi, 23 Mokgosi Makaba, d. 24 Diutluleng, d. 23 a. Thutlwa, d. ce. 1800 a. Mosomi, i a. AnD eae
ec. Mokgoba Mathome, d. 80 Mina Seeing ct 17 Matlala b. Pilane Motsepe : Malekutu, k. 26 Matsapane Mutle Mamogale b. Kgare Seitlamo Matlapeng Ikaneng, d. 02 Sebego, d. 37 Mokgatla b. Tawane, d. 51 b. Thapuse, d. 93 b. son Molale, d.
d. Seripa, reg. Kgadushe Kunwana isleras. Gly OF Mayotia c. Kgamanyane Seamoge, d. 88 Sekwati, d. 61 bro. Mathibe, d. 99 Matsobane Lerothodi c. Sekgome Legwale Gaborone Mokgosi,d. 17 Senthuthe Lenewe, d. 36 ; 1929
e. Seshigo Garant Waseueeaaat Paledi ; (bros. kd. by Tebele, c. 30) John More d. Kgame, ii Moewasele, k. 22 Molefi, hr. Seboko Gaseitsiwe, d. 89 a. Moilwe, d. 75 c, Montsiwe c. Gasebone, 93-6, 2-13 ce. son Hendrik
f NOtalanOTew LSS isos” Ae b d. Lenewe — Moemise Sekukuni, i, k. 80 son Salatiel Mutle Patule e. Sekgome, heir Sechele, d. 93 sonBathweng, d. 10 b. Sebogodi, d. 77 d. Wessels d. Sehume, 96-02
a, » Ke Alfred, ch. 17 Matou e. Isang, heir Alfred Mampuru, ii, k. 83 Solomon i Sebele, d. 11 sonSeepapico ¢. Ikalaheng, d. 94 e. Badirile e. Mosome, ii, d. 13
g Masanyane 1 Uru, i, ele, d. 2 c. Ikalaheng, d. 2 a: e. Mosome, ii, d.
tig Kgolokwe, d. 93 g. Sechele, d. 18 Gopane, d. 04 f. John, dep. 19 h. Koikoi o. Molekane {. Molatlhegi
g. Makveteng, d. 04 h. Sebele, ii Mokgatle a Lencwe g. Letlamoreng j. Moroka p. Makgwe
h. Kgoloane and cozz. ib Samuel k. Sefunelo, d. c. 30q. Matlabe
p Malekutu, d. 04 1. Moroke r. Shwedintle
k, Sekukuni, ii m. Tshepinare s. Ramolekana
1. Padutle n. Samuel
7 A —
@5 al eoeaule ?radipetsa kgwa?, 1750 dila ?, 1760
phaswa ? b thiba ?c. 1780 [swene (These from Rey. thametsu tshelaPhala
dimihanar cecele cecele® golopo N. Roberts) lalaMetlwa2 sepa®
nkwe ? fatshwana® tuba godiri gwasa
tladi 2 dima petlwana nala godimo? chologa
lat[IJhwa ?¢ gata tlakana> ngolo Pkewa tlakana® noga
dingwane ? chama >thulwa, c. 22 dikwa tlola ?, 1780 tlhomaThebe® gata (v. No. 8) no[a]naPelo 2*
hohu 2 firiP thamaga disa robaMetlwa gata betlwana eakaNtwa gagaP
hyana® rutla thulwa me-rileba kgwa] jwetshepe® firid... hiri (v. No. 8)? gwasa thubaPula
aa phaswae ngope a tuba _ tladi, 48 ? thula ? fitlha Kgosi thologa® noga begalaTladi
kgalwa di-thaha bdingwane, ¢. 35 pewelbe: ?, 46 evele adupe dalaMetlwa ?, 08 ? Powe’s noga hatane tshelaPhala throughout baRolong (ef. 16, f. 19) ?
rutla nkwe noga golopo kgoba’.. rema-kaCheka gaga tshelaPhala lau ?, 1820 ngana ngana
hula aE nanne, soswe®, kete-kete nkwe ; dthamaga ?, 63 butswana nkwe ?, 34 bbapoga tshelaPhala do. do. do. (v. 238-25), prob. before 1820 notoana abakgomo abaKgomo Shalini
ulaca lat{I]hwa °ganelwa, ngana tlakana, dima ngane ?, 69 thamaga utlwaKgang teane kodisa ntofaJ¢, metsaPhala,28 ntwa jakgomo (hachane)
ceretla (teketa) dingwane tuku (i), dima : gasa ?, 65 kwa?, 73 nganab opaThebe penapena chame ? tlhware® lauP caga no[a]naPelo, 11, 34 do. ntwa (“gou-kom”’), 36
ica *ohu ) ntsho (fenya Ramphelana, 63 tlhaba tuba ?, 77 lobela pulana btlakana sitaPheho ; ngope aoe abaKgomo IhabaTau gataKgomo, 38, or jaKgomo™ abaNkwe
jaNku rutla § z fatlha, 68 thamaga, pondo dingwana koba Sets gasace lomaKgomo do.,° c. 30 do. alpofhi gataKgomo tlhomaThebe, 44, or eakaThebe ? gwasa® (gaeKgomo)
ntshoP phaswa Tele sonyaP tlhakanad, 74 cecele [e]dimas ?, 81 pota fatlha, 68 dingwana lekaNtwac eganelwa, 38 noga lekaNtwad ?, 30 gaiKkgomo thopaKgomo, 51 rooiBaaitye ilwa ngana
au ntsho di-thaha Se Sue eee cee dkoba, isaKoma, 80 galwa tkoba ?, 85 edima echedi Kgari rata Kgosi, 40 nanne, 42 rema tsholaTau tlholaDibe nokoana Clokwana, 54 gata, thibaKgomo ? senyat? ? nare law
pulana pulana dingwana Gites ea pee tshelae jaNko, lalaMetlwa (lalla- tshela, 88 tuku?, 89 naila ntsho kgalwa suba folosad, 4 ganaTsatsi ganelwa® tshelaT'au gataKgomo osaTlou tlotlaKgosi, 57 busa® busa ?ngope jaTaub
kgalwad kealwa 8 iain cues 0 ts'epe 50.0 Lira), -Abilwe, 84f. kwa ?, 89 merileba yulana tsosa, 56 thuba) tsa’ Kgang tsa’ Kgosi? eakaThebe® ribaNtoa (kubaN.) disaKgomo, i, 61 fenya ? lokwanaP, 75 jaKgomo
niwanac, 73 eae tladie ne u tshela koba ntwane, lwelaMotse, 91{. Ime-rileba, 93 Stlhwana, 93 loto jakgomo lwelaMotse, 64 tlholaKgang, gwasa jaPoho lokwana ganena disaKgomo ii, 63 jaPoho ? gata fikang, 79 (notwane)
et{ladi, 83 tladi ratwa (lathwa), 76 ee 9 = a hlana ngana = tuba, 96 Mnkwe, 97 koba, 94 hlaba rutla emelwa, 07 canaMokgwa, 66 goiwa tlhware losaNkwe tlhoare rooiBaaitjc
bitsi, 93 dltoercienne, 94° : l i ” 20) sete ; leslie hohu aparaNkwe, kuka ‘ Dngana ?, 00 shela alola, 70 ntshwabis Portuguese tladi isaNtwae ?, 65 abaNtwa sitaKgosi tsetse4, 69 alosa gang, 82 lo-haha
athulae, 99 Sohu, 98” baRAMOKGOPA oe 8 dingwane, 04 bitsi dingwana HE, 55 (WO), kgwaliba, 03 gasa ?, 04 tlakana ?, 99 tlakana dima gana Aoketsa, 74 rata Kgomo CeakaThata, 78 eakaKgang, 74 tlotla Kgosi godu, 72 tlakaMokgwat, 85 d(henya Kgosi)
erut[lja, 09 Tutla rutla, 03 Shae ee Ane 4 gohu dima petlwana, 07 swene ?, OT tladi naila kgomoca lwelaKgosi, 81 fganaKgomo eakaNtwae tlotlaKgan, 75 lwelaMotse, 75 isaNtwa dihaMotse petu, 78 f. disalkkgomo®, 89 bmogolodi, 85
phaswa hacia, OF y ae dingwana, 0 naila tlakana4 cecelet Il 7 golopo, 10 tuba ?, 11 me-rileba, 03 pulana thamaga ganelwa - NelaKgosi ; eakaThata, 82 isaKgosi nesaPula tlhaselwae, 85 tlhaselwa® fitlhaKgosi, 93 ganelwa
1a LEE CE ten 4 : kgau, 08 fatshwana, 15 naila, 13 kgalwa ngana, 08 sonya, 08 ngana4, 09 thiba phatshwa ?, 87 rataKgomo, 85 fitlhaKgosi . henyaKgosi doanelwa, 98 sita ganelwa, 97 nosi, tlatsa!, 00 sitaKgosi, 96
cess elbielnennlisiye), alas etlhwana!, 12 2 as i ; sie kgobaMotse, 89 letaNtwa, 88 tsie, 90-2 eakaPula eakaPula, 87 ganelwa ?, 85 eakaNtwa chubaTladi ceakaThata, O4 gasa letaKgosi etheoga, 06 tlotlaKgosi, 02 £
nkwenyana fngana, 18 tladi, 19 tladi thulwa, 17 Inkwe, 15 jane@, 96 tlogela, Sip tlhomaThebe, 96 aparaNkwe, 00 lwelaMotse, 98 ; sole, 93 ; mokaoe eakaKgosi, OT = caka or apaPula tlhware, 09 thibaKgosi d ;
dswene, 99 kakasetlaoka, OL etshelaPhala, 96 ganaMokgwa, 96 [tsaakgang]abaTonatona yaChupa lwelakgosi ?, 10 gapela Skgare, 08 fpula, 12 henyaKgosi¢
chamaga, 02 tsapa, O4 tlogela, 04 Komponi, 07 eakaThata lau, OL eakaNtwa, 02 eakaThata gaenaKgang flaKgang’, 14 phutha, 16 ganelwa, 16 letal gost, 14
(naila, 06) thubaNtwa, 11 shotlaKgosi, 11 sotlaKgosi?, 10 ‘lwelaNaha, 08 bfitlhaKgosi, 04 ganaMokgwa etaPene ? betlaKgang eakaThata
sokola, 15 rataKgosi, 15 lwelaMotse, 15 f. tlholaDibe eakaKgang, 01 f. siea-ka-Lotlhaka lata Kgosi (anon.), 18
tswaKgotla, 17 ilaMotse ? lwelaKgosi, 16 German, 14 fenyaKgosi noko
lataKgosi, 19 Perekisi *
h(sitaKgosi, 19) disa Kgosi sepala
* T imagine that P. is the name of the present Inspector of Native Reserves, i.c. Purchase. (Note by the Rey. E. Silk, who collected this list.) The word usually means ‘‘ peaches” (W. A. N.). Mr. Silk also mentions that one legaekgomo was living in Noy. 1920. ‘That regiment was therefore no doubt circumcised well within the nineteenth century, even as late as 1840, Watshelaphala before 1820, and Manoga (cp. baNgwaketae and baRolong) near 1800.
NV.B.—Regiments in list 27 without brackets are of the baMaidi.
All except the italicised names are those of Ganyesa (Rolong ?).
( 253 )
SESUTO AND SECHWANA PRAISES.
By the Rev. Professor W. A. Norton, M.A., B.Litt.
If we allow a wider meaning to literature, and make the term cover all
art-forms of the spoken language, we at once obtain a whole range of such.
Open, for example, Carl Meinhof’s volume on the poetry of the African at
the title-page, and you will find the fairy tale, myth, saga, epic, hymns
and embryo drama, proverb and riddle and song, enumerated as examples
of what I mean. If these examples fall mostly below the literary level, in
the way of bulk, of our European poets and dramatists, the samie can by
no means be said (in the matter of finish) in the case of many of the royal
praises, delivered at dawn from the edge of the acropolis rock of some chief
in his praiser’s stentor voice, and re-echoed among the boulders which
strew the stad or astu. I had never realised the full meaning of the Greek
city state, with its central town and citadel, and but scattered villages
beside, through wide territory, till I saw the stads of the chieftains Sebele
and Lenchwe in Bechwanaland. The acropolis of the latter is an enormous
pile of rocks, like Pelion on Ossa, approached by a most narrow winding
path, tangled with bush.
To turn back from such chiefs to their hors literature (if I have success-
fully made good its claim to the term), of which they are in their kind, like
Alcinous of old, the patrons, let me illustrate the lithoko (or praises), which
to my mind have something of the heroic type of Homer’s epic. To test
this statement one needs, of course, to have an appreciation of them in the
original : let me give first an attempt to render the spirit of them in English
verse, and I will choose the famous story of Lethole, the material for which
was gathered by my old friend the Resident Commissioner of Bechwanaland
in the days when he was still in Basutoland. Lethole, chief of the
Makhoakhoa in Basutoland (early nineteenth century), had hired Zulus
to help him attack the brother of the famous chieftainess Mantatise. He
refused them their share of the spoil, but they caught him later and sen-
tenced him to death. He begged them first to let him sing his own praises
for the last time, and they stood listening.
254 Transactions of the Royal Society of South Africa.
Spellbound stood his foes
Listening to the death-marked warrior :
Off, off he goes,
Like an arrow from the bow-string,
Shoots amid the throng,
As they stand aghast a moment,
Then raise a song:
They a song far other-sounding,
Snarl of cheated beast :
Start the men, and dart the heavy
Spears, seeking feast.
Fast they fly, but he flies faster
Over field and fell,
Yet one fleeter than the others,
With dart so fell,
Reaches him, all but outracing
That tremendous shot :
Reaches and his back transpierces :
Blood outbursts hot :
Down it streams, but he pants onward,
Coughing out red spume,
Recking not his ebbing vigour,
Counts but the room
That he sets between the bloodhounds
And his flying feet :
Once he turns—far off they follow,
Less fleet ... less fleet...
So he staggers down the burn-bank
To the water’s edge,
Shelter’d from those grim pursuers,
Hid in the sedge.
For a moment he emerges
On the opposing bank :
Once again his strength he urges—
Slow, slow he sank,
Pouring out his latest blood-stream
As the warriors came,
Singing with last breath the praises
Of his doughty name.
(Southern Cross, May 1915.)
. Khomo e
Sesuto and Sechwana Praises.
255
No. 1. Lethole’s song was kindly supplied to me by Mr. J. C. Macgregor,
C.M.G., Resident Commissioner of Bechwanaland, when A.C. of Leribe.
It runs as follows :—
Lethole le thunyang,
Le thunyang khomong tsa batho,
La re ho thunya, la apesa pelesa.
Hlabeli, tlola Rakhabane a bone ;
A bone, ha ho qhalangwa menaila.
Khwahlan’a maroba
Mokhaloli-qenatsan’a maroba,
Mokhaloli ba o hlobile sesela,
O ts’o tjekela o le feela.
Hlabeli tswetse ea se itja mohlana.
Bashanyana ba llela sekaqa.
Ha lla Posho le Lenyolosa.
Mohlankana oa ghoba nku ka letolo,
Hlabeli,
tshwana ea ngwan’a
Lechesa
K tsamaile ea sehlaka matsika
Mor’a Lechesa kolobe li ea jana,
Kolobe li ea jana Bolaoaneng,
Ka mona ka ha likubu matshaneng.
Lefuma (-o 2) le kwana haMahemane,
Ha rwesa mekoli khomo e tehehali ;
Thokwana ea rwala shwahla mola-
leng.
L. the Dust (archaic for Lerole),
which shoots up among people’s
cattle,
when it shoots up, it covers the
beasts.
