Corresponding author: Robert W. Scotland (
Academic editor: Sandra Knapp
A global revision of
Wood JRI, Williams BRM, Mitchell TC, Carine MA, Harris DJ, Scotland RW (2015) A foundation monograph of
The approach adopted in this study arises from a consideration of what taxonomists should be focussed on in the 21st century. Our knowledge of flowering plant diversity comes primarily from taxonomy that has accumulated piecemeal through studies that are geographically restricted. A limitation of this approach to the taxonomy of any widespread or sizeable group is that species usually comprise a combination of restricted endemic species, just over half of all species of flowering plant are single country endemics (
In contrast to the geographical focus of floristic projects, monographic studies at a global level have been undertaken by individual botanists or teams of botanists (
To speed up the taxonomic process while retaining the extensive specimen sampling of a global monographic treatment we believe it is necessary to focus on species delimitation at the expense of other aspects of the traditional monograph. This can be achieved as long as the pragmatic and heuristic nature of taxonomy is appreciated. In other words, for many groups we can rapidly assess specimen level variation, write keys and delimit species without devoting large amounts of time to infra-specific levels of variation and hybridisation and similar issues. Our approach, however, is quite different from many traditional flora projects as we have made full use of electronic tools and technologies, DNA sequencing and basic phylogeny estimates for the group. We use the term “Foundation Monograph” for our approach.
This monograph of
We faced two severe constraints in preparing the
For species delimitation and the preparation of descriptions we examined specimens and made use of the available literature. Dissection of flowers and fruiting capsules was carried out where possible, but care was taken not to damage types or unusual material and capsules are rare or unknown in many species. In species delimitation we gave additional weight to authors who had extensive experience in the genus and knowledge of the species in the field as we were acutely aware that our field knowledge was limited to certain geographical areas. Thus, we have followed the treatment of (
Our desire to produce a concise account of the genus in a limited time has of necessity resulted in the exclusion of certain elements common to many monographs. We have accepted subspecies and the more significant varieties but have accounted for all the recognised infraspecific taxa scattered through the literature down to the level of variety. Forms and subvarieties are not accounted for and are only cited when also treated as the basionym of a taxon of higher rank. For reasons of budget and time we have only seen and cited representative specimens of each species. To cite and map all known collections of each species would be an impossible task within our constraints. A glance at the
The use of the name
Prior to beginning this monograph of
Species of
The European species of
We have treated 190 species of
Molecular data formed an integral component of our approach. Silica gel dried (10%) and herbarium (90%) material were used to sample 130 species of
Molecular data were used in conjunction with morphology to resolve species delimitation in a number of complexes and to inform decisions regarding infrageneric classification and the systematic order used in the taxonomic treatment (
At the species level, molecular data supports the distinction of a number of geographically separate but morphologically similar species pairs such as
The phylogenetic and biogeographic implications of the results are discussed elsewhere (
Summary cladogram showing major groupings resolved in
Within Clade X, there is a strong geographical signal with Australasian (Clade E) and South American (Clade F) clades of
Clade Y is restricted to the Eurosiberian region and three major subgroups are resolved within it. The first (Clade I) comprises species that are often fastigiate in habit and have flowers that are not aggregated into heads. The second (Clade J) comprises species that are typically subshrubs with sericeous leaves, pubescent seeds and often a spiny habit. The third clade (Clade K) comprises species with leaves that are typically tomentose and flowers that are borne in heads.
Whilst some attempt has been made above to correlate morphological traits with the clades resolved in the molecular analysis, it should be noted that the fit is imperfect. We have been unable to identify unique, un-reversed morphological synapomorphies for any of the groups highlighted in Figure
As noted above, there were pragmatic grounds for excluding
The molecular results have been used to establish a framework for the linear sequence of species adopted in the taxonomic account. Efforts have been made to establish an order that maximises morphological similarity between adjacent species within the constraints of relationships inferred using molecular data. Species that are not sampled for molecular data were placed on the basis of their morphology. Within major groups, the molecular sequence has occasionally been abandoned so as to place species which are known to hybridise next to each other or to place species with close morphological similarity together. The sequence used is as follows:
The species concept is problematic in
Notes are provided to indicate where plants intermediate between two species are known to occur. The status of these intermediates is rarely known with certainty. Hybrids are only well documented in a few cases (
An important and perhaps surprising problem in species delimitation is that a considerable number of species are only known from a handful of collections and in several cases from the type only. Moreover, many specimens are inadequate. Only in a few cases do we know what the rootstock is like and in many species the capsule and seeds are unknown. This makes any taxonomy based on capsule and seed characters difficult to formulate and, where it has been attempted, difficult to use or evaluate, as in the case of the Australian species. Good field observations are also lacking in many cases. We do not know the flower colour or the potential height of many species as these details are often not recorded.
Very few new species are described in this monograph. This is mainly a reflection of the fact that
Habits of the four new species described in this treatment.
For the reasons expressed in the previous paragraphs it should be emphasised that the status of a number of species remains uncertain until further collections and more detailed study in the field is possible. We are not completely convinced that
At the infraspecific level, we have utilised the ranks of subspecies and variety:
During our studies we have noted a range of characters of use in species delimitation. Their value is discussed below and we have provided lists of distinctive features, which may be of use in identification. It should be noted that the lists are not always exhaustive as information is not available for all species and not all useful characters are clearly defined.
Habit. Species of
All southern hemisphere species are perennial trailing or twining herbs with cordate, hastate or sagittate, petiolate leaves with the partial exceptions of
Annual herbs:
Erect (or at least ascending) herbs:
Trailing or twining herbaceous perennials, which may be woody below, with hastate, sagittate or truncate leaves:
Lianas:
Unarmed non-twining, non-fastigiate shrubs:
Erect unarmed undershrubs with stiff, thin, woody, subfastigiate branches and (usually) few, relatively small leaves:
Low cushion plants with branched woody rootstock:
Spiny undershrubs can be divided into three subgroups:
Only the old lower branchlets spinescent:
All (or most) branches spinescent:
Branches and peduncles (where present) spinescent, sterile lateral spines (formed from abnormal sterile peduncles?) also present:
Underground parts. These are poorly known.
Leaf and stem indumentum. There is much variation in indumentum, with the majority of species hairy in some form.
Leaves velvety-tomentose:
Leaves densely sericeous/canescent:
Very finely sericeous and often somewhat glabrescent:
Villous-tomentose perennials (dense longish hairs), characteristically woody at the base but with more or less herbaceous branches (always associated with flowers in heads):
Plants glabrous or nearly so (several of these are sometimes partially puberulent):
Sepal indumentum strikingly different from that of stem and leaves:
Leaves and bracts. In this treatment, no great distinction is generally made between bracts and leaves. In the great majority of cases the leaves which subtend flowering peduncles (i.e. bracts) are almost identical to the lower stem leaves and differ only in their progressively smaller size towards the apex of the stem. Only in a few cases, principally those species where the inflorescence appears terminal, have we made a clear distinction between leaves and bracts.
In terms of leaf shape and molecular phylogeny the genus can be divided into two main natural groups, which are mostly easily separated apart from a few Somali or Socotran species. The first group (Species 1–106) has leaves with cordate, truncate, sagittate or hastate leaf bases and clearly demarcated petioles; basal leaf auricles are frequent. Leaves may be entire, undulate, crenate, dentate or sinuate-lobed. Included are a small number of petiolate species with rounded to broadly cuneate leaf bases, which include
Amongst unusual leaf features are
Leaves abruptly contracted to a subsessile base:
Twining/trailing plants with strongly dimorphic leaves include most Australasian species and the following but the list is not exhaustive:
Leaf auricles bifurcate or otherwise divided:
Leaves equilaterally triangular in form:
Inflorescence. Inflorescence structure is quite diverse in the genus and is of taxonomic importance. Flowers are arranged in axillary cymes but this structure is not always obvious. Most commonly, cymes are clearly pedunculate with paired bracteoles at the branching point(s). In this account the peduncle is measured from the base where it arises from the main stem to this branching point. The term pedicel is used above this point. Pedicels are often very variable in length on a single inflorescence especially when the cyme has a clearly monochasial structure. Bracteoles are mostly small, sometimes caducous and relatively unimportant in distinguishing species.
Inflorescence terminal:
Axillary pedicellate flowers with peduncle absent or very short:
Flowers sessile, solitary or paired:
Inflorescence paniculate in form:
Inflorescence racemose in form:
Peduncles paired (at least sometimes):
Sepals. The calyx consists of five separate overlapping sepals, which are commonly somewhat similar in size and shape but often slightly unequal. The two outer sepals are nearly identical; the middle is commonly asymmetric with two halves unequal and the inner pair similar to each other. All sepals may be scarious marginally, but the inner sepals often have wider scarious margins and are less hirsute than the outer pair. The sepal margins are entire or sometimes slightly undulate. In several species, the lower part of the sepal is more or less colourless and contrasts with the distinct green apical portion. In general the size and relative size of the inner and outer sepals, sepal shape, texture and indumentum are all of taxonomic importance. Unusual sepal structures include:
Outer sepals noticeably shorter than inner sepals:
Outer sepals conspicuously larger than inner sepals:
Sepals conspicuously accrescent:
Sepals very lax (not appressed to base of corolla):
Sepals spathulate (and reflexed):
Sepals rectangular:
Sepals with a conspicuously different coloured apex:
Sepals mostly > 10 mm long:
Sepals all very short, < 5 mm long:
Corolla. The corolla of all
Corolla often or always deeply lobed:
Corolla yellow (or cream):
Corolla blue or bluish:
Midpetaline bands glabrous:
Stamens. Stamens are unequal in length. The main taxonomic character of interest is in the filaments. Sessile or very shortly stipitate glands are present on the lower expanded part of the filaments in all annual species, and trailing and twining species from the Mediterranean and Central Asian regions. They are absent from species with cuneate or attenuate leaf bases and apparently from the twining and trailing species from the southern hemisphere.
Style and stigma. The length of the undivided portion of the style is provided in the descriptions but this may well be more variable than the dimensions given as we have not generally examined the style of many examples of each species. More significant is the relative length of the undivided portion of the style to that of the stigmas. In a number of species, the stigmas are almost as long as the undivided style although they are usually much shorter. The stigmas are weakly exserted from the corolla in a number of species (
In most species the stigmas are linear-filiform and co-extensive with the style arm. In a small group of Socotran species previously placed in
Stigmas 3:
Stigmas not coequal with style arm:
Stigmas thickened, oblong to ellipsoid in form, much shorter than style:
Stigmas equalling or longer than style:
Style and ovary indumentum. The indumentum of the ovary, ripening capsule and style is of considerable taxonomic importance. Style indumentum correlates closely with that of the ovary but there are cases where the ovary is hirsute and the style glabrous (
Ovary indumentum has been used extensively in species delimitation by Sa’ad, Rechinger and others. Indeed (
Style sometimes hirsute immediately below stigmas:
Ovary (and usually the capsule) hirsute at least at apex:
Capsule and seeds. The fruit is a capsule and is ovoid, subglobose or somewhat ellipsoid and acuminate in shape. It has the same indumentum as the ovary. The base of the style is persistent in some species. The dehiscence is loculicidal. The capsule is basically bilocular and 4-seeded with trigonous seeds. However, unilocular capsules and single seeds occur quite frequently by abortion and may sometimes be species specific. The shape of the seed is more or less ellipsoidal if only one seed is present. Seeds may be glabrous or variously hairy. The surface may be smooth, reticulate or tuberculate.
Despite the great variety of fruit characters there are severe practical limitations in their use for taxonomic purposes, particularly in the herbarium. Most specimens are collected in flower and capsules and ripe seeds are often missing. Still more serious is the fact that for a large number of species the capsule and seeds are unknown. Given the small number of fruiting specimens available it is often impossible to be certain whether single seeded or 2–4-seeded capsules are species specific or the result of chance abortion. Our observations of seed ornamentation do not always agree with those of other authors and it is not always easy to be sure whether this is the result of natural variation, wrong identification or observation using different strengths of magnification. Consequently, caution should be exercised in relying on distinctions based solely on seed characters. Some generalisations include:
Many fastigiate species are 1-seeded (
All herbaceous petiolate species have glabrous seeds, the seeds commonly being tuberculate or rugose, sometimes more or less reticulate with raised wavy lines as in several African species
Capsules and seeds are usually found in abundance on annual species.
Capsules and seeds are usually present on herbaceous petiolate species.
The following character lists may prove useful although they are not necessarily exhaustive:
Peduncles reflexed in fruit (most common in annual and Australian species):
Seeds hirsute:
Seeds smooth, glabrous:
Keys for the identification of
South America
North America
Australia
New Zealand
Southern Africa (Botswana, Lesotho, Namibia, South Africa, Swaziland)
Tropical Africa (Sahel south to Angola, Mozambique and Zimbabwe, including Socotra and Madagascar)
North Africa (Morocco, Algeria, Tunisia, Libya, Egypt, Mauritania and Niger)
Atlantic Islands (Azores, Canaries, Cape Verde, Madeira)
Europe (Flora Europaea area)
Levant (Turkey, Cyprus, Syria, Lebanon, Palestine/Israel, Jordan)
Arabian Peninsular (including Socotra and Kuwait)
Indo-Iranian Region (Afghanistan, Bhutan, India, Iran, Iraq, Nepal, Pakistan)
Former Soviet Union
East Asia (Burma/Myanmar, China, Japan, Korea, Mongolia, Eastern Siberia).
