The Vegetation of Socotra

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706

0.B. POPOV: THE VEGETATION O F SOCOTRA

The vegetation of Socotra. By G. B. POPOV,


Desert Locust Survey, Nairobi.
(Communicated by Dr B. P. Uverov, C.M.G., F.R.S., F.L.S.)

(With Plates 5S59 and 1 Map


[Read 24 May 19571

INT~ODUCTION
The existing literature on the vegetation of Socotra is rather scanty. The island received
much attention from the collectors a t the end of last and early in this century, when
several British and German expeditions visited the island. Important contributions
were made to the flora, chiefly by Balfour (1888), Forbes et al. (1903) and Vierhapper
(1907), but these works only contain general brief descriptions of the physiognomy of the
vegetation without an attempt at the classification of vegetation communities or types.
Similar brief references are also found in Engler (1910),and Wettstein (1906),and the
available information was used by Pichi-Sermolli (1955) in his review of the plant
ecology of arid and semi-arid zones of East Africa.
The present author had an opportunity in 1953 to spend a few weeks on Socotra in
connexion with control of the Desert Locust. A general ecological survey of the island
was carried oyt in order to investigate its possible suitability as a habitat of the locust.
An account of locusts will be published elsewhere, and B list of Saltatorial Orthoptera of
Socotra, with zoogeographical analysis of the fauna, is in the course of publication
(Popov & Uvarov, in press).
The plants collected during the visit were first examined by Mr B. Verdcourt of the
East African Herbarium, Nairobi, where the duplicates were retained. The original
specimens were later presented to the British Museum (Natural History) for final
determination by Dr G. Taylor, Miss D. Hillcoat and Mr A. W. Exell. The names of plants
referred to in this paper were subsequently checked by Mr J. B. Gillett of Royal Botanic
Gardens, Kew, who also compiled the list on pp. 717-20. The author wishes to offer his
sincere thanks to the above and to Dr B. P. Uvarov for editing this paper.
PHYSICAL GEOGRAPHY A N D SOILS

The coastal plains. The southern coastal plain of Naukad is the largest on Socotra,
being some 30 miles long and up to 5 miles wide. To a large extent it i s covered with
packed gravels, stones and coarse sand, with an overlying mantle of loose sand. The loose
sand belt runs in a ribbon along the widest part of the plain, locally developing into windblown crescentic dunes up to 7 m.high (PI. 55, fig. 1). This sand has been largely formed
through the decomposition of granites, while the beach sands, formed all along this plain,
are mainly of shell and coral origin.
The watercourses traverse the plain along single, well-defined beds of boulders and
rock slabs, without extensive fanning out or leaving finer deposits. The banks are hardly
ever more than 1 m. high, in parts exposing recent conglomerates and raised coral
beaches.
Clay and silt deposits are found along some depressions, such as at Mahallas, where the
soil is somewhat saline, or in very small quantities along the minor watercourses which,
on being blocked by sands, spill and dry up over the plain.
Elsewhere, the southern coastal plain is narrow, and much of the southern coast
presents spectacular abrupt cliffs and bluffs, rising steeply to the plateau above. These
cliffs are honeycombed with caves and fissures and there is some water seepage, which
forms tufa encrustations and deposits in many p&rts of the cliffs.

G. B. POPOV: THE VEGETATION OF SOCOTRA


ZI

707

708

G. B. POPOV: TEE VEGETATION Or SOCOTRA

Between Ras Shoab and Katanahan there is a narrow inland basin at Nait separated
from the sea by a line of dunes, but filling up at spring tides. The water gradually evaporates, leaving a salt deposit, which is the main source of salt on Socotra. Similar basins are
said to exist in other parts of the island, as at Ras Bishuri, near Kalansiya.
The northern coast is composed of a number of smaller coastal plains, separated from
each other by rocky headlands. Kalansiya, Goba (Karma) and Hadibo plains are open
to the sea and correspond to the bays of the same name. Eastwards from Houlaf to
Momi, the lowland plains are separated from the sea by ranges of hiUs and divided by
watersheds into series of small river basins reaching the sea through narrow gorges; Kam,
Moabbadh and Khor-garieh are the major ones. From Hamadera to Ras Momi, the coast
is again formed by precipitous cliffs.
Much of the soil of the northern coastal plains, like that of Naukad, is composed of
packed gravel, stones and coarse sand, with a small admixture of clays. There are some
deposits of soft sand, on the Goba plain as thin sheets or hummocks under vegetation,
but at Houlaf reaching an unexpected thickness, covering the Houlaf promontory on the
one hand, and encroaching on to the seaward slope of Hawari hill to a height of 100 m.
or so, on the other (Pl. 55, fig. 2). It might be more than a coincidence that Acacia
edgeworthii forms an association in this area, but is rare elsewhere.
A feature common to the greater majority of the northward draining watercourses of
Socotra is the formation of estuaries, separated from the sea by sand-bars, which are
broken only by the heavy floods. The beds of rivers are covered by stones and boulders
most of the way across the plain, but h e r soils become deposited a t the estuaries, where
agriculture is practised. Even here the depth of the soil is so slight, that it is not sufficient
to take the full roots of the date palm, which on reaching the rock bed begin to arch and
bulge above the soil level.
The large river basin of Zahr was only seen from the air. No large sand deposits were
noticed, and in general appearance this area was similar to the coastal plain at Kalansiya.
The vegetation is probably akin to that of the coastal zones and not of the limestone
plateau proper.
The limestone plateau. This forms the greatest area of Socotra and varies in height
between 300 and 700 m., but in places, such as Reiged, rising to 900. Within the confines
of the limestone plateau, soil deposits are nowhere great. This area is much dissected by
gullies and ravines, and the surface worn smooth or pot-marked by erosion. Little
pockets of fhe, grey clayey soil occur in the hollows and between rock crevices, but
locally larger shallow soil deposits become accumulated in depressions,where the gradient
is slight. The largest of these were seen a t Homhil, on the summit of Reiged and the
plateau of Momi.
The granite highlands. The decomposition of granites, of which the Hagghier Massif is
built, has led to the formation of rich red soils. These have become accumulated in the
valleys and the plains and even along the less steep slopes, to form the deepest and most
fertile soil deposits on Socotra.

CLIMATE
The climate of Socotra is tempered considerably by the north-east and the south-west
monsoons and is less torrid than that of the adjacent mainlands. The temperatures
recorded by me in spring, 1953, on the coast a t Hadibo were: January, mean 24,
maximum 30, minimum 15.5; February, mean 25, maximum 31, minimum 16.5; March,
mean 26.5, maximum 32.2, minimum 18.3" C. The relative humidity during this period
varied from 75 to 90yo in the morning and from 55 to 75 yo a t midday. Occasionally
even higher humidity was recorded and morning dew was of frequent occurrence.
Higher temperatures were experienced in some of the valleys in the lowlands, and they
were usually associated with lower humidity. The highest recorded temperature waa
35" C. At higher altitudes, particularly in the Hagghier highlands, temperatures were

709

0. B. POPOV: THE VEGETATION OF SOCOTRA

noticeably lower, particularly at night and a minimum of 13.5" C. was recorded a t


Adho Demalu (lo00 m.) in late March. Frost, however, is unknown to the inhabitants.
May is said to be the hottest month, when heat and humidity rise during the period
of calms between the monsoons. Later, the heat is relieved by the south-west monsoon,
but the high winds are dry and cause desiccation, seldom bringing rain. September is
another period of calm airs and high humidity, which ends with the onset of the northeast monsoon, blowing until early May and bringing the main rain to Socotra, with the
highest precipitation in November and December.
Table 1 is compiled from rainfall figures recorded by the Royal Air Force Station a t
Ras Karma, the only locality on Socotra where regular weather observations have been
maintained. These figures can be regarded as representative of the amount of rainfall
that can be expected in the drier parts of the island. Rainfall is said to be greater in the
eastern part of Socotra than the western and, judging by the vegetation, it is probably
higher still in the highlands generally and in the Hagghier particularly where, in addition
to the rain, frequent mists and dews occur.
Table 1. Rainfall records for Ras K a m (in mm.)
1943
1944
1945

Jan.
tr.
4.8
3-6

Feb.

