NationalLibraryOfScotland/medical-history-of-british-india huggingface dataset:
Medical Officers of the Army of India.
65
TABLE II.—Distribution of Laccadive Plants—continued.
NAME.
I n d i a .
C e y l o n .
N i c o b a r s .
A n d a m a n s .
B u r m a .
M a l a c c a .
M a l a y A r c h i p e l a g o .
A u s t r a l i a .
P o l n e s i a .
A m e r i c a .
A f r i c a .
M a u r i t i u s .
K e e l i n g I s l a n d s .
C h a g o s I s l a n d s .
Stachytarpheta indica
.
x x
x
x
x
x
x
...
. .
x
.
.
.
x
Premna integrifolia .
.
x
x
x
x
x
x
x
x
x
... ..
x
...
. . .
Leucas aspera.. .
x
. . .
. . . ...
x
. . .
x
...
. . .
...
...
x
...
. . .
Boerhaavia repens
.
.
x
x
x
x
x
x
x
x
x
...
x
x
x ...
Aerua lanata..
.
x
x
x
x
x
x
x
... . . . ... X
x
...
. . .
Achyranthes aspera .
.
x
x
x
x
x
x
x
x
x
x
x
x
x
...
Hernandia peltata
.
.
x
x
x
x
x
x
x
x
x
...
x
x
...
. . .
Euphorbia Atoto
.
.
x
x
x
x
x
x
x
x
x
... ...
... ...
. . .
Euphorbia pilulifera.
.
x
x
x
x
x
x
x
x
x
x
x
x
... x
†Phyllanthus Emblica
. x
x
x
x
x
x
x
... ... ...
... ... ... ...
Phyllanthus maderaspatensis x
x
...
...
... ... ...
x
... ...
x
x
... x
Acalypha indica.
.
x
x
x
x
x
x
x
... ...
... x
x
... ...
†Ricinus communis. .
x
x
x
x
x
x
x
x
x
x
x
x
... ...
Ficus bengalensis.
.
x
x
. . . ...
x x x
...
. . . ... ... ... ...
. . .
†Artocarpus incisa.
.
x
x
. . .
... . . . x
x
x
x
x
x
...
... . . .
†Musa sapientum.
.
x x
x
x
x
x
x
x
x
x
x
x ...
. . .
†Agave vivipara (?). . x
x
. . . ...
x
x
x
x
x
x
x
x ... ...
†Dioscorea sativa.
.
x
x
x
x
x
x
x
x
x
x
x
x ... ...
Gloriosa superba. .
x x x x x x
. . . ...
. . . ... ... ......
. . .
†Areca catechu.
.
x x
x
x
x
x
x
x
x
x
x
... ... ...
†Cocos nucifera.
.
x
x
x
x
x
x
x
x
x
x
x
...
x
. . .
Pandanus odoratissimus .
x
x
x
x
x
x
x
x
x
x
x
x
... ...
†Colocasia antiquorum
.
x
x
x
x
x
x
x
x
x
...
x
x
... ...
Cyperus arenarius. .
x x
. . . ...
x
. . .
. . . ...
. . . ... ... ... ...
. . .
Cyperus pennatus.
.
x
x
x
x
x
x
x
x
x
... x
x
... ...
Oplismenus compositus
.
x
x
x
x
x
x
x
x
x
x
x
x
...
. . .
Setaria verticillata.
.
x
x
x
x
x
x
x
x
x
x
x
...
... . . .
Spinifex squarrosus .
.
x
x
x
x
x
x
x
...
. . . ...
. . . ......
. . .
Andropogon contortus. .
x
x
x
x
x
x
x
x
x
x
x
x
... ...
Apluda aristata. .
x x
. . . ...
x x x
...
. . . ... ... ......
. . .
Cynodon dactylon (?)
.
x
x
x
x
x
x
x
x
x
x
X
X
...
. . .
Eleusine ægyptiaca .
.
x
x
x
x
x
x
x
x
x
x
x
... ... . . .
†Eleusine Coracana(?)