H. (2.e. L.) leap, that R. may see,
may see when the vanguard have
been dispersed.
A strong man of the maroba (L.’s
circumcision mates).
M. (a bird white and black) [tail ;
you singing bird, they pulled off your
you sing without it (7.e. have no
pride).
H. as a cow just calved ate the after-
birth.
the boys cried for a lump of it,
there cried P. and L.
the young lad H. drives sheep by the
lightning,
the black cow of Lechesa’s child
went and beat the M. (7.e. Maroba ?)
Son of L. the pigs eat each other.
They eat each other at B.,
Here in these little lakes in the
hippos’ place.
Poverty (? wealth) is at M.’s:
Bunches of beads are tied on a cow
(? a maiden).
The fawn-coloured cow had plenty
(of beads) on her neck.
_ To end the Sesuto part of my paper, I give No. 2, an ordinary impro-
visation of a South Mosuto boy praising himself (Mogapunyana of
Vereeniging, son of Masopha, a Motloung of Basutoland) :
I am the brindled one of Ramatsikitlinyane,
The child of the wife of Khwali :
Take the ox’s horn, and pluck it earthward,
Let the lads run with it.
They said: “ The black kine are not lean,
But in summer they trouble us ever.”
I picked the bat off the willow tree,
I trod in a slippery place and slid :
I came forth carrying the men’s shields in a pile:
256
Transactions of the Royal Society of South Africa.
I slaughtered the lamb, and divided it among the elders.
But the lamb choked one of the aged ones... .
Then he took the dipper to the stream,
That he might drink and swallow. |
(Contributed by Legabo of Cape Town.)
The original is as follows :
Ke nna Thamanyane ea Ramatsikitlinyane (i.e. leopard)
ngwana wa mosali wa Kewali :
nka lenaka o le kgolele fatshe,
bashemane ba le nke motiitil.
Ba itse “* Tse ts’wana ha li ote
empa lehlabula li hlole li re khathatsa.”’
Ka thwalla maborokoane fateng sa Moluane,
ka hata lereli ka thella:
ka tswa ke sikere lithebe tsa banna sekwankwetla :
ka hlaba konyana, ka ea ka e abela magheku :
eaba konyana e khama leqheku ;
eaba le kakalatsa liho ho ea nokeng,
ho ea theosetsa nama.
No. 3. A seTaung piece:
Lesa, buabua lechabana :
Mosali o lebetse Lekete RaMpioane,
O lolobetse a ea tlung.
A ba a tsoha ka meso a le hleka.
Lesa, u hlomme thebe thoteng :
Bashanyana, le se ke la e ts’oara,
Ke thebe ea molisana Selemo.
Lelothoane, mphahle, ke ee holimo,
Ke ee bona likhomo tsa mokan’a me.
Ka kena kahare, ka otla lekaba
Le maoto a meleka.
Se ntshoere, Phakoe-e-tshoenyana
Ea selemong ha Motsetse.
Nonyana ena, ho thoe’ng, ke Ra-
Potieti ?
Ke re, ke eromme mollo ka morung :
E ne e fihla e qhonokha molomony-
ana.
L., tell a varied tale :
the woman hath forgot Lekete, the
father of Mpioane,
She goes straight to the house :
but she rose early and bright (pure).
L., you have planted the shield on
the hill :
Lads, touch it not,
it is the shield of Spring the herdboy.
Salvifolia, let me brush past thee and
Bo Wp
to see the cattle of my mate.
I entered within, I smote the ox (to
be slaughtered at his marriage)
with a crooked shin.
It seized me, little white hawk
at the precipice (of the Viervoet) at
M.’s place. (M. was headman of
the village at Orange Springs, near
Modderpoort, O.F.8., and grand-
father of Enea, now dead, who
supplied the piece).
Why is this bird named R. ?
Then I sent it for fire to the forest.
It was pursing up its sulky little
mouth.
Sesuto and Sechwana Praises. 257
Though these are not a chief’s praises, | insert them as another example
of the poetic effort which is constantly being put forward by native bards,
professional and otherwise.
I will now pass to the praises of the Bakgatla chiefs, and begin with those
of Kgamanyane, the father of Lencwe, and those of his father and grand-
father. This was that Kgamanyane who was sjamboked before his people
by Herklas Malan, the veld-kornet of Rustenburg, as related to me by the
widow of the chief, daughter of the great Moshesh. I find myself wondering
whether the veld-kornet’s friends commemorated his work as worthily
from the literary point of view as the people of the man he sjamboked.
At any rate, the incident should remind us that, as there was another side
to the heavy-handed official of the “* Bushlice ”—for such was the elegant
soubriquet which these praises fasten upon the steadfast foretrekkers
(doubtless they had others for the rooinek, and even the missionary)—so
there was another side to the ow kerel or schelm to whom, as he thought, he
was administering due chastisement, for complaining, on behalf of his
people, of the hardness of the tasks they were set to. It will be remembered
that the Boers demanded labour from the natives in return for the protection
of their better arms against the raids of Moselekatse. Hence the difference
of opinion. To the honour of the Dutch Church be it said that, to this day,
even the independent Bakgatla of Bechwanaland retain their mission
among them, in spite of their disagreement with the Dutch of the Transvaal
as their political overlords.
The Praises of Kgamanyane the sjamboked run thus :
INO: 4:
Kgapetla e tlhaba-tlhaban thata a
Masoswe a Kgosih,
o sere o tlhaba ea ekete o tlaletswe,
etswe le metlhef, o se ke o tlallwa.
Makgalimotse o sefega sa nkwe,
o kgeleli ea tau,
mogats’a Likolo a Ramontsana-tlou.
Tuba tub&é ea Makhinya-khinya-
khinya,
eare he le tsamaea,
tlhako se kgopa se senwe sa.
lekhinya.
Lekobela le tsoga kwa lefiolen.
Sharp-plercing shard of the royal
Masosoe (kg.’s own regiment),
smite not as though in anxious
uncertainty,
perchance sometime thou’rt anxious:
be not so.
Fast-striding M. with the leopard-
crest (or breast),
with the lion’s sear,
Spouse of L., Ramontsana-tlou’s
daughter ;
(praise-name of Kg.)
When he walks,
One shoe strikes against the other
of Lekhinya :
but the supple knee-bender rises
from the knee.
258
Kwa seleteh go iwana o monana,
go legora le leletsa Molefi le rago-
Mojalefa.
E ne e-na le Ramolibe tshukulu,
tshukulu ea ga ka le Ramorontse,
eare e fitlha fa Mankwe, ea tlhapa,
ea ja taka, ea boea ka matsogo,
ea ba ea boea ka likgokono,
tshukulu ea ga ka le Ramorontse.
Magoapitlana, fiwana-mahulu wa ga
Kegafela,
fwana mahulu, tshaba tlhaga: O le
man ?
O le lifako li nkwela gale,
tlhagana tsa marallana.
Motho, ke tshaba wa selepe ;
wa molamu ga ke mo tshabe :
a ka re “ tou,” ka hunyela,
ka hunyella legapen.
Transactions of the Royal Society of South Africa.
Where the good land is, there’s a
new (born) child (7c. Keven
Pollo), )
there is the hedge to _ protect
Molefi and the father of the Heir
(2.e. R’lane).
(Kg.)
The rhinoceros
Ramolobe,
my rhinoceros and Ramorontse’s.
When it arrived at Mankwe, it swam,
it ate lime, which came up its arms,
Yes, it came even to his elbows,
(the elbows) of my rhinoceros and
78:
was with
Walker like the tortoise, Nwana-
mahulu of Kgafela,
Child of the tortoise, shun the dry
grass, for who are you ?
You are the hail which has fallen on
me longsyne :
You are the small dry grass of the
hills.
‘* Man, I fear him of the axe.
him of the kerrie I fear not :
he may smite, but I pullin my head :
I pull my head into my shell!”
Nearly every line is crammed with allusions like the lines of a Greek
chorus or a Hebrew prophet, and those who cannot follow much of them
must allow for their effect.
even in my rough version.
Benjamin in Genesis :
Judah is a lion’s whelp :
I hope, however, something of it may be given
It is very much that of the blessing of Judah or
From the prey, my son, thou art gone up:
He stooped down, he couched as a lion,
And as a lioness: who shall rouse him up ?
Benjamin is a wolf that raveneth :
In the morning he shall devour the prey,
And at even, he shall divide the spoil :
or that of Joseph in Deuteronomy :
The firstling of his bullock, majesty is his :
And his horns are the horns of the wild ox:
With them he shall push the peoples, all of them.
Sesuto and Sechwana Praises. 259
No. 5. I add the praise of Pheto, grandfather of Kgamanyane :
Ke Ramopyane a baKgatla, Iam R. of the Bakgatla,
ke tshukulu ea ga Mphetei': keeme. the rhinoceros of Mpheteng, let me
be!
Pheto a Molefe, ke ratwe ke batho. _P. son of Molefe, I am loved by none
(ke ratwa ke mme Moliane) : (save by my mother M.).
ke tshukulu ea ga Mphetef, ke eme. I am the rhinoceros of Mpheteng.
Ge le sa mphete, le-tlhola-lillo, Let me be; if you do not pass me
by, bringer of lamentation.
ke tlholla bomaeno go beolwa. I do make your mothers to be shorn
for grief.
The last line refers to the quaint custom by which bereaved women
shave the head, as in Bible times, except for a tuft or tasselling of hair-
strings from the middle of the crown, having at a short distance the effect
of a football cap. The Praise concludes :
Ramopyane, o se be o tshwaetse : R. maybe thou hast made an end
(of a wounded man).
kgosi ga e tshwaele, e nna fela : Tis not for a chief to do so: he lets
be:
(Repeat these two lines, adding ga Raseakanyo =at R.’s place [’tis so].
These of Pheto’s son Pilane complete the series of three generations :
No. 6:
It is Pilane, the rock of the anvil stone, the slippery one :
Those who touch it will leave their fingers,
And that Mabine has proved.
I am Pilane that makes to war (ke P. a malosa)
The beast of Kopong and the wood of Litlhotlhe.
(Selo se mo K...L.)
(2) The common Folk are poking at his hole and annoying him :
The beast has been annoyed by Mmamorogoana of the Mabulisa (vassals
of the baKgatla).
He is thrusting forth his head, he pecks savagely,
And returns to his place Kopong....
(1
~—
The original runs :
(1) Ke Pilane, ke tlhapa la ntswe pilwane, ke letlhapa
le le boreleli Pilane: ba le tshware, ba tla
tlog[el]Ja menwana: le tshwerwe ke Mabine, a tloga meno.
(For the rest, v. Dico, p. 77.)
Ba lintlha ba nntse ba se gwaisa, se gwaisitswe
ke Mmamorogwana oa BaMabolisa, se ne se ntsha tlhogo, se ba
thObonya, se boea se boela kafa Kopon,
kafa Kopoopooi, kafa Litlhotlhe. Tau makunya a ga
ba-Phalaphala, etc., as in Mr. Wookey’s Dico, p. 78.
(2
“~~
260 Transactions of the Royal Society of South Africa.
The last lines (from Tau . . .) may be rendered :—
The roaring lion of the flood of peoples,
Descending from the smooth rocks of the monkeys,
Roaring and hastening to divide the spoil.
When he arrived at Selitlwe,
He roared, and then silenced his voice,
This lion of the Hyenas.
(Pilane’s circumcision regiment were called the Hyenas, maFiri.)
After this my informant adds :
Nwaketse babeli bale ba kgopywa ba wa, ba kgopywa
ke ese ke be gaufi: etlare go le ke atamela Pilane,
rumo le tla nwa mali a Setate, ntsi e tla nwa
bobete, Pilane, segen ga Nwaketse a Morara ntwet.
This means:
Those two BaNwaketse are trembling and fall before
I could be near. Now, when I Pilane approach,
the spear shall drink Setate’s blood, and I Pilane,
as a fly, shall drink his gore, in a lake at the
place of Nwaketse, descendant of Morara—in battle
(in which Pheto has sent his son P. to aid the
Bakwena. Setate is a Newaketse).
No. 7. Sekgatla Praises by Seatile Meshake Lebotse, praising himself :
Ke rile ka bopyoa senare, ka ba ka bopyoa setau, moroa kgosi ;
When I was formed, I was formed with the nature of the wild ox and the
lion, a King’s son ;
phatlana ea nna mokgeletsanyana, meritsana ea ea le litomego :
my forehead has a groove, with the hair following the cupping-mark :
ke palame kota ka bala motse ka re: “ Se ga se motse, motsana.
I climbed a tree-stump, I took stock of the Stad, and said: This is no
town, but a village :
Ntoa e ka tla nasuta, e ka suta: ke apere metja, ke ila lerole
War may come nigh, yea, come nigh.” I put on my kilts, I shun the dust
ka makutung, ke sa gate lefatshe, ke ila le motho a nkgoma
from the feet ; nor do I kick up the earth, I shun the touch of men
ke feta, ke be ke tsitsibane, ke be ke batle go omana,
in passing, I condemn, and am apt to curse,
potepote le motho, ke mmolaee, ke mo tseela thebe.
and to chase one in and out, that I may slay him, and may take from him
his shield.
Sesuto and Sechwana Praises. 261
Lithebe, re ne re sa li pege, re ne re li kgoagetsa feela
The shields, we never hang them up, but just within reach we hitch them,
re li pega mo mopiping. Ke nna mosimane oa Mokongoana oa Mamokoena
oa Seroka.
hanging upon the pipe tree. I am the youth of M. of Mamokoena’s house,
the daughter of 8.
(The penultimate sentence means the weapons are always ready.)
No. 8. Setoutoe’s Praises of Senelo, his grandfather, who went with the
Mafirt regiment to Ngwato (being himself a lelima), saying :
Ke e’o bona koa bana ba ea teng. I am going to see where they are going.
Mamalima o ja ka legetla, o ja ka perepe ea legetla. Moth’o itaea ka
patla oa
Mamalima eats with a shell, she eats with a big spoon of shell. A man
hits with
Malima, o othibosetse chaba, li tla be li go lemoga, baSeame ka baRatlhagana.
the stick, he of the M. (regt.) he prevents the enemies coming to gaze at him,
both baS. and baR. (in B.P.).
Ba re, ““ A! merafe ea Boroa e bilioa fela, e le mapshega: A! e bilioa fela.
They say that the nations to the South (from Tvl.) aresimply cowards. Ah!
(b) Ga le ’mone a Molefe (7.e. Senelo) : ga le mmone, o bapile le lona :
You do not see M.’s son, he is by your side:
a ea tshoga ka e le kgoelo e kgoeloga botlhatsana :
it happened, as he darted from the bush,
Mogale a phofa tsa kgaka, Mogale a liphofa,
the hero of the kgaka’s wings, the guinea-fowl feathers suit him well,
li ea mo tshoanelatshoanetse, mogats’a Nketso a Kgetse. (Note the rhyme.)
the husband of N., child of Kg. (Senelo killed his cousin, also a hero : there-
fore called Kgoelo the darter. Both were leaders of the Matlhako.)
We pass from Lencwe’s people to the Bakgatla ba Makau, between
Rustenberg and Pretoria. Moewise (b. c. 1845, father of the present chief)
and his brother Molikwe were sons of the old chief Leamoge, called in
Dutch, for good reason, Sjambok ! (1828-88).
No. 9. Sereto sa Seamoge :
Kgomo e tshopha ke golile, I am an ox with spots and am grown,
lilekana, ke golile, ke luma makg- the spots are equal, I am so big, I
wata roar wherever I go (or hoarsely).