1 | Erect cerrado perennial with woody xylopodium; leaves sessile or subsessile |
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– | Twining, trailing or prostrate herbs of varied habitats; leaves distinctly petiolate |
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2 | Outer sepals 3–6 mm long |
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– | Outer sepals 6–14 mm long |
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3 | corolla <1 cm long; outer sepals 5–6 mm long |
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– | corolla 2–3 cm long; outer sepals 3–4.5 mm long |
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4 | Leaves deeply palmatisect or pinnatisect, the segments usually very fine |
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– | Leaves entire, weakly lobed, dentate or crenate, the basal auricles entire or bifid, never palmatisect or pinnatisect |
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5 | Outer sepals 10–14 mm long; corolla 2.5–4 cm long |
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– | Outer sepals 7–10 mm long; corolla 1–3 cm long |
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6 | Prostrate plants; leaves < 3 cm long, usually much less; flowers solitary (rarely paired) |
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– | Prostrate or twining plants; leaves mostly >3 cm long; flowers in 1–many-flowered cymes |
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7 | Leaves suborbicular, glabrous or nearly so; corolla with glabrous midpetaline bands |
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– | Leaves ovate-deltoid, pubescent; corolla with hirsute midpetaline bands |
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8 | Leaves entire or very shallowly lobed (Chile) |
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– | Leaves incised-dentate (Peru) |
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9 | Corolla (1.5-)2–3 cm long, pink; leaf auricles usually bifid |
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– | Corolla 1–2.5 cm long but, if more than 1.5 cm corolla cream; leaf auricles entire, rarely bifid and, if so, corolla < 1.5 cm long |
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10 | Leaves linear oblong, glabrous, petiole < 6 mm long, flowers always solitary |
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– | Leaves ovate, deltoid or strap-shaped, pubescent or hirsute, petiole > 5 mm long, flowers 1–many |
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11 | Ovary and capsule apically pilose; plant commonly white-pilose |
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– | Ovary and capsule glabrous; plant variously hairy to subglabrous |
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12 | Corolla 1.6–2.5 cm long, cream; flowering stems slender, c. 1–1.5 mm thick |
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– | Corolla 1–1.8 cm long, white or pink; flowering stems relatively stout, 2–3 mm thick |
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13 | Leaves usually 4–5 times as long as broad, puberulent, the auricles sometimes bifid |
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– | Leaves ovate-deltoid, 2–3 times as long as broad, usually hirsute, the hairs more or less spreading, the auricles never bifid |
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14 | Inflorescence of (1-)3–7-flowered cymes; peduncles 1.5–12 cm long; corolla pinkish |
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– | Flowers solitary or paired; peduncles 1–3(-6) cm long; corolla white |
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1 | Annual. Leaves narrowly oblong-oblanceolate with a long petiole-like base; corolla 5–6 mm long |
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– | Perennial. Leaves various but never narrowly oblong-oblanceolate with a long petiole-like base; corolla more than 10 mm long |
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2 | Corolla pink, 1.8–4.5 cm long, leaves strongly dimorphic, the upper leaves deeply incised (naturalised in California) |
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– | Corolla < 2.5 cm long but, if longer, white; leaves not dimorphic; upper stem leaves not deeply incised |
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3 | Outer sepals 3–4.5 cm long; corolla 3–4 times longer than calyx, usually pink |
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– | Outer sepals 6–11 mm long; corolla mostly about twice as long as calyx but, if much more, white |
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4 | Sepals mostly 9–11 mm long; corolla > 2 cm long |
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– | Sepals < 8 mm long; corolla < 1.8 cm long |
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5 | Leaves and stem white-tomentellous; leaves abaxially with prominent raised veins |
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– | Leaves and stem not white-tomentellous; leaves lacking prominent raised veins |
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6 | Leaves ovate-deltoid, neither lobed nor deeply incised, auricles simple; outer sepals narrowed to base; peduncles with 1–5 flowers |
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– | Leaves deeply lobed or incised and/or auricles deeply bifid; outer sepals often truncate to auriculate at base; peduncles with 1–2 flowers, rarely more |
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7 | Corolla 1.3–1.8 cm long; leaf margin incised-dentate; sepals reddish-brown |
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– | Corolla 1–1.5 cm long; leaf margin usually entire or undulate; sepals not reddish-brown |
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1 | Corolla more than 1.5 cm long |
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– | Corolla less than 1.5 cm long |
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2 | Leaves usually strongly dimorphic and/or with narrowly linear segments; sepals > 4 mm long |
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– | Leaves not or only weakly dimorphic, the segments never linear; sepals < 4.5 cm long |
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3 | Sepals less than 3 mm long |
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– | Sepals more than 4 mm long, often much more |
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4 | Fruiting peduncles straight or sinuate, never recurved |
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– | Fruiting pedicels recurved |
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5 | Leaves with central lobe entire or undulate; basal auricles distinct, not intergrading with sinuate-margined central lobe; corolla 1–1.5 cm long |
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– | Leaves with central lobe dentate, sinuate or dissected, the basal auricles variously lobed or dentate |
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6 | Flowers usually solitary, peduncles solitary; seeds winged; corolla 7–9 mm long |
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– | Flowers usually in small axillary cymes, peduncles sometimes paired in leaf axils; seeds unwinged; corolla 9–15 mm long |
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7 | Corolla < 10 mm long; stems stout, coarsely hairy |
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– | Corolla > 12 mm long; stems relatively slender, softly pubescent or glabrous; corolla 1.2–1.5 mm |
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8 | Leaves sericeous-tomentose with appressed hairs, basal lobes usually not prominent |
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– | Leaves glabrous to roughly pubescent with spreading hairs; basal lobes usually prominent |
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9 | Peduncles very short, < 12 mm long |
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– | Peduncles > 12 mm long |
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10 | Sepals glabrous to sparsely hairy |
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– | Sepals pubescent, often densely so |
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11 | Seeds winged; corolla 7–9 mm long |
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– | Seeds unwinged; corolla 9–12 mm long |
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1 | Leaves dimorphic on the same plant, some being ovate-deltoid in form, others being deeply laciniate with filiform lobes |
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– | Leaves more or less uniform in shape, generally ovate to suborbicular in form |
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2 | Sepals 6–8 mm long; peduncles not reflexed in fruit |
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– | Sepals 4–6 mm long; peduncles reflexed in fruit |
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3 | Corolla > 2 cm long |
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– | Corolla < 2 cm long |
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4 | Corolla relatively small, < 14 mm long; plant subglabrous with only a few hairs, these especially on petioles |
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– | Corolla 15–19 mm long; plant pubescent |
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1 | Flowers sessile; corolla scarcely exceeding calyx; calyx strongly accrescent in fruit |
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– | Flowers pedicellate and/or pedunculate; corolla much exceeding calyx; calyx not markedly accrescent in fruit |
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2 | Calyx < 6 mm long |
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– | Calyx 6–15 mm long |
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3 | Corolla 2–4 times longer than the calyx; leaves usually < 2 cm long |
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– | Corolla only slightly exceeding calyx; leaves usually > 2 cm long |
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4 | Leaves ovate-deltoid, sagittate, never lobed |
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– | Leaves usually lobed or segmented but, if entire, linear-oblong |
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5 | Plant completely glabrous, even on the exterior of the corolla; flowers pedunculate |
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– | Plant pubescent; flowers pedicellate but peduncles absent or very short |
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6 | Peduncles short or absent, flowers solitary |
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– | Peduncles always present, short or long, 1–5-flowered |
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7 | Leaves with 5–9 linear or filiform lobes; corolla lobes obtuse or rounded |
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– | Leaves entire or 3–5-fid, lobes not markedly narrow; corolla lobes acute |
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8 | Leaves always entire, silvery-sericeous, margins not inrolled; sepals acute |
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– | Leaves usually palmately-lobed, rarely entire, brownish-villous, margins inrolled; sepals obtuse |
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9 | Corolla < 15 mm long |
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– | Corolla > 15 mm long |
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10 | Leaves broadly to narrowly triangular to ovate, the base truncate, sagittate or hastate but auricles not lobed |
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– | Leaves palmately-lobed with the central lobe much longer than the auricles, which are usually bilobed |
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11 | Flowers solitary; petioles short; plant decumbent to erect, rarely twining; corolla indistinctly lobed |
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– | Flowers in cymes of 1–6; petioles to 6 cm, plant usually twining; corolla lobed |
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12 | Central lobe of leaf broad, coarsely serrate to pinnatisect; plant roughly hairy |
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– | Central lobe of leaf linear-oblong (rarely broad), entire; plant glabrous to finely pubescent |
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13 | Leaves linear with hastate base |
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– | Leaves various, usually pinnately to palmately lobed or triangular-ovate |
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14 | Sepals obtuse; corolla shallowly lobed |
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– | Sepals acute; corolla deeply lobed |
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15 | Leaves unlobed, entire or crenate |
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– | Leaves palmately or pinnately lobed |
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16 | Corolla < 2 (-2.5)cm long; calyx < 10 mm long |
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– | Corolla 2–3. 5 cm long; calyx > 10 mm long |
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17 | Sepals abruptly narrowed into an apiculate point |
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– | Sepals rounded to acute but not abruptly narrowed |
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18 | Central lobe of leaf sinuate; flowers usually 2 or more |
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– | Leaf entire; flowers usually solitary |
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19 | Leaves pinnately nerved with sinuous-margined central lobe |
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– | Leaves palmately lobed or palmatifid |
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1 | Undershrub with woody spinescent branches |
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– | Plant unarmed although stems sometimes stiff and woody |
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2 | Leaves abruptly narrowed at base; flowers in subsessile clusters |
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– | Leaves cuneate at base, flowers solitary or paired, not pilose |
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3 | Stems and leaves hirsute; flower clusters usually 2–6-flowered; corolla > 12 mm long; bracteoles 3–5 mm wide |
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– | Stems and leaves glabrous to thinly pubescent; flower clusters 1(-2)-flowered; corolla 8–10 mm long; bracteoles 1–2 mm wide |
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4 | Flowers shortly pedicellate, solitary or paired; stigmas linear (Somalia) |
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– | Flowers sessile, solitary; stigmas clavate (Socotra) |
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5 | Ovary glabrous; sepals broadly obovate-elliptic |
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– | Ovary hirsute; sepals oblong-lanceolate |
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6 | Annual herb; plant entirely herbaceous |
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– | Perennial herb or undershrub |
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7 | Lamina ovate, abruptly narrowed into a distinct petiole |
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– | Lamina oblong, lanceolate or oblong (rarely ovate), sessile or narrowed at base with petiole not clearly demarcated |
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8 | Leaves crenate; sepals obtuse; corolla pink |
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– | Leaves entire; sepals acute; corolla blue |
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9 | Flowers 3–6 in sessile axillary clusters |
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– | Flowers up to 3 in pedunculate cymes |
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10 | Corolla large, 2.5–4 cm in length; mountain liana |
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– | Corolla < 2 cm in length; low herb if occurring on mountains |
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11 | Flowers arranged in few- to many-flowered heads, these pedunculate or sessile, the flower bases often concealed by bracts |
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– | Flowers solitary or in lax cymes with pedicels clearly developed, flower bases usually easily visible |
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12 | Leaves linear, < 5 mm long. Undershrub |
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– | Leaves oblong or lanceolate, > 10 cm long. Herbs or undershrubs |
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13 | Bracts and calyx glabrous |
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– | Bracts and calyx variously hirsute |
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14 | Flower heads subsessile; peduncles < 5 mm long |
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– | Flower heads distinctly pedunculate with peduncles > 10 mm long |
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15 | Corolla pale pink or white; woody below |
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– | Corolla blue; herbaceous |
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16 | Leaves abruptly narrowed into a short petiole up to 6 mm long; sepals uniform in colour |
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– | Leaves gradually narrowed to base; sepals bicoloured, pale basally, green at the apex |
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17 | Leaves obovate, pubescent, up to 10 mm wide; sepals 5–7 mm long |
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– | Leaves oblong-oblanceolate, sericeous, < 6mm wide; sepals 8–9 mm long |
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18 | Leaves linear, cuneate at base; bracts linear, appressed to flower head |
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– | Leaves suborbicular, ovate, lanceolate or oblong; bracts neither linear nor appressed to flower head |
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19 | Leaves suborbicular; plant densely covered in brownish sericeous hairs |
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– | Leaves lanceolate, ovate or oblong; plant not sericeous |
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20 | Leaves usually basally cordate; bracts and sepals villous with brownish hairs; flowers usually whitish |
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– | Leaves basally truncate to subhastate; bracts and sepals pubescent; corolla blue |
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21 | Leaves sessile, the base of the lamina attenuate at the base |
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– | Leaves petiolate, lamina hastate or sagittate, well demarcated from the petiole |
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22 | Flowers solitary, sessile; stigmas clavate, shorter than the style arm |
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– | Flowers grouped into cymes or, if solitary, pedunculate; stigmas linear |
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23 | Inflorescence racemose in form with very shortly pedunculate cymes |
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– | Inflorescence paniculate or cymose; flowers with long peduncles |
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24 | Herbaceous plant; leaves with sinuate or undulate margins; basal rosette present |
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– | Herbaceous plant becoming woody with age; leaves entire; basal rosette absent |
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25 | Basal leaves villous, ephemeral; bracts linear |
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– | Basal leaves sericeous, persistent; bracts oblong to oblanceolate |
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26 | Corolla 2–5 times as long as calyx; sepals <4.5 mm long |
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– | Corolla up to twice as long as calyx; sepals usually > 5 mm long |
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27 | Leaves crenate; fruiting peduncle reflexed |
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– | Leaves not crenate; fruiting pedicels not reflexed |
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28 | Erect or twining plant; leaves sericeous with prominent veining on abaxial surface (Zimbabwe) |
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– | Trailing or twining herb, veining on abaxial surface of leaf not prominent |
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29 | Flowers in 1–7-flowered cymes; peduncles 1.5–12 cm long; corolla < 1.2 cm long |
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– | Flowers usually solitary; peduncles 1–3(-6) cm long; corolla > 1.2 cm long |
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30 | Central lobe of leaves coarsely dentate; stems and leaves roughly hirsute |
|
– | Central lobe of leaf entire to undulate; stems and leaves farinose to softly pubescent |
|
31 | Leaves ovate to triangular; auricles not bifurcate |
|
– | Leaves oblong or strap-shaped; auricles commonly bifurcate |