.
x x
. . . ...
x
. . .
. . . ...
. . . ...
x
......
. . .
Lepturus repens.
.
x x
. . ...
. . .
. . .
. . .
x x
...
x
x
...
Nephrodium molle.
.
x
x
x
x
x
x
x
x
x
x
x
x ... ...
Nephrolepis tuberosa
.
x
x
x
x
x
x
x
x
x
x
x
x
... ...
The cultivated species are not taken into account in the calculations given
below. Even when they are excluded, we find that 18 species extend to Africa,
America, Australia, and Polynesia, as well as to Asia ; four species extend to all the
great geographical divisions except Polynesia ; one to all the divisions except
Australia. Three that occur in Asia, Africa, and America are excluded both
from Australia and from Polynesia.
On the other hand, 11 that are common to all the other divisions are absent
from the New World, and no fewer than seven of these belong to the coast flora
K
66
Scientific Memoirs by
which is characterized by ocean-distribution. 1 Two others ( Vernonia and the
herbaceous Phyllanthus) are excluded from Polynesia also, but both of these are
simple weeds of cultivation. One coast species that extends both to Australia
and Polynesia ( Euphorbia Atoto ) does not reach America, and at the same timé
appears to be absent from the African coast. Five species extend from Asia to
Africa only. One of them ( Peristrophe bicalyculata ) is a very common Indian
weed that extends into Assam and into Burma, but, though it appears in the
Laccadive islands, it has not hitherto been reported from Ceylon. Another
(Launea pinnatifida ) is a coast species much less extensively distributed than
species of this class usually are ; it appears to be confined to the coasts of India
and Ceylon, to Mauritius, and to the eastern coast of Africa. 2. Three species that
are characteristic of Continental Asia appear in Mauritius. These are Tylophora
asthmatica, Barleria cristata, and Leucas aspera ; their distribution as weeds
probably accounts for the presence there of all three. 3 The other 15 are con-
fined to Asia. 4
The following table shows more compactly the constitution of the Laccadive
flora :—
TABLE III.—Constituents of the Laccadive Flora.
Distributional Features.
No. of species.
Per cent.
Cultivated species .
.
.
.
.
.
.
.
17
21¼%
Cosmopolitan in the Tropics
.
.
.
.
.
.
8
22½%
Almost Cosmopolitan: not in Polynesia
.
. .
.
4
5%
Ditto
ditto
: not in Australia. . . .
1
1 ¼
Nearly
ditto
: not in Australia nor in Polynesia
.
3
3¾%
Tropics of old world and Polynesia. . . . .
.
1 1
13¾%
Tropics of old world alone
.
.
•
.
•
.
2
2½%
Asia, Australia and Polynesia only
.
.
.
.
.
1
1 ¼ %
Continental Asia and Continental Africa
.
.
.
.
5
6¼%
Ditto
and Mauritius
.
.
.
.
3
3¾%
Confined to Asia
.
. . . . . .
.
15
1 8 ¾ %
8o
100 %
1 These sea-coast species not extended to the new world are Guettarda speciosa, Scœvola Koenigii,
Tournefortia argentea, Hernandia peltata, Premna integrifolia, Cyperus pennatus a n d
L e p t u r u s r e p e n s .
But to these have to be added Morinda bracteata, Launea pinnatifida, Euphorbia Atoto, Cyperus arenarius.
and Spinifex squarrosus, which are circumscribed in their distribution even within the eastern hemisphere.
And, besides these, Thespesia populnea, which is so common on tropical shores in the old world, is only
known in the West Indies as an introduced and naturalized species. Altogether therefore 13 of the
Laccadive sea-coast species are plants confined to the old world.
2E l e u s i n e C o r a c a n a , o n e o f t h e c u l t i v a t e d s p e c i e s , h a s a l s o t h i s l i m i t e d A s i a t i c a n d A f r i c a n
distribution.
3 Moringa pterygosperma, a cultivated species, is also limited to Asia and Mauritius.
4 Phyllanthus Emblica, which is cultivated, is also confined to Asia.
Heritiera littoralis.