I insert the interpretations given locally, even when I am not satisfied
with them. I also preserve spelling as given, which is often an index to
dialectical phonetic.
20
262
No. 10. Molikwe’s Praises:
Phekola ea sebata,
moloko oa banna, ga te itse go leka :
fwana oa Matlhali ka tsa bogale,
ba nchupetsa mankwe :
nwana kanthe kewali kwa bosa eabo
ga li fele.
ke phuphutla thamaga,
moshemane o tswetswen le Moemise.
Phata ea likewali ka gobala,
phatagali ikgwal ka gobala.
Nos il:
Mo Thubatse motho o tswanh mo
tlhake[h ]
motl[h]ak’a morota wa masebutla sa
limo :
motho wa bolabola, molohwana wa
maitsibolla :
o seba ga le mmone, go sa le [le-]
phetha a le tjeki.
Transactions of the Royal Society of South Africa.
Charm of a beast,
I belong to virile men, I know not
to attempt :
a child of the M. regiment that came
with valour,
a man that can fight with tigers,
child that is like a partridge in the
morning when it trips along.
I prod the red and white ox (after
which one of the regts. was called).
the boy that was born with M.
I was wounded (but still went on,
with my wound, fighting).
striped with great wounds like a
partridge.
There at the Steelpoort River is the
man that comes forth from the
reeds, the reedbed
of the kloof of him who can face
giants :
the man talks and talks, the little
mouth that talks first :
he reproaches, but you do not see
him, he is like a bead so small
(you do not see it on the ground).
We now turn to the North Bakwena, and first we have the praises of
Moewasele II (killed c. 1820).
Now i2F
Pholoholo ea Bo-tlhapa-tlou, ea noka ea Metse-motlhaba, ke e tsetlha,
The beast of the elephant-bath of the river of troubled waters is yellow,
ee metse e li matla: e tlhotse e gwelelwa ke lichaba, e gwelelwa,
whose mane is stiff: the livelong day it is shrieked at by the tribes,
e phamotse maralu.
it seized the milch kine.
a fha lira h roba magolelo,
when the foes crush the herbs,
fha temofi ea ga Ramatokwane ?
in the garden of R. ?
Nna le boPolile re bo-eben :
I. and P.’s men are in a quandary
Lira tsa ga Kgari linyana ?
Are the foes of K. (old Kg. of Nwato) few ?
Sesuto and Sechwana Praises. 263
re lebalanye :
and forget each other (? lebelanye, look at each other).
fhala le mo pine? :
only you are in the dance:
pelo go setse go utlwagala
in the heart there still is heard
mogatlhare wa ga Tsamaa-gotlhe,
the chatterer who goes everywhere,
Raseitlhamo aKwena
Seitlhamo’s sire and Kwena’s son (2.e. Mocwasele IT.).
No. 13. Praises of Sechele I, son of Mocwasele II (a plea for the
geception of the baHurutse, baKgatla, etc., when fleeing from the Boers
mm 1852) :
Re na le motho, o Neokencoke : We have a man N. (the Subtle, 7.e.
Sethele).
Kare fa lekwalo le tla le befile, If there come a letter with bad talk,
Le eene a kwale la gagwe, a re: He also writes his letter, and says :
Tsafe]a, o le ise, Masholobotlho, Take it and carry, M. (S8.’s old
servant).
Batho se bao: a ga ba bone barwa Here are people: don’t they see
rrabo BaHurutse 2 their friends the BaH. ?
Batho ba ea re[ng] bafa] noa kofi, (There are) people who, when they
drink coffee,
Ba ba nee morogo ba o nwe. give them (baH.) dregs to drink.
Shupegetsa ba-bina-kgabo botshelo, Show mercy to the monkey-dancers
(ad.).
Ba tle ba tshele ka wena : That they may live by thee !
Ba go thele * madi a kgofa (ba fete To bring on thee blood of the Bush-
mogocoana), louse (z.e. Boer)—Let them pass
the cupping bowl.
EH bee re dimo a sale a ya wena nosi. Let the cannibals (¢d.) eat thee up
alone.
N.B.—The baTlokwa, baHurutse, bagaManana (sometimes ranked with
the baKgatla), bagaMalete, bagaMokibilu, and baTlhako all took refuge
with the baKwena under Sechele from the Boers.
No. 14. Lesike’s Praises of Sebele I, fighting the Bamangwato under
Macheng (soon after 1860), at Kgama’s request :
Kwena matuduanye wa ga Sechele, The diver crocodile of Sechele (fr. of
Sebele),
Matuduanye wa ga rraSebogiso, The diver of Sebogiso’s sire,
* Thela, seKwena dialect for tshela, to pour (broad e; with shut e, means to live =
phela).
264 Transactions of the Royal Society of South Africa.
o tlhakantse batho le Matebele,
Kare ba lebile ditima-modimo,
A bo a e-tla a tlhaola Matebele,
A bolafe]a, a ba latsa mmetha :
Ka ba ka tlhoma ke re “‘ Kwena”’:
Ka khutla, ka bua se sele, ka re:
Tsankole, pholoholo,
Ka setlhabela-manong,
Manong a a kwa Mabele-a-podi,
A re a fhofha, a fhofha, a khubidu,
A ya thata ea ga Phometsi-a-kwena,
Ka ga matuduanye wa ga Sechele,”’
Matuduanye wa ga rraSebogiso.
Merachwanyana ea Serowe,
Masilo a ntshitse matlhwana a mo
lebile :
‘““ Kgama e tla bona kae botshelo ?”’
K kile ea raka Kwena letsibogo :
Moseya ole o mfhera,
O thata:
Ba tla tshwarwa ke mfhera-mangole.
Ke Nao-dintlha wa ga Mocwasele,
Nao-ditelele wa kgosi,
Mosii mosii wa dira,
Mokgarametsege Malope (? Molope)
Metseng ea batho
Fha gare Mahutagane le Kurwe.
Confounded the folk, even the
Matebele,
When they beheld the Ditima-
Modimo (? regiment).
He came and cut down the Matebele,.
Killed them, laid them in heaps :
And then, I had said “*‘ Kwena ”’ :
I stopped and spake otherwise :
“ Quick little jumper of a steen-
bok,
The beast the vultures stab,
The vultures of Teats-of-the-goat.
(Mtn.),
The red vultures, which fly, fly,
Which eat the power of the Unafraid
(Prover of Leviathan),
Of the diver of 8.
The diver of S.’s sire.”’
O ye small tribes of 8. (Kgama’s.
present stad),
M. gazed on him with all his eyes :
‘** The Hartebeest, how will it live 2 ”’
It crossed the river before the Kwena
came :
The further side it is very narrow:
“his hard:
They will be caught by the knees.
I am the active one of M.,
The long-footed one of the chief.
And overtaker of the foe:
The forward Malope (little bird with
long tail).
In the cities of men,
Among the narrow places and at K.
(Both in the east of the Bakwena country).
Batho ba tshabile tladi ea makau,
Ba tshabile tladi ea boMashamatse, *
Pholoholo ea boGafiwaiiwe (Seb.’s
sister).
E se gadima lwa bo-Motshwane a
Mmaitle,
La Makatapitse a ga Botshabelo.t
The people feared the lightning of
the youths.
They feared the lightning of the men
of Mashamatse.
The beast of Gafiwafiwe’s folk :
"Tis the lightning of the men of
Motshwane (son of Mmatle).
Of the Makatapitse regiment of the
Stronghold.
* Elder brother of Baruti, Sebele’s cousin. Most of the names are of members of
the Bakwena royal family.
+ Sechele’s stad as a city of refuge for the tribes mentioned in No. 13, note.
Sesuto and Sechwana Praises.
Se dule se nakaletse selemela,
S’wa godimo, soo Kgosidintsi
Le Seadiiwana a Mocwasele:
Se o'le, go sa ntse go itebetswe,
Fha gare ga mebilana ea batho
Ha ga Mangwato le ea Bokalaka.
Le-fhera-batho la boKgabo le
Keakge,
La dikgosi: dikgosi boMotswane a
Mmatle
Bokgaimena la Legoyane :
Nfhere wa dipholoholo le batho,
O kile o kile wa patika boTlhankane.
Bagamangwato, lo tlhasetswe eng 2
Lo tseneletswe ke’ng ?
Maemong lo tseneletswe ?
Ke le-tlhanega batho, ke kolwane la
Matlhorangwe :
Sebele ke lekatapitse la
Tumagole :
Ke tlhorufiwa ea boKgari, a Sechele.
(2? loo)
O e-tla, le-bola[e]a, le go rebola, wa
ga Sechele :
O e-tla: a bolafela, a ba a lesa:
Ke rafela, gobo a lesitse Moatswae,
A lesitse MorwaKgomo, a reboga.
Kwena, mogolole, ke ea go leboga,
Ka go bolaea le go rebola :
Kgosi o jafhile seng, wa lesa Ma-
thubi ?
Kgosi e busa ka bana ba merafhe !
No. 15. Another Praise of Sebele:
Mochacha mogakatsa-mala,
More mojewa-o-botlhoko :
O ka jewa ke monna a sa 0 itse,
O ka mo tshwara, wa mo kgobola
mmele,
mere ea tla ea fela, re ee pele;
265
The Pleiads are out and shine,
They fall from above, they are on
K.’s side, .
On Seadingwana’s,
daughter :
They fell unexpected *
Among the streets of men
In Mangwato and Karangaland.
The disappointer of Kgabo and
Keakge’s men,
Of the chiefs, the chiefs of the party
of Motshwane, Mmatle’ sons,
Of Kgaimena and Legojane ;
One who broke both beasts and men,
and once pressed the’ folk of
Tlhankane.
O ye Mangwatos, what attacked you?
Ye were approached by what ?
In your steads invaded ?
"Tis the Leaver-of-men-dead, ’tis
the youth of the man that sits
but little.
S. is a lethubantwa of Tumagole,
Mocwasele’s
He is the rained-on grass of Kgari’s
men (son of Sechele).
He comes, the slayer, yet to deliver
even Sechele’s son :
He comes: he slew, again he spared.
I say it, in that he spared Moatswae
(a Mongwato),
He spared Moatswae (son of K.), and
delivered him.
Kwena, my elder brother, I thank
thee,
For slaying and for sparing !
How well didst thou, chief, when
thou sparedst M.
The Chief rules by the sons of the
nations (spared by him).
Bitter herb, which poisoneth within,
Physick bitter in the taking :
Should it be eaten by one unknowing,
It should seize him, bruising his
body,
And medicine would not suffice,
though to hand,
* Ts this passage connected with Donati’s comet, 1858-9 ?
266
ha fela boshokwe le bomakgoro-
metso,
Kgomo e lela kwa Phapane,
e lela e re “ Sebele we!”
e tlholetse bo-Ngwato go senyeha.
Me a thutlwa tse ke tsa molapo wa
Sehoka ?
a di neilwe Phoka ?
Ga di tshwarwe,
le ha kgomo tsa bangwe di tla tshaba,
di tlogele marole mo tlotleng.
Transactions of the Royal Society of South Africa.
Neither boshokwe nor bomakgoro-
metso.
The ox lows at Phapane (Hill).*
It lows and says “ Sebele.”’
It caused boNgwato to be de-
stroyed. (Oxen are the usual
native casus belli.)
Are yon giraffes those of Sefoka’
vley ?
Have they been given to Phoka ?
Nay, they are not to be caught,
Even if the cattle of others flee,
And leave their calves in the ruins.
No. 16. Yet another Praise of Sebele :
Mashweu-shweu ao a ya (ao)
tlou tsa Mothathe, masepela phalo :
Tlou l kile tsa etela Sebele, eo o
Sechele ;
o tsetsoe[g]o fhenya lichaba;
re raea ka a fhetlha Mokmakanenh ;
le ga maNwato, o tla go fhitlha :
me eare fha kgomo tsabo li gapiloe,
a bile, a samile cheka,
Mogats’ a Motshipi.
Those white men are eating up
The elephants of M. They are those
who outstrip in their going,
The elephants paid a visit to Sebele,
Sechele’s son.
He was born to rout the nations.
We mean because he pierced to M.
and at maNegwato shall he arrive :
and when their cattle are taken, _
he made a war-axe his pillow,
that spouse of M. (? Sebele I.’s wife).
No. 17. Leboko la ga Mpelege (by himself on his deeds in the Kwena-
Kgatla War, ’75-9) :
Batho ba yelwe ke Hauhau-koma :
Ba yelwe ke tlhone rea tlhabana.
Ka erile re se na ho tlhabana ntwa,
ra ba ra tlhonama ra nna fela.
Tloga, o ba abe, morwa Mocwasele,
o ba abe,
O ba abe le makau :
sefofatlele se leka macomane.
O pelo-kgale, fwana wa ga Mmopi :
O pelo-kgale, o tla digela batho.
Ke mmonye ka ntwa ea ha Kgafela.
O digetse banna ba le bantsi :
Tsela-e-mang o re, leke a mo etsa,
A tla a tsena mo ganong g’a dilo, g’a
di-ya-batho.
(Ke khutla foo Mpelege.)
Men are eaten by Hauhau (7.e. with a
crackling of bones)—’tis a secret :
They are eaten by sadness as we fight.
When we had fought our war,
we were sad and sat still.
Arise, O son of Mocwasele,
and marshal them with the youths:
aswift one (z.e. Mpelege) tries the race.
He is patient, the child of Mmopi :
He is patient, he shall lay men low.
I have seen him in the war with the
Kgafela.
Surely he hath laid low many men :
T. saith he will try to do likewise,
and get him into the mouth of the
beasts, of the man-eaters.
(The words of Mpelege are ended.)
* Close to Pilane siding.
( 267 )
NOTE ON A DETERMINANT WITH FACTORS LIKE THOSE
OF THE DIFFERENCE-PRODUCT.
By Sir Tuomas Murr, F.R.S.
1. The determinant in question, when of the 37¢ and 4°" order, is
1 a+b ab 1 a+tb+e abtac+be abc
1 b-+e be 1 b+etd be+tbd-+cd bed
| tl c+d cd\, 1 c+td-+e cdtce+de cde
1 dtetf detdftef def |,
the scheme of construction being readily grasped when the scheme for
the first row and the scheme for the second column have been observed.
Clearly it is such that when of the n™ order the number of variables
occurring in any row is n—1, and the total number of variables is 2(n—1).
2. Recalling the effect produced on a determinant by reversing the
order of its rows, we see that the function under consideration, F say,
is at most altered only in sign when the order of its variables is reversed :
for example,
F(a bcd) = —Fidc ba),
KRabedef)— Efedcbd a).
3. Performing on the 4-lne determinant the operations
TOW ,—IOW,, LOW,—FroOWs, LOW;—IowW,
we obtain
Fiabcdef) =—|a—d (b-+c)(a—d) be(a—d)
b—e (c+d)(b—e) cd(b—e)
c—f a \(c—f) de(c—f)
= —(a—d)( c—f). F(bede),
= (d—a)(e—b ieee F(b cde),
where it will be observed that the three factors on the right are got by,
as it were, subtracting
from the row d e f
the row a 6 e,
and the F on the right from that on the left by removing from the latter
its first and last variables.
268 Transactions of the Royal Society of South Afreca.
Exactly similarly it is found that
F(abcdef gh) = (e—a)( f—b)(g—c)(h—d) . F(be def),
and we thus finally reach
F(abedef gh) = (e—a)( f—b)(g—o(h—@)
- (e—b)( f—o)(g—4@)
(eof —d)
. (e—d).
4. The marked resemblance of this development to that of the alternant
[eahbl ct deie= ||
cannot but strike the student, the factors in both cases being all differences,
and the number of them in the one case being the same as in the other.