Mar.

0.1
2.1
3.5

tr.
31.4

Nil

Apr.
tr.
tr.
Nil

May
tr.
Nil

June
Nil
Nil

4.3

91.0

July
Nil
0.06
tr.

Aug.

Sept.

Oct.

Nov.

Dec.

tr.

3.2

Nil
Nil

89.6
60.2
5.5

75.2
76.6

4.0

3.8
4.1
21.3

Nil

VEGETATION
I n the following pages the major vegetation communities w i l l be described in turn, then
ascribed to the nearest vegetation type outlined by Pichi-Sermolli (1955). There are two
fundamental difficulties in fitting the vegetation communities of Socotra into this general
classification. The first is due to our comparative lack of knowledge of the climatic and
other factors affecting the vegetation of Socotra, which serve as the basis for the major
divisions ofthe vegetation types in other areas. The second is due to the peculiar structure
of some of the vegetation communities on Socotra, particularly from the point of view
of their physiognomy. Thus the Croton-short grass community, so prevalent on Socotra,
has no real equivalent in East Africa, and the rock communities do not reach the same
development there as they do on Socotra. On the other hand, the Acacia-short grw
savannah, so common in East Africa, is not represented on Socotra. The classificationhas
therefore not always been obvious, and the Socotran communities have been referred to
the one or the other type on the basis of a comparison and the personal acquaintance of
the author with the vegetation communities in other parts of East Africa.
The basis used by Pichi-Sermolli for the division of the vegetation of East Africa into
two major zones-the arid and the semi-arid-has chiefly been the aridity index.
I am in general agreement with Pichi-Sermolli, who regards the greatest part of Socotra
as being within the arid zone, with a narrow semi-arid belt in the upper part of the
Hagghier range, but he overstressed the aridity of the island, a point which w i l l be made
clear in the following description.
(1) The co&stalp h i m

Maritime and halophytic communities


Avicennia nzarinu mangroves occur as local narrow belts a t a number of points along
the coasts of Socotra. One of these was seen at Gubbet Nait, between Ras Katanahan
and Ras Shoab-the locality noted for its salt deposits (p. 708). Here, the salt flats are
fringed with a woody salt-bush, Arthrocnemum sp. ?, followed inland by a sward of coarse,
spiky salt-grass, Heleochloa dura.

710

Q.

B. POPOV: THE VEOETA!PION OF SOCOTRA

The beach dunes along the southern coastal plain of Naukad are fixed seawards by the
same salt-gram (Pl. 55, fig. 3) and landwards by sea-lavender, Linzon&unz spp., and in
some places by another salt-loving woody herb, AtP.ipkx.stoch%i forma sohtrana;
these three plants appear to be frequently associated with sands of coralline or shelly
origin.
Tamarix sobtram is evidently less halophytic than the foregoing species, as it often
grows away from the coast in less saline, sometimes gypseous soils. It is most common
at Khor Garieh, where it forms a narrow belt along the upper end of the inlet, which is
evidently subject to tidal inflows. Elsewhere it is occasional; either as a subdominant
within the Linwnium association or aa a rock-plant in some of the inland valleys and
gorges, usually on tufa deposits.
Suaeda naonoica is another member of the halophytic coastal communities of the
Naukad plain, but is really common only at Mahallaa, where it fringes the palm groves,
on low-lying silty soil. The wiry grass, Ae1urop-m sp., and the beautiful, red-flowered,
spreading convolvulaceous herb, I p m e u pes-cupae, occur in small colonies on the
beaches, but the rush Juncus arabicus, which commonly grows in the brackish estuaries,
can also be found in the marshes in some inland areaa, where water is quite fresh.
Some other plants such as Sporobolus spieatus, F q m i a socotrana, ZygophyUum sp.,
Heliotropium sp., Aerva microphylla and a few others are also tolerant of salt to a greater
or lesser extent, though probably not dependent on it. These are common members of
mixed xerophytic communitieson the coastal cliffs, growing nearest to the edge of the sea.
The above-mentioned communities can be directly ascribed to Pichi-Sermollistype 1,
Maritime vegetation (arid zone).

The sand belt


On the southern coastal plain of Naukad many of the dunes are completely devoid of
vegetation (Pl. 55, fig. l), while others support a thin growth of plants (Pl. 55, fig. 4 ;
P1. 56, fig. 5). No species of plant is evidently con6ned to this area, though some,
e.g. Calotropis prmra-the
large-leaved Asclepiad, are more common here than elsewhere on the island. The perennial plants probably tap the gravel soil below and as such
cannot be regarded as truly representative of the sand belt. Among these are Indigofera
sp.nov.,l I. oblongqolia, Ziziphus sp., Salsola fwskdii, Panicum rigidum, CorchomEs
depressus, Cucumis prophetarum and the above-mentioned Calotropis procera, which are
equally common on the adjoining gravel plain free of surface sand. The numerous annual
herbs and ephemerals are probably more repremntative of sandy soils,where they grow
in greater abundance than on the gravel plains. At the time of the visit (March),most
of these were already paat their prime and the following were recorded as common:
Fqoniu paulayana, Portulaca quudriJida, Dactylocteniurn aristatum, Phyllanthw sp.,
Hdiotropium sp., Ageratum wnymides, Boerhavia repens, Cenchrus cilia&, Enneapogon sp.
and ZygophyUurn simplex. As is common in the sandy areas of the adjacent mainlands
the cornunity is characterized by nanophanerophytes and ephemerals (Pl. 56, fig. 5).
Along the northern coast, only a t Ras Houlaf are the sand deposits deep enough to
affect the vegetation. Acacia edgeworthii forms a consociation here, whilst elsewhere it is
occasional. The dominant annuals, also largely conked to the sand, being Dactyloctenium
aristaturn and a species of Cyperw. The majority of other common plants, however, also
occur on the adjoining sand-free,gravelly and stony slopes of Hawari. Of these the most
noteworthy are Indigofera cordifolia, I. nephromrp, I. colutea, S a r a latqolia,
Aerva sp., Crotalaria sp., Tribulw terrestris, Lindenbergia sokutrana, Ruellia sp. and
Arnebia hispidissima (Pl. 55, fig. 2).
intricata Boiss. in the sense of Bdfour, Bot. Socotra, Tram. R. Soc. E d i d . 31, 74 (1888).not of
Boissier, Flora 07ientdis 2, 190 (1872). To be described shortly by J. B. Gillett.

G. B. POPOV

Journ. Linn. SOC. Bot. Vol. LV, PI. 55

G. B. POPOV

Journ. Linn. SOC. Bot. Vol. LV, PI. 56

Figs. 3-8. Explanation of Plate on p. 720.

Q.