|
32 | Plant densely pubescent to subtomentose; leaf margins shallowly lobed; outer sepals 9–11 mm long |
|
– | Plant thinly to densely pubescent: leaf margin usually entire; sepals 6–8 mm long |
|
1 | Annual herbs; plants slender, herbaceous, never rhizomatous or woody at the base |
|
– | Perennial herbs or undershrubs, usually robust, the base woody or, if herbaceous, rootstock rhizomatous |
|
2 | Flowers densely clustered, peduncles and pedicels, absent or, if present, very short |
|
– | Flowers solitary or in lax cymes |
|
3 | Flower 3–6 in axillary clusters; corolla pale pink (Sahara) |
|
– | Flower clusters terminal (formed from the uppermost leaf axils); corolla blue |
|
4 | Corolla c. 1 cm long; ovary and capsule hirsute; some flowers usually present in uppermost leaf axils (Mediterranean) |
|
– | Corolla 1.5–2.5 cm long; ovary and capsule glabrous; flowers all terminal (Morocco) |
|
5 | Leaves petiolate; leaf blade abruptly narrowed to a truncate or cordate base |
|
– | Leaves clearly sessile or leaf blade gradually narrowed to base |
|
6 | Flowers blue; leaves entire, fruiting peduncle not strongly recurved |
|
– | Flowers pinkish, leaves crenate; peduncle strongly recurved in fruit |
|
7 | Pedicel absent, bracteole adjacent to calyx |
|
– | Pedicel present, bracteole distant from calyx |
|
8 | Corolla 7–10 mm long, entirely blue |
|
– | Corolla 14–40 mm long, usually blue, white and yellow banded |
|
9 | Capsule pubescent; sepals with distinct, different coloured lower and upper portions, pubescent ( |
|
– | Capsule glabrous; sepals without distinct upper and lower areas, glabrous to pubescent |
|
10 | Upper portion of sepals acute to acuminate, longer than basal portion |
|
– | Upper portion of sepals obtuse to acute, shorter than or equalling the basal portion |
|
11 | Plant with spinescent branches, at least below |
|
– | Plant unarmed |
|
12 | Plant subglabrous to densely sericeous (Morocco and Algeria); flowers solitary or in ebracteate clusters |
|
– | Plant densely pilose to tomentose (Egypt); flowers in bracteate heads |
|
13 | Leaves < 15 mm long, abruptly narrowed at base; all branches spinescent |
|
– | Leaves 1–3 cm long, tapered at base; only the old basal, often leafless branches spinescent |
|
14 | Leaves attenuate at base and lacking a distinct petiole; plants never twining |
|
– | Leaves hastate, sagittate or (less commonly oblong), abruptly narrowed into a distinct petiole; plants twining or not |
|
15 | Mature stems woody and divaricately branched |
|
– | Mature stems not woody except below, not divaricately branched |
|
16 | Stems and leaves with spreading hairs |
|
– | Stems and leaves appressed hairy and more or less sericeous with silvery hairs |
|
17 | Sepals with a pale lower portion and green apex; flowers separate or in few-flowered clusters; leaves mostly pilose or pubescent, sometimes subtomentose |
|
– | Sepals uniform in colour; flowers in dense heads; leaves densely tomentose (Sinai) |
|
18 | Ovary and capsule hirsute; corolla c. 2 cm long |
|
– | Ovary and capsule glabrous; corolla <1.5 cm long |
|
19 | Sepals oblong–oblanceolate, acute; inflorescence lax with some flowers separate |
|
– | Sepals lanceolate or ovate, acuminate; flowers clustered into heads |
|
20 | Leaf margin undulate; plants apparently prostrate |
|
– | Leaf margin entire; plants ascending to erect |
|
21 | Cushion plant; flowers solitary or paired, very shortly peduncled |
|
– | Plants not cushion–forming; flowering stems mostly > 5 cm long, flowers in lax terminal groups |
|
22 | Upper surface of leaves glabrous (Morocco) |
|
– | Upper surface of leaves at least thinly pubescent (widespread) |
|
23 | Plant usually < 25 cm high; extreme base of leaf widened and scarious; sepals sericeous and spreading pilose |
|
– | Plant usually 20–50 cm high; extreme base of leaf not widened and scarious; sepals with spreading hairs only |
|
24 | Leaves almost completely entire, occasionally lobed at base |
|
– | Leaves undulate, dentate, sinuate–lobed or incised |
|
25 | Inner sepals longer and more prominent than outer sepals; plant completely glabrous |
|
– | Inner sepals equalling or shorter than the outer sepals; plant hirsute or glabrous |
|
26 | Corolla yellow, 3–4 cm long; outer sepals oblong–obovate without recurved apex (East Mediterranean) |
|
– | Corolla pink < 2.3 cm long, outer sepals spathulate with reflexed apex (Algeria) |
|
27 | Stem base herbaceous; plant usually glabrous to adpressed pubescent |
|
– | Stem base woody, plant pubescent, often densely so |
|
28 | Sepals < 5 mm long |
|
– | Sepals > 6 mm long |
|
29 | Corolla yellow or yellowish; stigma much shorter than style; petioles all very short, more or less 1 mm |
|
– | Corolla blue or white; stigma and style more or less equal or stigma only slightly shorter; petioles > 2 mm |
|
30 | Leaves more than twice as long as broad, usually acute; bracteoles 0.5 mm wide |
|
– | Leaves less than twice as long as broad, rounded; bracteoles 1–3.5 mm wide |
|
31 | Flowers in compact pilose axillary heads; stigma clavate |
|
– | Flowers solitary or in lax cymes or, if clustered, at apex of stem; stigma linear |
|
32 | Sepals < 5 mm long; peduncle deflexed in fruit; slender trailing herb |
|
– | Sepals > 5 mm long; peduncles not deflexed in fruit; plant moderately robust |
|
33 | Flowers clustered at the apex of a peduncle–like stem; stigmas commonly three |
|
– | Flowers not clustered, solitary to several; stigmas 2 |
|
34 | Perennial herbs with herbaceous base; flowers 1–several; peduncle not suppressed |
|
– | Woody based plants from Morroco; flowers solitary, peduncle commonly short |
|
35 | Leaves undulate to sinuate, not dimorphic |
|
– | Leaves (or some of them) deeply incised, commonly dimorphic with basal leaves differing markedly from upper stem leaves |
|
36 | Flowers solitary; leaves truncate (NW Africa) |
|
– | Flowers up to 5; leaves cordate (East Mediterranean) |
|
37 | Corolla yellow |
|
– | Corolla pink or white |
|
38 | Leaves sericeous, the segments of the upper leaves linear |
|
– | Leaves pubescent to pilose, the segments of the upper lanceolate to oblong–elliptic |
|
39 | Corolla purple, the centre white usually with 5 purple spots; peduncles absent; sepals < 5 mm long |
|
– | Corolla pink or white, lacking a purple-spotted centre; peduncles almost always present; sepals > 4.5 mm long |
|
40 | Calyx lanceolate in outline; sepals lanceolate to ovate |
|
– | Calyx oblong in outline; sepals obovate, obtuse to truncate |
|
1 | Leaves sessile; shrubs or woody-based herbs |
|
– | Leaves abruptly narrowed into a distinct petiole; herbs or shrubs |
|
2 | Ovary glabrous; leaves not sericeous; flowers of sessile or pedunculate clasters; plant herbaceous with a woody base (Cape Verde) |
|
– | Ovary usually hirsute; leaves sericeous; flowers varied but not in sessile or pedunculate clusters; undershrubs or shrubs (Canaries) |
|
3 | Prostrate hummock-forming undershrub with spinescent branches |
|
– | Erect shrubs or herbs, the branches not spinescent |
|
4 | Leaves oblong, > 0.5 cm wide; inflorescence terminal, paniculate, many-flowered |
|
– | Leaves filiform to linear < 0.5 cm wide; inflorescence unbranched or sparingly branched, axillary and terminal, few-flowered (< 10) |
|
5 | Annual herbs, neither twining nor trailing; corolla blue |
|
– | Perennial herbs, lianas or undershrubs, twining or trailing the basal parts often woody |
|
6 | Leaves sessile, tapered at base; corolla 3-coloured |
|
– | Leaves abruptly narrowed into a distinct petiole; corolla entirely blue ( |
|
7 | Pedicels absent, bracteole appressed to base of calyx, filiform to more or less lanceolate |
|
– | Pedicels borne at least 5 mm below calyx, always filiform |
|
8 | Calyx < 5 mm long, plant entirely herbaceous |
|
– | Calyx > 5 mm long, base of plant woody, rarely entirely herbaceous |
|
9 | Corolla 10–15 mm long; plant entirely herbaceous (Azores) |
|
– | Corolla > 15 mm long; plant woody at base (Madeira, Canaries) |
|
10 | Leaves dimorphic, upper leaves deeply segmented |
|
– | All leaves similar, upper leaves not deeply segmented |
|
11 | Leaves strongly hirsute |
|
– | Leaves glabrous or nearly so |
|
12 | Leaves oblong-ovate, densely villous beneath |
|
– | Leaves elliptic, lanceolate, ovate or oblong, finely pubescent to tomentellous beneath |
|
13 | Leaves ovate, outer sepals 10–13 mm long |
|
– | Leaves linear-lanceolate to oblong, sepals 6–9 mm long |
|
14 | Cymes long-pedunculate; leaves large 4–11 cm long (Madeira) |
|
– | Cymes borne on peduncles < 2 cm long; leaves < 6 cm long |
|
15 | Leaves < 4 × 1.5 cm wide; cymes 1–2-flowered (Gran Canaria) |
|
– | Leaves mostly > 4 × 1.5 cm; cymes 1–6-flowered |
|
16 | Sepals 9–10 mm long; corolla white with pink midpetaline bands (Lanzarote) |
|
– | Sepals 5 mm long; corolla bluish (Tenerife, La Gomera) |
|
1 | Leaves abruptly narrowed at the base into a distinct petiole |
|
– | Leaves gradually narrowed at the base, lacking a distinct petiole |
|
2 | Leaves cuneate or truncate at the base |
|
– | Leaves sagittate or hastate at the base |
|
3 | Robust plant, densely lanate; sepals obtuse |
|
– | Slender plant, pubescent; sepals acute to acuminate |
|
4 | Annual herb, corolla 7–12 mm long ( |
|
– | Perennial herb, corolla 1.5–2 cm long |
|
5 | Pedicels absent, bracteole appressed to base of calyx, filiform to more or less lanceolate |
|
– | Pedicels borne at least 5 mm below calyx, always filiform |
|
6 | Pedicels 0–3 mm long; leaves ovate to suborbicular |
|
– | Pedicels 3–12 mm; leaves lanceolate to oblong, often falcate |
|
7 | Calyx with mostly short appressed hairs |
|
– | Calyx with spreading hairs only |
|
8 | Leaves strongly dimorphic, the upper leaves deeply divided ( |
|
– | Leaves not strongly dimorphic; the upper leaves not deeply divided |
|
9 | Leaves sericeous beneath; leaf segments narrow, linear to oblong |
|
– | Leaves with spreading, often slightly asperous hairs; leaf segments mostly broad |
|
10 | Sepals < 5 mm long |
|
– | Sepals > 5 mm long |
|
11 | Flowers very small; capsule pubescent, borne on recurved peduncles |
|
– | Flowers usually 15–25 mm long; capsule glabrous, not recurved |
|
12 | Corolla < 17 mm long. Portugal |
|
– | Corolla 25–45 mm long. Eastern Europe |
|
13 | Liana; leaves oblong and elliptic, retuse, glabrescent; corolla 15–17 mm long |
|
– | Twining or decumbent perennial herb with herbaceous stems; leaves usually triangular, acute, farinose or pubescent; corolla 10–15 mm long |
|
14 | Plant glabrous; flowers yellow; sepals emarginate and apiculate |
|
– | Plant pubescent; flowers usually pink; sepals acute or acuminate |
|
15 | Annual herbs; plants entirely herbaceous |
|
– | Perennial plants, at least the basal portions and rootstock woody |
|
16 | Flowers sessile or nearly so, peduncle and pedicel shorter than calyx |
|
– | Flowers distinctly pedunculate, the peduncle and pedicel several times longer than the calyx |
|
17 | Corolla 7–10 mm long, entirely blue |
|
– | Corolla 14–40 mm long, usually blue, white and yellow banded |
|
18 | Capsule pubescent; sepals with distinct, different coloured lower and upper portions, pubescent ( |
|
– | Capsule glabrous; sepals without distinct upper and lower areas, glabrous to pubescent |
|
19 | Upper portion of sepals acute to acuminate, longer than basal portion |
|
– | Upper prtion of sepals obtuse to acute, shorter than or equalling the basal portion |
|
20 | Cushion plants with prostrate stems with or without short flowering stems |
|
– | Plants not cushion forming; flowering stems at least 5 cm high |
|
21 | Flowering stems absent or extremely short |
|
– | Short but distinct flowering stems present |
|
22 | Leaves glabrous above, midrib only distinct |
|
– | Leaves sericeous above, lateral veins distinct ( |
|
23 | Indumentum of sepals more or less spreading and distinct from that of the leaves (Spain) |
|
– | Indumentum of leaves and sepals similar (Balkans) |
|
24 | Lateral veins distinct (Bulgaria) |
|
– | Lateral veins not distinct (widespread). |
|
25 | Mature stems woody and divaricately branched |
|
– | Mature stems not woody, or only so below, not divaricately branched |
|
26 | Plants silvery-sericeous, the hairs appressed |
|
– | Plants densely pubescent to pilose, some hairs conspicuously spreading |
|
27 | Outer sepals more or less cordate at the base, conspicuously gibbous |
|
– | Outer sepals neither cordate basally nor gibbous |
|
28 | Flowers in dense heads overtopped by bracts, which form a kind of involucre |
|
– | Flowers in lax terminal groups, the pedicels usually obvious |
|
29 | Ovary and capsule glabrous (Spain and France) |
|
– | Ovary and capsule hirsute |
|
30 | Stem leaves distant, few; sepals all acuminate (Caucasus) |
|
– | Stem leaves imbricate, numerous, some sepals obtuse (Adriatic region) |
|
31 | Cliff plant with long pendent sems |
|
– | Plant of open slopes, decumbent to erect |
|
32 | Plant usually < 25 cm high; extreme base of leaf widened and scarious |
|
– | Plant usually 20–50 cm high; extreme base of leaf not widened and scarious |
|
33 | Inflorescence with flowers clustered at apex of stem |
|
– | Inflorescence lax |
|
34 | Sepals densely pilose |
|
– | Sepals with scattered spreading hairs |
|
1 | Annual herbs; plants slender, herbaceous, never rhizomatous or woody at the base |
|
– | Perennial herbs or undershrubs, usually robust, the base woody or, if herbaceous, rootstock rhizomatous |
|
2 | Flowers densely clustered at the apex, peduncles and pedicels, absent or, if present, very short |
|
– | Flowers solitary or in lax cymes |
|
3 | Sepals terminating in a prominent mucro 1.5–3 mm long |
|
– | Sepals acute to obtuse, lacking a distinct terminal mucro |
|
4 | Leaves petiolate; leaf blade abruptly narrowed to a truncate or cordate base |
|
– | Leaves clearly sessile or leaf blade gradually narrowed to base |
|
5 | Flowers blue; leaves entire, fruiting peduncle not strongly recurved |
|
– | Flowers pinkish, leaves crenate; peduncle strongly recurved in fruit |
|
6 | Corolla 7–10 mm long, entirely blue |
|
– | Corolla 14–40 mm long, usually blue, white and yellow banded |
|
7 | Leaves distinctly petiolate, base of lamina hastate, sagittate, cordate, rounded or very broadly cuneate |
|
– | Leaves sessile or base of lamina tapering at base |
|
8 | Inner sepals conspicuously longer than outer sepals; plant glabrous (including midpetaline bands); corolla yellow |
|
– | Inner sepals equalling or shorter than outer sepals; plant hirsute at least on the midpetaline bands; corolla pink or white (yellowish only in |
|
9 | Rigidly erect, divaricately branched plant |
|
– | Trailing plant, stems not divaricately branched |
|
10 | Flowers in axillary, pedunculate, pilose heads |
|
– | Flowers solitary or in lax cymes, never arranged in dense pilose heads |
|
11 | Leaf base broadly cuneate to rounded; undershrub with tomentose leaves and solitary white flowers |
|
– | Leaf base hastate, sagittate or truncate; trailing or twining herbs, only slightly woody at base; flowers or not, commonly pinkish |
|
12 | Sepals < 6 mm long; leaves never deeply incised or lobed |
|
– | Sepals > 6 mm long; upper leaves often incised or dentate |
|
13 | Leaf margin entire; fruiting peduncles not recurved |
|
– | Leaf margin strongly crenate; fruiting pedicels recurved |
|
14 | Sepals < 4.5 mm long, scarious-margined |
|
– | Sepals 5.5–6 mm long, margins not scarious |
|
15 | Sepals 10–15 mm long, leaf margin entire to obscurely undulate |
|
– | Sepals 6–10 mm long, leaf margin crenate dentate to incised |
|
16 | Leaves dimorphic, at least the upper ones incised; ovary glabrous; corolla pink or white |
|
– | Leaves not dimorphic nor upper leaves incised except sometimes in |
|
17 | Leaves sericeous, the segments of the upper leaves linear |
|
– | Leaves pubescent to pilose, the segments of the upper lanceolate to oblong-ellipti |
|
18 | Leaves glabrous with a ciliate margin |
|
– | Leaves densely pubescent to tomentose |
|
19 | Corolla yellow; leaves commonly dimorphic |
|
– | Corolla pink or white, leaves never dimorphic |
|
20 | Ovary glabrous; sepals obovate to broadly elliptic, leaves sinuate, coarsely pubescent |
|
– | Ovary hirsute (? rarely glabrous); sepals ovate to elliptic; leaves undulate to crenate, softly tomentose |
|
21 | Leaves with spreading hairs; sepals acute; corolla white to pale pink |
|
– | Leaves uniformly short-tomentose; sepals apiculate; corolla pink |
|
22 | Flowers several in axillary heads; leaves, stem and sepals densely villous |
|
– | Flowers in terminal heads (sometimes a few axillary also) or not in head-like structures; stem and leaves glabrous, pubescent, pilose or sericeous |
|
23 | Bracts ovate, cordate, up to 3 cm wide; leaves reticulate below |
|
– | Bracts oblong-elliptic or lanceolate, up to 1.5 cm wide; leaves not reticulate below |
|
24 | Flower heads sessile or subsessile |
|
– | Flower heads pedunculate |
|
25 | Lower, old stems spinescent, stems ascending, < 40 cm long |
|
– | Plant unarmed, stems procumbent, > 40 cm long |
|
26 | Bracts linear to lanceolate, < 0.5 cm wide |
|
– | Bracts oblong-elliptic or lanceolate 0.5–1.5 cm wide |
|
27 | Plants with branched stems forming a lax open inflorescence |
|
– | Plants with compact terminal inflorescences or cushion plants, never forming a much branched open inflorescence |
|
28 | Corolla white, c. 1 cm long; stem glabrous |
|
– | Corolla pink or with pink midpetaline bands, > 1 cm long, stem appressed hairy to pilose |
|
29 | Ovary glabrous |
|
– | Ovary hirsute |
|
30 | Corolla 1–1.5 cm long; stems flexible, herbaceous, pilose |
|
– | Corolla 2–2.5 cm long; stems stiff and woody, appressed pubescent |
|
31 | Sepals at apex abruptly narrowed and mucronate |
|
– | Sepals gradually narrowed to an acute or acuminate apex |
|
32 | Stems and leaves densely spreading pilose; leaves all oblong (Gaziantep region) |
|
– | Stems and leaves pubescent or thinly pilose; leaves variously shaped, often oblanceolate-spathulate near the base of the stem, rarely oblong |
|
33 | Leaves and stem adpressed-sericeous |
|
– | Leaves and stem with spreading hairs (often also sericeous) or more or less glabrous |
|
34 | Sepals with a conspicuous pouch near base |
|
– | Sepals lacking a conspicuous pouch near base |
|
35 | Sepals < 10 × 8 mm |
|
– | Sepals > 11 × 11 mm |
|
36 | Dwarf cushion-forming shrublet |
|
– | Low perennial with woody base and distinct ascending or erect stems |
|
37 | Sepals with conspicuous spreading hairs; leaves with forked central vein |
|
– | Sepals with appressed hairs; leaves with one simple central vein |
|
38 | Plant usually < 25 cm high; extreme base of leaf widened and scarious; sepals sericeous and spreading pilose |
|
– | Plant usually 20–50 cm high; extreme base of leaf not widened and scarious; sepals with spreading hairs only |
|
39 | Flowers clustered, usually at apex of peduncle-like stem; plants cushion-forming or not |
|
– | Flowers solitary or clearly separate in a lax terminal cyme; plants strictly cushion forming |
|
40 | Outer sepals bicoloured with a pale base and green apex |
|
– | Outer sepals uniformly coloured |
|
41 | Corolla < 1.5 cm long |
|
– | Corolla > 1.5 cm long |
|
42 | Stems, leaves, sepals and ovary glabrous or nearly so |
|
– | Stems, leaves, sepals and ovary conspicuously pilose |
|
43 | Plant < 15 cm high; sepals with long caudate apex |
|
– | Plant usually > 25 cm high; sepals acute to acuminate |
|
44 | Plant with conspicuous spreading hairs; corolla pink |
|
– | Plant subglabrous or thinly appressed pubescent; corolla white or pale pink |
|
1 | Plants annual; all parts of the plant herbaceous |
|
– | Plants perennial; plants usually woody below, but, if entirely herbaceous, with perennial rhizomatous roots |
|
2 | Leaves crenate; sepals obtuse; corolla pink |
|
– | Leaves entire; sepals acute; corolla blue |
|
3 | Pedicels absent, bracteole appressed to base of calyx, filiform to more or less lanceolate |
|
– | Pedicels borne at least 5 mm below calyx, always filiform |
|
4 | Trailing or twining herbs with leaves abruptly narrowed at the base into a distinct petiole; plants not with woody stems nor flowers arranged in head-like clusters |
|
– | Herbs or shrubs, never twining or trailing, leaves gradually narrowed at the base, lacking a distinct petiole but if petiolate, stems woody or flowers in head-like clusters |
|
5 | Corolla > 15 mm long; flowers usually solitary |
|
– | Corolla < 12 mm long; flowers 1–5 in axillary cymes |
|
6 | Sepals < 4.5 mm long; plant usually glabrescent |
|
– | Sepals > 6 mm long; plant pubescent |
|
7 | Leaves crenate; fruiting peduncles deflexed |
|
– | Leaves not crenate, entire apart from (sometimes) forked auricles or weakly sinuate margins; fruiting peduncles not deflexed |
|
8 | Central leaf lobe ovate to triangular in outline; basal auricles not forked; plant often twining |
|
– | Central leaf lobe linear-oblong in outline; basal auricles sometimes forked; plant usually trailing |
|
9 | Leaves with lamina abruptly narrowed at base and clearly separate from the (sometimes very short) petiole; stigmas clavate or at least thickened upwards |
|
– | Leaves sessile or with lamina attenuate at base with no distinct petiole; stigmas various |
|
10 | Flowers in hirsute heads, the hairs spreading and somewhat concealing the calyx |
|
– | Flowers solitary; sepals glabrous or sericeous, easily visible (Oman) |