Terminalia Catappa.
Cocos nucifera.
Pandanus (2 species).
Barringtonia speciosa.
Calophyllum inophyllum.
68
Scientific Memoirs by
the vegetation on this island was destroyed by the great eruption of 1883, and
the factor of human interference in its refloration is eliminated, as it is quite un
- v i s i t e d b y m a n a n d i s q u i t e u n i n h a b i t a b l e .
On the shore Dr. Treub found seeds or fruits of—
Growing there he found young plants of
—
Erythrina (species).
Calophyllum inophyllum.
Cerbera Odallam.
Scævola Kœnigii.
Ipomœa biloba.
Hernandia peltata.
Cyperaceæ (2 species).
Gymnothrix elegans.
The last-named species is clearly a wind-conveyed grass; the others are as
clearly all sea-conveyed species. On the mountain proper he found Scœvola
Kœnigii again, also Tournefortia argentea, four palpably wind-conveyed Com
- p o s i t œ ,
w i t h t w o g r a s s e s , e l e v e n f e r n s , a n d t w o m o s s e s , — a l l e v i d e n t l y ( e x c e p t
Scœvola and Tournefortia) wind-dispersed. The general similarity between the
coast list and the coast section of the Laccadive list is very striking and be
-comes all the more so when we consider that Krakatau is only 21 miles from
Java, and 10 and 20 miles from Siebesie and Sumatra respectively, and has land
on every quarter, while Anderut, the nearest island of the Laccadive group to
the mainland, is 130 miles from the Indian coast, and the archipelago, as a whole,
has land on one side only.
While the species that constitute the coast flora are the first to arrive on
newly emerged coral islands, they form the section of the flora least liable to
subsequent alteration. Thus, while Dr. Alcock has brought to our notice 16
species that were not seen by Mr. Hume, none of these belong to the coast flora,
while three—the ferns and Cynanchum—are almost certainly wind-conveyed
species, and three—Vitis, Oldenlandia and Herpestis —are almost certainly
bird-introduced. The others are all instances of plants intentionally or uninten
-tionally introduced by man.
The extremely low percentage of vascular cryptogams in the Laccadive list,
as compared with the percentage found by Dr. Treub on Krakatau l, deserves
remark. There the vascular cryptogams formed 46½ per cent. of the whole and
62 per cent. of the interior flora to which they properly belong. In the Lacca
- d i v e s , o n t h e c o n t r a r y , t h e y f o r m b u t 2 ½ p e r c e n t . o f t h e w h o l e , o r 3 p e r c e n t . o f
the inland flora.
The state of affairs on Krakatau is, as Dr. Treub shows, in accordance with
expectation and in agreement with the evidence afforded by mountainous vol
- c a n i c i s l a n d s e l s e w h e r e . T h e s e b e c o m e f i r s t s t o c k e d w i t h a w i n d - c o n v e y e d
cryptogamic flora which survives the more readily that its constituents are not
1
A n n . J a r d . B o t . B u i t e n z o r g , v i i , 2 1 8 .
Medical Officers of the Army of India.
69
highly specialized, and which prepares the way for a more highly specialized
phanerogamic flora. But the larger proportion of the phanerogams that do
reach such islands are ocean-distributed and littoral species that cannot ascend
beyond a certain height, those whose fruits or seeds are adapted for very dis-
tant wind-conveyance being comparatively few in number, and the proportion
of such fruits or seeds that escape from perishing in the ocean en route being
very small. Hence, in such islands the proportion of cryptogams to phanero-
gams always remains high, and is increased in a direct ratio with the distance
of the island from adjacent land, and in an inverse ratio with the suitability of
the surface of the island for the support of vegetable life.
The conditions that prevail in newly emerged coral islets are very different.
Their low elevation in itself diminishes the chances of wind-conveyed spores
alighting on them, and is in itself unfavourable for the germination of many of
the spores that do alight. Then, for those that have succeeded in alighting
and that have germinated the struggle for existence is intensified by the simul-
taneous arrival of another class of species, independently conveyed, more highly
specialized, with its individual species more largely represented, and for which
the same physical conditions are in the highest degree favourable. These
conditions appear sufficiently to explain the difference of the proportions in
Krakatau and in the Laccadives.