711

B. POPOV: TIIE VEGETATION OF SOCOTRA

Coarse sand, gravel and stony plaina


The &st vegetation zone inland from the sea is formed by the woody herb community
(nanophanerophytes). At Hadibo the dominant species is Indigofera sp. nov.,
frequently associated with other woody herbs such as Convolwulusfastigiatus, Campylanthus spinosus, the tussocky grass Hyprrhenia h i m , some annual herbs, Tephrosia
apolliwa, Aerva javanica, Farsetia hgii&quu and a carpet of ephemeral short grasses,
Aristida A c e m i o n i s and dlelamcenchris abyssinica (Pl. 56, fig. 6). Along the river basins
east of Houlaf, this community is but poorly represented, but some of its elements occur
within the shrub zone (see below). Between Ghadheb and Kalansiya, however, it is well
developed, reaching a width of up to 3 km. on the Goba plain.
Other common nanophanerophytes in this area are Bidens bitemzata, Aerva microphylla,
Dactyloctenium huckelii, Panicum rigidunz, Euphorbia spiralis, E . septemsuhta and a
dwarf Commiphora; each of these plants being sometimes dominant locally but rare or
even absent elsewhere (Pl. 56, fig. 7 ) .
I n addition, Aloe perryi and Euphorbia arbuacula form local isolated communities in
some areas, while occasional bushes of Salvadora persiea, Acacia edgeworthii and C&a
rotundifolia, usually growing close to the seashore, are conspicuous by their larger size.
Inland this belt is followed by the shrub zone dominated by Metred, Croton socotranus, which is easily the commonest shrub on Socotra. Sometimes it forms almost pure
stands, but is more often associated with other shrublets such as Plumpoda virgata,
Justicia rigida, Lycium sohtranum, Grewia erythraea, Ballochia amoena, Trichocalyx
orbiculatus, Dirichletia obovata, Asparagus africanua var. microcaps ; some succulents,
such as Aloe perryi and Carduma sowtrana; herbs, Aerva lanata, Clossonemu revoilii,
Cucumis jicifolius, Pedalium murex, Alzagallis arvensis, with the grass carpet formed
mainly by Aristida adscensionis and Melamwnchris abyssinica (Pl. 56, fig. 8).
This shrub zone is fairly wide, extending to the river basins and the gravelly foothills,
where it merges into the mixed woodlands on the lower slopes, or ends a t the foot of the
limestone cliffs. Its density is evidently governed by the humidity and the edaphic
factors, and some of the gravel plains along the coast support little or no vegetation.
The succession of plants leading to the climax formation of Croton shrub zone, has been
observed a t Ras Karma airfield and other parts of Socotra, where parts of the bush were
cleared for runways and roads. The first invaders were the ephemeral and annual herbs
dominated by Tephrosia apollinea, followed by such perennials as Corchorus depressus
and Cucumis prophetarum, then by perennial prostrate grass, Panicum rigidum, as a
sand-binder for small mounts of drifting sand, then bushes of Indigofera sp.nov. and last
of all Croton sowtranus. The succession has been slow due t o predation by goats and sheep
and the climax has far from re-established itselfin the eight years since the abandonment
of the station a t Ras Karma.
The vegetation communities and zones described above can safely be attributed to the
Subdesert shrub and grass type of Pichi-Sermolli, although the dominant Croton-short
grass community is not found on the East African mainland.
Minor hills and dry hill slopes
The Croton socotranus association frequently encroaches on the hill-sides and sometimes
remains dominant there. I n some localities, however, particularly at a height of
100-250 m. it becomes a t least partly replaced by the Commiphora community represented by such xerophilous elements as Comrniphora parvifolia, C. socotram, Aloe perryi,
Euphorbia nubica, E . spiralis, Barleria tetracantha, Blepharis spiculifolia, Cissus subaphylla
and others.
It is easier to regard this community as a transition between the subdesert shrub and
grass and the subdesert shrub with trees, rather than to attempt t o ascribe it to either.
JOURN. LI. SOC.-BOTANY,

VOL. LV

XH

712

Q.

B. POPOV: THE VEGETATION OF SOCOTBA

(2) The 1-p-u


The dopes of the limestone plateau are often sheer cws, but sometimes they are less
steep, composed of tumbled-down slabs and chunks of rock and shelves separated by
stony screes. Not only is the whole limestone block honeycombed with fissures, caves
and ledges, but the individual rocks are scoured with cracks and hollows, where particles
of soil accumulate, affording foothold for vegetation (Pl. 57, fig. 11).
These slopes are veritable rock gardens, surprising for the variety of their 00ra. There
is undoubtedly some gradation with altitude, but the limits were not rigid, particularly
for some species ; the decisive factor probably being the occurrence of mists, and the main
change becomes apparent a t about 500 m. The plants growing nearest the edge of the
sea have a.lready been mentioned. A short distance above, Dendrosicyos sowtrana, Cissus
subaphyllcl, Euphmbia arbu-scuh and Adenium sokotranurn combine to form a landscape
peculiar to Socotra (Pl. 57, fig. 9), together with some Euphmbia nubicu, E . spiralis,
Commicaryuus sp., Mmrma sowtrana, Withunia riebeckii, Cynanchum linifolum. In
addition, the Croton sowtranus community becomes established wherever soil deposits
are suEcient to permit its development.
The annual and perennial herbs are equally varied and abundant, and the commonest
recorded were Triclwdesma microcalyx, Pdiearia skphuwcurpa, P . diversifolia, Corchorus
erodioides, Lactuca r h y n c k r p a , Crotdaria leptocarrpa, Ipowweu blepharocephah,
Boerhavia repens, M e o k ~ g ominim, Convoldus q k r d u s , Oldenlandia pulvinata, and
the ferns Actinbpteris austrdis and Adiantum balf.u.ii.
At the height of 500 m. and above, rock-plants are represented by Dorstenia gigas,
Kleinia swttii, Ficus sowtrana, Boswellia sp., presenting an equally characteristic if
somewhat different appearance (Pl. 57, fig. 10). Other common elements within this
mixed community are BosweUia sp., Tetragoniapentandra,Adenium socotranum, Euphorbia
oblanceolata, Hibiscus swttii and some others (Pl. 57, fig. 11).The young seedlings usually
require a small quantity of soil but, once established, the plants appear to draw sustenance
from the bare rock.
Other plants, such as Psetdomussm& q d ~ e r aPolymrpaea
,
spicata, P . de'varicuta
and Haya obovata, grow suspended from ledges of rock. Yet others are only found growing
in pockets of dark rich soil, deep in the crevices of rocks; of these Begonia sowtrana,
Exmum afine, Peucdanum wrdatum and the fern Adiantum balfourii are the commonest. A greater number, however, show preference for screes, shelves and valleys,
where some soil becomes accumulated. Where soil deposits are poor, the vegetation is
characterized by succulents such aa Aloe perryi, Kdancluw farinacea, K . robu-sta and
some small woody herbs-Melhunia
murim%, Diceratella incana, Leucas urticifolia,
Helichrysum gracilipu, Teucrium sohtranum, in addition to the above-mentioned rock
plants and an occasional E u p h b i a arbuscuh. I n sheltered valleys, where soil deposits
are more abundant, sometimes fairly dense thickets become established; here the
commonest shrubs and trees are represented by Ruellia insignis, Ormcurpum caerukum,
Acacia pennivenia, Dicliptera effusa, Croton sulcif&us, Psiadia schweinfurthii, Vemnia
cockburniana, Rhus thyrsi)?ora, Ficus sowtrana, Justicia heterocarpa, Acridocarpus
sowtranus and Cissu-s quadranqdaris. These are bound by lianas and creepers, such aa
C. panicuhta, Tragia bdfourhna, Curroria decidua subsp. volubilis and Diosmea lamta.
The above description is based on the vegetation of Reiged, the 900 m. high limestone
bluff west of Hadibo. Although the majority of the plants mentioned were also found in
the other parts of the limestone plateau, the composition of the vegetation varied and
the dominants in one locality were sometimes rare or absent in others. The most noteworthy example is the Drmnu-Boszuellia community on the slopes of the Hamadera
hills a t Homhil (Pl. 57, fig. 12), chiefly composed of Dramena cinnubari,Bomellia ameero,
B. elcrngata, B. sowtrana, Aloe perryi, Adenium sokotranum and Mitolepis intricata. While
Dramena is more wmmon on the limestone slopes, B d l i a predominates on the plain

G. B. P O P O V

Journ. Linn. SOC. Bot. Vol. LV. PI. 57

G. B. POPOV

Journ. Linn. SOC. Bot. Vol. LV, PI. 58

Figs. 13-16. Explanation of Plate on p. 720.