|
11 | Flower heads pedunculate |
|
– | Flower heads sessile or nearly so |
|
12 | Leaves glabrous; stems woody, sometimes spinescent |
|
– | Leaves pubescent; stems herbaceous (except below), never spinescent |
|
13 | Spiny undershrub; leaves all alternate; flower clusters of up to 6 flowers |
|
– | Unarmed undershrub; leaves often opposite towards branch tips; flowers usually 1–2 together |
|
14 | Branches spinescent |
|
– | Branches not spiny although sometimes woody and rigid |
|
15 | Flowers solitary or clustered, sessile; sterile spinescent peduncles absent |
|
– | Flowers solitary or clustered borne on spinescent peduncles; sterile peduncles often present as spines (Oman) |
|
16 | Sepals addressed pubescent; stigma clavate, shorter than style arm (Socotra) |
|
– | Sepals with spreading hairs; stigmas linear, co-extensive with style arm |
|
17 | Ovary glabrous; sepals broadly obovate-elliptic |
|
– | Ovary hirsute; sepals oblong-lanceolate |
|
18 | Sepals long-pilose with woolly hairs (Oman) |
|
– | Sepals densely pubescent to tomentose but lacking long woolly hairs |
|
19 | Plant with long, slender, spine-tipped branches, short spinescent side shoots absent or very few |
|
– | Plant with stout spinescent primary branches and numerous short (< 4 cm long), usually stout lateral spine-like shoots |
|
20 | Leaves with rigid, acute apex, basal leaves not undulate; flowers usually in clusters of > 1, clusters elongating at maturity |
|
– | Leaves with soft obtuse to subacute apex, the basal leaves often undulate; flowers mostly solitary |
|
21 | Flowers arranged in sessile or pedunculate, pilose clusters |
|
– | Flowers arranged in a lax open inflorescence or sessile or shortly pedunculate along an elongate axis, not in pilose cluster |
|
22 | Nearly leafless subshrub with glabrous to adpressed pubescent stem and leaves; leaves minute, linear, < 5 mm long (Hadramaut) |
|
– | Leafy plants at least basally; leaves and stem pubescent, pilose or villous; leaves > 2 cm long |
|
23 | Sepals bicoloured, base colourless, apex greenish; ovary glabrous |
|
– | Sepals uniformly coloured green; ovary hirsute |
|
24 | Flowers usually more or less solitary, sometimes clustered; sepals oblong with an acute apex |
|
– | Flowers in heads, very rarely solitary; sepals gradually narrowed to an acute to long acuminate apex |
|
25 | Heads subsessile; dwarf mountain plant with stems < 10 cm high (Asir) |
|
– | Heads pedunculate; desert plant with stems usually >15 cm |
|
26 | Stems and leaves with long villous hairs; style pilose |
|
– | Stems and leaves shortly hairy; style glabrous or nearly so |
|
27 | Sepals bicoloured; base colourless, apex greenish |
|
– | Sepals uniformly green |
|
28 | Leaves sinuate margined; basal rosette persistent; plant entirely herbaceous (Abd ul Kuri Island) |
|
– | Leaf margins entire; basal rosette absent or ephemeral; plant usually woody at least below |
|
29 | Ovary hirsute; corolla deeply lobed; undershrub to 3 m with very rigid branches and peduncles arising at 90° to each other (Saudi Arabia) |
|
– | Ovary glabrous; corolla shallowly lobed; herbs or undershrubs to 50 cm, branching not as above |
|
30 | Flowers sessile or nearly so, forming a long narrow inflorescence |
|
– | Flowers borne on conspicuous, often rigid peduncles; inflorescence open |
|
31 | Flowers solitary (Socotra) |
|
– | Flowers in very shortly pedunculate cymes (Yemen) |
|
32 | Peduncles bearing monochasial cymes, inflorescence pubescent (Oman) |
|
– | Peduncles mostly bearing single flowers; inflorescence almost glabrous except sepals |
|
33 | Basal leaves villous, ephemeral; bracts linear |
|
– | Basal leaves sericeous, persistent; bracts oblong or oblanceolate |
|
1 | Plants annual; all parts of the plant herbaceous |
|
– | Plants perennial; plants usually woody below, but, if entirely herbaceous, with perennial rhizomatous roots |
|
2 | Lamina ovate, abruptly narrowed into a distinct petiole |
|
– | Leaves oblong, lanceolate or oblong (rarely ovate), sessile or narrowed at base with petiole not clearly demarcated |
|
3 | Leaves crenate; sepals obtuse; corolla pink |
|
– | Leaves entire; sepals acute; corolla blue |
|
4 | Flowers solitary, corolla blue |
|
– | Flowers clustered or grouped, very rarely solitary; corolla pinkish |
|
5 | Flowers 3–6 in sessile axillary clusters |
|
– | Flowers up to 3 in pedunculate cymes | |
6 | Sepals glabrous |
|
– | Sepals adpressed-pilose |
|
7 | Trailing or twining herbs with leaves abruptly narrowed at the base into a distinct petiole; plants not with woody stems nor flowers arranged in head-like clusters |
|
– | Herbs or shrubs, never twining or trailing, leaves gradually narrowed at the base, lacking a distinct petiole but if petiolate, stems woody or flowers in head-like clusters |
|
8 | Sepals < 5 mm long |
|
– | Sepals > 6 mm long |
|
9 | Leaves crenate; fruiting peduncles deflexed |
|
– | Leaves entire; fruiting peduncles not deflexed |
|
10 | Plant completely glabrous; corolla yellow |
|
– | Plant pubescent at least on the midpetaline bands and usually elsewhere; corolla variously coloured |
|
11 | Sepals rectangular in form (Afghanistan) |
|
– | Sepals variously shaped, never rectangular in form (India, Iran, Iraq) |
|
12 | Leaves entire, densely hirsute |
|
– | Leaves sinuate-margined, sparsely pubescent |
|
13 | Ovary and capsule hirsute; sepals bicoloured with distinct apical portion; leaves entire to undulate |
|
– | Ovary and capsule glabrous; sepals lacking a distinctly coloured apical portion; leaves undulate to sinuate or dentate |
|
14 | Corolla < 1.2 cm long, white or cream (India) |
|
– | Corolla 2.5–3.5 cm long, pink or purplish |
|
15 | Leaves distinctly petiolate, the lamina clearly separate from the petiole |
|
– | Leaves sessile or with lamina attenuate or cuneate at base with no distinct petiole |
|
16 | Flowers in pedunculate hirsute heads |
|
– | Flowers solitary |
|
17 | Leaves glabrous; stems woody, sometimes spinescent |
|
– | Leaves pubescent; stems herbaceous (except below), never spinescent |
|
18 | Spiny undershrub; leaves glabrous to finely sericeous, < 1 cm wide |
|
– | Unarmed undershrub; leaves tomentose, 1–3.5 cm wide |
|
19 | Plant spiny or with spinescent branches |
|
– | Plant unarmed, although branches sometimes rigid and hard |
|
20 | Flowers in a terminal cluster towards the apex of the stem |
|
– | Flowers mostly or entirely axillary |
|
21 | Ovary and style glabrous; primary branches only spinescent |
|
– | Ovary and style hirsute; lateral shoots spinescent as well as primary branches |
|
22 | Flowers borne on spinescent peduncles; sterile spines often also present |
|
– | Flowers sessile or nearly so; sterile spines usually absent |
|
23 | Outer sepals much shorter than the inner sepals |
|
– | Outer and inner sepals similar in length |
|
24 | Sepals 3–10 mm long |
|
– | Sepals 10–11 mm long |
|
25 | Sepals long-pilose |
|
– | Sepals adpressed pubescent |
|
26 | Sepals 4–5 mm long, thinly pilose; corolla 1.5–1.7 cm long |
|
– | Sepals 7–10 mm long, densely pilose; corolla 1.6–2.5 cm long |
|
27 | Sepals 12–15 mm long |
|
– | Sepals < 8 mm long |
|
28 | Plant with long, slender, spine-tipped branches, short spinescent side shoots absent or very few |
|
– | Plant with stout spinescent primary branches and numerous short (< 4 cm long), usually stout lateral spine-like shoots |
|
29 | Leaves with rigid, acute apex, basal leaves not undulate; flowers usually in clusters of > 1, clusters elongating at maturity |
|
– | Leaves with soft obtuse to subacute apex, the basal leaves often undulate; flowers mostly solitary |
|
30 | Flowers arranged in dense terminal or axillary heads or clusters |
|
– | Flowers variously arranged in a lax, branched inflorescence |
|
31 | Heads terminal (occasionally with a few flowers below the terminal head) |
|
– | Heads axillary, sometimes terminal as well |
|
32 | Plant with woody stems |
|
– | Plant with herbaceous stems, woody only at the base |
|
33 | Leaves with impressed veins, hairs dense but short; sepals < 10 mm long; Corolla < 1.2 cm long |
|
– | Leaves without impressed veins; sepals > 10 mm long; corolla > 1.5 cm long |
|
34 | Ovary and style glabrous; stigmas c. 6 mm long |
|
– | Ovary and style hairy; stigmas c. 3 mm long |
|
35 | Plant silvery-sericeous; inflorescence very lax with individual peduncles and pedicels clearly visible |
|
– | Plant not sericeous or, if somewhat so, inflorescence of dense heads with individual peduncles and pedicels not easily visible |
|
36 | Outer sepals with a conspicuous pouch; plant to 30 cm |
|
– | Outer sepals lacking a conspicuous pouch; plant usually < 10cm |
|
37 | Leaves linear, < 0.2 cm wide |
|
– | Leaves oblong-elliptic or oblanceolate, > 5 cm wide |
|
38 | Stem with spreading hairs |
|
– | Stem with appressed hairs |
|
39 | Heads solitary, strictly terminal |
|
– | Heads with 1–2 flower groups below terminal heads |
|
40 | Lower peduncles absent or < 0.5 cm long; heads sessile or nearly so |
|
– | Lower peduncles well-developed, > 1 cm long; heads mostly distinctly pedunculate |
|
41 | Cushion herbs from which arise erect flowering stems, ovary comose |
|
– | Plants not cushion forming, ovary glabrous or hirsute |
|
42 | Sepals bicoloured, the base pale, apex green; ovary glabrous |
|
– | Sepals of uniform colour; ovary hirsute at apex |
|
43 | Branches rigid and woody; leaves apiculate; corolla < 1.5 cm long; corolla < 1.5 cm long; flowers usually 1–3 |
|
– | Branches not noticeably rigid; leaves acute but not apiculate; corolla > 1.5 cm long; heads many-flowered |
|
44 | Ovary hirsute, at least at apex |
|
– | Ovary glabrous |
|
45 | Sepals 14–16 mm, ovate with a long aristate point, almost half its length |
|
– | Sepals 10–12 mm, lanceolate to ovate, acuminate but not long-aristate |
|
46 | Stems and leaves with long villous hairs; style pilose |
|
– | Stems and leaves shortly hairy; style glabrous or nearly so |
|
47 | Plant densely brown-velvety-tomentose; leaves reticulate |
|
– | Leaf indumentum not as above; leaves not reticulate |
|
48 | Stem stout, 4–5 mm wide; bracteoles elliptic, 4–5 mm wide, sepals obovate |
|
– | Stem relatively slender, < 3 mm wide; bracteoles lanceolate, 2–3 mm wide; sepals lanceolate |
|
49 | Sepals bicoloured; base colourless, apex greenish |
|
– | Sepals uniformly coloured green |
|
50 | Corolla 1.7–2.5 cm long, ovary and capsule hirsute |
|
– | Corolla 1–1.5 cm long; ovary and capsule glabrous |
|
51 | Flowers usually more or less solitary, sometimes laxly clustered, sepals oblong with an acute apex |
|
– | Flowers always in dense heads, sepals tapered to an acute to long acuminate apex |
|
52 | Only lower heads pedunculate; heads on upper part of stem sessile |
|
– | All heads distinctly pedunculate except perhaps the uppermost |
|
53 | Bracts < 3 × 1 cm, lanceolate (Iraq–Iran) |
|
– | Bracts mostly 3–4 × 1.2–2 cm, ovate (Afghanistan) |
|
54 | Sepals bicoloured, pale below with a green apex; plants with herbaceous stems, flowers somewhat clustered |
|
– | Sepals of one colour; plants commonly with woody rigid stems, few leaves and flowers well separated |
|
55 | Corolla < 1.5 cm long; ovary and capsule glabrous; sepals oblong-obovate |
|
– | Corolla 1.7–2.5 cm long; ovary and capsule hirsute; sepals ovate to lanceolate, acuminate |
|
56 | Sepals glabrous or nearly so |
|
– | Sepals pubescent, canescent or otherwise hirsute |
|
57 | Stems completely glabrous |
|
– | Stems adpressed pubescent |
|
58 | Corolla 1.7–2 cm long, pink |
|
– | Corolla < 1. 5 cm long, white or very pale pink |
|
59 | Sepals obovate, mucronate, c. 5 mm long |
|
– | Sepals oblong, acuminate, c. 7 mm long |
|
60 | Leaves filiform |
|
– | Leaves linear, oblong or oblanceolate |
|
61 | Sepals ovate |
|
– | Sepals obovate-elliptic |
|
62 | Sepals 6–7 × 3–4 mm; inflorescence narrow, few-flowered |
|
– | Sepals 4–5 × 2 mm; inflorescence commonly much branched and many flowered |
|
63 | Stems appressed hairy, finely sericeous to strigose |
|
– | Stems with spreading hairs at least below |
|
64 | Sepals tiny, suborbicular, c. 2 mm; plant divaricately branched |
|
– | Sepals > 3 mm long, longer than broad; plant not divaricately branched |
|
65 | Sepals obtuse to rounded; corolla deeply lobed; inflorescence much branched forming an intricate mass |
|
– | Sepals acute, acuminate or obtuse and mucronate, always terminating in a point. Corolla at most shallowly lobed; branching not so extensive as to form an intricate mass |
|
66 | Corolla 0.8–1 cm long; ovary pubescent |
|
– | Corolla > 1.2 cm long; ovary glabrous |
|
67 | Corolla pink |
|
– | Corolla white |
|
68 | Sepals lanceolate, elliptic or oblong, acuminate; stems not leafy, very rigid |
|
– | Sepals obovate, abruptly narrowed to a muconate apex; stems leafy and somewhat herbaceous |
|
69 | Stem and leaves white-sericeous |
|
– | Stem and leaves not white-sericeous |
|
70 | Branches slender, not very rigid; leaves lanceolate to ovate |
|
– | Branches short, stiff, relatively stout; leaves linear-oblong |
|
1 | Plant an annual herb; flowers solitary, blue |
|
– | Plant perennial, herbaceous or woody; flowers pink, white or yellow |
|
2 | Trailing or twining herbs with leaves abruptly narrowed at the base into a distinct petiole; plants not with woody stems nor flowers arranged in head-like clusters |
|
– | Herbs or shrubs, never twining or trailing, leaves gradually narrowed at the base, lacking a distinct petiole but if petiolate, stems woody |
|
3 | Sepals < 5 mm long |
|
– | Sepals > 5 mm long |
|
4 | Corolla < 2.8 cm long, pink, flowers usually solitary |
|
– | Corolla >2.8 cm long, white, yellowish or pink, flowers usually more than one |
|
5 | Leaves with an elongated strap-shaped central lobe |
|
– | Leaves triangular in form |
|
6 | Plant completely glabrous; inner sepals longer than outer sepals |
|
– | Plant hirsute; inner sepals equalling or shorter than outer sepals |
|
7 | Undershrubs with petiolate leaves, the lamina abruptly narrowed at base |
|
– | Herbs or undershrubs with sessile leaves or leaves gradually narrowed into an indistinct petiole |
|
8 | Spiny undershrub; leaves glabrous to finely sericeous, < 1 cm wide |
|
– | Unarmed undershrub; leaves tomentose, 1–3.5 cm wide |
|
9 | Plant spiny or with spinescent branches |
|
– | Plant unarmed, although branches sometimes rigid and hard |
|
10 | Flowers in a terminal inflorescence |
|
– | Flowers axillary |
|
11 | Branches all spinescent; lower leaves oblanceolate-obovate; flowers 1–several in a terminal cluster |
|
– | Only the old lower branches spinescent; lower leaves linear to narrowly oblanceolate; flowers in a terminal cyme |
|
12 | Flowers borne on spinescent peduncles; sterile spines often also present |
|
– | Flowers sessile or nearly so; sterile spines absent |
|
13 | Outer sepals glabrous, much larger than the inner sepals |
|
– | Outer sepals pubescent, equalling or smaller than the inner sepals |
|
14 | Flowers arranged in terminal heads or clusters, occasionally with a few flowers on the stem below the main cluster; stems herbaceous (if woody, see 134. |
|
– | Flowers variously arranged in lax, branched inflorescences, stems often woody |
|
15 | Leaves, stem and sepals all silvery-sericeous |
|
– | Leaves, stem or sepals with conspicuous spreading hairs, sometimes sericeous as well |
|
16 | Inflorescence a compact head, pedicels and peduncles not clearly visible |
|
– | Inflorescence lax, peduncles and pedicels easily visible |
|
17 | Outer sepals conspicuously pouched (Crimea); plant to 30 cm |
|
– | Outer sepals lacking a conspicuous pouch; plant rarely exceeding 15 cm |
|
18 | Stem with spreading hairs |
|
– | Stem with appressed hairs |
|
19 | Sepals with spreading hairs |
|
– | Sepals with appressed hairs |
|
20 | Low perennial with linear, sericeous leaves; flowers solitary |
|
– | Erect or ascending plants with stems usually > 10 cm tall; flowers mostly clustered |
|
21 | Sepals bicoloured, with pale base and green apex; plants with herbaceous stems and flowers somewhat clustered |
|
– | Sepals of one colour; plants commonly with woody rigid stems, few leaves and flowers well separated |
|
22 | Corolla < 1.5 cm long; ovary and capsule glabrous; sepals oblong-obovate |
|
– | Corolla 1.7–2.5 cm long; ovary and capsule hirsute; sepals ovate to lanceolate, acuminate |
|
23 | Sepals glabrous or nearly so |
|
– | Sepals pubescent, canescent or otherwise hirsute |
|
24 | Corolla pink; sepals 5–7 mm long; plant divaricately branched |
|
– | Corolla white; sepals 4–5 mm; plant not divaricately branched. |
|
25 | Sepals obovate, mucronate, c. 5 mm long |
|
– | Sepals oblong, acuminate, c. 7 mm long |
|
26 | Stems appressed pubescent, finely sericeous to strigose |
|
– | Stems densely sericeous, pubescent or pilose, some hairs spreading at least below |
|
27 | Sepals obtuse to rounded; corolla deeply lobed; inflorescence much branched forming an intricate mass |
|
– | Sepals acute, acuminate or obtuse and mucronate, always terminating in a point; corolla at most shallowly lobed; branching not so extensive as to form an intricate mass |
|
28 | Corolla white, 0.8–1 cm long; ovary pubescent |
|
– | Corolla pink, > 1. 2 cm long; ovary glabrous |
|
29 | Stem and leaves white-sericeous |
|
– | Stem and leaves not white-sericeous |
|
30 | Inflorescence of very dense, axillary clusters; lower leaves clearly oblanceolate; lower branches often somewhat spinescent |
|
– | Inflorescence scape-like, flowers1-several at apex of stem; leaves strictly linear, branches never spinescent |
|
31 | Branches slender, not very rigid |
|
– | Branches short, stiff, relatively stout |
|
32 | Leaves linear-lanceolate, up to 3 mm wide; sepals lanceolate, acuminate; stems subsericeous |
|
– | Leaves lanceolate to ovate, 3–15 mm wide; sepals often abruptly narrowed at apex; stems pubescent |
|
33 | Stems sparingly branched; corolla > 1.5 cm long, pink; ovary glabrous |
|
– | Stems much branched; corolla < 1.5 cm long, white or pinkish; ovary usually hirsute |
|
34 | Plant densely pubescent; leaves linear-oblanceolate |
|
– | Plant thinly pubescent; leaves linear-oblong |
|
1 | Leaves distinctly petiolate, the blade abruptly narrowed onto to the petiole; trailing or twining herbs |
|
– | Leaves lacking a distinct petiole, the blade narrowed at base; undershrubs or perennial herbs with a woody rootstock, neither twining nor trailing |
|
2 | Sepals < 4.5 mm long |
|
– | Sepals 5–7 mm long |
|
3 | Leaves glabrous or nearly so, margin entire |
|
– | Leaves densely pubescent to tomentose, margin undulate, crenate or dentate |
|
4 | Leaf base hastate, often with bifid auricles; sepals 7–10 mm long |
|
– | Leaf base truncate to subcordate with simple, poorly developed auricles; sepals 6–7 mm long |
|
5 | Unarmed perennials woody at base only |
|
– | Undershrubs with spinescent branchlets |
|
6 | Stems erect, branched; leaves glabrous or adpressed hairy beneath; ovary and capsule glabrous |
|
– | Stems prostrate to ascending but always low; leaves sericeous; ovary and capsule pubescent |
|
7 | Leaves oblong-oblanceolate, 5–25 mm wide; flowers 1–5 in compact cymes, 1.8–2.5 cm long |
|
– | Leaves linear to linear oblanceolate, < 5 mm wide; flowers usually solitary, 1–1.6 cm long |
|
8 | Flowers mostly axillary; outer sepals 4.5–6 mm long; stems herbaceous |
|
– | Flowers all terminal on the branches; outer sepals 6–7 mm long; stems somewhat woody and rigid |
|
9 | Sepals glabrous to thinly pubescent. Outer pair suborbicular, much wider than inner sepals |
|
– | Sepals hirsute, all similar in shape and size |
|
10 | Flowers clustered at apex of peduncle-like stem |
|
– | Flowers axillary |
|
11 | Prostrate cushion plant, the flowering branches without spines |
|
– | Erect undershrub, flowers borne on spinescent peduncles, usually (always?) with sterile stem spines towards the apex of the flowering shoots |
|
Accepted names are in bold italics. All names of specific, subspecific and varietal rank in the genus
Type specimens and their location are cited for all recognised taxa whether species, subspecies or varieties. We have lectotypfied species where we have seen appropriate material for lectotypification but have not lectotypified where there is doubt about the selection of a lectotype. A particular problem relates to the plants described from North Africa by Maire. The types of these species were supposed to have been deposited at the Université d’Alger (AL) and are cited for AL by
Wherever possible, at least one specimen is cited for every country where a species is known to occur. Occasionally, a literature record is cited and in a few cases of common species, no specimen is cited as the species is assumed to be present because of its wider distribution. Records requiring confirmation are indicated with a question mark (?). Although cited specimens are limited to those seen by the authors and are representative of the species, some effort has been made to select material that is either widely distributed or likely to be available in the country where the plant occurs. Unfortunately this has not always been possible. The herbaria where these specimens are found are not cited as we are not generally aware of where they are distributed. We have seen all collections in BM, E, K, LE, OXF, P and W and sporadic examples from other herbaria if material has been loaned or images were available online.