In concluding, the writer wishes to acknowledge his great indebtedness to
Dr. Alcock for much information concerning the present flora of the Lacca-
dives and to Dr. King for numerous valuable suggestions while analysing its
constituents.
On the Life History of a Himalayan Gymnosporangium
( G. CUNNINGHAMIANUM, NOV. SP. )
BY
SURGEON-MAJOR A. BARCLAY, M. B.
BENGAL MEDICAL SERVICE.
I had long suspected that the Aecidium on the wild pear ( PyrusPashia, Ham.),
which is in certain years very common in the neighbourhood of Simla (Western
Himalayas), was related to a Gymnosporangium on Cupressus torulosa, Don.,
and I noted the probable connection in a paper submitted to the Asiatic Society
of Bengal in 1887,1 and incidentally in a paper on Caeoma Smilacis in these
Memoirs, Part IV; but it was not until the spring of 1889 that I succeeded in
proving this by experiment.
The wild pear is abundant in Simla, but the cypress is not. The Aecidium
is consequently by no means common, except in the neighbourhood of affected
cypress trees. An occasional aecidial patch may, however, be found at a con
-siderable distance from any cypress tree.
In the following paper I propose first to describe the characters of the fungus
as it occurs on its two hosts, and then to pass on to a consideration of its title
to be considered a distinct species.
I. Teleutosporic stage on Cupressus.
The teleutospore beds are hemispherical dark brown compact bodies during
dry weather, and are found on the ultimate small branches, as well as on older
twigs of 4 to 5 mm. in diameter. Their appearance during dry weather is re
-presented in Fig. 1, Plate I. During very moist weather these beds swell up en
-ormously into gelatinous masses, and soon assume a yellow ochre colour, due to
a plenteous formation of sporidia. The swelled condition of the teleutospore
beds is shown on Fig. 2, Plate II. During heavy rain the gelatinous masses fall
off from the tree. Teleutospore beds may be found during the greater part of the
year. I have missed them only during August to November.
The mycelium producing these spores is perennial, and causes some slight,
but by no means conspicuous, thickening of the parts involved. It ramifies
between the cells of the bark tissue, and penetrates down to the central woody
tissue, but does not apparently enter the latter. Near spore beds the mycelium is
1
ournal of the Asiatic Society of Bengal, Vol. LVI, Part II, No. 3, 1887.
72
Scientific Memoirs by
extremely abundant, but, as we recede from this area the amount of mycelium
quickly diminishes. Where abundant near beds of teleutospores the hyphae con-
tain orange-red oil globules; but elsewhere the mycelium is colourless. It never
appears to enter the cells, and there are consequently no haustoria. The
stimulant effect of the mycelium on adjoining cells is shown in their increased
size. The bark cells, just below the outer sub-epidermal palisade-like cells, as
seen in transverse sections of the youngest twigs, are normally about 30μ in
diameter, whilst in areas pervaded by mycelium these present a diameter of
about 50μ .
The teleutospores are slender spindle-shaped yellow bodies, on long stalks
covered with a substance capable of swelling exceedingly when moistened (Fig. 5,
Plate II). Sometimes, when the spore beds are moist, and some spores are scrap-
ed off, the central axile stalk comes out of the gelatinous sheath as a very slender
stalk. In such cases there is a characteristic disc between the base of the spore
and the stalk (Fig. 9, Plate II). There is no appreciable constriction at the
septum. Each cell of the spore exhibits a central nucleolar space, or nucleus,
and near the septum each cell has two germ spores. The fresh spores, when
scraped off their beds and examined immediately in water, measure 75.6
X 2 5 . 2
μ .
They have a thin membrane, not usually thickened anywhere, and without
any external markings. Sometimes a slight thickening of the apex, with a short
blunt conical process, may be seen.