G. B. POPOV

Journ. Linn.

Fig. l i

SOC. Bot. Vol. LV, PI. 59

713

0. B. POPOV: WE VEQETA!t'ION OF SOCOTRA

where, in addition, a species of Lunnea, not seen elsewhere, forms a small shady grove.
The vegetation on the southern hills surrounding Homhil is rather different, forming
a mixed shrub thicket, the main components of which are Rhus thyrsifEwa, Dirichktia
venulosa and Punica protopunica.
Lannea aspleniifolia is a common member of the communities on the limestone slopes
inland from Kalansiya, but rare elsewhere, while the vegetation of the cliffs along the
southern coastal plain is characterized by a predominance of rock plants, chiefly shrubs,
such as Boswellia spp., Dyerophytum pendulum, Cyptolapis wbicularis, Cochlunthus

socotranus, Secamone sowtrana, Euphorbia oblanceolata,E . nubica, Hibiscus swttii, Barleria


spinma and Dorstenia gigas.
The altitudinal zonation is also variable; for example, along the northern coast,

Dorstenia gigas is seldom seen below 500 m., while along the southern coast it is commonly
encountered down to 100 m.
With the exception of the thickets found in the sheltered valleys, the communities of
the limestone plateau described so far can be ascribed to the subdesert shrub with trees
LW has been done by Pichi-Sermolli. The mixed thickets ofthis region, together with those
on the lower slopes of the Hagghier, on the other hand, may be referred t o the subdesert
bush and thicket (type 6 of Pichi-Sermolli).
By contrast with the slopes, the top of the main limestone plateau is remarkably arid.
The perennial vegetation is rather scanty, consisting of stunted bushes of Jatropha unicostata, Croton socotranus, Picus socotram, Adenium sokotranum, Aloe perryi and an
occasional Dracaena cinnubari. The annual vegetation is more abundant, being chiefly
composed of grasses-Hyparrhenia hirtu, Arthraxm lancifolius, Aristida funiculata,
Cymbopogon sp. and Pennisetum setaceum ; some herbs-Asphodelus tenuifolius, Digera
alternijolia, Achyranthes aspera and some others (Pl. 58, fig. 13).
Here again there is some difference in the plant composition between the various parts
of the island: thus the community populating the summit ofthe Hamadera hill is chiefly
characterized by low woody perennials such as Mitolepis intricata, the larger shrubs and
trees being absent. The watershed between the coastal valley of Kalansiya and the Goba
plain supports a thin growth of Jatropha uniwstatu and Adenium sokotranum, in parts
replaced by the Croton association.
The paucity of the vegetation on the plateau may not be so much due to the soil and
aridity, since many vegetation communities develop well on rocky slopes under even
more arid conditions, as to the high winds, particularly during the south-west monsoon.
It is significant that vegetation was found to be much better developed in ravines and
gorges where the adjoining cliffs afford some protection. Thus on Reiged, mixed thickets
have developed on the uppermost shelf, but are absent from the summit.
Parts of the limestone plateau are indeed so sparsely clothed with vegetation that one
feels tempted to ascribe them to the desert vegetation, as has been done by PichiSermolli on the information provided by Balfour (1888). However, the sparsity of the
vegetation here is mainly due to one outstanding climatic factor (wind) and not general
d a e r t conditions, and many of the species found here are prominent members of the
semi-arid vegetation elsewhere on Socotra. In all fairness, therefore, one should refer
the largest part of the limestone plateau t o the Subdesert shrub and grass type of PichiSermolli, with islands of less arid vegetation belonging to the Subdesert shrub with trees
and the Subdesert bush and thicket types.
(3) The granite massij of the Hagghier
Along the northern slopes, the coastal shrub-zone extends to the foothills, where it
merges with the mixed thickets composed of a variety of shrubs such as Jatropha unicostata, Dirichletia obovata, Cordia obovata, Cylista sp., Acridocarpm socotranus ; trees like
8krculia rivae, Aberia abyssinica and Tamarindus indica, and an undergrowth of herbs
and annuals such as Tewrium balfourii, Leucas neu$izeana, L. urticifolia, L. virgata,

xx-2

714

G. B. POPOV: THE VEGETATION OF SOCO"E.4

Jwtkia heterocarpa and many others mentioned for the shrub zone and the lower slopes
of limestone plateau (Pl. 58, fig. 14).
With an increase in altitude, and presumably rainfall, vegetation becomes denser;
many of the elements mentioned for the higher slopes of the limestone plateau such as
Rwllia insig.niS, Rhw thyr&@a, ownmwpum CaeFUleum, Ckrodendmmg&um are also
fairly common here, but some due to better edaphic conditions preeent a somewhat
different "pect. Thus, Fkw socotmna, which in the limestone belt grow as a stunted
rock shrub, here develops into a h e spreading tree.
There is no difficulty in ascribing these mixed thickets together with those of the
sheltered valleys of the limestone region to the subdesert bush md thicket type of PichiSermolli. These thickets gradually change to the evergreen thickets of the Hagghier
Highlands, which belong to the semi-arid vegetation zone.
The main change in vegetation becomes apparent at an altitude of over 750 m., where
many elements appear for the Srst time. The vegetation of the Hagghier Highlands may
be divided into three main types: the grasslands, the thickets and the association of
woody herbs. One might include the rock vegetation, but this community does not reach
the same development on granites, as on the limestone. The surface of rocks is covered
with lichens so thickly that the naturally pink granite appears grey from a distance.
The higher forms of flora, however, are restricted to the occasional fissures where succulents such as K&nchoe farinacea, K . roclm&i;folia,
Aloe per+, and some herbs such aa
Exmum a@m, 2. mruleum and Begonia socotrana find a foothold. However, there
appear to be no rock shrubs.
The grmshnda are to a large extent composed of The&
pwadrivaEvis, Hypwhenia
hirta and Arthramn hncqoliw (Pl. 58, fig. 15). They form the main pastures for the
Socotran cattle and cover a fairly large area of the watershed between the granite peaks.
The division between the grasslands and the thickets is a very sharp one.
The thick.& occupy the lower slopea of the highlands, but are absent from the steeper
peaks. Their specific composition is a rich and varied one, the following being the
commonest in order of their dominance: shrubs and small trees-Cep?u&croton sowtranus, Carissa eddis, Bum~shildebrandtii, Dodonaea viscosa, Ficus sowtrana, Indigofera solcotrana, RueUiu insignis, BosweUia amzero and E u p h b i a soeotrana; smaller
bushes--iliedyotis stellarioides, Hypoestes pdmwm, Ahphylus rAwiphyUus, together
with a tangled undergrowth of Cocculus balfourii, C i s w qu.udran&aris, the twining
Traqia baifourianu and E&kp8ds sp. and the Dioscorea h&
lianas (Pl. 58, fig. 16).
The community of the woody herbs develops on the stony slopes leading steeply from
the watershed, where it replacea the grasslands and the thickets. This is characterized by
such species as Euryqs socotranus, Hypericum torCuosum, H. mysorense and Aerva
revol'uta, with an h o s t Alpine appearance. Other common perennial and annual herbs
include Pluchea obovatu, P.aromatics, Pulhria vier&,
&idia socotrana, hillonia
puberula, lsatureja remota, Sigeshkia opientalk and Ldw munwpsis.
fiacuem cindmri is fairly common, though never as dominant as at Homhil; the
other plants noticeable for their appearance being the flat-topped Commiphora plunifrom and laxge-leaved amaryllid, Haemanthus grandifolius.
The higher peaks of the Hagghier, above 1200 m., were not visited, but according to
the eaxlier deacriptions, they are characterized by such woody herbs as Nirarathanznos
ma&foliw, Diehrmphlu chrysanthmifoliu and various species of Hdichrysum.
The vegetation of the Hagghier Highlands belongs to the semi-arid zone, and one can
attribute the T-Bpwhenia
grassland to the Open gradand and the thickets
to the Evergreen scrub (types 3 and 4) of Pichi-Sermolli. The nanophanerophytic
community of the Hagghier is less easy to classify. Climatically it belongs here, physiognomicdy to the Subdesert shrub and grass type of the arid zone. Perhaps it is best to
regard it as part of the Evergreen scrub type.
The southern slopes of the Hagghier are more arid than the northern, particularly at