Where a species is known from a few countries the country order in which specimens are cited is arbitrary but in cases where a species is known from many different countries the preferred order is as follows: European and Mediterranean countries are arranged from West to East beginning with the Atlantic Islands but southern African countries are arranged from South Africa northwards.
We have cited references to where all type specimens were published. We have not cited references to pages in standard floras unless they add to the information in the present monograph by providing additional descriptive material, illustrations or maps. However we have cited recent works where illustrations, paintings, drawings or photographs are provided as these are often a very useful aid to identification, capturing the appearance of a particular species in a way that words do not. We have cited references to relevant literature in the discussion of infraspecific variation and taxonomic problems.
Spiny or unarmed shrubs or subshrubs or prostrate or erect herbs, stems often twining or trailing. Leaves alternate (rarely subopposite), simple, sessile or petiolate. Flowers variously arranged, solitary or in various kinds of inflorescence, usually cymose in structure although reduced to heads, flower pairs or other arrangements; each flower subtended by a pair of small bracteoles; calyx of 5 free sepals, these usually entire, slightly to very unequal, usually of two similar outer sepals, two similar inner sepals and an asymmetric middle sepal whose two halves are dissimilar; corolla funnel-shaped with a spreading limb and a short glabrous basal tube, the limb with five hirsute external midpetaline bands which terminate in a tooth or lobe; stamens 5, included, inserted at the top of the basal tube, filaments unequal, the basal part slightly dilated, glabrous or minutely glandular, the glands sessile or shortly stipitate, anthers equal, oblong to oblong-sagittate, pollen tricolpate, more or less spherical, colpi long and broad, exine thick; ovary usually ovoid, less commonly globose or conical, hirsute or glabrous, the base with a distinct disc, bilocular, each locule with 2 ovules; styles glabrous or hirsute, filiform divided upwards into 2 (rarely 3) arms, stigmas coextensive with style arms (very rarely slightly shorter), linear or, rarely, thickened upwards and ellipsoid or clavate. Capsule bilocular or by abortion unilocular, the dehiscence loculicidal or from the base, 4-seeded or less by abortion; seeds hirsute or glabrous, smooth tuberculate or obscurely ridged, one side convex and the other flat (see Figure
Nearly all species are trailing or twining perennial herbs with flowers in pedunculate cymes (sometimes reduced to single flowers) arising from the axils of leaf-like bracts.
Plate “
Glabrous perennial herb with trailing or twining stems up to 2 m long. Leaves petiolate, 2.5–7 × 1.5–5 cm, deltoid, acute to acuminate, margin entire, base cordate but not cuneate onto the petiole, auriculate with auricles weakly 2 (– 3)-lobed, with one lobe larger than the other; petioles 0.5–4.5 cm long. Flowers 1–5 in pedunculate axillary cymes; peduncles 3.5–16 cm long; bracteoles 3–5 × 0.5–1 mm, linear to linear-lanceolate, acute; pedicels 8–11 mm, so inflorescence rather dense; outer sepals 6–7 × 5 mm, broadly oblong-obovate to rectangular, truncate and minutely mucronate, glabrous, scarious; inner sepals 7–11 × 4.5–6 mm; corolla 3–4 cm, pale yellow, undulate, midpetaline bands glabrous except for a few hairs near the apex; filaments with sessile glands below; ovary glabrous; style glabrous, divided 11–15 mm above the base, stigmas 3 mm. Capsule glabrous; seeds smooth (especially in Iraq) to tuberculate. [
East Mediterranean region to Crimea and Iraq: Greece (Aegean Islands only): Rhodes (
A very distinct, nearly completely glabrous species with a yellow corolla, acutely-angled deltoid leaves and the outer sepals much smaller than the inner sepals. Molecular studies (
TURKEY, Gaue Sber,
Perennial herb with tap root and stems somewhat woody below, similar in all details to
Northeast Turkey (
Molecular studies (
ALGERIA,
Glabrous, trailing perennial herb, the stems angular, reaching at least 75 cm. Leaves petiolate, 1.2–3 × 1–2.2 cm. ovate, apex acute to more or less rounded and mucronate, margin entire, base truncate (below) or cordate (above), the auricles small, triangular-acute, venation reticulate; petioles 1–2.5 cm. Flowers axillary, pedunculate, solitary; peduncles 3–9 cm, commonly flexuose in bud; bracteoles 3–9 × 0.25–0.5 mm, linear-oblanceolate; pedicels 0.5–3.5 cm; outer sepals 4–5 × 2.5–3 mm, spathulate, the apex abruptly widened above the oblong base, rounded, sparsely ciliate, commonly reflexed; inner sepals 5–6 × 2.5–3 mm, similar in shape but apex emarginate and not reflexed; corolla 1.7–2.3 cm long, pink, weakly lobed, midpetaline bands glabrous; filaments glandular below; ovary conical, glabrous; style glabrous, divided 5–6 mm above the base, stigmas 5–6 mm. Capsule glabrous, style persistent; seeds slightly rugose. [
Restricted to the Magreb of northwestern Africa: Algeria (
A very distinctive species because of its reflexed spathulate sepals and unusual ovate, truncate, reticulate-veined leaves.
“Europe” (lectotype LINN 218.1!. designated by
Perennial herb from an extensive creeping underground rootstock, branched at base with trailing or twining quadrangular stems to about 75 cm long, plant glabrous to sparsely hairy. Leaves petiolate, 1–7 × 0.5–4 cm, broadly to narrowly ovate-deltoid, obtuse or acute, mucronulate, margin entire or undulate, base hastate to sagittate with simple auricles; petioles 1–2.5 cm. Flowers 1–3 in axillary pedunculate cymes; peduncles 1–5 cm; bracteoles 2.5–3 mm, filiform; pedicels 0.6–20 mm; sepals 3.5–4.5 × 2.5–3.5 mm, obovate to oblong, obtuse to mucronulate, scarious-margined; corolla 1.5–2.5 cm long, white or pink, undulate but not lobed, midpetaline bands often dark pink, pubescent; filaments glandular below; ovary glabrous, style glabrous, divided 7–8 mm above base, stigmas 2.5 mm. Capsule glabrous; seeds tuberculate. [
A very common cosmopolitan weed of all temperate regions which also grows in upland regions throughout the tropics.
A very variable species especially in indumentum, leaf shape and flower colour, of which many forms and varieties have been described (
CHINA, cultivated plant grown from seed collected by Staunton at “Pechelee” (holotype BM001053866!).
Perennial herb with long decumbent stems from a central rootstock to at least 50 cm, glabrous or, on older parts, minutely scabridulous. Leaves petiolate, 3–5 cm long, formed of an oblong, acute, entire central lobe 2–4 mm wide, a broadly cuneate base and horizontally to weakly reflexed auricles, these mostly bifid with acute segments; petioles 4–7 mm. Flowers axillary, pedunculate, solitary; peduncles 3.2–4.5 cm, slighty flexuous; bracteoles 3 mm, linear-filiform; pedicels 4–8 mm; sepals 6–7 × 3.5–4 mm, obovate, obtuse and sometimes mucronate, glabrous, margins scarious, inner sepals slightly larger; corolla 2–2.8 cm long, pink, very shallowly lobed, the midpetaline bands extended as short teeth, nearly glabrous but with a few hairs near apex; filaments glandular below; ovary glabrous; style glabrous, divided 12–14 mm above base, stigmas 2.5–3.5 mm. Capsule glabrous, seeds glabrous, minutely tuberculate.
We recognise two subspecies:
Distinguished by the decumbent habit and distinctive strap-shaped leaves, the central lobe elongated.
Very common in northern China, Mongolia and Siberia becoming rare in Kazakhstan, where it is largely replaced by
A subsp. typo habitu suberecta et foliis triangularibus.
KAZAKHSTAN, “in rupestribus montium Tarbagatai ad torrentium Dschanybek”,
Distinguished by its suberect habit and triangular leaves, c. 3–5 × 1.5–4 cm.
Russia: Siberia (
GREECE, Parnassus, Lake Zouvala,
Trailing perennial herb with very slender stems 10–30 cm long, vegetative parts densely pubescent. Leaves petiolate, 0.5–1.3 × 0.3–1 cm, suborbicular to ovate with obtuse apex to deltoid with acute apex, margin undulate, base cordate to hastate; petioles 2–9 mm. Flowers solitary, pedunculate, axillary; peduncles 2–11 mm, strongly recurved in fruit; bracteoles 1–1.5 mm, linear; pedicels 2.5–6 mm; outer sepals 2–3 × 1.5–2 mm, oblong-elliptic, somewhat truncate at both ends, pubescent, margins scarious. Corolla 0.8–1 cm long, pink, unlobed, midpetaline bands pubescent; filaments glandular; ovary pilose; style glabrous, persistent, divided 3–3.5 mm above base, stigmas c. 1.5 mm. Capsule borne on a recurved peduncle, pilose; seeds glabrous, obscurely rugose. [
Greece (
A distinctive species, superficially resembling a diminutive
SAUDI ARABIA, Wadi Fatma,
Perennial (possibly sometimes annual) herb with trailing stems to at least 50 cm from a slender central tap root; stems glabrescent to pubescent. Leaves petiolate, 1.2–4.5 × 0.6–4 cm, ovate-deltoid, apex obtuse, margin sinuate, base auriculate and cordate; petioles 0.5–3.5 cm. Flowers 1(-3) borne on axillary peduncles; peduncles 7–30 mm, commonly recurved in fruit; bracteoles 2 mm, filiform; pedicels 3–5 mm; outer sepals 3–5 × 3–4 mm, obovate, rounded, glabrous, slightly concave; inner sepals slightly narrower, 2.5–3 mm wide; corolla 0.9–1.3 cm long, pink, distinctly lobed, midpetaline bands brownish, thinly pubescent; filaments glandular below; ovary glabrous; style glabrous, divided c. 2 mm above base, stigmas 1 mm. Capsule glabrous, strongly exserted from the sepals, recurved in fruit; seeds glabrous, smooth (not rugulose as stated by Sa’ad,1967: 226). [
A widespread Sahara-Sindian species, generally uncommon and very scattered in occurrence but most frequent in Egypt; usually a weed of sandy fields. “Mauretania” (
Very distinct species with sinuate leaves and pink, lobed corolla borne on a recurved peduncle. The leaves are sometimes exceptionally small.
CHINA, Yunnan, Dali,
Pubescent perennial herb with (probably) decumbent stems from a thickened rootstock, young growth brownish-tomentose; stems to 60 cm, probably reaching 1 m, Leaves shortly petiolate, 1.1–3.5 cm long, the central lobe 0.2–1 cm wide, linear or oblong, acute, margin entire, undulate, sinuate or more or less dentate, base hastate, the auricles simple or bifid, sometimes intergrading with sinuate leaf margin. Flowers 1–2, axillary, pedunculate; peduncles 1.5–4 cm; bracteoles 3–4 × 0.5 mm, linear or filiform; pedicels 6–15 mm long, straight to slightly bent; outer sepals 7–10 × 4–5 mm, ovate, acuminate, villous with ciliate margins; inner sepals similar but much less hairy; corolla 1.2–1.4 cm long, pink or white, unlobed, midpetaline bands pilose, extended as short teeth; ovary and style glabrous. Capsule glabrous; seeds nearly smooth, glabrous.
Endemic to SW China: Yunnan (
Apparently rare and localised and no recent collections seen.
MYANMAR/BURMA, Shan plateau,
Coarsely pilose perennial herb with decumbent stems from a thickened taproot; stems to 20 cm but probably much more. Leaves petiolate, 1–1.5 × 0.2–0.5 cm, ovate-deltoid, acute, margin sinuate-dentate, base truncate to subcordate, coarsely pilose; petioles 4–6 mm. Flowers axillary, solitary, pedunculate; peduncles 1–1.5 cm; bracteoles 1–2 mm, filiform; pedicels 4–7 mm; outer sepals 6–7 × 2–3 mm, oblong-oblanceolate, acute, pilose on dorsal surface; inner sepals similar but 5 × 3 mm, obovate, scarious-margined; corolla c. 1.3 cm long, colour unknown, apparently weakly lobed, midpetaline bands pilose; ovary glabrous; style glabrous, divided c. 2.5 mm above base; stigmas 2 mm. Capsule not known.
Myanmar (Burma). Only known from the type collection found at c. 1700 m.
This poorly known species might prove to be a variant of
INDIA, Tamil Nadu/Kerala, Nilgiri Hills,
Perennial scrambling and climbing herb to at least 50 cm, stems pubescent, the hairs reddish on young parts. Leaves petiolate, 2–8 × 2–6 cm, lanceolate to broadly ovate-deltoid, acute and mucronulate, margin variable, undulate to deeply dentate, base broadly cordate in outline but cuneate onto the petiole, auricles entire to deeply dentate, pubescent on both surfaces, especially on the veins beneath; petioles 1.5–3 cm. Flowers 1–2 (-3) in pedunculate, axillary cymes; peduncles often paired, 6–8 mm; bracteoles c. 1.25 mm, caducous, ovate, acuminate; pedicels 8–10 mm, more densely pubescent than peduncles; sepals 6–7 × 3–4 mm, outer sepals obovate-elliptic, abruptly narrowed at apex, apiculate, pubescent, inner sepals similar, obovate, mucronate, scarius-margined, subglabrous; corolla 10–12 mm, whire or cream, deeply lobed, mid-petaline bands terminating in a tuft of hairs; filaments glabrous; ovary glabrous, style glabrous, divided c. 5 mm above base, stigmas 1.5–2 mm, linear. Capsule glabrous, seeds glabrous.
Endemic to South India: Nilgiri and Palni (Pulney) Hills (
Like the two preceeding species, this is a geographically isolated species. Although quite variable, the leaves are often strongly dentate and the auricles lobed. The corolla is similar to that of the South American species
Cultivated plant grown in Chelsea Physic Garden from seed received from Paris (holotype BM001035798!).
Very variable trailing or twining perennial herb up to 1 m high, stems angled, vegetative parts always hirsute, thinly to densely pubescent, pilose or tomentose. Leaves petiolate, 2.5–8 × 2–6 cm, ovate, apex obtuse or acute, often mucronate, margin entire to undulate, base cordate and cuneate onto the petiole, usually auriculate, auricles rounded to acute, entire or dentate; petioles 0.5–1.5 (-6) cm. Flowers 1–3 (-8) in pedunculate, axillary cymes (often clearly dichasial); peduncles 2–14 cm, very variable from specimen to specimen; bracteoles filiform to linear or linear-oblanceolate, acute, 6–14 × 0.5–1.5 mm, pedicels 5 –15 mm; outer sepals 7–15 × 3–5 mm, oblong-elliptic, acute or acuminate, bicoloured, sometimes slightly constricted below triangular, slightly deflexed dark green apical portion, inner sepals scarious-margined 8–10 × 5–6 mm, shorter but broader; corolla 2.8–3.6 cm, white, cream, or pink, unlobed, midpetaline bands pilose, sometimes darker coloured; filaments glandular below; ovary pilose, style pilose, divided c. 9 mm above base, stigmas 3 mm. Capsule pilose; seeds papillose. [
Widely distributed from the eastern Mediterranean region east to the Caucasus and Iran: Greece (
A very variable species in indumentum, leaf shape, peduncle length, number and colour of flowers and size and shape of sepals Attempts have been made by
TURKEY, Merzivan,
Presumably trailing herb of unknown length; stems and leaves pubescent. Leaves similar to those of
Turkey. Only known from the type collection.
This species is not conspecific with
SYRIA,
Twining perennial herb, stems angled, glabrous. Leaves petiolate, 3–4 × 2–2.5 cm, ovate-deltoid, obtuse, margin undulate to crenate or weakly lobed, ciliate, base cordate and attenuate onto the petiole, beneath thinly pubescent. Flowers 1–3 in pedunculate axillary cymes; peduncles 4–14 cm, glabrous; bracteoles linear, acute, 6–8 × 1 mm, ciliate; pedicels 0.8–1 cm, thinly pilose with stiff spreading hairs; outer sepals 9–10 × 5–6 mm, oblong-obovate, slightly pandurate, abruptly constricted at apex into a mucro, the apical portion dark-coloured, pilose with stiff brown hairs; inner sepals glabrous, membranous; corolla 3.2 cm, yellow, unlobed, midpetaline bands thinly pilose towards the apex; filaments glandular below; ovary pilose; style thinly pilose, divided 5 mm above the base; stigmas 2 mm. Capsule and seeds not seen. [
A rare and very local species of the Syrian border with Turkey, known from a handful of collections: Turkey (?); Syria (“Latakia” fide
Resembling
SYRIA/IRAQ, Aleppo to Mosul,
Perennial herb with decumbent stems up to 1 m long from a central rootstock, vegetative parts pubescent with crisped, somewhat retrorse hairs, occasionally villous to subtomentose. Leaves petiolate, 1.5–6 × 1.5–5.5 cm, ovate-reniform, apex obtuse, margin undulate, crenate-dentate to sinuate-dentate, base cordate and cuneate onto the petiole; petioles 1–4.5 cm. Flowers 1–5 in pedunculate axillary cymes; peduncles 3–9 cm; bracteoles 3–8 mm, filiform; pedicels mostly 1–1.5 cm but sometimes longer resulting in a very lax inflorescence; outer sepals 6–8 × 4–5 mm, obovate or broadly oblong, obtuse, retuse or truncate and mucronate, scarious, pubescent, inner sepals membranous with a truncate base, glabrous or nearly so; corolla (1.5-)2.5–3.5 cm long, pink to purplish, unlobed, midpetaline bands thinly pilose; filaments glandular below; ovary glabrous or with a few apical hairs, style glabrous or sparsely pilose, divided 5 mm above base, stigmas 4 mm. Capsule glabrous; seeds glabrous, strongly tuberculate. [
We recognise two varieties which can distinguished by indumentum and floral characters:
Indumentum of leaves and stem puberulent to pubescent; corolla 2.5–3.5 cm long.
Eastern Mediterranean region east to Iran, growing as a weed, often in fallow fields: Turkey (
EGYPT,
Distinguished by its denser villous to tomentose indumentum combined with a smaller corolla about 1.5 cm long.