When placed in water the spores germinate very rapidly. In one cultivation
spores were placed in water at 9 A.M., and at 2 P.M. many had thrown out pro-
mycelia 60 to 80μ in length (April). I have looked carefully for the second kind
of spore, with short stalks, thin walls, lighter coloured, and much constricted at
the septum, as described by Dietel1 in most species investigated by him, but
have not found them. I could not find any second form of spore decidedly
different from the form I have described. At first each cell in germinating throws
out two commencing promycelia from the pores near the septum, the four form-
ing a cross. Only one promycelium from each cell, however, continues to de-
velope, the other remaining undeveloped (at least in water cultivations). All the
coloured contents of each cell of the spore wanders in mass into the very end
of the developing promycelium, and keeps to the end, however long the tube
may grow, leaving the proximal end of the tube quite colourless. Sometimes
in a water cultivation the tubes attain a very considerable length; but they are
usually short (Fig. 3, Plate II). The promycelial tube measures from 8 to 11 μ
in diameter: it is usually widest at the end, where the coloured contents have
accumulated. The end now invariably, so far as my observations extend, divides
into four compartments by four septa, the lowest just including the coloured
contents (Figs. 3, 4, Plate II). Stout sterigmata are formed from each cell,
1 Ueber das Vorkommen von Zweierlei Teleutosporen bei der Gattung Gymnosporangium, Hedwigea,
1889, Heft 2, p. 99.
Medical Officers of the Army of India.
73
and very frequently all these are on one side of the promycelium, i.e. in a row
one above another (Fig. 3, Plate II). This is not invariable, but usual. Each
sterigma bears a sporidium (Fig. 4, Plate II), and therefore four such are formed
by each promycelium. The sporidia are orange-red and oval, and measure from
15 X 9µ to 22 X 14µ. When all the four sporidia have been formed, the promy
-celium is almost colourless.
. When many spores are germinating in a very small quantity of water, only
those which have germinated earliest proceed to the normal development of
sporidia: those which germinate later (48 hours) usually exhibit the abnormal
process first described by Kienitz-Gerloff, and which I have elsewhere' described
as characteristic of Caeoma Smilacis, and a Uromyces on Solidago Virgaurea;
that is to say, after the end of the promycelial tube has divided into four cells,
the cells round off and become detached (Fig. 8, Plate II). This abnormal
growth is not confined to spores of any peculiar form, for, as I have already
noted, there is only one kind of spore. I have not seen such detached cells
germinate.
In connection with the germination of these spores I may draw attention to
another curious and interesting variation. In water cultivations I have occa
- s i o n a l l y s e e n t h e c o n d i t i o n r e p r e s e n t e d i n F i g . 1 0 , p l a t e I I , i n w h i c h a t e r m i n a l
cell of the promycelium has developed a very long slender tube, in every respect
resembling the germ tube of a sporidium. Such growth obviously suggests the
suppression of sporidial formation.
The sporidium germinates very quickly, after ripening (Fig. 6 c, Plate II).
The formation of secondary sporidia in water cultivations is not uncommon.
II. Æcidial stage on Pyrus Pashia, Ham.
This stage of the life history of the fungus I have already described
in the Journal of the Asiatic Society of Bengal, 2 but, in order to complete the
present paper I will describe it again. It is remarkable that this member only of
the several species of allied fruit-trees occurring in this region harbours the parasite.
I have frequently looked in vain for it on other species. Another noteworthy
feature is that, whilst in some years the Æcidium occurs abundantly, in others
it is rarely found. As the teleutospores are formed abundantly every year, this
can only be accounted for on the assumption that certain well-defined atmo
-spheric conditions must concur to enable the sporidia to attack the leaves.
In 1885 the Æcidium was very common; in 1886 it was remarkably rare; in
1887 it was again fairly abundant, though not nearly so common as in 1885; and
now, in 1889, it is again common (I was absent from India during 1888). Mr.
W. L. Dallas, Assistant Meteorological Reporter to the Government of India
1
T h e s e M e m o i r s , P a r t I V .