Q. B. POPOV: THE VEQETATION

715

OF SOCOTEA

lower altitudes, where mixed thickets soon become replaced by the Croton shrub, or
Boswellia--succulent communities, which in the southern part of Socotra extend to a
greater altitude than in the northern. Aloe forbesii, in this area, almost completely
replaces A. p w y i .
(4) Riverim communities
The majority of the streams on Socotra are sporadic, carrying water only during and
for a short time after rains. In their upper reaches, particularly, the vegetation fringing
these runnels is hardly different from that of the adjoining slopes. There are, however,
a number of springs, streams and pools and some river estuaries, which retain water for
a greater part of the year and thus encourage the formation of plant communities not
represented, or at least not so fully developed elsewhere. The following are the most
noteworthy.
( a ) The semi-aquaticcommunity represented by Juncwr arabicus consociation, which
forms along most estuaries, as well aa along some of the stagnant pools and springs in
the interior, where in addition it includes the tall sedge Cladium m r i s c u s .
The shallower parts and edges of sluggish streams are lined with Lythrum hyssopifolia, Phyla rwdijbra and with the aquatic graas, Paapalidium gemidurn, growing in
deeper water. In sheltered parts along the edge of streams, ferns such as Pteris vittata
and Adianbm balfmrii are common. The upper parts of estuaries, where water begins
to stagnate, are carpeted with a low spongy growth of sedges and grasses of the genera
Fimbristylis, Cyperus, Erapostis and Paapdidium.
( b ) The date palm, Phoenix dactylifera, which is the main cultivated plant on the
island, is planted along the streams, where water-table was high and wherever soil
permits this. In addition, the streams with the water at or near the surface are lined
with groves of Ficus salicifolia, Euclea sp., Cordia obtusa, Acridocurpus socotranus, and
Buxw hildebrandtii with an occasionallarger tree such aa Sterculia rivae, Lunma ornifolia,
Tamarindus indica and Ziziphus sp., the latter usually associated with drier conditions.
There are also many herbs, particularly common along these watercourses: Leucas
virgata, Ocimum hadienae, Hypoestes sobtram, Cassia tora, Heliotropium zeylanicum,
Hibiscus sidiformis, Aerva javanica, Pedulium murex and Ver?wnia cineraacem being the
most prominent, while the ephemeral grasses are represented by Aristida adacemionis,
Chloris barbata, Cenchrus setigerus, Rhynchdytrum microstaehyunz, Apluda mutica and
Heteropogon contortus (Pl. 59, fig. 17).
(c) The drier, sandy stretches of main watercourses, where surface water is absent
most of the year, such aa the one at Hodaf, are characterized by Ziziphus sp., Ochra&nus baccatus, and such herbs aa Aerva javanica, Tephrosia apllinea, C l e m sp. and
Ruta graveolens.
The above description is based on the Hadibo plain, but is generally true of the northern
coast aa a whole. The valleys of the southward draining rivers, which cross the Naukad
plain, were found to be larger and more arid and the change of the vegetation more
gradual. The southern riverine valleys of the Hagghier contain groves of the wild orange
Citrus aurantium,A c r h w socotranus,Peperovmiare$exa, Ziziphus sp. and Tamarindus
trees together with an occasional Borassus palm. The lower reaches support Ziziphus
sp. and the much-branching smaller Ziziphus sp., the latter persisting all the way to
the coastal plain, where the sandy dry bed of rivers is, in addition, dotted with bushes of
Aerva microphylla, Indigofera sp.nov. and Dadyhtenium hackelii much more densely
than the adjoining plain (Pl. 59, fig. 18).
( 5 ) Agriculture
Most of the flood waters quickly drain off Socotra, but a small quantity becomes
absorbed and later reappears as small streams and springs. The granite range is particularly water-bearing and many streams draining from this range are said to flow for about
xx-3

716

a. B. POPOV:

THE VEGETATION OF SOCOTEA

eight months in a year, while permanent springs abound in this region. The limestone
plateau is much more arid, particularly a t the western end of the island, but even here
there are a number of permanent springs, while small amounts of rain wafer, which
collect in hollows, may sufice for human consumption for up to 3 months after rain.
On the southern coast, water is scarce, limited to one or two springs at the base of cliffs
and several wells of brackish water. On the northern coast, the estuaries of larger rivers
retain water for most of the year and water-table is high at Hadibo, Ghadheb and
Kalansiya, where nearly every household has its well. On the Goba plain, however,
water is brackish.
Agriculture is practised on a smali scale. Date gardens exist along most estuaries and
river basins on the north mast, while on the southern, there is an extensive plantation
at Mahallas. Where soil is inadequate for the growth of the plant, additional soil is piled
up against the roots, within a circular stone wall, giving the palms the appearance of
growing in pots.
Vegetable gazdens, irrigated from wells, exist at Hadibo and one or two other villages
along the north coast. Sweet potatoes, pumpkins, cotton, tomatoes, pepper and some
other vegetables are grown for household use.
There are also one or two small fields of finger millet (Eleusinecwacana) a t Moabbadh,
irrigated from the adjoining stream. These are said to be the property of the Sultan and
are probably still in the experimental stage.
Wild orange trees (Citrus aurantium) are found in abundance in the valleys of the
southern Hagghier. The plant may have been originally introduced, but appears to be
growing wild now. The pods of Tamrindus indica, the berries of Cordia sp. and some
other indigenous plants such as G h s m m a revoilii (fruit)and Dwswrea h m h (tubers)
are eaten.
Although most of the population of Socotra is pastoral, and stock breeding is undertaken as keenly as on the adjoining SomalilandPeninsula, overgrazingdoes not seem to be
a problem on Socotra. This is almost certainly due to a balance of stock population
maintained by natural causes in the absence of veterinary services, rather than to any
control by the inhabitants. I n any w e , the recent census shows no notable increase in
either human or stock population since the end of the last century, or even compared
with the figures given by early travellers.