Scattered over the range of the species. Examples seen include
PALESTINE/ISRAEL,
Perennial herb with trailing or twining stems from a woody base 0.4–1 m long; stem and vegetative parts adpressed tomentellous. Leaves petiolate, somewhat dimorphic; lower leaves 3–3.5 × 2–3 cm, broadly to narrowly ovate, acute, margin crenate, base broadly cordate and cuneate onto the petiole; middle and upper leaves with an acute triangular central lobe 3–5 × 0.4–0.6 cm, the margin entire to sinuate, basal auricles deeply lobed with many acute lobes; petioles 0.3–2.3 cm, diminishing in length upwards. Flowers 1–3 in pedunculate axillary cymes; peduncles 1.5–5.5 cm; bracteoles 2–4 mm long, filiform; pedicels 2–8 mm, frequently recurved; outer sepals 8–10 × 4–5 mm, obovate, obtuse, densely pubescent; inner sepals c. 1 mm shorter, obovate-elliptic, rounded and crenate at apex, scarious; corolla 2.2–3 cm long, yellow, unlobed, midpetaline bands shortly pubescent near apex; filaments glandular below; ovary pubescent; style glabrous, divided 8 mm above base, stigmas 2–3 mm. Capsule apically pubescent; seeds verruculose. [
Lebanon south to Sinai: Palestine/Israel (
Resembles
TURKEY, Ankara,
Perennial herb with decumbent or prostrate stems spreading from a central tap root and reaching 50 cm, vegetative parts softly tomentose. Leaves petiolate, 1.5–4 × 1–3 cm, ovate to ovate-triangular, apex acute to mucronate, margin undulate, sinuate or, above, weakly 5-lobed, base cordate and shortly attenuate onto the petiole, veins prominent below; petioles 0.5–1.4 cm. Flowers 1–2 in pedunculate, axillary cymes; peduncles 0.8–2.5 cm; bracteoles 2 –4 mm, linear to filiform; pedicels 0.5–1.4 cm; outer sepals 7–10 × 5–8 mm, broadly ovate, rounded and mucronate to acute, somewhat convex, tomentose, greyish, inner sepals c. 7 × 5 mm. glabrous, membranous; corolla 2.6–3 cm long, deep pink, unlobed, midpetaline bands adpressed pilose, terminating in a tooth; filaments glandular below; ovary pilose (or fide
Almost restricted to Turkey: Turkey (
The small ovate-triangular leaves and the softly tomentose indumentum help to identify this species. It might be confused with some forms of
TURKEY, Marash,
Similar in overall morphology to
Endemic to Turkey. Previously known only from the type collection but rediscovered in 2008 (
LEBANON, between Hasbey and Rasheiya,
Annual herb, mostly branched at base, with decumbent or ascending stems to c. 30 cm, vegetative parts thinly pubescent. Leaves petiolate, 2–4(-5) × 1.5–3 cm, ovate or reniform, apex rounded, margin entire or undulate, base weakly auriculate, cordate and cuneate onto the petiole; petioles up to 10 cm on basal leaves but mostly 2–3 cm on cauline leaves. Flowers solitary, axillary, pedunculate, becoming congested upwards; bracts resembling small leaves, but sometimes deeply palmately lobed with acute lobes; peduncles 0.5–5 cm, elongating and reflexing in fruit; bracteoles 3–4 mm, filiform to linear-lanceolate; pedicels 0.3–1.5 cm; sepals 3–6 × 3–4 mm, broadly oblong-obovate, prominently mucronate, stiffly hirsute with spreading hairs; corolla 1.5–2(-2.8) cm, pink or pinkish purple, unlobed, midpetaline bands pilose; filaments glandular below; ovary glabrous; style glabrous, divided c. 4 mm above base, stigmas c. 1.5 mm. [
Eastern Mediterranean, apparently especially common in Cyprus: Cyprus (
The retuse, strongly apiculate sepals, reflexed fruiting peduncles and annual habit are distinctive.
MOROCCO, Oued Cherrat,
Perennial herb with stout somewhat woody rootstock from which arise various short decumbent, subglabrous, pubescent to pilose stems to 50 cm. Leaves petiolate, 0. 8–3.2 × 7.5–3.5 cm, ovate-deltoid or reniform, rounded to obtuse, margin undulate to coarsely serrate, base cordate, usually shortly and softly tomentose-sericeous but sometimes with longer hairs, occasionally subglabrous; petioles 0.5–0 7 cm, often flexuose. Flowers solitary, borne on axillary peduncles; peduncles 0–1 cm; bracteoles 2.5–7 mm, filiform; pedicels 1–8.5 cm, commonly flexuose and somewhat deflexed in fruit; calyx in flower clearly longer than broad, sepals 4.5–9 × 3–6 cm, lanceolate to linear-oblong, acute to apiculate forming a narrow calyx, the inner sepals broader; corolla 2.2–3(-4) cm long, pink with a darker centre, midpetaline bands sericeous near apex; filaments glandular below; ovary glabrous; style glabrous, divided c. 7 mm above base; stigmas 3–4 mm. Capsule glabrous; seeds finely tuberculate. [
Distinguished from
Leaves glabrous above
Morocco (Only known from the type).
Based on
Leaves sericeous. The long flexuose pedicels are also very distinct. Much more common than the type variety.
Endemic to Morocco where it usually grows on schists: Central Rif (
MOROCCO, Demnate,
Perennial herb with relatively slender rootstock (c. 3 mm wide) from which emerge various short decumbent or ascending stems 5–15 (-20) cm long, vegetative parts pubescent. Leaves petiolate, 2–4 × 1–3 cm, dimorphic, lower leaves ovate-deltoid, obtuse, margin undulate to dentate, base truncate to shallowly cordate and shortly cuneate on the petiole; upper leaves somewhat smaller, strongly dentate, apex acute; petioles 1–5 cm. Flowers solitary on axillary peduncles; peduncles 0.3–2.5 cm long, very variable in length; bracteoles 4–6 mm, filiform; pedicels 1–2.5 cm, commonly flexuose; calyx in flower about as long as broad, outer sepals 4.5–7 × 3.5–6 mm, oblong-obovate, mucronate, adpressed pubescent; inner sepals c. 7 × 5 mm, broadly obovate, mucronate, soon scarious; corolla 2.6–3.3 cm, white or pink, unlobed, midpetaline band terminating in a point, nearly glabrous (slightly scabrous); filaments glandular below; ovary glabrous; style glabrous, divided 9 mm above base; stigmas 4 mm. Capsule glabrous; seeds finely tuberculate. [
Endemic to Morocco: High Atlas (
Similar to
It appears that the sheet with the original collection in Maire’s herbarium was cut in two and part deposited at Montpelier. This part is selected as the lectotype. The other part of the sheet may be at AL. We have not been able to trace type material of
MOROCCO,
Perennial herb from a stout tap root with decumbent stems to 30 cm, vegetative parts pilose. Leaves petiolate, 0.7–2.8 × 0. 5–3.3 cm, dimorphic, lower leaves ovate-deltoid, obtuse, margin crenate, base cordate to truncate, upper leaves deltoid, apex acute, margin incised-lobed, base cordate; petioles 1–3 cm, flexuose. Flowers solitary, borne on axillary pedicels; peduncles absent; bracteoles 2–4 mm, filiform; pedicels 3–35 mm, becoming strongly recurved in fruit; sepals 2.5–5.5 × 2.5–5.5, ovate to obovate, acute or obtuse and apiculate; corolla 1.7–3 cm long, purple with cream centre and (usually) dark purple marks around throat, midpetaline bands pilose towards apex; filaments glandular below; ovary glabrous; style glabrous, divided 5–9 mm above base; stigmas c. 4 mm. Capsule glabrous; seeds finely tuberculate.
Endemic to the Western Rif in Morocco (
Distinguished from the
Southern Europe, (lectotype LINN 218.26!, designated by
Trailing or twining perennial herb with slender creeping rootstock; stems terete, to 2 m long; vegetative parts thinly pilose to densely sericeous-tomentose. Leaves petiolate, strongly dimorphic; lower leaves 2–4 × 1–3 cm, ovate-deltoid, apex apiculate, acute or obtuse, margin irregularly crenate, base cordate and shortly cuneate; upper leaves slightly larger, to 6 × 6 cm, similar in outline but deeply sinuate-lobed to 3–5-partite with narrowly oblong, entire to coarsely dentate segments; petioles 1–5 cm long. Flowers axillary, pedunculate, solitary or in a dichasial cyme with up to 4 flowers; peduncles 3–10 cm; bracteoles 3–12 mm, filiform, linear to narrowly linear-lanceolate; pedicels 5–13 mm; outer sepals 5–9 × 4–6 mm, variable in shape, elliptic to obovate, acute to obtuse, glabrous to hirsute, margin often scarious, undulate, inner sepals slightly broader with broad scarious margins, often basally auriculate; corolla (1.6-)1.8–4.5 cm long (very variable in size), pink (rarely white), very weakly lobed, midpetaline bands darker, shortly pubescent; filaments glandular below; ovary glabrous; style glabrous, divided 5–10 mm above base; stigmas 2. 5–5 mm, relatively stout. Capsule glabrous; seeds glabrous, obscurely tuberculate. [
A very variable species in indumentum, sepal form and flower size, this reflected in the extensive synonymy below. Two subspecies, sometimes treated as separate species, can usually be distinguished although intermediates occur occasionally, for example
Variably hirsute but hairs not appressed and sericeous. Leaves variously dissected but lobes not linear. [
Madeira (
Based
Plant softly sericeous in all parts. Leaves finely dissected with narrow linear lobes. [
Mostly East Mediterranean region with scattered records in the west: Turkey (
The species as a whole is widely distributed around the Mediterranean extending to Madeira and the Canary Islands, where it may be introduced.
This species is morphologically very similar to
This group is exceptionally complex and there are few clearly demarcated species. Although
TANZANIA, Kilimanjaro,
Vigorous twining herb with stems reaching 2 m, vegetative parts varying from densely hirsute to subglabrous. Leaves petiolate, 3–8.5 × 1.8–6 cm, ovate-deltoid, acute, margin entire or obscurely crenate, base cordate (rarely hastate); petioles 7–30 mm. Flowers in many-flowered, axillary, pedunculate, bracteate heads; peduncles 1–13 cm long; bracteoles 5–10 × 3–7.5 mm, ovate, acute, scarious, tardily caducous; pedicels very short, 2–4 mm long; outer sepals 9–13 × 5–8 mm, broadly ovate, acute to apiculate, villous, becoming scarious; corolla 2.5–3(-4) cm long, very wide at the mouth, white, purplish or pink with a dark centre, unlobed, midpetaline bands pubescent; ovary glabrous; style glabrous, divided c. 7 mm above base; stigmas 2–2.5 mm, linear, slightly shorter than the style arm. Capsule glabrous; seeds glabrous, the surface with wavy, white-topped ridges. [
Mountains of eastern Africa from 1800 to 3500 m:. Ethiopia (
A very distinctive afromontane species because of its many-flowered capitate inflorescence but variable in indumentum, plants from Ethiopia commonly less hirsute and with slightly larger corollas than those from further south.
SOUTH AFRICA, Cape, without collection data (holotype G, not seen).
Perennial herb, usually coarsely brown-pubescent on all vegetative parts, occasionally glabresent or white-pubescent; stems to 1.5 m, climbing or prostrate, sometimes woody towards the base. Leaves petiolate, 1.5–5 × 1.5–4 cm, very variable in form but always palmately veined, oblong, ovate or reniform in outline, sometimes unlobed with coarsely crenate to laciniate margins, often deeply palmately lobed or palmatisect, base more or less hastate, apex acute or obtuse; petioles 0.4–1.5 cm. Flowers solitary or arranged in few-flowered cymes, peduncles 4–10 cm; pedicels 4–15 mm; bracteoles 4–7 mm, filiform to linear; outer sepals 8–13 × 5–8 mm, oblong-ovate, acute or acuminate, often scarious-margined; corolla 2–3.2 cm, white, unlobed, fimbriate, the midpetaline bands thinly pilose, terminating in a small tooth; ovary glabrous; style glabrous, divided 5–6 mm above base, stigmas 1.5–2 × 0.5–0.75 mm, very narrowly ellipsoid. Capsule glabrous; seeds glabrous, smooth except for the obscurely rugose angles. [
South Africa: Eastern and Western Cape (
Recognised by the large unlobed, often fimbriate, white corolla combined with palmately-veined leaves and very short, thick stigmas. Some specimens are almost indistinguishable from the Mediterranean
Plants from Namaqualand (northwestern Cape) have a shorter calyx (6–7 mm long) and smaller corolla (12–16 mm) and have been treated as a distinct species,
SOUTH AFRICA, Western Cape, George near Zivarte valley,
Perennial herb, glabrous to thinly pubescent in all vegetative parts; rootstock thick; stems to 3 m, prostrate or climbing. Leaves petiolate, very narrowly hastate, the central lobe 2–6 × 0.1–0.6 cm, linear to oblong, auricles very small, 0.2–1.2 × 0.1–0.4 cm, usually bifid, apex acute, margin entire; petioles 0.3–1.4 cm. Flowers axillary, pedunculate, usually paired (rarely solitary), peduncles 3–8(-14) cm; bracteoles 3–4 mm, subulate to narrowly lanceolate; pedicels 2–11 mm; outer sepals (5-)6–9 × (4-)6–8 mm, obovate, the apex truncate or rounded and often mucronate, margins scarious; corolla (1.2-)1.5–2.5(-3) cm long, white or pinkish, shallowly lobed, the midpetaline bands thinly pubescent terminating in teeth; ovary glabrous; style glabrous, divided 15 mm above base; stigmas ellipsoid, c 1 × 0.75 mm. Capsule glabrous; seeds glabrous, rugose with pallid ridges. [
South Africa: along the southern fringes of Western and Eastern Cape (
Recognised by the very narrow, hastate leaves combined with rounded, scarious-margined sepals, 2-flowered peduncles and relatively large corolla. The distinctive stigma suggests a close affinity with
SOUTH AFRICA, Western Cape,
Very similar to
South Africa (Western Cape, especially in the Clanwilliam area) and Namibia centred on Namaqualand (
Molecular studies (
We have only cited the isotypes we have seen. The isotype at W appears to represent
SOUTH AFRICA, Cape.
Perennial herb, thinly pubescent in all vegetative parts; rootstock thin, woody; stems to 70 cm, prostrate. Leaves petiolate, relatively small, lanceolate to ovate in outline, the central lobe 1– 5 × 0.3–0.8, cm, oblong to lanceolate, dentate, pinnatisect to pinnatifid, characteristically cordate-deltoid, auricles prominent, usually bifurcate, 0.3–1.5 cm, usually dentate, apex acute or obtuse; petioles 3–12 mm. Flowers solitary (rarely paired), axillary, pedunculate, peduncles 0.8–3 cm; bracteoles 3–7 mm, subulate to linear; pedicels 3–12 mm; outer sepals 9–12 × 4–6 mm, ovate, acute to acuminate, inner sepals scarioius, pubescent only in central vertical lines; corolla 1.6–2.8 cm, white or pink, very shallowly lobed, the midpetaline bands pubescent, terminating in a tooth; ovary glabrous; style glabrous, divided c. 7 mm above base; stigmas 4 mm, linear. Capsule glabrous; seeds smooth. [
South Africa: Eastern Cape, KwaZulu-Natal, Free State, North West, Gauteng, Mpumalanga, Limpopo (
Distinguished by its essentially pinnately-nerved central leaf lobe. It seems close to
SOUTH AFRICA, Northern Cape,
Completely glabrous perennial herb with decumbent to ascending stems to 30 cm long from a central taproot. Leaves shortly petiolate, 1–2 (-3) cm long, very variable in form on the same plant and between plants, sometimes linear with a hastate base and minute auricles (
South Africa except KwaZulu-Natal (
Distinct for being completely glabrous with small, delicate leaves and short obovate, rounded to slightly fimbriate sepals.
SOUTH AFRICA, Free State,
Perennial herb with woody taproot from which spread numerous stems to 60 cm, plant covered in adpressed brown to silvery hairs. Leaves 1–2.5 × 0.5–2 cm, lanceolate to ovate in outline, variable in form from pinnatisect to palmately 5-lobed, often with the terminal lobe much longer and deeply toothed and the basal lobes bifid, base truncate to shallowly cordate; petioles 1–5 mm long. Flowers solitary, axillary, pedicellate but not pedunculate (rarely peduncle to 1mm); bracteoles 1–2 mm long, subulate; pedicels 2–6(-10) mm, outer sepals 4–5(-6) × 2–3 mm, ovate to oblong-elliptic, acute; corolla 7–10 mm long, pink or white, shallowly lobed, midpetaline bands pubescent with brown hairs; ovary glabrous; style glabrous, divided 2.5 mm above base; stigmas 2.5 mm, slightly widened upwards. Capsule glabrous; seeds glabrous, smooth but muricate on angles. [
South Africa except KwaZulu-Natal (
Distinguished by the solitary, pedicellate flowers and near absence of peduncles combined with the very small calyx, the sepals usually about 5 mm long and thinly covered in brownish hairs. The inflorescence is similar to that of
There are specimens apparently intermediate with
SOUTH AFRICA, Eastern Cape,
Perennial herb similar in facies to
South Africa, almost endemic to the Cape (
Distinguished from
NAMIBIA,
Densely hispid-pilose perennial with prostrate/trailing stems from a central taproot to 70 cm; hairs rusty-brown in colour. Leaves petiolate, 1–3.5 (-5) × 0.5–2.5 cm, ovate in outline, deeply pinnatisect, abruptly narrowed and cuneate onto the petiole; petioles 1–8 mm. Flowers 1–2, axillary, subsessile; peduncles to c. 0.3 cm; pedicels 0; bracts filiform, 5–9 × 0.5 mm; outer sepals broadly ovate with a long caudate apex, c. 7–8 mm at anthesis, accrescent to 12–13 mm, becoming somewhat scarious, the margin crisped; corolla 10–12 mm long, deeply lobed for c. 4 mm, nearly concealed by calyx, white with pilose midpetaline bands; ovary glabrous, divided c. 4 mm above base; stigmas 2 mm, linear. Capsule glabrous; seeds glabrous, rugose. [
Namibia (
Very distinct because of the subsessile flowers and accrescent calyx, which almost conceals the corolla.