We are thus led to expect that the said alternant is derivable from
F(a, 6,. . ., h) by specialization ; and such is readily found and shown
to be the case. For, returning to the development at the close of the
preceding paragraph, and putting
Gull O10 26
we obtain
(e—a)(a—b)(b—c)(c—d)
(e—b)(a—c)(b—d)
(e—-c)(a—d)
1.€. (—1)® . (b—a)(c—b)(c—a)(d—c)(d—b)(d—a)(e—-d)(e—c)(e—b)(e—a),
so that our result is
Fabcdeabe) =| ab ed? e | = Glade de).
5. An interesting alternative mode of proving this last equality is to
show that
Ei(@ 0.60 era) bie). 4).@9 be cd? es ia bec ceales
Taking for shortness’ sake determinants of the 4" order we have
1 a+tb+e ab+ac+be abe a —@ a —l
1 b+e+d be+bd+ed bed b> —b b —Il
| 1 ctdta ced+teatda cda| : |c® —ce ec —I1
|1 d+a+b da+db+ab dab d? —d @ 1
(d—a)(d—b)(d—c)
(a= 0) (@—c)(a0)
c : (b—c)(b—d)(b—a)
(c—d)(c—a)(c—6)
=-.| abe d 2;
Determinant with Factors like those of the Difference-Product. 269
and consequently the multiplicand
== ——| 0° bt cd? |.
It is also just worth noting that the one determinant can be directly
transformed into the other, the first step of the process being to perform
on F (abcda b) the operation
col,—(a+b-+c-+d) col.
6. Besides the specialization
Jeed, 05, 0c
there are a number of others resembling it although of less interest : for,
without causing the function to vanish, we may put
jl —-¢ OL e,
gy — 1d Of Diote,
i — Gor bor Cc ore;
for example,
F(a bc deeee) = (e—a)(e—b)?(e—c)3(e—d)*.
7. Of far greater interest, however, is a specialization of a different
kind, namely, that which leads to a new determinantal representation
for the product of the binomial sums of a, b, ¢, . . . It is well known that
for representing such a product as
(a+6)(a+e)(a-+d)
. (b+0)(b+d)
. (c+d)
there is no form analogous to the alternant, and that in fact when the
said product does occur in determinantal analysis it appears as a quotient,
namely,
pa’ 2°62 d® | a? bt? a2 |
8. Going back to the result of §3 and altering the signs of the second
triad of variables, we obtain
F(a, b,c, —d, —e, — f) = (—d —a)(—e —b)(—f —e
or
(ri aoe) co) aC)
(Co)
= (—1)® . (d-+a)(e-+b)( f+)
(d-+-b)(e-++c)
(de) ;
and similarly for all other orders of F. In other words, By altering the
270 Transactions of the Royal Society of South Africa.
signs of the second n—1 variables in F(a, b, c, . . .) all the differences in the
factorial development of F are changed into sums.
9. From this as an immediate consequence we have
Kia, b, C, UE — a, ==) ay (d+-a)(a--6b)(b-+¢)
. (@+6)(a+c)
. (d+e)
== product of the binomial sums of
Gin ley On ye
and similarly for every other such product. We can thus formulate the
theorem that Lf in the new determinantal equivalent for the difference- product
of n quantities we alter the signs of the second n—1 variables in the functional
symbol, the resulting determinant rs equal to the product of the binomial sums
of the said n quantities : for example, from the equality of § 4
F(a, 6, c, d, e; a, b,c) =| a? bared? 7.)
we deduce the companion equality
F(a, b, c, d, —e, —a, —b, —c) = | a® b ct d® && | — | a® Bb! c? a e* |.
10. There is an entirely different way of viewing the product of binomial
sums, namely, as the eliminant of a pair of equations in x. Taking, for
example, the equations
(a+b+c+d+e)a* + (abe+ ... +cde)u? + abede = oF
a + (ab+... +de)a? + (abed+ ...+bcde) = 0
where the coefficients are sums of combinations of a, b, c, d, e, it 1s not
difficult to show that their eliminant is the 10-factor product
(at+b)(at+c) ... (d+e) . ; <2 SC)
One sure basis for proof lies in the fact that, if we multiply both sides of
the second equation by x and perform addition we obtain
(x--a)(x-+-b)(x+e)(a-+d)(x-+e) = 0,
and that the substitution of any one of the five roots of this derived equation
in the two original equations changes their left-hand members into the
product of four of the factors of E. Another basis of proof is the fact
that if in the original equations we put one of the ten factors equal to 0,
say a+b = 0, the equations are transformable into
(ba + ws e
e-+d-e
(x?-++ab)(x?-+cd-+ce+de) =0
and are thus seen to be consistent.
Determinant with Factors like those of the Difference-Product. 271
The latter mode is less readily generalizable than the former. To give
an indication of the line it takes, it may be stated that in the case of 7
variables the original equations
(a+...+g)a® + (abe+...+efg)x* + (abcde+ ...+cedefg)x* + abcdefg =
a + (ab+...+fg)z* + (abed+...+defg)x? + (abcdef+...+bedefg) =
are transformed into
(x?--ab) { (cdefg) x*-+-(cdefy) gx? + (cdefg) wae
(a2-+ab) { w+ (defy) 2? +(cdefg)4} = Of,
where, be it remarked as a digression in passing, the trinomial factors in
the left-hand members are exactly similar to the left-hand members of the
pair of equations with which we started this paragraph.
11. As a natural consequence of the result here foreshadowed we are
led to consider the dialytic eliminant
Doe 5.
i eee
Dy 53 95
1 a Or
where
5, stands for a+b+c+d-+e,
De aa abtact ... de,
and so on.
Multiplying it column-wise by unity in the form
1 : oi
—e 1
Pure cae 1 :
ee - —é IL |
and then multiplying row-wise the resulting product by unity in the form
1
e 1
et e2 il :
e® e* e2 il
we reach the result
ean
I 4, 4,
4, 4. '
1 4, e4+5,e?+5,|,
272 Transactions of the Royal Society of South Africa.
the elements taking the simple form here given by reason of the equality
5m — COm_1 te? m—2— on — 4 ine
Since the last element
e4#+5,e7+5, = e*+(e4,+4,)e?+(e4,+4,)
= e4-+ e34,-+¢674,+e4,+4,
= (e+a)(e+b)(e+e)(e+d),
and its cofactor is our dialytic eliminant of the next lower order, it is clear
that we shall have finally
5y 53 5s, - | = (e+a)(e+b)(e+c)(e+d)
as Ons . (d+a)(d+-b)(d-+c)
5, 5s 5s . (c-+a)(e+b)
1) 53) 5) . (b+a),
as was expected.
12. A less pleasing but more quickly effective mode of proof would be
to attest the existence of any one of the linear factors, say d+-e, by showing
that the putting of d+e equal to 0 in the determinant causes the latter
to vanish. Or, again, we might perform the operation
col, — de.col, + d?e?. col, — de coly,
when we should find that the 4th column as thus altered contains the factor
d—+e.*
RoNDEBOSCH, S8.A.,
Ath May 1922.
* Apparently the first to observe this peculiar bigradient was Mr. A. M. Nesbitt.
See Educ. Times, Ivii (1904), p. 490.
—
( 273 )
COLOUR AND CHEMICAL CONSTITUTION.
Part XVII.—THE Azo DyEs AND OTHER Monocyctiic CoLours.
By James Morr.
Some preliminary work on the azo dyes appears in Part XII of this.
work, pp. 215-216, accompanied by the remark that the problem appeared
hopeless of solution. Since then, by examining all the related simpler
substances, I have discovered a method of calculating their colours, and
thus uniting them to the monocyclic class already discussed in Part XIII.
The examination of the simpler substances could only be done by
photography. Other colour chemists will therefore join me in thanking
Mr. E. R. Grills (of Hortor’s Ltd., Johannesburg), who presented me with
a diffraction-grating specially mounted on a thin cover-glass such as is
used in microscopy : this instrument enabled me to photograph down to
A 320 and thus find the absorption of many colourless substances which
required to be examined in order to discover the theory of the azo dyes.
The fact that I had to use a glass (not quartz) lens prevented a complete
investigation further down in the ultra-violet. This I hope to attack
later on.
Remarkable to say, the very first photograph I took with the new
apparatus gave a discovery. Alkaline paraoxyazobenzene in water was
the substance the spectrum of which was photographed, and the negative
showed not one broad band at A 420, as previously supposed from visual
observation (Tuck, J.C.S., London, 1907, diagram on p. 450, and Moir,
loc. cit.), but two distinct although overlapping bands at AA 433 and 395.
Further investigation showed that all the members of the family show this
phenomenon: sometimes, as in acid butter-yellow, both bands are very
distinct, and sometimes they are so overlapped that only a faint luminosity
appears in the middle and the band looks at first like a single very broad
one; in such cases the estimate of the centres of the two bands is quite
uncertain, being in doubt by more than five units.
Again, all the substances are indicators and change colour if made
acid or alkaline, thus giving three kinds of spectrum according to the
274 Transactions of the Royal Society of South Africa.
reaction of the water in which they are examined. As all the absorption-
bands are double, each substance has six bands in all, and it is the object
of this paper to calculate all the six bands of each substance.
This leads to the result that each of the six bands corresponds to a
different chemical constitution, or at least to a different orbital motion of
an electron round the molecule.
As examples, p-oxyazobenzene has AA 433 and 395 when alkaline,
AA 490 and 463 when acid, and AA about 340 and 315 when neutral, the
corresponding figures for butter-yellow being AA 490+460 and 543-+508
and about 405+375. Taking the former example, the six chemical consti-
tutions are respectively : Ph. NH. NOH.C,H,0’, Ph.NOH.NH.C,H,0O’,
Ph. NHCl: N.C,H,0H, Ph ..Ne@NHC!. C,H,OH, Ph. NEY Nene:
Ph: N:>N 7 C,H,OH,
SOLUTION OF THE PROBLEM OF THE Oxy-COMPOUNDS.
Let phenate-ion, C,H,O’ have A 287 in water.*
Let the a-methylene-interposition-colour-factor, -CH,-, be 0-92.
Let the a-1mino- * e. . -NH-, be 1-09.
Let the B-methylene- : cA 4 bemietbs
Let the 6-imino- ie - be 1-22.
Let the a-oxidation-factor be. é é ; >» =O:
Let the p- cf 2. . 1-035;
Let the a-phenyl- substitution. fneter he . Pbsar
Let the B- _,, f Ror aes . + 1-03;
Then we have all the data for calculating the bands of the oxy-com-
pounds of azobenzene, azomethine, stilbene, benzaldehyde, benzhydrol,
benzophenone, diphenylamine, triphenylcarbinol, phenylhydroxylamine, etc.
The method of calculation is, as before, factorial. All the relative
factors taken from the formula of the substance are multiplied together,
the result being the wave-length of the band-centre corresponding to that
formula. For example, Ph NH NOH C,H,0’ gives :—
287 (phenate) 1-09 (a-imino) x 1:10 (a-oxidation) x 1-22 (f-imino) x 1-03
(B-phenyl), which equals 432-4, the decimal part having no significance on
account of the factors being correct only within one-half per cent.
In order to simplify the subsequent calculations, some of the factors
may be multiplied together to begin with, thus giving useful subsidiary
factors.
Thus : a-carbinol, CHOHa, = 0:921:10 =1-01
p- 3 ,CHOHS) = 1-15 x1-0351—= Jat
* Observation agrees with this.
+ As found before in Part XIII, A, of table on p. 36.
Colour and Chemical Constitution. 275
a-oximino, NOHa, =1:09x1:10 =1-20
See ee NOMS al 221-0302 — 1-26
Bpenzyleh@ He —— 15 < 1-050 185
Bb-Ph . CHOH = 1-19X 1-03 = 1-23
f-anilino, PhNH =the Gl Os 255
f-Ph . NOH = 209 Ob, dO
The above example, one of the bands of alkaline p-oxyazobenzene,
now becomes: 287 (phenate) x 1-20 (a-oximino) x 1-255 (f-anilino)=4321.
The other band, from the formula Ph NOH NH C,H,0’ is got from:
2871-09 (a-imino) x1:30 (6-phenyloximino)=407. The former agrees
exactly with observation, but the latter is a little too high. The reason
why the spacing and overlapping of the pair of bands varies has not been
discovered.
We will now proceed to calculate all the oxy-compounds, beginning with
the simplest.
1. The a-CH, group put into phenate gives p-cresolate :
A 264 (or 287 x 0-92).
and Ewbank).
calculated
Not observed in water, A about 285 in alcohol (Baly
2. The a-CH, group taken out of quinomethane gives ionised quinol:
calculated A 3154 (or 290—0-92) for HOC,H,0’: alternatively phenate
xX oxidation-factor 1-1: not observed.
The following table exhibits the rest of the results :—
Name.
p-oxybenzaldehyde .
p-oxy benzophenone
p-oxybenzylalcohol .
-oxybenzhydrol
Fuchsone :
p-oxydiphenylmethane
p-oxytriphenylmethane
-arainophenol
Quinone-imine
Quinone-anile
p-oxydiphenylamine
Quinone-oxime
-p-oxyazobenzene A .
5 B.
p-oxybenzalaniline A
$s B
p-benzalaminophenol A
9 B
p-oxystilbene A
B
2?
Basis of Calculation.
Quinol x CHOHa
Foregoing x Pha
Phenate x CHOHa
Foregoing x Pha
Foregoing again x Pha
Cresolate x Pha .
Foregoing x Pha
Phenate x NHa .
» ™%*NOHa
Foregoing x Pha
Aminophenol x Pha
Quinol x NOHa . : :
Phenate x NOHa x PhNHf
<x NHa x PhNOH#
<x CHOHa x PhNH
x CH,a x PhNOH
< NOHa x PhCH,
<x NHa x PhCcHOHS
x CH,a x PhCcHOH
x CHOHa x PhCH..f
Calcu-
lated A.
319
362
290
329
374
300
340
313
344
390
355
379
432
407
364
343
407
384
323
344
Observed A.
330 (alc.).
Agrees visually.
See Part XIII.
99
380 (alc.).
About 355 (ether).
Agrees visually.
397(NaOH aqueous).
433.
395.
About 375.
About 345.
400.
About 375.
_ 329 not
separated.
276 Transactions of the Royal Society of South Africa.
In addition the monocyclic phthaleins may be calculated, using the
CO,H factor of Part XIII: the results are the same as in Part XIII and
agree with observation. Other unobserved but calculated substances are :
PhCH .CH . C,H,0’ (cale. A356), Ph. N.N.C,Hj0° (calemaeeeae
No No
Ph,N . CHOH . C,H,0’ (calc. A 375), and Ph,C: CH. C,H,0’ (calc. A 339
double). Almost all the possible substances have thus been considered.
Note.—The bands of the acid solutions of the oxyazo and oxyazomethine
dyes are calculated * by multiplying the above-calculated (alkaline phase)
bands by the factor 1-18, and the bands of the neutral phase by multiplying
by a factor which is about 0-8. All the six bands of each substance are thus.
calculated.
Monocycitic AMINO-COMPOUNDS.
The calculation of these is simply made from the oxy-compounds by
multiplying by the factor 1-18 for acid solutions, the factor for ae.
ING
For neutral solutions the factor ox about 1-05.
TABLE OF CALCULATED AMINO-COMPOUNDS (ACID).
Name. Calculated A. Observed A.
p-toluidine salt : ‘ . 2641-18 = 312
p-aminobenzaldehyde ate 377 Agrees visually.
p-aminobenzophenone ; : 427 Probably exceptional.