(6) Weeds of cultivatim


Many of the weeds have probably been introduced relatively recently, as suggested by
their abundance in the cultivations in the vicinity of villages and their relative rarity
elsewhere. The commonest noted were Argemme qnexicana, Portulaca oleracea, A i m
mnariense, Trianthma penhndra, Psoraleu wrylijolia, Datura metel, Solanum
coagukcns,Cassia h a , Cassia sp., Ricinus unnmunis and some others. These are common
both in the palm-gardensand the small vegetable gardens near the villages. Other weeds
are often also common members of the coastal and some other communities on the
island, and a few of them are endemic to Socotra. Of these, Hdiotropium dentaturn,
ZygophyUum simp&, Achyranth aspera, Digera alternifolia, Indigofera tinctoria and
Sida mats are the most noteworthy.
SUMMARY

Our information on the extent and the distribution of the vegetation types on Socotra
referable to the arid zone may be summarized as follows:
1. Maritime vegetation is present over small areas along the coast of Socotra. The
observed communities have been described and similar, or slightly different, ones are
believed to exist to a limited extent in other parts of the island which were not seen, such
as near Kalansiya and a t Shoab and possibly elsewhere.

G . B. POPOV: THE VEGETATION OF SOCOTBA

717

2. Deszrt vegetation. No vegetation community can truly be ascribed to this type.


There are limited gravel and sand stretches on the coastal plain, where the paucity of the
vegetation is due to edaphic aridity, and parts of the limestone plateau, where perennial
growth is reduced by winds, but the existing vegetation can be described as a debwed
form of other plant communities reaching the climax under more favourable conditions.
3. Subdesert shrub and grass covers most of the island, i.e. the whole of the southern
coast, most of the northern coast and the western and central parts of the limestone
plateau.
4. Subdesert shrub with trees is another well-represented vegetation type, comprising
the rock communities of the limestone slopes, the BosweUia-Drc,cuenu community of
Homhil and possibly other similarly sheltered parts of the limestone plateau.
5 . Subdesert scrub is not represented, a t least in its typical composition. On the basis
of the vegetation cover one might possibly regard the denser Croton formations aa
belonging to this type, but this would be atypical in relation to the communities
characteristic of this type.
6 . Subdesert bush and thicket. The mixed thickets of the lower slopes of the Hagghier
and some of the sheltered valleys within the limestone region, such as the Goahal gorge
can only be ascribed to this type, even though somewhat atypical.
7. Xerophikvus open woocuand is not represented in its typical composition.
8. Vegetation of sites where water is present includes numerous riverine communities
and the cultivations of Socotra.
9. Vegetation of semi-arid type. The vegetation of the semi-arid zone is confined to the
highlands of the Hagghier, above the height of 850 m., where only two vegetation types
can be recognized, the Open grassland and the Evergreen scrub.

Species referred to in G. B. Popovs paper, The Vegetation of Socotra


by J. B. Gillett, Royal Botanic Gardens, Kew.
Families in the sequence of Bentham and Hooker, Genera Plantarum
Taxa marked with an asterisk are believed t o be endemic t o Socotra, those with
two asterisks belong to genera endemic to Socotra (in view of our imperfect botanical
knowledge of the Horn of Africa these statements must be considered as provisional).
Menispermaceae
*Cocculua balfourii Schweinf. ex Balf. f.

Papaveraceae
Argenwne m x i c a n a P.
Cruciferae
*Diceratella incana Balf. f.
Farsetia longisiliqua Decne.

Capparidaceae
Cadaba rotundifolia Forsk.
Cleorne sp.
Maesocotrana (Schweinf.) Gilg

Resedaceae
Ochradenua baccatua Del.
Flacourtiaceae
Aberia abyssinica clos

Caryophyllaceae
**Haya obovatu Balf. r.
*Polycarpaea divaricatu Balf. f.
P . spicata Wight ex Am.
Portulacaceae
Portulaca oleracea L.
P . quadri;fida L.

Tamaricaceae
Tamurix aokotrana Vierh.
Hypericaceae
Hypericum mysorense Wight & Am.
*H. tortuosum Balf. f.
Malvaceae
*Hibiscus scottii Balf. f.
H . siclifomis Baill. ( H . t e m t w r (c8V.)

Mast. not of Cav.)


Si&a ovata Forsk.
Sterculiaceae
Melhania muricata Balf. f.
Sterculia rivae (K. Schum.) Chiov.
Tiliaceae
Corchorus depressus (L.) Chistens.
*C. erodwides Balf. f.
G e e erythraea Schweinf.

Malpighiaceae
*Acridocarpua socotranwr Oliv.
Zygophyllaceae
*FagoniapaulayarmWagn. & Vierh.ex Vierh.
F . sowtrana (Balf. f.) Schweinf.

G . B. POPOV: THE VEGETATION OF SOCOTBA

717

2. Deszrt vegetation. No vegetation community can truly be ascribed to this type.


There are limited gravel and sand stretches on the coastal plain, where the paucity of the
vegetation is due to edaphic aridity, and parts of the limestone plateau, where perennial
growth is reduced by winds, but the existing vegetation can be described as a debwed
form of other plant communities reaching the climax under more favourable conditions.
3. Subdesert shrub and grass covers most of the island, i.e. the whole of the southern
coast, most of the northern coast and the western and central parts of the limestone
plateau.
4. Subdesert shrub with trees is another well-represented vegetation type, comprising
the rock communities of the limestone slopes, the BosweUia-Drc,cuenu community of
Homhil and possibly other similarly sheltered parts of the limestone plateau.
5 . Subdesert scrub is not represented, a t least in its typical composition. On the basis
of the vegetation cover one might possibly regard the denser Croton formations aa
belonging to this type, but this would be atypical in relation to the communities
characteristic of this type.
6 . Subdesert bush and thicket. The mixed thickets of the lower slopes of the Hagghier
and some of the sheltered valleys within the limestone region, such as the Goahal gorge
can only be ascribed to this type, even though somewhat atypical.
7. Xerophikvus open woocuand is not represented in its typical composition.
8. Vegetation of sites where water is present includes numerous riverine communities
and the cultivations of Socotra.
9. Vegetation of semi-arid type. The vegetation of the semi-arid zone is confined to the
highlands of the Hagghier, above the height of 850 m., where only two vegetation types
can be recognized, the Open grassland and the Evergreen scrub.

Species referred to in G. B. Popovs paper, The Vegetation of Socotra


by J. B. Gillett, Royal Botanic Gardens, Kew.
Families in the sequence of Bentham and Hooker, Genera Plantarum
Taxa marked with an asterisk are believed t o be endemic t o Socotra, those with
two asterisks belong to genera endemic to Socotra (in view of our imperfect botanical
knowledge of the Horn of Africa these statements must be considered as provisional).
Menispermaceae
*Cocculua balfourii Schweinf. ex Balf. f.

Papaveraceae
Argenwne m x i c a n a P.
Cruciferae
*Diceratella incana Balf. f.
Farsetia longisiliqua Decne.

Capparidaceae
Cadaba rotundifolia Forsk.
Cleorne sp.
Maesocotrana (Schweinf.) Gilg

Resedaceae
Ochradenua baccatua Del.
Flacourtiaceae
Aberia abyssinica clos

Caryophyllaceae
**Haya obovatu Balf. r.
*Polycarpaea divaricatu Balf. f.
P . spicata Wight ex Am.
Portulacaceae
Portulaca oleracea L.
P . quadri;fida L.