Plate 4065 in Curtis, Botanical Magazine 70 (1844); epitype (designated here): SOUTH AFRICA, North West Province, Gauteng, Magaliesberg,
Perennial herb with all vegetative parts tomentose with brown or grey hairs; rootstock stout, woody; stems 20–100 cm long, decumbent and trailing to erect, occasionally apparently rambling over shrubs, often woody towards the base. Leaves subsessile or shortly petiolate, 1–2.5 × 0.1–1.5 cm, narrowly oblong with or without basal auricles to palmately 5-fid (
South Africa (
Usually easily recognised by the densely tomentose indumentum combined with revolute leaf margins. The calyx and corolla are similar in size to that of
ZIMBABWE, Gweru,
Perennial herb, all vegetative parts covered in appressed sericeous hairs; rootstock woody, very stout, apparently horizontally spreading; stems erect or ascending, rarely rambling over shrubs, 20–80 cm high. Leaves shortly petiolate, 0.8–3 × 0.2–2 cm, oblong to obovate, apex acute to apiculate, margin entire to crenate, not revolute, base truncate to cordate, prominently veined especially on the lower surface; petioles 0.5–3 mm. Flowers solitary, axillary, pedunculate; peduncles (0.1-)0.3–2.5 cm; bracteoles 3–5 mm, linear; pedicels 2–10 mm; outer sepals 8–10 × 3–6 mm, broadly to narrowly ovate, tapered to an apiculate apex; corolla 16–20 mm long, white or pale pink, shallowly lobed, the midpetaline bands pubescent, terminating in teeth; ovary glabrous, finely acuminate; style glabrous, divided 5 mm above base, stigmas 5 mm, linear. Capsule glabrous seeds smooth.
Endemic to Zimbabwe, gowing in grassland on serpentine deposits, 1270–1700 m. (
Somewhat variable in habit but readily recognised by the broad oblong-obovate leaves, silvery sericeous indumentum, acute sepals and larger corollas.
Affine
ZIMBABWE, Salisbury [Harare], a weed, 29 June 1927,
Perennial herb, all vegetative parts pubescent with somewhat asperous, sometimes rufous hairs; rootstock a woody taproot; stems prostrate or twining, up to 2 m long. Leaves petiolate, 3–6 × 0.5–2.5 cm, variable in shape, ovate-deltoid, auriculate, sometimes the auricles lobed, the central lobe commonly oblong, apex acute, the margins undulate, sinuate-dentate to pinnatisect, base hastate; petioles 3–30 mm. Flowers 1–6 together (very rarely all solitary on the same plant) in axillary pedunculate cymes; peduncles 10–35 mm; pedicels 2–15 mm, bracts 2.5–4 mm, linear; sepals very unequal, outer sepals 6–8 × 4–5 mm, ovate to elliptic, acute; inner sepals 4–6 × 3–4 mm, nearly glabrous, apiculate; corolla 9–12 mm long, white or pale pink, lobed, the midpetaline bands pubescent, terminating in prominent teeth; ovary glabrous; style glabrous, divided c. 3 mm above base, stigmas 2.5 mm, linear. Capsule glabrous; seeds glabrous, smooth. [
South Africa (
This species was treated as
A cultivated plant (
ETHIOPIA,
Perennial herb, all vegetative parts similarly obscurely puberulent to pubescent; rootstock a woody taproot; stems prostrate or twining, up to 2 m long. Leaves petiolate, (0.5-) 2–10 × 0.2–4 cm, variable in shape, narrowly deltoid in outline, auriculate with the basal auricles simple or, more commonly bifurcate, the central lobe oblong to oblong-lanceolate, much longer than the auricles, apex acute to apiculate, margin entire to undulate, base commonly more or less truncate and briefly cuneate onto the petiole, leaves near base of stem often with a broader central lobe than those near apex; petioles 5–25 mm. Flowers (1-) 2–6 together (very rarely solitary) in compact axillary, pedunculate cymes; peduncles 8–35 mm; bracteoles 2–5 mm, linear or linear-lanceolate; pedicels 1–10 mm, outer sepals 5–6 (-7) × 2–3 mm, lanceolate to ovate, acute, usually pubescent, inner sepals up to 5 mm wide, suborbicular, apiculate, margins scarious, glabrous or pubescent on the midrib only; corolla 7–12 mm long, white or pink, lobed, the midpetaline bands pubescent, terminating in prominent teeth; ovary glabrous; style glabrous, divided 3–4 mm above the base; stigmas c. 3 mm. Capsule glabrous; seeds glabrous, smooth. [
South Africa (
The type of
Cultivated plant grown at Uppsala (lectotype LINN 218.6!, designated by
Perennial herb, appressed pubescent to farinose in all vegetative parts, especially the younger stems; rootstock not known; stems to c. 1 m, twining or prostrate. Leaves petiolate, 3–9 × 2–6 cm, characteristically cordate-deltoid, auricles usually acute, apex acute to acuminate, margin entire, undulate or serrate; petioles 1–4.5 cm. Flowers 1–6 in axillary pedunculate cymes, peduncles 1.5–5 cm; bracteoles 1–2 mm, subulate; pedicels 1–15 mm; outer sepals 6–8 × 3–5 mm, lanceolate, ovate or elliptic, acute the apex often slightly reflexed, pubescent, inner sepals suborbicular with scarious margins, glabrous; corolla 10–15 mm, white or pinkish, lobed, the midpetaline bands pubescent; ovary glabrous; style glabrous, divided 4 mm above base, stigmas 1–1.5 mm, linear. Capsule glabrous; seeds glabrous, rugose. [
South Africa (
Usually readily recognised by the triangular-ovate, shortly pubescent to farinose, very acute leaves and small, deeply lobed corolla. However, occasionally plants are seen in which the leaves are ovate or sinuately lobed to more or less palmatisect, particularly in South Africa (
SOUTH AFRICA, Cape,
Very variable perennial herb, the vegetative parts usually thinly to densely pubescent, very rarely glabrous; stems decumbent, trailing, rambling or ascending usually < 60 cm long; rootstock a stout taproot. Leaves petiolate, sometimes dimorphic with ovate-deltoid (below) and narrowly lanceolate leaves on the same plant, 1–2.8 (-5.5) × (0.1-) 0.3–1.4 cm, ovate-deltoid to narrowly lanceolate, apex acute, margin entire (very rarely undulate), base sagittate or hastate, the basal auricles not bifid, varying greatly in width; petioles 2–4 (-7) mm. Flowers axillary, pedunculate, 1(-2); peduncles (3-) 6–33 mm; bracteoles 2–3 mm, linear to linear-lanceolate; pedicels 2–5 (-12) mm, outer sepals 5.5–8 × 4–5 mm, ovate, broadly oblong to obovate, acute to obtuse, the apex often somewhat bent outwards, glabrous or pubescent, inner sepals glabrous; corolla (1-) 1.2–1.7 cm. pink or white, shallowly lobed, midpetaline bands pubescent, terminating in a tooth; ovary glabrous, style glabrous, divided 3–4 mm above base; stigmas 2–4 mm, linear. Capsule glabrous; seeds glabrous, minutely rugose. [
South Africa (
Distinguished from related species by the solitary pedunculate flowers, the corolla typically 1.2–1.7 cm long. The leaves are essentially ovate-deltoid, becoming linear in some cases, although often very narrowly so and the basal auricles are simple. The petioles are often very short.
The type of
Although our concept of
TANZANIA, N. of Lake Nyasa,
Prostrate perennial herb, densely tomentose on all vegetative parts, often brownish when dry. Leaves petiolate, 1.2–5 × 0.5–2.5 cm, variable in shape, ovate, lanceolate-deltoid or, most commonly oblong, margin undulate to crenate, base hastate to sagittate; petioles 3–6 (-20) mm. Flowers solitary, axillary, pedunculate; peduncles solitary or, occasionally, paired, 1.5–4 cm, often arching; bracteoles 5–7 mm, linear; pedicels 3–8 (-15) mm; outer sepals 9–11 × 5 mm, ovate, acute to shortly acuminate, densely hairy; corolla (1.3-)1.5–1.8 cm long, white, unlobed, midpetaline bands pubescent, terminating in a tooth; ovary glabrous. Capsule glabrous; seeds glabrous, nearly smooth but somewhat rugose on the angles. [
Malawi (
We have widened the concept of this species from that of
As with many other African species, intermediates with other species are found. These have the same indumentum, relatively large corollas and sepals of
SOUTH AFRICA, Eastern Cape,
Perennial herb, densely brownish or whitish villous in all vegetative parts; rootstock not known; stems to 60 cm, slender, twining or (?) prostrate. Leaves petiolate, 2–4 × 0.7–1.2 cm, deltoid with cordate, hastate or sagittate base, apex acute, margin undulate or crenate; petioles 4–7 (-12) mm. Flowers axillary, pedunculate, solitary or paired, peduncles 1.5–2 (-6) cm; bracteoles 6–7 mm, linear; pedicels 3–6 (-10) mm; outer sepals 8 × 4 mm, ovate, abruptly narrowed above the middle to an acuminate apex; corolla 16–21 mm long, white, shallowly lobed, the midpetaline bands densely pilose, terminating in a tooth; ovary glabrous; style glabrous, divided 8–10 mm above the base, stigmas 2.5 mm linear. Capsule glabrous; seeds glabrous, obscurely rugose with pallid ridges, not puberulous as stated by
South Africa: Eastern Cape (
Distinguished by the relatively slender, twining stems, dense indumentum, hastate or sagittate leaves and abruptly acuminate sepals.
SOUTH AFRICA, KwaZulu-Natal, Pietmaritzburg,
Perennial herb, densely hirsute with brownish hairs in all vegetative parts; rootstock woody; stems to c. 1 m, apparently trailing (rarely climbing), relatively stout. Leaves petiolate, 1–6 × 0.8–4 cm, ovate-deltoid, simple, apex acute, margin undulate to irregularly dentate, base cordate; petioles 5–10 (-15) mm. Flowers 1–5; peduncles 2–6.5 cm; pedicels 3–6 (-15) mm; bracteoles 6–12 × 1–2 mm, linear, narrowly lanceolate or narrowly oblanceolate; outer sepals 14–18 × 7–9 mm, broadly to narrowly ovate, obtuse or acute, margin undulate or crenate; corolla 15–30 mm, white or cream, shallowly lobed, the lobes broadly triangular, acute, c. 5 mm long, the midpetaline bands densely pilose; ovary glabrous, acuminate; style glabrous, divided 5–8 mm above base, stigmas 6 mm, linear. Capsule glabrous; seeds tuberculate. [
South Africa: centred on KwaZula-Natal extending to Eastern Cape, Free State, Lesotho, Swaziland and Northern Province (
Distinguished by its tomentose, entire, ovate to oblong, cordate hirsute leaves. Plants described as
SOUTH AFRICA, KwaZulu-Natal, Mooi River,
Perennial herb, shortly pilose with stiff spreading hairs in all vegetative parts; rootstock not known; stems to at least 80 cm, apparently trailing. Leaves petiolate, very narrowly hastate, the central lobe 2 -5 × 0.2–0.4 cm, linear-lanceolate, basal auricles 3 -4 mm long, bifid, apex acute to apiculate, margin entire; petioles 5–16 mm. Flowers axillary, pedunculate, 1 (–2), peduncles 3–5.5 cm; bracteoles 6–10 mm, linear; pedicels 3–8 mm, noticeably more densely hirsute than peduncles; outer sepals 14–18 × 8 mm, ovate, long acuminate, margin undulate, inner sepals distinctly shorter; corolla 25–30 mm, yellow-green, deeply lobed, the lobes triangular, acuminate, c. 10 mm long, the midpetaline bands densely pilose, terminating in the apex of the lobes; ovary glabrous, style glabrous, divided 6–10 mm above base, stigmas 3–5 mm.
South Africa: KwaZulu-Natal (
Included by
Apart from two anomalous species (
The first two species treated here,
The taxonomy of the American species is difficult as can be appreciated by the synonomies listed under many species, the same infraspecific entity being placed variously under different species. However, we believe that
MEXICO, León,
Pubescent perennial herb from a stout tap root; stems decumbent or trailing to at least 1 m. Leaf blade very variable in size and form, 1.5–4 (-6.5) × 1–2.5 cm, most commonly with a narrow linear-ligulate central lobe much longer than the small lobed or bifurcate auricles, sometimes palmatisect, sometimes broadly ovate-deltoid, auriculate, usually densely and finely pubescent, apex acute, base cordate, margin entire, undulate or (rarely) crenate-serrate; petioles 0.5–2.5 cm. Flowers 1–3 in pedunculate, axillary cymes; peduncles 1.5–9 cm; bracteoles 1.5–2.5 × 1 mm, linear-lanceolate; pedicels 2–9 (-17) mm; outer sepals 6–8(-12) mm, narrowly elliptic, truncate to auriculate at base, margin entire to crenate, apex truncate and mucronate to acute; corolla 1.4–1.8(-3.0) cm long, white, white with dark centre or pink, shallowly lobed; midpetaline bands pubescent, terminating in a mucro; filaments eglandular; ovary glabrous; style glabrous or pubescent just below the stigmas, somewhat persistent, divided 5–7 mm above base; stigmas 2 mm, weakly exserted. Capsule glabrous, seeds minutely rugose. [
A very variable plant in many respects. However the vast majority of specimens have small leaves with a narrow linear-ligulate central lobe and short bifurcate or otherwise lobed auricles. In most plants the outer sepals are abruptly narrowed to auriculate at the base, but in many specimens including the type, they are gradually narrowed to the base. Plants are usually densely pubescent. The recognition of the following varieties only account for some of the great variation seen in this species.
Flowers relatively small; sepals 6 – 8 mm long; corolla 1.4–1.8(2.3) cm long, usually pink.
Mexico: north and central (
A var, typo floribus grandioribus, sepalis 11 – 12 mm longis, corolla 2.5–3 cm longa, plerumque alba, in centro atropurpurea.
UNITED STATES OF AMERICA, Texas, New Braunfels,
Distinguished by it is larger flowers; sepals 11–12 mm long; corolla 2.5–3 cm long, usually white but often with a dark centre.
United States, Texas: widely distributed but not common in at least seven Texan counties (
Intermediates with
UNITED STATES OF AMERICA, Texas,
Trailing or twining herb with stems at least 60 cm long from a central rootstock, the vegetative parts densely pubescent to whitish-tomentellous. Leaves petiolate, 2–5 × 1.3–3 cm, ovate-deltoid to broadly oblong, obtuse or acute, margin undulate to incised-dentate, base shallowly cordate and cuneate onto the petiole, auricles simples or toothed, veins very prominent on lower surface; petioles 1–2.5 cm. Flowers 1(-2) borne on long axillary peduncles; peduncles 3–5 cm, often bent at apex; bracteoles 1–2 mm, minute, linear-lanceolate; pedicels 3–14 mm; sepals 9–12 × 5 mm, broadly oblong, apex rounded to emarginate and mucronate, base truncate to somewhat auriculate; corolla 2.5–3 cm long, white, usually with a maroon centre, unlobed, midpetaline bands pilose, terminating in a tooth; ovary glabrous; style divided c. 8–10 mm above base, glabrous or, just below the stigmas, pubescent; stigmas 2 mm, weakly exserted. Capsule glabrous; seeds glabrous, smooth, black. [
Endemic to Texas in the United States of America: restricted to Holocene sands in Brooks and Hidalgo Counties (
A recently described species which requires further study. It may prove only to be an unusually distinct form of
Based on
Thinly to densely pubescent herb from a thick rootstock, sometimes sericeous on young parts, but more or less glabrescent; stems trailing (rarely twining), up to 2.5 m long. Leaves petiolate, 2–8 × 2–6 cm, very variable in form, usually linear or oblong with prominent elongate bifurcate basal auricles, but occasionally ovate-deltoid to suborbicular with rounded auricles, apex usually acute, margin entire or undulate, base cordate to truncate; petioles 0.5–3.5 cm. Flowers 1–2 (-3), axillary, pedunculate; peduncles 2–4 (-6.5) cm; bracteoles 2–4 mm, lanceolate; pedicels 5–10 mm; outer sepals 7–9 × 5–7 mm, elliptic, obtuse, mucronate; corolla 1.5–2.5 cm long, pink, very shallowly lobed with slightly fimbriate margins, midpetaline bands dark, pilose, terminating in a pilose mucro; ovary glabrous; style divided 6–10 mm above base, stigmas 1.5–3.5 mm, cylindrical to linear, unusually variable. Capsule glabrous; seeds rugose. [
Endemic to central Chile from Antofagasta south to Santiago (
This polymorphic species is usually easily distinguished from all other South American species by the leaves with bifurcate auricles combined with pink corollas usually around 2–2.5 cm long. However, some specimens from Coquimbo (
ARGENTINA, Pampas de Buenos Aires,
Finely adpressed-pubescent herb from a thick rootstock, sometimes sericeous on young parts; stems trailing, up to 2 m long, 1–3 mm in diameter. Leaves petiolate, 3–6 (-11) × 1.2–3 (-6) cm, very variable in shape but usually oblong-lanceolate or strap-shaped with pronounced (rarely bifurcate) auricles, occasionally ovate-deltoid, characteristically 5 times as long as broad, apex usually obtuse, mucronate, margin undulate to crenate or serrate, base hastate to cordate; petiole 5–40 mm. Flowers (1-) 2–5 in axillary, pedunculate cymes; peduncles 0.6–4 (-6) cm; bracteoles 2–4 mm, narrowly lanceolate; pedicels 3–12 mm; outer sepals 6–8 × 3–5 mm, elliptic, acute; corolla 1–1.5 cm long, pink, shallowly lobed, midpetaline bands pubescent terminating in teeth; ovary glabrous, style divided 3–6 mm above base; stigmas c. 1.5 mm, narrowly ellipsoid. Capsule glabrous; seeds smooth to indistinctly tuberculate. [
Argentina (
The adpressed pubescent, lanceolate to strap-shaped leaves combined with the small corolla are characteristic. Flower size and usually leaf shape serve to distinguish it from
CHILE, Coquimbo,
Glabrous or puberulent herb from a deep rootstock. Stems 30(-50) cm long, numerous, trailing. Leaf blade 0.6–2 × 0.4–1.6 cm, ovate-deltoid; base truncate and briefly cuneate onto the petiole, auricles not well-developed; apex obtuse and mucronate or acute; margin entire; petiole 4–8 (-12) mm. Flowers solitary (rarely paired), axillary, pedunculate; peduncles 10–18 mm; bracteoles 2–6 mm, linear; pedicels 2–4 mm; outer sepals 7–10 × 5–7 mm, elliptic, obtuse; corolla (1-) 1.5 (-2) cm long, pink, shallowly lobed, midpetaline bands pilose, terminating in small teeth; ovary glabrous; style divided 4–5 mm above base, stigmas 2–3 mm. Capsule glabrous; seeds smooth but with slightly muricate angles. [
Central Chile (
This is an Andean species variable in indumentum with a superficial resemblance to
ARGENTINA, Chaco,
Finely pubescent trailing or twining herb from a thick rootstock, stems up to 1 m long. Leaves petiolate, 1–5 × 0.5–2 cm, ovate-deltoid, auriculate, apex obtuse and mucronate, margin weakly crenate, base cordate; petioles 3–13 mm. Flowers 1–4 in axillary, pedunculate cymes; peduncles 1–3 (-8.5) cm; bracteoles 1.5–2.5 mm, narrowly ovate; pedicels 5–15 mm; outer sepals 4–6 × 4–5 mm, broadly elliptic to obovate, obtuse, inner sepals truncate; corolla 0.7–0.8 cm long, pale pink, shallowly lobed, midpetaline bands pilose in the upper half terminating in small teeth; ovary glabrous, acuminate; style glabrous, divided 3.5–4 mm above base; stigmas 1.5 mm. Capsule glabrous; seeds strongly tuberculate. [
Endemic to Argentina: Corrientes and Chaco (
A local endemic with a small corolla growing on sand deposits in river valleys.