Aminobenzylalcohol . : : 342
leigh eae : : 388 | suka et
Fuchsonium salts. : : 440 430.
Aminotriphenylmethane _. : 401
p-phenylenediamine . K 370
Quinone-diimine : : 405 About 370 (ether).
Aminodiphenylamine ; 420
Quinone-phenyldiimine ; 460 488 and 450.
p-nitrosoaniline ; : : 448 410 vague.
Aniline yellow A : ; 512 520.
as B , 480 485.
Aminobenzalaniline A : : 430 About 440.
be B : 5 405 About 415.
Benzalphenylenediamine A : 480 | 465 broad, probably
Bes B : 453 J double.
p-aminostilbene A. : : 381
® Baan 4.05
Amino-azoxybenzene : ‘ 527
* See the Table at the end for the observations.
Colour and Chemical Constitution. 261
Monocyciic DIMETHYLAMINO-COMPOUNDS.
ee NMe,HA ,
The multiplying factor for cine is 1-25, 2.e. for acid solutions. For
the neutral phases the factor is 1-13, 2.e. the bands of a neutral dimethylamino
substance are practically the same as those of the corresponding oxy sub-
stance examined in acid solution (see previous page).
TABLE OF CALCULATED DIMETHYLAMINO-COMPOUNDS (ACID).
Name. Calculated A. Observed A.
Dimethyl-p-toluidine : : ; 330
p-dimethylaminobenzaldehyde . : 399 About 380 and 355.
Dimethylaminobenzophenone . e 452 Probably exceptional.
Dimethylaminobenzhydrol : : 411 About 405.
Dimethylaminobenzyl alcohol . : 363
Dimethylfuchsonium salts . : : 466 460.
Dimethylaminotriphenylmethane : 425
Dimethyl-p-phenylenediamine . : 392
Dimethyl-quinonediimine . : : 429 About 400.
Dimethylaminodiphenylamine . : 445
Dimethylphenylquinonediimine . : 488
Nitrosodimethylaniline é ; : 474 457,
Butter-yellow A ; ; : ‘ 542 543.
53 B : : : : 509 508.
Dimethylaminobenzalaniline A. 2 508 513.
a Ba: : 480 482.
Benzaldimethylphenylenediamine A. 455 460.
Bae: 430 427.
Dimethylaminostilbene A . : : 430
33 Bay : t 403 375.
Dimethylaminoazoxybenzene : 559
Dimethylamino-stilbeneoxide ‘ 445
Dimethylamino-benzophenoneanile 524
There are still many blanks in the observations, it being almost impos-
sible for one investigator to repeat all the work which has stretched over
the last thirty-five years ; but the calculations cover nearly all the possible
monocyclic compounds of this class.
Quinone Compounds.—Quinone in water has a band at A455 or 460
(centre vague), but cannot be observed in alkaline solution, 7.e. as
HO.O.C,H,: O’ on account of oxidation. Assuming, however, that
the water solution is ionised, we calculate the a-oxo-interposition factor to
be about 457—3154 (quinol) or 1-45.
Quinonedioxime (in excess NaOH, as it is a very weak acid) has a sharp,
strong band at A366, and quinonemonoxime in alkali a band which is
21
278 Transactions of the Royal Society of South Africa.
roughly intermediate between those of quinone and its dioxime. This
gives a ratio :O0//: NOH of 1:15 experimentally, whereas the ratio
—O—/—NOH— is 1-45+1-2 or 1-21. Alternatively, quinonedioxime
calculated from HO—NOH - C,H, - NOH - H, or from NO- C,H, : HNOH,
should have A 412 instead of A 366, so that it is necessary to make some
distinction between the two groups attached to the benzene ring.
The old terms chromophore and chromogen involve two elements and a
double bond and are now out of date, since, as I have shown in the first
paper of this series, colour may arise merely from the attachment of two
eroups to a ring, the whole being oxidised.
I propose the new term hapton * for the second element and its attach-
ments, reserving the old term auxochrome for OH, NH,, NHAIk, and NAIk,
restrictively. The hapton is intended to be a general term for ©, N,
O,,or 8;
Such a substance as sodium phenate, consisting of a ring and an auxo-
chrome but no hapton, has not a strong absorption visible in great dilution
(say N/740,000) as the true colouring-matters have. Technically it is a
coloured substance with an absorption-band in the ultra-violet. Its penta-
bromo substitution-product is yellow, the ‘loading’ having raised the
absorption-band from A 287 to about A 365, but the absorption-band is
still not strong.
Now let a hapton be added, giving in the unoxidised condition
CH,C,H,0’, NH, - C,H,:- 0’, HO-C,H,- 0’, SH: C,.H,-~ O.-aihesersimll
remain coloured substances, not colouring-matters. They are leuco-com-
pounds, with only shallow absorption.
Parahydroxybenzaldehyde is the simplest true colouring-matter giving
a strong, sharp absorption-band even in high dilution. We thus require
for a colouring-matter a ring, an auxochrome, and a hapton, and the whole
must be oxidised. It is best to adopt an agnostic attitude as to the nature
of the oxidation. The formula HO-C,H,-CHO refers to solid neutral
hydroxybenzaldehyde, but the coloured aqueous alkaline solution may
be O'- C,H,: CHO, 0:C,H,: CHO’, O-C,H,-~CHO’ and half a dozen
| |
other formule involving the valencies of the ring itself.
Apparently the relation of the hapton to the auxochrome may be either
ortho, meta, or para, but the bands of meta-compounds appear to be less
pronounced.
The term dye is restricted to colouring-matters which stick to fabrics
or mordants, e.g. phenolphthalein is not a dye.
* Cf. haptophore in bacteriology.
Colour and Chemical Constitution.
MISCELLANEOUS OBSERVATIONS ON AZO-DYES AND AZOMETHINES.
1. Ortho-hydroxyazobenzene (alkali) AA 441 and 390.
Ds ” c (acid) A 480 broad single.
3. Para- os 3 2475 broad = AA 491+460 in
weaker acid.
4. CH,-der. of (1) = benzeneazo-p-cresol (alkali) A 493 broad single.
DSO, K-,, » = s p-phenolsulphonate 4455 ,, ee
6. 2—-4-6-tribromobenzeneazophenol (alkali) AA 430-+384.
7. Benzeneazo-o-bromophenol a Ad 436+ 390.
8. 2-nitrobenzeneazophenol = Ad 487+ 440.
9, 4- x5 A 490 broad single.
10. 4-nitrobenzeneazo-a-naphthol ie A576 broad = AA 602-552.
11. Foregoing in alcoholic alkali A598, =AA 623-572.
12. No. 9 in Bs Ph A 502 » =A) 540+490.
13. 4-sulphobenzeneazophenol (alkali) A 440 and ? 400.
14. 4-bromo-__,, 35 35 Ah 445+395.
15. 4-ethoxy-_,, ,. a Ah 447-+390.
16. 4-sulphobenzeneazo-o-cresol ,, / 463 single.
eigen 3 min Pee Sa ore A475 ,,
18. Benzeneazoguaiacol 3 A, 460 broad single.
19. Benzeneazothymol - AA 455+410.
20. a -a-naphthol ‘ A 496 broad single.
21. 4-sulphobenzeneazo-a-naphthol (alkali) AO13; 55 Bs
22. Benzeneazo-tetrahydro-a-_,, BA Ad 470 +430.
23. he -catechol . AA 480-4430.
24. 4-diphenylylazophenol ss : A 451.
25. Diphenyltetrazo-p-phenolsulphonate (alkali) i 475 broad single.
26. 4-NO,C,H,CH : NC,H,OH fs A455 ,, 33
27. 4-NO,C,H,N : CHC,H,0H He A about 410.
28. 4-NO,C,H,CH : NC,,H,OHa - AA 560+-505 in alcohol.
29. 4-NO,C,H,N : CHC,,H,OHa eet Ad 510 (+545 faint) in alcohol.
30. 4-diphenylylazonaphthionic acid (in HCl) A 575 broad single.
31. Congo-red (in HCl) A 580 ss a
279
From these we infer (1) that the ordinary substitutions have about the
same small effect as in the dicyclic series : paranitro and naphthyl have the
largest factors = 1:16 and 1-19 respectively ; (2) the benzidine dyes are
practically the same as the diphenylyl dyes, 7.e. only half the molecule acts.
* T have to thank Professor G. 'T. Morgan for this specimen (J.C.8., 1922, p. 5).
ae
( 281 )
FUNGI OF THE STELLENBOSCH DISTRICT AND
IMMEDIATE VICINITY.
By Dr. P. A. vAN DER BuJL.
(From the Department of Phytopathology and Mycology, University of
Stellenbosch.)
No separate list of the fungi occurring in the Stellenbosch district has
so far been published.
For some years Miss A. V. Duthie has been collecting principally the
Myxomycetes and the Higher Basidiomycetes occurring in the district,
and J am indebted to her for kindly placing her collection at my disposal
and for allowing me to embody it in the present paper.
The only way we can hope to learn the distribution of the South African
fungi is to study closely those occurring over small areas, and to extend
such work throughout the Union. More lasting benefit is to be derived
from the close study of the fungi occurring over limited areas than collecting
a few here, there, and everywhere.
In this connection Miss A. Pegler of Kentani rendered valuable service
to the study of South African Mycology by giving close attention to the
fungi occurring in her district.
In 1898 J. Medley Wood published a list of the fungi occurring in Natal.
Incomplete as this list is, it is nevertheless of great assistance to anyone
working on fungi in Natal. |
In the present list the genera and species are listed alphabetically
under the families to which they belong, and the host is also often stated.
The Agariceae are omitted from this list. They will require a few seasons
to work up satisfactorily, and will be dealt with later. In the case of
_ specimens not collected by myself the name of the collector is always
given. The list contains 98 genera, 148 identified species, and 20 fungi
not specifically identified. As the study of the fungi of the district pro-
gresses the number will no doubt be considerably increased.
With the co-operation of students and friends it is hoped to work up
the fungus flora of other districts as well, and to publish these from time
to time.
282 Transactions of the Royal Society of South Africa.
MyYXOMYCETES.
Plasmodiophorales.
Plasmodiophora brassicae, Wor., Brassica oleracea.
Myxogastrales.
Ceratiomyxaceae.
Ceratiomyxa-fruticulosa, Macbr. Decayed wood. A. V. Duthie, June
LOU:
Physaraceae.
Badhamia utricularis, Berk. Dead wood. A. V. Duthie, June 1917.
Diachaea leucopoda, Rost. Decayed leaves. A. V. Duthie, Aug. -1916.
Diderma effusum, Morg. A. V. Duthie.
Fuligo septica, Gmelin. Decayed wood.
Leocarpus fragilis, Rost. Dead needles of Pinus. A. V. Duthie, Aug.
196:
Physarum auriscalpum, Cke. Dead needles of Pinus. A. V. Duthie,
Ae iG.
Physarum pusillum, Lister. Dead stem, Solanaceae. A. V. Duthie.
Physarum sinuosum, Weinm. A. V. Duthie.
Physarum vernum, Somm. A. V. Duthie.
Didynuaceae.
Didymium difforme, Duby. A. V. Duthie.
Didymium nigripes, var. xanthopus, Lister. A. V. Duthie.
Siemonitaceae.
Comatricha nigra (Pers.), Schroet. A. V. Duthie.
Lamproderma scintillans, Morgan. On moss. A. V. Duthie.
Stemonitis splendens, Rost. Dead wood. A. V. Duthie.
Stemonitis splendens, Rost., var. Webberi, Lister. A. V. Duthie.
Heterodermaceae.
Cribraria argillacea, Pers. Decayed stump, Pinus. A. V. Duthie,
May 1917.
Dictydium cancellatum, Macbr. A. V. Duthie.
Reticularvaceae.
Dictydiaethalium plumbeum, Rost. Dead wood. A. V. Duthie.
Reticularia lycoperdon, Bull. Dead wood. A. V. Duthie.
Lycogalaceae.
Lycogala epidendrum, Fr. Dead wood. A. V. Duthie.
Lycogala flavo-fuscum, Rost. A. V. Duthie.
Fungi of the Stellenbosch District. 283
Arcyriaceae.
Arcyria cinerea (Bull.), Pers., Forma digitata. Dead wood. A. V. Duthie.
Arcyria denudata, Sheldon. Dead wood. A. V. Duthie. —
Arcyria nutans, Grév. Dead wood. A. V. Duthie.
Arcyria Oerstedtii, Rost. Dead wood. A. V. Duthie.
SCHIZOMYCETES.
Actinomyces scabies (Thax.), Guss., Solanum tuberosum.
Bacterium nectarophilum, Doidge, Pirus communis.
Bacterium tumefaciens, Sm., Pirus malus.
Pseudomonas pruni, Sm., Prunus persica, Nov. 1921; Prunus domestica
(Prunes), Jan. 1922.
PHYCOMYCETES.
Albuginaceae.
Albugo candida, Lér., Cochlearia Armourica, Dec. 1921; Raphanus
raphinastrum.
Entomophthoraceae.
Empusa Grylli (Fres.), Nowakowsk1.
Empusa muscae, F. Cohn.
ASCOMYCETES.
Exoascaceae.
Taphrina aurea, Fr., Populus pyramidalis, Dec. 1921.
Taphrina deformans (Fcl.), Tul., Prunus persica (Peach and nectarines).
Helvellaceae.
Morchella conica, Pers. On ground. A. V. Duthie.
Pezizaceae.
Lachnea hemisphaerica (Wigg.), Gill. A. V. Duthie.
Lachnea sp. A. V. Duthie.
Peziza spp. Two large species and several smaller ones. A. V. Duthie.
Mollisvaceae.
Aspergillus niger, Bref., Alium cepa. Nov. 1921.
Erysiphaceae.
The fungi of this family were only found in their conidial fructifications.
Their identification was hence based on the hosts on which they occurred.
Erysiphe cichoracearum, D. C., Cucumis pepo., Nicotiana tabacum.
Erysiphe polygon, D. C., Pisum sativum. Nov. 1921.
284 Transactions of the Royal Society of South Africa.
Podosphaera leucotricha (Ell. and Ev.), Salmon Pirus malus. Nov.
1921.
Sphaerotheca pannosa (Wallr.), Lev., Rosa sp. Nov. 1921.
Uncinula necator (Schwein.), Burr, Vitis vinifera.
Hy pocreaceae.
Claviceps paspali, 8. & H., Paspalum. Feb. 1922.
Claviceps sp., Pennisetum macruorum.
Dothidiaceae.
Phyllachora cynodontis (Sacc.), Niess., Cynodon dactylon. Feb. 1922.
Phyllachora melianthi (Thuem.), Sacc., Melianthus major. Oct. 1922.
Phyllachora sp., Pennisetum macruorum. March 1922..
Coryneliaceae.
Corynelia uberata, Ach., Podocarpus. Oct. 1921.
Mycosphaerellaceae.
Mycosphaerella fragariae (Tul.), Linn., Fragaria vesca.
Pleosporaceae.
Venturia inaequalis (Cke.), Aderh. (Fusicladium stage), Pirus malus.
Venturia pirina, Aderh. (Fusicladium stage), Pirus communis.
Melanconidaceae.
Calospora arausiaca (Fab.), Sacc., Quercus sp.
A ylarvaceae.
Hypoxylon sp. Old wood. A. V. Duthie.
Xylaria sp. Old wood. A. V. Duthie.
BASIDIOMYCETES.
Ustilagvnaceae.
Ustilago avenae (Pers.), Jens., Avena sativa. Nov. 1921.
Ustilago bromivora (Tul.), F. v. M., Bromus unioloides. Oct. 1921.