Tamaricaceae
Tamurix aokotrana Vierh.
Hypericaceae
Hypericum mysorense Wight & Am.
*H. tortuosum Balf. f.
Malvaceae
*Hibiscus scottii Balf. f.
H . siclifomis Baill. ( H . t e m t w r (c8V.)

Mast. not of Cav.)


Si&a ovata Forsk.
Sterculiaceae
Melhania muricata Balf. f.
Sterculia rivae (K. Schum.) Chiov.
Tiliaceae
Corchorus depressus (L.) Chistens.
*C. erodwides Balf. f.
G e e erythraea Schweinf.

Malpighiaceae
*Acridocarpua socotranwr Oliv.
Zygophyllaceae
*FagoniapaulayarmWagn. & Vierh.ex Vierh.
F . sowtrana (Balf. f.) Schweinf.

718

-0.B. POPOV:

THE VEQETATION OF SOCOTBA

Zygophyllaceae (wnt.)
Tribulue twrsstris L.
ZygophyUum &n&x L.
Rutacese
Ruta gravwkm L.
Citrcls mrantium L.
Buraera.ceae
*BosweUia ~umeeroBalf. f.
*B. ebngata Balf. f.
*B. 8oCOi%KMU&Balf. f.
C m p m w f & (Bslf.f.) -1.
*C. p h n i f m (Balf.f.) Engl.
eowmna (Belf.f.) Engl.
Rhemnaceaa
Zizyphua sp.
Vitaceae
* C k m pun(Balf.f.) Planch.
qua&L.
*C. uubqvhyllu (Balf. f.) Planch.
Sepindweee
D0dona;ea viawsa (L.) Jwq.
*AlbpbyLua rhuuiphyllua Mf.f.

*c.

c.

*Lanneuq h G f o l i a (Balf.f.) Engl.


*L. orn$oZia (Belf.f.) Engl.
*Rhw thyrsiijlore Balf. f .
Leguminosae: Pspiliomtse

cram lt?ptouwpa Balf. f .

c y l k h Sp.
Indigofera wlutm (Burm. f.) Merr.
I. wrdifolia Heyne ex Roth
I. m p h m r p a Balf. f.
I . obbngifolia Forsk.
*I.aokotrana Vierh.
I . tinetoria L.
*Loworro1Eop8iB Balf. f.
Medimgo mimima L.
Ornwmrpum caertJeum Balf. f.
P 8 0 r h W9yc%fO&J L.
T e p h r o h apollineu (-1.)
Link.
Caesalpinioideaa
C a a h tura L.
Tamm-indua indic0 L.
Mimosoideaa
Acacia edgeuwthii T.Andera. (A. sowtmna
Balf. f.)
*A.pennivenia Balf. f.
Crassulaceae
*Kahne?we farinacm Balf. f.
*K. robuatu Balf. f.
K. rotundifolia Haw.
Lythr&CeaS
Lythmm hyaao@fo&z L.
PUnim*Punica potopu7aic0 Balf. f.
Cucurbitaceae
Cucumkjkifoliua A. Rich.
C. prophtarum L.
**Dend.ro&yoa soc~tranaBalf. f.

Begoniaceaa
*Begonia s o c ~ & a n aHook. f.
Aizoaceas
Aizom canarkme L.
*Tetregoniapentandm Bslf. f.

TAnthema penta;ndra L.
Umbelliferae
* * N k r a t h ~ w a.8a&foliua
a
Balf. f .
*Peucedanum cmdatum Balf. f.
Rubiweae
D .w.d k t i a obovata Balf. f.
*D. wn&sa Balf. f.
* # a h & pube&
Balf. f.
*HedyotiS atelkBalf. f.
*O,!ddandia pubinafu (Balf. f.) Vierh.
**Pluwpodu virgata Balf. f.

(Balf. f.)
WeRlh8ITl
Compositse
Ageratum conyzoides L.
Bidens W m d a (Lour.)Merril & SherfT
Dichmcephula &rysanthem@olia (Bl.) DC.
*Euryopam t r a n u a Balf. f.
Heliohryeclm
Oliv. & Hiern.
*Kkinia ecoti% (Balf. f.) Chiov.
+La;ctuccr rhyndwmrpa Balf. f.
* P l u c k aromatic0 Balf. f.
*P. obovatcr Balf. f.
*Peia&io achweinfurthii BaH. f.
*Pdkwia divemifolia Balf. f .
*P. atepha7wmrpa Balf. f.
*P. viemeoides Balf. f.
Sigeabkia orienta2is L.
Ver?wniacimraacem Sch. Bip.
* V. cockhrniana Balf. f.
Plumbrrginsceae
*Oyemphytum penddum (Balf. f.) Kuntze
Sp.
Primulaceae
AnagaUis amen& L.
Salvadoraceaa
s
u persica L.
Ebenweae
E u c h ap.
Apocynmeaa
*A&niumaokotranum Vierh.
C&sa eddk (Forsk.) Vahl
Asclepiadaceae
Calotropia p c e r a (Ait.) Ait. f.
Carallurn eocotrana (Balf. f.) N.E. Br.
**Cochlcmthuaaocotranua Balf. f.
Cryptokpia m~ularischiov.
Curroria decidua Planch. ex Benth. sap.
volubilis (Balf. f.) Bullock
* C p n e h u m linifolium (Balf. f.) Bullock
comb.nov. (Viwtoxkum linifolium Balf.
f. in Proc. R. SOC.Edinb. 12, 79 (1884))
Ectudiop& sp. (aensu Bdf. f.)
c f b a a a m a rev&& Franch.
*P8f3UdO?nU&W43dU @$E?'tZ

0.B. POPOV: THE VEGETATION OF SOCOTRA

*+Mitole@ intPicarla Balf. f.


*Secamne socotranu Balf. f.
Gentianma
*Emcum aflne Balf. f.
*E. CaeruIeUllz Balf. f.
Boreginaceae
Arnebia hiepidkeinut (Lehm.) DC.
*Cordia obowcta Balf. f.
*C. o b t m Bdf. f.
*Helwtl.qpium dentaturn Belf. f.
H . zeylanicum (Bunn. f.) Lam.
*Trichodesma microdyx Balf. f.
Convolvulaceae
Conuolvulw,faatigiatua Roxb.
C. gbmeratua Choisy
Ipomoeu bkpbrosepak Hochet. ex A. Riah.
I . p c a p m (L.) Roth
SeclcEeralatifoliaHochat.&Steud.exHochst.
Sol&naceae
Datura metel L.
*Lycium sokotranum wagn. & Vierh.
Sohnum waguhm Forsk.
* Withania riebeckii Schweinf. ex Balf. f.
Scrophulariacem
*Campylanthua s p i n o m Balf. f.
*Lindenbergia sohtrarm Vierh.
Pedaliaceae
Pedalium murex L.
Acanthaceae
**BaUochia a m e n u Balf. f.
Bark& apinosa Hook. ex Nees
*B. tetracantha Balf. f.
*Blepha& S p i c d i f O h Balf. f.
*Dicliptera effaa Balf. f.
*Hypoestes pubescens Balf. f.
*H. aokotrana Vierh.
Juaticia hetwocarpa T . Andera.
*J.rig&
Balf. f.
*Ruellia imignis Balf. f.
+Trichocalyx orbiculata Balf. f.
Verbemceae
Avicennia marina (Forsk.) Vierh.
*Clero&ndrum galeatum Balf. f.
Phyla nodifira (L.) Greene
Labiatae
Leucm neuflizeana Courb.
L. urticiifolia (Vahl) Benth.
*L. virgata Balf. f.
Ocimum hadieme Forsk.
*Satureja r e m t a (Balf. f.) Vierh.
*Teucrium balfourii Vierh.
*T.sokotranum Vierh.
Nyctaginaceae
Boerhavia repens L.
Commicarpua sp.
Amaranthaceae
Achyranthes apsera L.
Aerva javanica (Bunn. f.) Jw. ex Schult.
A. lanuta (L.) Juss. ex Schult.