2
V o l . _ L V I , P a r t I I , N o . 3 , 1 8 8 7 .
L
76
Scientific Memoirs by
blue, the spermatia assume a brick-red colour. The spermatia are large and
oval, measuring 8x4µ. On several occasions I placed these spermatia in weak
solutions of sugar, but never observed any yeast-like germination.
This completes the description of the fungus, and I now proceed to de
- s c r i b e b r i e f l y t h e e x p e r i m e n t s I m a d e t o e s t a b l i s h t h e c o n n e c t i o n b e t w e e n t h e t w o
phases of existence of the parasite.
During March and April I frequently applied the sporidia obtained from
the teleutospores on Cupressus to the leaves of Pyrus, and in most cases saw
spermogonia developed in ten to fourteen days. The exact method of procedure
was as follows. Small twigs of Pyrus were cut off under water, and then placed
for a day or two under a glass shade in my laboratory, means being taken to
keep the air within moist. In the meanwhile some dry teleutospore beds were
scraped off from Cupressus, and put into a watch-glass containing water, the
whole being kept in a moist atmosphere. In a very short time these beds swelled
up extraordinarily into light tremulous gelatinous masses, and in 24 hours were
covered wi t h l i ght yel l ow spori di a. These spori di a were t hen t ransferred wi t h
a sterilized glass rod to the leaves on the twigs of Pyrus under the glass shade,
and left there until the earliest sign of attack was apparent. They were then
put into an ordinary gardener's forcing-box. If kept too long under a glass
shade, the leaves soon withered. Even in the forcing-box the leaves did not
remain alive long enough for the production of æcidia, although numerous char
-acteristic spermogonial patches were produced.
As I found it impossible to keep such twigs long enough alive to allow of
the formation of æcidia, I tied some twigs of Cypress, on which there were
teleutospore beds, on to three bushes of Pyrus in my garden, after the wet
weather had set in. On each bush, after about a fortnight, many of the newly
unfolded leaves exhibited bright orange-red spermogonial patches, and these
went on to the formati on of æci di a about a month l ater. The photograph,
reproduced in Fig. 1, Plate 3, represents a twig taken from one of these bushes
in my garden.
These experiments show quite conclusively that the Pyrus œcidium is
related to the teleutospores which are borne on Cupressus torulosa.
With regard to the geographical distribution of this Gymnosporangium, I am
sorry to say I have but little information. During an excursion into the interior of
the Himalayas I found the Æcidium many miles from Simla. My friend Dr. D. D.
Cunningham informs me that he found what appears to be the same æcidium,
at Almora in the Kumaun Hills in 1874, and he has kindly placed the drawings
he made of i t at my di sposal (Fi gs. 13, Pl ate II, and 2, Pl ate III). I am tol d, not
however with certainty, that Cupressus torulosa occurs there. After so long a
time Dr. Cunningham could not himself remember whether any of these trees
7 8 Scientific Memoirs by Medical Officers of the Army of India.
differences. This species has two forms of teleutospore, with four pores to each
c e l l ; i t h a s l o n g e r æ c i d i a , o p e n i n g i n a d i f f e r e n t m a n n e r , a n d t h e æ c i d i o s p o r e s
are somewhat smaller than in the Himalayan species. The Himalayan species is
therefore distinct from all these.
I have also carefully compared the Himalayan species with Farlow's
description of the American species,1 and feel persuaded that it is distinct from
any one of them. There can be no doubt whatever that it is quite distinct
from G. Ellisii (Berk.) and G. biseptatum, Ellis, both of which are characterized
by the teleutospores being usually divided into more than two cells, and both of
which, curiously enough, are the only other Gymnosporangia borne on species
of Cupressus ( C. thyoides ).
A new name is therefore wanted for the Himalayan species, and I propose
calling it Gymnosporangium Cunning hamianum, after my accomplished friend
Dr. D. D. Cunningham, F.R.S.
1 The Gymnosporangia, or Cedar Apples, of the United States: W. G. Farlow, 1880.