Ustilago hordei (Pers.), K. & 8., Hordeum vulgare. Sept. 1921.
Ustilago maydis (D. C.), Tul., Zea Mays.
Ustilago nuda (Jens.), K. & S., Hordeum vulgare. Oct. 1921.
Tilletiaceae.
Tilletia laevis, Ruhn., Triticum vulgare.
Urocystis tritici (Bjerk.), Wint., Triticum vulgare. Oct. 1921.
Melampsoraceae.
Melampsora tremulae, Tul., Populus sp. Sept. 1921.
Fungi of the Stellenbosch District. 285
Pucciniaceae.
Aecidium resinicolum, var. tumefaciens, Wint., Raphnia angulata.
Octet 9it.
Phragmidium subcorticum, Wint., Rosa sp. Nov. 1921.
Phragmidium violaceum (Schultz), Wint., Rubus Vitifolius.
Puccinia chrysanthemi, Roze., Chrysanthemum sp.
Puccinia graminis, Pers., Avena sativa, Hordeum sativum, Triticum
vulgare.
Puccinia granularis, K. & Cke., Pelargonium sp.
Puccinia hypochaeridis, Oud., Hypochaeris radicata. Oct. 1921.
Puccinia malvacearum, Mont., Malva parviflora. Oct. 1921. Mathiola sp.
Puccinia maydis, Bereng., Zea Mays. Feb. 1922.
Puccinia phragmites (Schum.), Koern., Phragmites communis.
Uredo Fici, Cast., Ficus sp.
Uromyces betae (Pers.), Kuhn., Beta vulgaris sp. 1921.
Uromyces cyperi, P. Henn., Cyperus sp. Oct. 1921.
Uromyces fabae (Pers.), de B., Vicia Faba. Oct. 1921.
Uromyces romuleae, n. sp., P. v.d. B. (MSS. name), Romulea rosea.
Uromyces striatus, Schroet., Medicago sativa.
Tranzschelia punctata (Pers.), Arth., Amygdalus communis; Prunus
armeniaca ; Prunus domestica (Prunes); Prunus persica.
Tremellaceae.
Exidia purpureo-cinerea, Kalch. Dead branch, Quercus, Sept. 1921.
Old log, Oct. 1921.
Tremella sp. A. V. Duthie.
Tremella microspora, Lloyd. A. V. Duthie.
Dacromycetaceae.
Dacromyces deliquescens (Bull.), Duby. Old log. <A. V. Duthie.
Hormomyces aurantiacus, Bon. Logs, Quercus. A. V. Duthie.
Thelephoraceae.
Stereum hirsutum (Willd.), Pers. Common saprophyte on logs of
Quercus and other plants.
Stereum purpureum, Pers. Old logs Quercus. Aug. 1922.
Stereum vellereum, Berk. Old logs Quercus.
Thelephora intybacea (Pers.), Fr. In plantations, Pinus.
Thelephéra penicillata, Lloyd. A. V. Duthie.
Poly poraceae.
Lenzites betulina, Linn. Stump, Quercus. Aug. 1921.
Lenzites Quineensis, Fr. (form of above). Old logs. Oct. 1921.
Merulius lachrymans, Facq. A. V. Duthie.
286 Transactions of the Royal Society of South Africa.
Trametes protea, Berk. Stump and logs of Pinus, Populus, and Quercus.
Trametes trabea, Otth. Logs of Pinus.
Polyporus adustus, Willd. Stumps Quercus. Aug. 1921.
Polyporus conchatus, Lloyd. Stump Populus.
Polyporus occidentalis, Klotz. Old logs. A. V. Duthie.
Polyporus ochraceus, Pers. Old logs. A. V. Duthie.
Polyporus rusticus, Lloyd. Log Pinus.
Polyporus sanguineus (Linn.), Fr. Old logs. A. V. Duthie.
Polyporus sulphureus (Bull.), Fr. Wound parasite of Quercus. Burned
stump of Kucalyptus. |
Polyporus versicolor (Linn.), Fr. Wound parasite of Prunus persica.
Common saprophyte on various old logs.
Polyporus zonatus, Fr. A. V. Duthie.
Boletaceae.
Boletus edulis, Bull. On ground.
Boletus, n. sp. (Herb. No. 508). On ground.
Agaricaceae.
Amanita muscaria (Linn.), Pers. On ground. A. V. Duthie.
Amanita phalloides (Fr.), Quél. On ground. A. V. Duthie.
Lentinus Lepideus, Fr. Stump of Pinus.
Schizophyllum commune Fr. Wound parasite on Prunus persica and
Acacia saligna. Common saprophyte on logs of various trees.
Phallaceae.
Anthurus Mac Owani, Marl. On ground. A. V. Duthie.
Phallus sp. On ground. A. V. Duthie.
Hymenogastraceae.
Rhizopogon luteolus, Fr. On ground. A. V. Duthie.
Rhizopogon rubescens, Tul. On ground.
Rhizopogon sp. On ground. A. V. Duthie.
Hysterangvaceae.
Protubera africana, Lloyd. On ground. A. V. Duthie.
Lycoperdaceae.
Calvatia lilacina, Berk. On ground. A. V. Duthie.
Catastoma magnum. On ground. A. V. Duthie. ®
Geaster plicatilis, Berk. On ground.
Lycoperdon pratense, Pers. On ground. <A. V. Duthie.
Nidulariaceae.
Cyathus vernicosus (Bull.), D.C. On ground. A. V. Duthie.
Fungi of the Stellenbosch District. 287
Podaxaceae.
Podaxon carcinomalis, Fr. On termite nest. A. V. Duthie. .
Sclerodermataceae.
Arachnion album, Schw. On ground. A. V. Duthie.
Arachnion scleroderma, Lloyd. On ground. A. V. Duthie.
Polysaccum crassipes, Fr. On ground under Kucalyptus trees.
Scleroderma cepa, Pers. On ground. A. V. Duthie.
Scleroderma laeve, Lév. On ground. A. V. Duthie.
Scleroderma flavidum, Ellis. On ground. A. V. Duthie.
Scleroderma tenerum, Berk. On ground. A. V. Duthie.
Tulostomataceae.
Tulostoma cyclophorum, Lloyd. On ground. A. V. Duthie.
Tulostoma sp. (a small plant). On ground. A. V. Duthie.
DEUTEROMYCETES.
Sphaeriordaceae.
Actinonema rosae (Lib.), Fr., Rosa sp. Nov. 1921.
Ascochyta pisi, Lib., Pisum sativum. Nov. 1921.
Phyllosticta richardiae, Hals., Zantedaschia aethiopica. Oct. 1921
Phyllosticta violae, Desm., Viola odorata. Nov. 1921.
Septoria graminum, Desm., Triticum vulgare. Sept.1921.
Septoria lycopersica, Speg., Solanum lycopersicum. Feb. 1922.
Septoria pelargonii, Syd., Pelargonium sp. C. K. Brain.
Septoria pisi, Went., Pisum sativum.
Septoria rubi, West., Rubus vitifolius. M. M. Radloff. Oct. 1921.
Septoria sp., Silene gallica.
Sphaeropsis malorum, Peck, Cydonia vulgaris ; Pirus malus.
Melancomaceae.
Gloeosporium ampelophagum, Sacc., Vitis vinifera. Dec. 1921.
Pestalozia sp., Pirus malus. Oct. 1921.
Septogloeum arachidis, Rac., Arachis hypogea. Feb. 1922.
Septogloeum mori, Lév., Morus sp. March 1922.
Monilraceae.
Botrytis cinerea, Pers., Vitis vinifera. March 1921.
Botrytis sp., Lathyrus odorata.
Oidiopsis taurica, Lév., Tropaeolum major. Feb. 1922.
Oidium sp., Euphorbia sp. Nov. 1921.
Oidium sp., Quercus sp. Nov. 1921.
288 Transactions of the Royal Society of South Africa.
Dematiaceae.
Alternaria solani (HE. & M.), Jones & Grout, Solanum tuberosum, Nov.
1921. Solanum lycopersici, Feb. 1922.
Cercospora protearum, Cke., Leucospermum conocarpum.
Cladosporium zeae, Peck., Zea Mays. Feb. 1922.
Coniothecium macowanil, Sacc., Protea grandiflora. June 1913.
Macrosporium cladosporioides, Desm., Beta vulgaris. Feb. 1922.
Macrosporium iridis, C. & E., Irissp. Feb. 1922.
Macrosporium maydis, C. & E., Zea Mays. Feb. 1922.
Macrosporium sp., Tree tomato. Dec. 1921.
Macrosporium sp., Eriobotryon japonica. Feb. 1922.
Macrosporium sp., Populus sp. March 1922.
Mucedinaceae.
Rhinotrichum rubiginosum, Fr. <A. V. Duthie.
Stilbaceae.
Isaria sp., Gonipterus scutellatus. C. K. Brain.
MyYcELIUM STERILE.
Rhizoctonia, Solanum tuberosum.
( 289 )
OBSERVATIONS ON THE PROTECTIVE ACTION
OF NORMAL SERUM IN EXPERIMENTAL INFECTION WITH
BACILLUS DIPHTHERIAE.
By tad) Mackin, MD. ChB, DP H.
(From the Department of Bacteriology, University of Cape Town.)
In carrying out test inoculations in guinea-pigs with B. diphtheriae
and diphtheroid bacilli for purposes of identification, where control animals
were injected with the particular strain plus a certain amount of diphtheria
antitoxin, it was noted that these animals were protected against lethal doses
of B. diphtheriae by normal horse serum as well as by the specific immune
serum. Though this observation did not involve any quantitative com-
parisons between the effect of the normal and the immune serum, the fact that
normal horse serum should act protectively in the same way as the serum of
a specifically immunised animal seemed of considerable importance, and
suggested the further investigation of the phenomenon. This normal-serum
effect is also of general interest in relation to “ non-specific therapy,”
where alien protein (e.g. animal serum, bacterial protein, etc.), injected
parenterally, is found to exert a favourable influence on the course of various
infections 1n a non-specific manner.
With a view, therefore, to analysing the effect, a series of experiments
were carried out which fully established the original observation, and yielded
further information of some theoretic and practical interest.
The protective and curative action of normal horse serum in B. diph-
theriae infection and intoxication has also been studied by other workers.
Kolle and Schlossberger concluded that in guinea-pigs infected with
B. diphtheriae normal horse serum had a limited curative effect, which was
in no way comparable with that of an antitoxic serum, and in the case of
animals injected with diphtheria toxin had a certain “ delaying ”’ action
but was not definitely curative. They stated that the effect was ‘“ non-
specific, resistance-increasing, stimulating,’ but did not offer any further
explanation of their results. Kraus and Sordelli claimed that normal horse
serum contains antitoxin to diphtheria toxin, and in virtue of this exerts a
‘‘ preventive’ effect against B. diphtheriae and its toxin. Cowie and
Greenthal showed that 1 c.c. of normal horse serum injected subcutaneously
290
Transactions of the Royal Society of South Africa.
and intravenously in guinea-pigs simultaneously with diphtheria toxin
always protects against 1 M.L.D., and may neutralise even 8 M.L.D.; they
showed that the activity of the serum depends on the proteins and not on
the alcohol-soluble fraction.
entirely due to a natural diphtheria antitoxin.
They regarded the protective power as not
In the writer’s original experiments guinea-pigs of approximately
350 grms. weight were injected subcutaneously with 2 c.c. of a 2-days
bouillon culture of virulent strains of B. diphtheriae, and at the same time
5-10 c.c. of normal horse serum were injected subcutaneously at a different
site from that of inoculation; these animals survived, while animals of
similar weight, injected with the same amount of the same cultures,
died within 24 to 48 hours, showing at autopsy the characteristic effects of
B. diphtheriae inoculations.
normal horse serum gave the same results.
Various strains and different specimens of
“For the detailed investigation, particular strains of B. diphtheriae isolated
from typical cases of throat diphtheria were used.
DEGREE OF PROTECTIVE ACTION OF NoRMAL HorRSE SERUM.
In order to ascertain quantitatively the degree of protection conferred
by normal serum, guinea-pigs of approximately equal weight were injected
subcutaneously with a fixed quantity of a B. diphtheriae bouillon culture,
and at the same time with varying amounts of horse serum; the minimum
lethal dose of the culture was also estimated in parallel experiment (Table I).
Thus 2 c.c. of serum protected against at least 12 lethal doses of the
culture (7.e. 1 c.c. serum against 6 M.L.D.).
TABLE I,
Guinea-pigs approximately 400 grms. weight injected subcutaneously in
flank with 3 c.c. of a 2-days bouillon culture of B. diphtheriae, “strain 3”’ ;
varying amounts of normal horse serum injected subcutaneously in back.
1. B. diphtheriae -+-6’c.c.
a
=
Seer Ea HS ue a ee
+5 ¢.c.
+4 c.c.
+3 6.6.
+2 G.c.
+1] ¢.¢.
+0-5 c.c.
+0-25 ¢.¢.
+0-1 c.c.
ho serum
serum
survived.
29
ele)
99
39
died 6th day.
psn Ab ay
Ae ORG ia ce
a) Abies
Be TP Aitoll
Observations on the Protective Action of Normal Serum. 291
Minimum LEetHAL Dost ESTIMATION.
Guinea-pigs approximately 400 grms. weight: same culture injected
subcutaneously.
1. B. diphtheriae bouillon culture, 1-5¢.c. . . died 2nd day.
2. es iss is IEOVG Car on ; oe cmicee a,
3. > 5 3 Orb Cee: eo At hie a,
A, - Ee = O25: Cie, are as LN ee
5, st ie es Ole. . . survived.
6. a - ss O:05c6.c.
die)
PROPHYLACTIC AS OPPOSED TO CURATIVE EFFECT.
It was found that normal serum, though protective or prophylactic, was
not curative; thus, to elicit the antagonistic effect, the serum had to be
injected without delay after the introduction of the infecting organisms ;
if an interval of two hours or longer was allowed to elapse before the serum
was injected, even when five times the protective dose was used, the animals
died (Table II).
THERMOSTABILITY OF THE ACTIVE SUBSTANCE IN THE SERUM.
In thermostability the active principle of the serum corresponds gener-
ally to antitoxin and other antibodies. Horse serum heated for one hour
at 57° C. was still protective, but was inactivated by exposure at 70° C.
for half an hour and at higher temperatures (Table III). The thermo-
stability at 57° C. excluded the possibility of the active substance being of
complement nature.
TAsiE ts
Guinea-pigs approximately 400 grms. weight injected subcutaneously
in flank with 3 ¢.c. 2-days bouillon culture of B. diphtheriae, “strain 3” ;
10 c.c. normal horse serum injected subcutaneously in back at varying
intervals after injection of the culture.
1. B. diphtheriae+serum, simultaneously . . survived.
2. a +.,, after 2hours . . died 3rd day.
3 53 snes Sie, acess 7 end.
+ a + ,, Sa) poe tee 2 ae
5 no serum. Sura 04! ae
TaBLe III.
Guinea-pigs approximately 450 grms. weight injected subcutaneously
in flank with 3 c.c. of a 2-days bouillon culture of B. diphtheriae,
292
Transactions of the Royal Society of South Africa.
“strain 2”; normal horse serum injected subcutaneously in back at
same time.
1. B. diphtheriae+10 c.c. unheated serum . survived.
Vp ng no serum . died 2nd day.
3. b. +10 c.c. serum, 57° C., 1 hr. . survived:
*4, ss +50 ¢c.c. of a serum dilution,
=):
ZO:
1 part in 5 of normal saline,
heated at 70° C., 4 hr. . died 2nd day.