719

A. microphylla Moq.
*A. revoluta Balf. f.
Digera altemifolia (L.)Aachera.
Chenopodiaceae
Arthmmum sp.
Atripkx stock& Boies. forma mkotrana
(Vierh.) Vierh.
Sdaola forakulii Schweinf.
Suaeda monoica Forsk.
Thymedleaceae
*&id& sowtrana (Balf. f.) Gilg
Buxeceae
B u x u hildebrandtii Baill.
Euphorbiaceaa
*Cephaloeroton sowtranua Balf. f.
*Croton socrotranua Balf. f.
*C. &+uctua Balf. f.
*Euphurbia arbuaula Balf. f.
E. nubica N.E. Br.
*E. oblanceolala Balf. f.
*E. septemsulcato Vierh.
* E . socotrana Bdf. f.
*E. spiralis Balf. f.
*Jatrophu unicostda Balf. f.
PhyUanthua sp.
Ricinua wmmunk L.
*Tragia balfou&na Gillett nom.nov. (T.
dioica Balf. f., Proc. R.Soo.E d i d . 12.96
(1884), non Sond., LinlMea 23,109 (1860))
Moraceae
*Dorstenia gigas Schweinf. ex Balf. f.
Fkua d i c i j o l i a Vahl
*F.aocotrana Balf. f.
Amaryllidaceae
*Haemanthus grandifoliua Balf. f.
Dioscoreaceae
*DiosCorea; lanuta Bdf. f.
Liliaoeae
*Aloeforbaeii Balf. f.
* A . perryi Bak.
*Asparapa a f k n u a Lam. var. microcarpua
Balf. f.
Asph~deluatenuifoliua Cav.
*fiacuena cinnubari Balf. f.
Juncaceae
Juncua arabicua (Asch. & Buch.) Adameon
Palmaa
Bwaaeus sp.

Phoenix dactylifera L.
Cyperaceae
C2adium ma&cua R. Br.
Cyperus sp.
Fimbri8tyliS sp.
Graminern
Aeluropua sp.
AplwEa mutica L.
Aristida adscensionis L.
A. f u n i c h Trin.& Rupr.
Arthraxon lancifoliw (Trin.)
Hochst.

720

U. B. POPOV:

THE VEUETATION OF SOCOTRA

Gramineae (cont.)
Cenchma ciliaria L.
C . eetiqem Vehl
Chloris barbata Sw.
cymbopgcm sp.
DactyEoctenium ariataturn Link.
* D . hackelii Wagn & Vierh. ex Vierh.
Eleusine wrwm (L.) Gaertn.
Enneapgcm sp.
Eragroath sp.
Heleochloa dura B o k .
Heteropogorr contortue (L.)Beauv. ex Roem.
& Schult.

Hypawhenia hirta (L.) Stapf.


Melunomnchria aby&nku (R.Br. ex Free.)
Hochst.
*Panicumrigidurn Balf. f.
Pmpalidium geminaturn (Forsk.) Stapf.
Pennisetum setaceurn (Forsk.)Chiov.
*Rhyn&dytrmn rnicroatachyum Balf. f.
Sporobolus apic&ua (Vahl)Kunth
The&
quudrivalvia (L. f.) Kuntze
Filiceg
Actinbpteria australis (L. f.) Lk.
*Adianturn Wfmhi Bak.
Pkh8 V'il%Zi% L.

REFERENCES

I. B., 1888. Botany of Socotra. Trans. R. SOC.Edinb. 31, 1-446.


BALFOUR,
ENGLER,
A., 1910. Die PponZmwelt A f r i h , imbesmzdwe seiner tropkchn Gebkte. 1030 pp. Leipzig.
FORBES,
H. 0. et al., 1903. The. N a . t u d iktory of Socotra and Abd-el-Kuri, 598 pp. Liverpooli
PICHI-SERMOLLI,
R. E. G., 1955. Tropical East Africa (Ethiopia, Somaliland, Kenya, Tangan-).
Arid Zone Research. 6 , 302-360. Plant Ecol., UNESCO, Paris.
VIERHAPPER,F., 1907. Kenntnk der Flora Siidarabiens und der Inseln Sokotra, &mh8 und Adb el
Kuri. Denhhr. A M . Wise. Wien, 1, 321-490.
WEITSTEIN,
R.. 1906. Sokotra. In Kersten Schenk, V e g e t d - b d d ~ , 3 ( 5 ) , 25-30.
EXPLANATION O F PLATES 55-59
PLATE

55

Fig. 1. Bare, crescentic dunes overlying stony floor of the southern coastel plain of Naukad. The
absence of vegetation is due to edaphic aridity.
Fig. 2. Sand deposits at Houlaf, northern coast. Aeocia edgezuo7thii forms 8 community here, but is
rere elsewhere.
Fig. 3. Sward of salt-grass,HeZeoch dara, on the beech dun- at Naukad. In the background, the
belt of the sea-lavender,Limonium app. which develops farther inland from the salt-grw.
Fig. 4. Sometimes the vegetation on the southern coastal plain is very sparse. Low crescentic dunes
with a few shrubs of Caotr+ p m a .
PLATE
56
Fig. 5. At the base of the hills the dunes are fixed by a community of woody herbs and ephemerals.
Fig. 6 . Indigofem sp.nov.4ort grass esaociation on the coastal plain a t Hadibo.
Fig. 7. Another woody herb community: the Dactyloctsnium hmMi&Eqlwrbia spiralis association
on the Gobe plain.
Fig. 8. The shrub zone; Croton socotranus--short grass association on Hadibo plain and at Moabbadh.

PLATE
57
Fig. 9. Community of rock plants on the lower s l o p of the Reiged. ( a ) Dendrosicyos sowtram,
( b ) Cisszls subaphyk and (c) Euphorbia ar6uscplla.
Fig. 10. Community of rock plants on the higher s l o p of Reiged. ( a ) Kleinia scottii, ( b ) Ficw
socotrana, (0) Dorstenia gigas, ( d ) BoaweUia sp.
Fig. 11. Rock vegetation on the middle s l o p of the Reiged; the dominantsare succulents and shrubs.
Fig. 12. A grove of &-ma
cinnabari on the s l o p of the Hamadera hill a t Homhil.
PLATE 58
Fig. 13. A typical view of the summit of the limestone plateau, with stunted shrubs (Jatropha unicoetata) and ephemeral%
Fig. 14. Mixed thickets on the lower slopes of the Hagghier.
Fig. 15. Grasslands on the Hsgghier mountains; dominants The&
q u a d r i d & and Hypawhenko
hirta.

Fig. 16. Mixed evergreen thickets in the Hagghier highlands.


PLATE 59
Fig. 17. F k w salkifolko, Cordia sp. and other plauts which together with the date palm form fairly
dense groves along the rivers, which carry water for most of the year.
Fig. 18. Dry watercourse cros~@the Naukad plain. Vegetation is denser, but of the m e composition as on the adjoining plain.

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