+60 c.c. of a serum dilution,
1 part in 5 of normal saline,
heated at 90° C.. $ hr. . died 2nd day.
+50 ¢.c. of a serum dilution, -
1 part in 5 of normal saline,
heated at 100° C. for 5
minutes . . died -in 24 hours,
showing more mark-
ed lesions than other
animals ; marked
haemorrhagic in-
flammatory oedema
at site of inoculation,
with marked
haemorrhages in
suprarenals, lungs,
and intestinal wall.
_EXPERIMENTS WITH RABBITS.
Using rabbits as the test animal, horse serum was found to exert the same
protective action against B. diphtheriae as in the case of guinea-pigs ; the
particular effect was not therefore limited to B. diphtheriae infection in
one species of animal only. Table IV exemplifies one of these experiments.
TABLE [V.
Rabbits of equal weight injected subcutaneously with 4 c.c. of a 3-days
bouillon culture of B. diphtheriae, “ strain 3.”
1. +20 c.c. normal horse serum injected
subcutaneously at different site . survived.
2. Noserum . . died 3rd day.
* The serum was diluted to prevent coagulation on heating.
Observations on the Protective Action of Normal Serum. 293
PROTECTIVE ACTION OF THE SERUM OF DIFFERENT ANIMALS.
- The serum from various animal species (ox, sheep, man, rabbit, cat) was
found to confer the same protection as horse serum ; in the experiments of
other workers on this subject the phenomenon has been studied from the
point of view apparently of its being related to horse serum only ; the effect
is, however, not characteristic of the serum of only one species of animal
(Table V). .
The question then arose as to whether the serum of one guinea-pig in-
jected parenterally into another guinea-pig would protect the latter against
B. diphtheriae infection. To ascertain this a number of experiments were
carried out, but with variable results: in some cases complete protection
was obtained ; in others, while the infected animals succumbed to the dose
given, there was a distinct delaying of the lethal effect (Table VI). Thus
it was apparent that though protection by normal serum from an alien
species was the general rule, the serum of the same species was not without
a similar effect.
TABLE V.
Guinea-pigs approximately 450 grms. weight injected subcutaneously in
flank with 3 c.c. of a 2-days bouillon culture of B. diphtheriae, “strain 2” ;
serum of various animals injected subcutaneously in back at same time.
1. B. diphtheriae+10 ¢.c. ox serum . . survived.
i +10 c.c. sheep’s serum. ; -
a + 8c.c. human serum . : ad
+10 c.c. cat’s serum -
- + 6c.c. rabbit’s serum . ; -
. +. 3.0. 4, ee oo 1C)) fs
ys +-10 c.c. horse serum re
8. : no serum. ; . died 2nd day.
5 tees ee ens
TABLE VI.
Guinea-pigs approximately 400 grms. weight injected subcutaneously
with 3 c.c. 2-days bouillon culture of B. diphtheriae ; serum from other
guinea-pigs injected subcutaneously at a different site.
1. (a) B. diphtheriae, “ strain 2 ’’+6 c.c. serum : . died 6th day.
(b) control (no serum injected) . ye ieee
2. (a) B. diphtheriae, ‘“ strain 2’’+-8 c.c. serum (55° C.) . survived.
(b) control . died 2nd day.
22
294 Transactions of the Royal Society of South Africa.
3. (a) B. diphtheriae, “ strain 2”’°+10 c.c. serum (from 2
animals; sera pooled) died 19th day.
(6) control Sly, Danica
4. (a) B. diphtheriae, “‘ strain 3”+5 c.c. serum (from 2
animals), and 5 c.c.
following day . . survived.
(b) control : ; . died 2nd day.
5. (a) B. diphtheriae, “strain3”+10c.c. serum. . survived. —
(6) control ; . died 2nd day.
6. (a) B. diphtheriae, “ strain 3 ’-++10 ¢.c. serum > bth Fee
(6) control : : . yg, eae ie
7. (a) B. diphtheriae, “ strain 3 +20 ¢.c. serum 5; othe
(b) control : : : : ‘ : : .- TS 2ndare
8. (a) B. diphtheriae, “‘ strain3”+10c.c. serum. . survived.
(b) control ; ; : : : : : . died 3rd day.
It is a phenomenon of considerable interest that the serum of one in-
dividual of an animal species, which is highly susceptible to experimental
B. diphtheriae infection, should be capable of affording some protection,
when injected parenterally in certain amounts, to another individual of the
same species infected with the particular organism.
EXPERIMENTS WITH OTHER PROTEINS ;
BACTERICIDAL EXPERIMENTS,
Experiments were carried out with egg-albumen, milk, and peptone to
determine whether the parenteral injection of these proteins would confer
any protection as in the case of the serum proteins. The results were
uniformly negative.
In vitro bactericidal tests were also made to ascertain whether horse
serum (in varying amounts), along with fresh guinea-pig’s serum as comple-
ment, exerted a specific bactericidal action on B. diphtheriae, but no such
effect could be demonstrated, thus excluding the possibility of the pro-
tection being due to a normal bactericidal immune body in horse serum.
EXPERIMENTS WITH DIPHTHERIA TOXIN.
Experiments were then carried out to ascertain whether the protective
effect was due to a natural antitoxic action on the part of the serum injected,
though it seemed difficult to explain on this basis the action of guinea-pig’s
serum in the case of infection in the same species.
A powerful toxin was prepared from B. diphtheriae, “ strain 3.” It was
found that 10 c.c. of horse serum was protective against 10 M.L.D. of the
Observations on the Protective Action of Normal Serum. 295
toxin (1 c.c. against 1 M.L.D.), whereas 1-5 c.c. of diphtheria antitoxin,
representing 1000 immunity units, protected against 10,000 M.L.D. The
minimum lethal dose was estimated in parallel experiments along with
the protective tests (Table VII).
A similar series of experiments were carried out in which the pooled
serum of three guinea-pigs was substituted for horse serum (Table VIII).
The parenteral injection of 10 ¢.c. of guinea-pig’s serum had apparently
no protective action in guinea-pigs against B. diphtheria toxin.
TABLE VII.
Guinea-pigs 250 grms. weight injected subcutaneously in flank with
varying doses of diphtheria toxin ; at the same time injected subcutaneously
in back with 10 c.c. normal horse serum. }
1. Toxin 0-1 ¢.c. 10 ¢.c. serum . . died 2nd day.
2 Pe OLOG Gre. ie : ; Oe Pete as
ow eee, 0-01 c.c.- As , ; ve eb lieges 3
4. ,, 0-005 c.c.+ e ; oes otMe as.
5 » 0:001 c.c.+ . . survived.
6 », 0:0005 c.c.+ s , s
7 sree | c.c. +1000 immunity units
diphtheria antitoxin i
M.L.D. Estimation.
1. Toxin 0-1 CiC new: ; . died 2nd day.
2 » 0-01 Chess: ; oy 2 3;
Ser OOM CrCn ee é 741 hee
fe 00005) Ieee. eb; 2. hw,
5 pee 000) celia a LOL ea
6 ; 0:00005 e.c. .. ; . survived.
7 0-00001 c.c. Ms
M.L.D.—0-0001 c.c.
Norte.—The small doses of toxin 0:05 c.c.—0-00001 c.c. were measured
by preparing decimal dilutions of the toxin 1 : 10, 1 : 100, 1 : 1000, 1 : 10,000,
1 : 100,000 and injecting the appropriate quantities of these dilutions.
Tasie VIII.
Guinea-pigs 250 grms. weight injected subcutaneously in flank with
varying doses of diphtheria toxin; at the same time 10 c.c. of guinea-pig’s
serum (from three animals, pooled) injected subcutaneously in back.
296 Transactions of the Royal Society of South Africa.
1. Toxn)0-00L)e:c. 10 c-exserum ie . died 2nd day
2. ,, O-0001 c.c.+ sp : : NAPADOIL ich
8. ,, 0-00005 c.c.+ - : . survived.
4 » 0:00001 c.c.+ v : as
Lethal Dose Estimation.
12 Loxin,0-00l. tcc : . “died 2nd day:
Dee es OOOO M ere nile: ; : mots ee
3. 3g (000005; e2e3,— . . survived.
4, ,, 000001 cc. . : : Fe
THE OCCURRENCE OF LocaL LESIONS IN ANIMALS SURVIVING AFTER
PROTECTION BY SERUM.
An interesting feature of these experiments was the occurrence of a
marked local lesion in animals surviving after protection by normal serum
against B. diphtheriae or its toxin.
After two to three days a well-marked subcutaneous indurated swelling
with surrounding inflammatory oedema developed at the site of injection
of the organisms or toxin; the overlying skin then became necrotic and
sloughed, leaving a punched-out ulcer with a grey sloughing base; the
lesion attained its full development in eight to ten days, and then slowly
healed ; in one instance the lesion involved the deeper tissues, and led to
perforation of the whole abdominal wall.
It was found that diphtheria antitoxin prevented the occurrence of the
local lesion as well as the general effects.
This type of lesion occurred in rabbits protected by serum as well as
in the case of guinea-pigs.
In the infection experiments no diphtheria bacilli could be detected in
the ulcers after sloughing of the skin, either by microscopic examination or
culture. On the other hand, in experiments where the injection of guinea-
pig serum delayed the lethal effect of B. diphtheriae for several days (e.g.
eight days, nineteen days, v. Table VI), and similar sloughing ulcers de-
veloped at the site of inoculation, B. diphtheriae were present in the lesion.
Thus the parenteral injection of normal serum, though protecting against
the lethal result of infection with B. diphtheriae, had no influence on its
local toxic action. It was apparent also that B. diphtheriae did not persist
in the tissues of animals fully protected by serum. ,
In unprotected animals the local lesion of the type described is not
commonly met with, as animals infected with a sufficient dose die within
a few days; for the full development of the local effect a period of eight to
Observations on the Protective Action of Normal Serum. ASHE
ten days is required, and it was apparent that the occurrence of this .
particular lesion in protected animals was simply dependent on the survival
of the animal. In rabbits, which are less susceptible to experimental
B. diphtheriae infection and resist larger doses than guinea-pigs, it has
been noted that sub-lethal doses may produce similar marked local lesions
apart from any serum protection.
DISCUSSION.
These observations represent a further contribution to the study of
non-specific immunisation by parenteral injection of normal serum.
While it is possible that in the case of horse serum a natural antitoxin
may contribute to the protection noted, and it is of importance that the
active substance in the serum corresponds to antibodies in thermostability,
the action of the serum in bringing about some other antagonistic effect
must be considered. This effect apparently does not depend on the trans-
ference of bactericidal complement or a natural bactericidal antibody ;
the serum is protective against infection apart from intoxication ; thus the
organisms are absent from the local lesion. It might be claimed, however,
that the antitoxic factor contributes to the elimination of the bacteria by
rendering them more susceptible to phagocytosis through neutralisation
of their toxin. It is to be noted also that normal serum is not antitoxic as
regards the local toxic effects, whereas diphtheria antitoxin protects animals
against local as well as general effects. The fact that the parenteral in-
jection of the serum of one guinea-pig into another exerts a “ delaying ”
effect in diphtheria infections and even complete protection, though this
species has little natural immunity to B. diphtheriae and no natural anti-
toxin can be demonstrated in the serum, clearly indicates that some
mechanism other than that of a natural antitoxin or other antisubstance
transferred with the serum plays the important part in increasing the
resistance of the animal.
These experiments are therefore of interest in their bearing on the general
question of “‘ non-specific immunity,” and provide a further illustration of
how the resistance of the tissues to a particular organism may be enhanced
by the parenteral injection of normal serum. The results, however, clearly
exclude the possibility that normal horse serum could be as effective a
therapeutic agent as a potent diphtheria antitoxin.
CONCLUSIONS.
1. In guinea-pigs experimentally infected with B. diphtheriae, normal
serum (horse, ox, sheep, cat, human), injected subcutaneously at the same
time as the inoculation, exerts a definite protective action.
298 Transactions of the Royal Society of South Africa.
2. Two c.c. of normal horse serum may protect in this way against
12 M.L.D. of a B. diphtheriae culture.
3. No protection occurs if the serum injection is delayed for two hours.
after the inoculation.
4. The activity of the serum persists at 57° C., but is lost at 70° C. and
higher temperatures.
5. The serum of one guinea-pig injected subcutaneously into another is
either protective to the latter experimentally infected with B. diphtheriae,
or at least exerts a definite delaying effect on the course of the infection.
6. Normal horse serum is also similarly protective in guinea-pigs in-
jected with diphtheria toxin ; 10 c.c. of serum may protect against 10 M.L.D.
7. The serum of one guinea-pig is not protective in the case of another
injected with diphtheria toxin.
8. In the case of animals surviving after protection by normal serum, a
marked local lesion develops at the site of inoculation.
REFERENCES.
KouLE and ScHLOSSBERGER.—Med. Klinik, 1919 (also quoted by Kraus and Sordelli).
Kravs and SorDE.xi.—Zeit. f. Immunitats., Teil i, Orig. 31, 2, 1921, 107.
Cowles and GREENTHAL.—Journ. Med. Research, 1921, 42, 261.
( 299 )
ON SOME UPPER BEAUFORT THERAPSIDA.
By 8S. H. Haventon, B.A., D.Sc.
(Published by permission. of the Hon. the Minister for Mines and
Industries.)
(With Plate XIII and two Text-figures.)
CYNIDIOGNATHUS LONGICEPS, gen. et sp. nov.
The specimen which forms the type of this new genus was collected on
the farm Vaalbank, Albert District, at the foot of the northern slope of the
Dreunberg during the course of the geological survey of the area carried out
in 1921. It consists of an almost perfect skull with part of the lower jaw,
and some fragmentary limb-bones, but the skull only will be described here.
The principal measurements are as follows :—
Maximum length : ; ) oo) oumannn
Dorsal length in mid-line . 330 mm.
Maximum width . : . 240 mm.
Width at canines : ’ © 80mm.
Interorbital width * , | oll nam,
Snout to front of orbit . : . 180 mm.
Orbital diameter . : > O0emm:
Snout to back of pterygoid flange . . 200 mm.
Maximum height SoD eam,
Premaxilla.—The two bones are separated by well-marked suture.
Each bone carries 4 incisors, but at the alveolar edge the facial portion is
fairly short. It extends backwards in its lower portion to form the lower
border of the external nares, but does not meet the nasal behind the nostril.
Above the nostril each premaxilla sends backwards a narrow tapering
process to the level of the back of the outer nares, the two processes lying
between the front parts of the nasals.
The palatal portion of the premaxilla is partly seen, although the ventral
surface has not been fully displayed. On the ventral surface, however, the
300 Transactions of the Royal Society of South Africa.
premaxilla passes backwards to the level of the back of the canine, appearing
in its posterior part as a narrow splint of bone lying medial to the maxilla.
The dorsal surface has been almost completely developed. Anteriorly the
bone forms the floor and outer side of the nasal opening—no ossified inter-
nasal septum being present. Near the front of the nasal floor on each side
there is a rounded foramen, and just anterior to this the septomaxilla lies
on the premaxilla. Behind the septomaxilla the lateral portion of the
premaxilla is a thin inclined plate of bone lying on the maxilla and passing
back to the level of the back of the canine. Medial to this plate there is a
deep channel which is flanked medially by an upstanding process, somewhat
swollen above, which extends from the level of the last incisor to the level
of the front of the canine. This palatine process forms the inner wall of
the premaxilla and is separated from its neighbour by a deep narrow channel,
which is partly divided by a longitudinal thin bony plate not reaching down
to the secondary palate. This palatine process of the premaxilla does not
seem to be separate from the main body of the bone. The bone is some-
what cracked, and the cracks are filled with red iron oxide; but the cracks
are so unsymmetrical, and no one of them can be matched by a counterpart
on the other side, that none can be considered as a suture dividing off the
premaxilla from a prevomer.
Broom has mentioned and figured the occurrence of such processes in
Lycochampsa ferox and Watson in Gomphognathus polyphagus, but no
adequate description has been given of either.
Septomaxilla.—This bone is only preserved in the right nostril. It lies
on the floor and against the lateral portion of the premaxilla and is almost
completely separated by that bone from the maxilla, only meeting the latter
at its upper corner. The front lower half of the bone is a very thin lamina,
concave from above (called the pars horizontalis of the os narialis by Wegner,
1922). Half-way up it thickens considerably and sends inwards a fairly
thin turbinal process (processus intrafenestralis of Wegner) which almost
reaches the mid-line of the skull. The turbinal process lies wholly in front
of the palatine process of the premaxilla. On the floor of the nostril there
is a foramen between the front of the septomaxilla and the narial portion of
the premaxilla. The septomaxillary foramen is very small.
Mazxilla.—The maxilla carries 1 large canine and 10 molars. The molar
series occupies a length of 103 mm. Only the sockets of the molars are
preserved. These are longer than wide, and begin almost directly behind the
canine.’ The last tooth is very small.
Lachrymal.—This bone has contact with the prefrontal, nasal, maxilla,
and jugal. In front of the orbit it is excavate as in many Gorgonopsids
and is bounded behind by a pronounced ridge which forms here the antorbital
border. At the bottom of the excavated portion is the lachrymal foramen
On some Upper Beaufort Therapsida. 301
which forms the entrance to a short duct running through the bone whose
posterior end opens into the orbit.
The other bones of the face and of the top of the skull are very similar
to those of Cynognathus platyceps, and their shapes and relationships are
adequately displayed in the figure.
** Vomer.”’—In section anterior to the orbit the vomer is broad, thin,
slightly concave downwards, with a ventral median keel and a dorsal median
groove flanked by thin upstanding walls. When traced forward in longi-
tudinal section the horizontal plate gradually disappears, the vertical plate
becomes higher and its ventral por-
tion swells, so that at the anterior
end, between the palatine processes
of the premaxille, the median plate is
sharp-pointed above and rounded
below. At the anterior end the
bone-mass is certainly single; along
its length there may possibly be a
suture separating an upper portion
from a lower; but the median sep-
tum, except in its lower swollen por-
tion, 1s very thin and much cracked,
so that it is not possible to be absol-
utely certain of the presence of a
suture. If a suture zs present, then
the bone between the premaxille is Cynidiognathus longiceps.—Htn.
the vomer. and the bone which roofs ‘=tegular section across snout. Dorsal
eos: ; view. Left septomaxilla not.preserved.
the pharynx is the parasphenoid; or
the former is the fused prevomers and the latter the vomer. It is certain
that the bone which roofs the pharynx forms the upper part of the
median septum at least; and that the condition is different from that
figured by Broom in Diademodon, in that no mesethmoid forms the upper
part of the septum. On the other hand, if the bone is single, the changes
which take place in it when traced forward from the pharynx are the
gradual loss of the horizontal septa, the thickening of the ventral part of
the vertical plate, and the complete or almost complete fusion of the walls
of the dorsal groove.
Posterior to the line of fracture in front of the orbits, the
becomes more arched with a ventral median keel and passes rapidly down-
wards and backwards, forming the front part of the groove in the palate ;
the hinder portion of this groove is formed by the pterygoids. Broom and
Watson have both figured this region in Diademodon, and have shown the
vomer as forming the whole of this vaulted area of the palate, and Watson
‘
‘vomer ”’
302 Transactions of the Royal Society of South Africa.
has further figured a large vomer in this position in some Gorgonopsia. I
have elsewhere expressed my inability to see a vomer separating the
pterygoids in Gorgonopsian skulls which I have examined, and the evidence
afforded by this Cynodont skull seems to re-open the whole question of the
identity of the bone which forms the median septum of the anterior part of
the skull and the roof of the front part of the pharynx.
Is it the anterior prolongation of the parasphenoid ? or is it the fused
prevomers of the Therocephalia ? Its position in front of, and slightly
flanked by, the pterygoids, and between the palatines, would seem to
homologise it with the prevomers; and until further evidence of its actual
connection with the parasphenoid is obtained, it would seem best to consider
it as the fused prevomers.
Basicranium.—The basicranial region is similar to that of Cynognathus
in general plan, but there are significant points of difference. The pterygoid
is large, the vomer extends back at least to the front of the median bar ;
and posteriorly the pterygoids stop some distance in advance of the triangular
basisphenoid plate and are separated from each other by an anterior pro-
longation of the basisphenoid along the median bar. Laterally the posterior
portion of the pterygoid is overlapped by the epipterygoid, which has a long
articulation with the basisphenoid. Posteriorly the epipterygoid is con-
siderably shortened. It has the same relations with the openings in the
brain-case as in other Cynodonts, but it does not extend outwards to
articulate with the quadrate, being separated from that bone by a consider-
able gap and lying as a thin lamina on the mass of the pro-otic. Watson
describes a somewhat similar condition in Protacmon brachyrhinus, but there
the epipterygoid terminates posteriorly in a thickened margin. Anteriorly
in our form there is a foramen between the pterygoid and epipterygoid,
which continues forward as a groove along the dorsal surface of the pterygoid.
This is the opening seen by Broom in Gomphognathus, and considered by him
to be the internal carotid foramen.
The basioccipital forms most of the floor of the brain-case behind the
sella turcica, extending from the condyles to the dorsum selle, where a
fracture shows it lying on the pro-otics as a thin plate of bone with triangular
section. It forms the inner boundary of the internal auditory opening and
is perforated on each side by two foramina for the exit of the branches of
the XIIth nerve.
Seen in dorsal view the basisphenoid forms the floor of the sella turcica
into which the carotid foramina enter. The carotid arteries enter the
ventral surface of the basisphenoid far back, and thus pass nearly horizon-
tally forwards through the bone. The side walls of the sella turcica are
formed by the anterior superior processes of the pro-otics, which bones lie
here between the basisphenoid and basioccipital. ~ Anteriorly to the sella
On some Upper Beaufort Therapsida. 303
turcica the basisphenoid is continued forwards as a median bar overlapped
laterally by the epipterygoids and then by the pterygoids. The dorsal
ridge of this median bar is formed of a thin, high plate.
Cynidiognathus longiceps—Htn.
Photograph of palatal view of type skull.
As in Diademodon the foramen jugulare lies just anterior to the occipital
condyle and faces downwards ; while the fenestra vestibuli looks forwards
and outwards, being separated from the foramen jululare by a thin vertical
plate of the periotic mass. The bone above the fenestra, instead of having
a vertical face, as in Diademodon, is bent horizontally to form an eave
304 Transactions of the Royal Society of South Africa.
overhanging the fenestra. This eave is deeply notched on its anterior edge.
The inner border of the notch is formed by the posterior end of the epiptery-
goid, the remainder of the border by the pro-otic. This notch must represent
the pterygo-paroccipital foramen of Diademodon, the retraction of the
epipterygoid from the quadrate giving rise to the open form. The foramen
for the exit of the VIIth nerve is larger and nearer the pituitary fossa than
in Diademodon.
The periotic (pro-otic) forms most of the side wall of the brain-case
behind the dorsum selle, articulating behind with the fused exoccipital
and supraoccipital, the latter forming the posterior wall of the canal for the
TXth-XIth nerves.
The paroccipital process is quadrangular in section, and not triangular
as in Diademodon.
Quadrate.—-Another feature of interest lies in the quadrate mass. The
quadrate is a plate of bone with a slightly hollowed anterior surface, and
lies on the anterior face of the squamosal, being furnished with an outer
flange which is clasped by the squamosal. It has no pterygoid wing. The
quadrato-jugal is lateral to the quadrate. It is bifid in character, consisting
of a larger anterior vertical thin plate and a smaller posterior one which
clasp the lower edge of the squamosal between them. There is probably
a quadrate foramen between the quadrate and the quadrato-jugal. A
somewhat similar quadrate complex exists in Protacmon, but there the
quadrato-jugal spreads inwards between the quadrate and squamosal.
Discussion of the relationship of this type to other Cynodonts is deferred
until a more complete account of the fossils of the Upper Beaufort Beds is
prepared. It is sufficient here to point to some of the advanced features
which this form possesses—the complete loss of the quadrate ramus of the
pterygoid, the gap between the quadrate and the epipterygoid, loss of a
pterygoid process separating the palatine and vomer, and extreme flattening
of the basicranium—all logical results from the evolutionary tendencies
traced in the Theriodontia, as pointed out recently by Watson.
The absence of an internasal process of the premaxilla and the usurpation
of the functions of prevomers by palatine processes of the premaxille—
functioning as supports to Jacobson’s cartilage—are interesting features
which are distinctly mammalian in character. The former is paralleled
in a large skull of Cynognathus trom Winnaarsbaaken, Albert District, now
in the South African Museum, in which the premaxille form a pillar in front
of the nostrils reaching upwards and backwards to the nasals.
On some Upper Beaufort Therapsida. 305
Cynidiognathus (%) broom, n. sp.
1911. Broom, Cynognathus berryi, Proc. Zool. Soc., pl. xlvi, figs. 1, 2.
The occiput and outer view of the bones of the brain-case of an incomplete
skull in the South African Museum collection (Cat. No. 1056) were figured
by Broom as Cynognathus berryr. Recent examination of the specimen has
shown that it possesses 10 molars, and cannot therefore belong to the genus
Cynognathus. In view of the general agreement between the two forms,
as far as their features can be compared, I have tentatively placed it in the
newly-erected genus.
Comparison with Broom’s figures shows that the division between the
two condyles is not quite strong enough in the drawing he gave. Further,
it seems doubtful whether the epipterygoid passed back to meet the quadrate
as described by Broom. As actually preserved, the suture between the
pro-otic and epipterygoid is rather more vertical below the big foramen than
in his figure and the broken lower edge of the plate shows an interdigitation
between the two bones. Unfortunately, the lower outer corner of the
palate figured by Broom no longer appears on the specimen, so that it is
impossible to settle the point definitely.
No suture can be seen between the exoccipital and paroccipital; the
features of the region are mainly as in Watson’s description of Diademodon.
The post-temporal fossa is above the level of the foramen magnum. The
fenestra ovalis lies lower than in Diademodon.
The basisphenoid and basioccipital in ventral view are somewhat
hollowed out, not flat, and in the posterior part there is a well-marked,
narrow, shallow, median keel flanked by two narrow elongated grooves
which pass forward well beyond the level of the tubera basisphenoidalia.
There is a pronounced occipital boss on the upper border of the foramen
magnum.
The series of 10 molars occupy a length of 85 mm. The first is immedi-
ately behind the canine. In Cynognathus and Lycochampsa there is diastema
between the canine and the first molar, which is relatively largerin the latter.
The frontal is more lozenge-shaped than in Cynognathus crateronotus,
reaching to within 4 or 5 mm. of the orbital border.
The probable length of the skull is 260-270 mm.; the interorbital
width is 52 mm., and the width at the canines 58 mm.
Atlurosuchus brownt, Broom.
1906. Broom, Trans. Phil. Soc. 8. Afr., xvi, 4, p. 376, pl. x.
Broom has described all the visible features of the top of the weathered
skull and of the postcranial skeleton. Since the type came into the posses-
306 Transactions of the Royal Society of South Africa.
sion of the South African Museum (Cat. No. 5875) it has been possible to
display the general features of the palate and basicranium, and to show
that the form must be classed among the Bauriamorpha.
The palatal aspect is similar to that of Bauria, except that it is probable
that the secondary palate is more fully formed than in that genus. The
septum dividing the posterior nares was figured by Broom. There is a
large suborbital fossa, bounded posteriorly by the pterygoid and laterally
by a bone which expands forwards and is probably the ectopterygoid.
The bone substance is very soft and sutures are usually indistinguishable
in a sandstone matrix such as this fossil possesses. There is a large cordate
interpterygoid vacuity.
The quadrate ramus of the pterygoid meets the quadrate, and between
it and the paroccipital there is a very considerable space.
The epipterygoid is a flat bone, fairly narrow in the middle, but expanded
at each end, resting on the quadrate ramus of the pterygoid and touching
the parietal. It is far more like the epipterygoid of the Therocephalia than
that of the Cynodontia.
The basisphenoid is broad, and apparently extends in the middle line
to the back of the interpterygoid vacuity. Near the front it is pierced by a
single circular foramen, probably the foramen for the carotids. It forms,
as usual, part of the border of the fenestra ovale, which is closed by the
stapes. The stapes are dumb-bell shaped bones, the outer end lying between
the paroccipital, quadrate, articular, and pterygoid.
The paroccipital is shallow—possibly on account of the post-mortem
flattening of the skull—but laterally is broad from back to front ; its under
surface is provided with a broad, shallow groove. Its inner end forms
the border of the large foramen jugulare, whilst medial and slightly posterior
to the latter is a smaller opening for the exit of the XIIth nerve.
No other details are visible, but enough is seen to place this form in the
Bauriamorpha, to which the ill-defined features of the upper surface of the
skull would also assign it.
ADDENDUM.
Since the above description of Cynidiognathus longiceps was drawn up,
Dr Broom has informed me in a verbal communication that the bone hitherto
known as basisphenoid in Therapsids (and so described in this paper) is,
in his opinion, the parasphenoid. In an immature Gorgonopsian skull from
the top of the Cistecephalus zone, whose basicranial region he has sectioned,
he finds the basisphenoid existing as two small ossifications lying in front
of the basioccipital—quite distinct from it and from the bone which sends
back a thin plate to underlie the basioccipital and which is anteriorly clasped
by the pterygoids, extending forwards above them as a median septum bone
On some Upper Beaufort Therapsida. 307
—the parasphenoid or vomer. In later life the basisphenoidal ossifications
fuse with the posterior portion of this parasphenoid.
According to this view, only the upper posterior portion of the bone
called basisphenoid in such figures as the one published by the writer as a
section through the brain-case of Alopecognathus minor (Ann. 8.A. Mus., xii,
p. 211), should be known by that name; the remainder is the true para-
sphenoid.
EXPLANATION OF PLATE.
Cynidiognathus longiceps, gen. et sp. nov.
1. Top view of type skull.
2. Side view of type skull.
3. Ventral view of basicranium.
4. Dorsal view of floor of brain-case.
5. Anterior view of left quadrate mass.
6. Ventral view of same.
Cynidiognathus broomi, sp. nov.
7. Side view of type skull.
8. Top view of type skull.
Atlurosuchus browni, Broom.
9. Dorsal view of horizontal section through type skull.
B.0O., Basioccipital ; B.S., Basisphenoid ; d.s., dorsum selle; Ep.pt., Epipterygoid ;
inner vest., inner vestibule of ear; J., Jugal; L., Lachrymal; Mx., Maxilla; N., Nasal ;
P.Mx., Premaxilla; P.O. (Pro. Ot.), Pro-otic; Patu., Palatine; Par. Occ., Paroccipital ;
Po.O., Postorbital; Pr.F., Prefrontal; Pr., Pterygoid; Q., Quadrate; Q.J., Quadrato-
jugal; Smx., Septomaxilla ; Sq., Squamosal ; St., Stapes; Tas., Tabulare ; Vo., Vomer.
Trans. Roy. Soc. 8. Afr. Vol. X. Plate XIII.
Neill & Co., Lid.
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