URUGUAY, Montevideo,
Very variable perennial herb with numerous, often branched, trailing stems from a stout central rootstock, most commonly glabrous, sometimes thinly pubescent and rarely white-tomentose. Leaves petiolate, 1–3(-5) × 1–3(-5) cm, very variable in form but always deeply divided, usually profoundly palmatisect or pinnatisect with narrow laciniate segments, rarely with broader segments; apex obtuse or acute; margin undulate or entire; petioles 3–15 mm. Flowers 1–2(-3), axillary, pedunculate; peduncles 1–3(-5) cm; bracteoles 2–3.5 mm; pedicels 2–10 mm; outer sepals 6–9 × 5–6 mm, elliptic to obovate, margins scarious; corolla 1–2 cm long, white or white with purple centre, lobed with acute apices, exterior glabrous to thinly pilose in correlation with overall plant indumentum, midpetaline bands present or absent but if present dark violet, pilose; ovary glabrous; style glabrous, divided 5–11 mm above base; stigmas 1.5–2 mm. Capsule glabrous; seeds smooth, black. [
Argentina (
An extremely variable species easily recognised by its deeply divided leaves and white flowers, which are occasionally with dark violet midpetaline bands. The following specimens are outstanding and could each constitute a distinct taxon, but which I hesitate to recognise in the absence of any matching material:
Two specimens from Bolivia (
PERU, Huara,
Trailing or (less commonly) twining herb from a thickened woody rootstock c. 1 cm thick, all vegetative parts grey-tomentose. Stems up to 1 m long, apparently more slender in twining plants, numerous. Leaves petiolate, 2–6.5 × 0.5–3 cm, ovate to ovate-deltoid, the auricles not well-developed, apex acute to obtuse, margin undulate to irregularly dentate, base cordate; petioles 5–12 (-22) mm. Flowers 1–2 (-3), axillary, pedunculate; peduncles 1–3 (-6)cm, often shorter than the leaves; bracteoles 2–4 mm, linear-lanceolate; pedicels 5–12 mm; outer sepals 7–10 × 4–6 mm, (narrowly) elliptic, usually acute; corolla 1.4–1.8 cm long, white, shallowly lobed, midpetaline bands extended into mucros, tomentose; ovary conical, 1.5–2 mm, glabrous or apically pilose, style divided 5–7 mm above base, glabrous except immediately above ovary, stigmas 2.5–3 mm. Capsule glabrous or apically pilose; seeds smooth.
We recognise two subspecies based on ovary and capsule indumentum:
Ovary and capsule completely glabrous.
The principal or only subspecies in Ecuador and Peru extending south into Bolivia: Ecuador (
The name
Based on
Ovary and capsule apically pilose. [
The only subspecies present in the Southern Cone extending north to Bolivia and Brazil: Chile (
PERU, Junin, Huancayo,
Herb apparently from a deep rootstock, glabrous or with a few scattered hairs on vegetative parts, stems trailing, 5–15 cm long. Leaves petiolate, 0.6–1.6 × 0.5–1.3 cm, ovate, apex more or less rounded, margin strongly crenate, base truncate; petioles 3–8 mm. Flowers 1(-2), axillary, pedunculate; peduncles 1.2–1.6 cm; bracteoles 2.5–4 mm, oblong; pedicels 2–5 mm; outer sepals 6–8 × 4.5–6 mm, obovate-elliptic, concave, scarious, emarginate, inner sepals similar emarginated or mucronulate; corolla 1.2–1.5 cm long, white to pale pink with a dark centre, deeply lobed, midpetaline bands glabrous terminating in a small tooth; ovary glabrous, style divided 5–7 mm above base, stigmas c. 2 mm. Capsule glabrous; seeds smooth, glabrous.
Peru: Junin, Cusco (
A distinctive nearly glabrous species with small ovate, basally truncate, crenate leaves.
Based on
Perennial herb from a tap root, thinly pubescent on all vegetative parts. Stems 15-40 cm long, trailing. Leaf blade1–2.3 × 0.6–1.2 cm, ovate-deltoid; base cordate and briefly cuneate onto the petiole, auricles prominent; apex shortly mucronate; margin incised-dentate; petiole 3–5 mm. Flowers solitary (rarely paired), axillary, pedunculate; peduncles 8–14 mm; bracteoles 1–2 mm, filiform; pedicels 3–5 mm; outer sepals 6–7 × 4mm, ovate, obtuse; inner sepals 6-7 × 6-7mm, suborbicular, rounded, slightly scarious; corolla 1.5–1.6 cm long, white, lobed, midpetaline bands pubescent, terminating in triangular teeth; ovary glabrous; style divided 4–5 mm above base, stigmas 2.5 mm. Capsule glabrous; seeds smooth, glabrous.
Moquegua (
A poorly-known species growing at low altitudes in Peru but easily recognised by its incised-dentate leaves. Morphologically it would appear to lie between
PERU, Huanuco
Pubescent to densely hirsute herb; stems twining up to 3 m high. Leaves petiolate, 3–8 × 1–4 cm, ovate-deltoid, strongly auriculate, usually large, apex usually obtuse and mucronate, margin undulate to sinuate, base broadly cordate to hastate with midrib area cuneate onto petiole; petioles 7–15 mm. Flowers (1-) 3–7 in compact axillary, pedunculate umbellate cymes; peduncles 1.5–12 cm; bracteoles 2–5 mm, narrowly lanceolate; pedicels 2–12 mm, apparently accrescent after anthesis; outer sepals 6–6.5 × 3.5–5 cm, elliptic, obtuse or acute; corolla 1.1–1.5 cm long, white to pink, deeply lobed, midpetaline bands brownish, pilose, terminating in a mucro; ovary glabrous; style divided c. 7 mm above base; stigmas 3 mm, more or less included. Capsule glabrous; seeds smooth. [
We recognise two subspecies, which are distinct through most of their range, but intergrade in parts of northern Argentina (
Distinguished by the more numerous flowers (there are nearly always some cymes with >3 flowers), the relatively short pedicels, the cymes usually forming rather tight umbellate clusters, and the smaller, usually pinkish, lobed corolla.
Amphitropical, Andes and southern Brazilian highlands in South America; United States of America and Mexico in North America: Ecuador (
In South America this species appears to be distinctly montane in distribution being limited to the Andes and the higher mountains of southeastern Brazil. Specimens from Andean Bolivia, Peru and Ecuador are very consistent in habit. Its status in North America is uncertain. The leaves are often more strictly triangular and more coarsely dentate than in South American plants but some specimens such as
Based on
Distinguished from
Northern Argentina (
According to the image in
There is at least one collection from Paraguay (
ARGENTINA, Corrientes, Ituzaingó,
Densely pubescent or tomentose trailing or twining herb from a thickened rootstock, stems to at least 1 m long, relatively stout. Leaves petiolate, 2–7(-10) × 0.7–2.8 cm, lanceolate-oblong with pronounced basal auricles, these sometimes lobed, apex acute and mucronate, margin entire to weakly undulate, base hastate; petioles 3–10(-15) mm. Flowers solitary (very rarely paired), axillary, pedunculate; peduncles 2–5 cm; bracteoles 2–5 mm, linear-lanceolate; pedicels 10–15 mm; outer sepals 10–13 × 5–6.5 mm, elliptic, obtuse; corolla 2.5–4 cm long, white with dark centre, shallowly lobed, midpetaline bands pilose, extended into teeth; ovary glabrous, style glabrous, divided c. 16 mm above base, stigmas 2–3 mm, cylindrical. Capsule glabrous; seeds smooth to finely tuberculate. [
Argentina: Corrientes and Misiones (
An apparently very distinctive local endemic characterised by its large, usually solitary flowers and densely hairy indumentum. However, a number of specimens indicate that it is more variable than O’Donell supposed.
BRAZIL, Rio Grande do Sul,
Prostrate perennial with woody base, the stems winged, twining at the apices. Leaves shortly petiolate, 2–6 × 0.2–0.4 cm, linear-oblong, base sagittate, apex acute, glabrous; petiole 2–6 mm. Flowers solitary; peduncles 25–60 mm, shortly winged; bracteoles 3–5 nn, lanceolate, deciduous; pedicels 8–12 mm; outer sepals 6–8 × 3–4 mm, obovate, truncate, mucronulate, glabrous or canescent, the margins ciliolate; inner sepals similar but with scarious margins; corolla 1.8–2.2 cm long, white, glabrous, midpetaline bands sericeous; ovary glabrous, subglobose; style divided 6–9 mm above base; stigmas 3–4 mm. Capsule glabrous, apiculate; seeds black, glabrous.
Brazil (Rio Grande do Sul, Paraná).
Resembling
Based on
Usually erect but occasionally decumbent to ascending cerrado perennial with woody xylopodium, vegetative parts villous with long spreading cobwebby hairs; stems several, erect, herbaceous, 20–30(-50) cm high. Leaves sessile or very shortly petiolate, 1.5–4.5 × 0.8–1.5 cm, ovate to oblong-ovate; base rounded, truncate to subcordate or sagittate; apex acute or (above) apiculate; margin entire; petiole 0(-2) mm. Flowers solitary (rarely paired); peduncles 20–50 mm; bracteoles 3–5 × 0.5–1 mm, linear-lanceolate, finely acuminate; pedicels 5–10 mm; outer sepals 10–15 × 5–7 mm, ovate, acuminate; corolla 3.2–4 cm long, white, unlobed, midpetaline bands pink, long-pilose, terminating in a mucro; ovary glabrous; style glabrous, divided 8–12 mm above base, persistent in fruit; stigmas 3 mm. Capsule glabrous, unusually large, c. 1 cm diameter; seeds smooth. [
Paraguay (
Unique species because of its adaptation to the cerrado biome. The erect habit, xylopodium, woolly, often cobwebby indumentum, subsessile leaves and large corolla all render it distinct from other American species.
Herbaceous trailing or twining species, relatively slender compared with species from other regions. Leaves petiolate, with the lamina narrowed to a sagitate or hastate base, commonly dimorphic or even trimorphic; basal leaves often simple, stem leaves often lobed with narrow segments. The ovary, style and capsule are always glabrous. Although
The taxonomy of Australasian species is difficult. Many species are superficially similar especially when young or showing only one leaf form, and the taxonomy is often based on the direction of the fruiting peduncle and seed sculpture so non-fruiting specimens and incomplete specimens can be impossible to name. In many herbaria, all species from this region were once filed under the name
Australian species are recorded as adventives or naturalised in other countries including the British Isles (
AUSTRALIA, South Australia,
Perennial herb from a central taproot with trailing stems to 1 m, plant adpressed pubescent to glabrescent. Leaves petiolate, 1–2 × 0.4–05 cm, lanceolate-deltoid, acute, margin entire to slightly undulate, base shallowly cordate and auriculate, the auricles entire or bifid; petioles 3–8 mm long. Flowers solitary, pedunculate, axillary, becoming recurved in fruit; peduncles mostly 1–2 cm long; bracteoles filiform, 1–1.5 mm; pedicels 3–12 mm; outer sepals 2–3(-4) × 2–3 mm, obovate to broadly elliptic, rounded and minutely apiculate, scarious, glabrous or thinly appressed pubescent; inner sepals similar; corolla 5–7 mm long, white or pink, very shallowly lobed, midpetaline bands almost glabrous except for a few hairs at apex; ovary glabrous; style glabrous, divided c. 2 mm above base, stigmas c. 1.5–2 mm. Capsule glabrous; seeds coarsely and irregularly tuberculate. [
Australia: South Australia and adjacent New South Wales (
Very distinctive because of the tiny calyx, lanceolate-deltoid leaves with basal auricles and the distinctive seed ornamentation.
AUSTRALIA, Queensland,
Perennial herb with trailing or twining stems to at least 50 cm, plant thinly pubescent to glabrescent. Leaves petiolate, dimorphic; petioles 2–10 mm; lowermost leaves 2.5–4 × 0.6–1.6 cm, deltoid, obtuse and finely mucronate, entire, base truncate and briefly cuneate onto the petiole, auricles absent; middle leaves similar but base more or less cordate, auricles present, often bifid or tridentate, the central lobe longer and narrower; middle and upper leaves with a narrowly linear-lanceolate central lobe 3–6 × 0.1–0. 6 cm, the basal auricles more or less reflexed so base sagittate, inconspicuous, bifid or trifid, 3–5 mm long, segments usually very narrow. Flowers solitary (very rarely paired), pedunculate, axillary; peduncles 1.2–3.5(-5.5) cm, recurved in fruit; bracteoles 1–2.5 mm, filiform; pedicels 4–12 mm; sepals 4–6 × 2.5–3 mm, obovate or elliptic, rounded and mucronate, scarious-margined, glabrous or pubescent on dorsal surface near apex; corolla 0.7–1.6 cm long, pink, shallowly lobed with triangular lobes, midpetaline bands pilose; ovary glabrous; style glabrous, divided 3–4 mm above base, stigmas 1.5–2 mm. Capsule glabrous; seeds with prominent wavy tubercles. [
Australia: Northern Territory, Queensland and New South Wales (
Very immature plants could be confused with
AUSTRALIA, South Coast, Bay 10 (Port Lincoln)
Perennial herb with twining or (occasionally) trailing stems to at least 50 cm, plant adpressed pubescent to more or less strigose. Leaves petiolate, not strongly dimorphic, 2.1–7.5 × 1–1.5 cm, narrowly deltoid, basally truncate and shortly cuneate onto the petiole, the central lobe linear, oblong, lanceolate or oblanceolate, acute, entire, 2–6 mm wide, basal auricles always present, 2–10 mm long, usually simple, occasionally bifurcate or toothed; petioles 6–10 (-20) mm. Flowers solitary or paired (rarely 3), pedunculate, axillary; peduncles mostly 1–3.5 cm long, not recurved in fruit; bracteoles 1.5–2.5 mm long, filiform; pedicels 4–10 mm; sepals 4.5–6 × 3–4.5 mm, broadly elliptic to obovate, rounded and mucronate at apex, margin somewhat scarious, dorsal surface pubescent; corolla 1–1.8 cm long, pink, lobed with broadly triangular lobes, midpetaline bands pilose towards apex; ovary glabrous; style glabrous, divided 4–6 mm above base, stigmas c. 2 mm. Capsule glabrous; seeds nearly smooth with obscure tubercles. [
Widespread in Australia, except the east coast, but most abundant in South Australia and Western Australia (
The usually very obviously twining stems with adpressed indumentum and narrowly deltoid auriculate leaves and straight peduncles serve to distinguish this species. The seeds are only obscurely tuberculate unlike those of
Robert Brown did not cite either a precise location or specimen in the protologue so Johnson was wrong to cite the Port Lincoln collection as holotype as there is another syntype mounted on the same sheet from a different location. In order to avoid future uncertainty we are formally designating the Port Lincoln collection at BM as lectotype. There is an isolectotype at Kew.
SOUTH AUSTRALIA, Cudnaka [Kanyaka],
Perennial herb with trailing stems to 1 m, vegetative parts covered in long, whitish appressed velvety hairs, densely so on young growth. Leaves dimorphic: leaves of non-flowering shoots long-petiolate, 1–2 × 0.6–1.5 cm, ovate-deltoid, rounded, margin serrate, base truncate (very rarely slightly auriculate) and shortly cuneate onto the petiole, petiole 1–2 cm; leaves of flowering shoots 1–1.5(-2.5) × 0.5–1.4 cm, ovate-deltoid in outline but deeply toothed and incised towards the base; petioles 0.2–0.8 cm. Flowers solitary, pedunculate, axillary; peduncles 0.3–0.8 (-1.2 )cm, becoming recurved in fruit; bracteoles linear c. 1 mm; pedicels 1–3 mm; outer sepals 4 × 3.5 mm, elliptic, apiculate, the point recurved, sericeous; inner sepals similar but with fewer hairs; corolla 5–6 mm long, white or pink, lobed with somewhat triangular lobes, midpetaline bands sericeous; ovary glabrous, style glabrous, divided 2–2.5 mm above base; stigmas c. 1.5 mm. Capsule glabrous; seeds slightly winged and with low irregular sinuate ridges. [
Australia: Southern South Australia and adjacent parts of Victoria and New South Wales (
This species shares with
AUSTRALIA, South Australia,
Perennial herb similar in general habit and distinctive features to
Australia: northeastern South Australia and adjacent Queensland (
AUSTRALIA, Western Australia, between Ashburton and Grey Rivers,
Perennial herb from a taproot with trailing (sometimes twining) stems to at least 75 cm, vegetative parts pubescent with adpressed or spreading hairs. Leaves petiolate, very plastic in shape, more or less trimorphic; petioles 0.3–3 cm, diminishing in length upwards; lowermost leaves 1–3 × 0.6–1.5 cm, deltoid, acute, margin undulate to crenate, base truncate and shortly cuneate onto the petiole; lower leaves sometimes distinct, 3–4 × 3–4 cm, deeply laciniate; middle leaves with elongate oblong undulate- or crenate-margined middle lobe, 2.5–3.5 × 0.4–0.8 cm and deeply lobed, more or less laciniate auricles; upper leaves with a very narrowly oblong central lobe 2–3.5 × 0.2–0.5 cm, two slightly shorter ascending lateral lobes and smaller bifurcate basal lobes. Flowers solitary or paired, pedunculate, axillary, not recurved in fruit; peduncles 0.6–3 cm; bracteoles 1–2 mm, filiform; pedicels 3–8 mm; sepals 4–5 × 2.5–4 mm, obovate or elliptic, acute or rounded, mucronate, pubescent; corolla 7–9 mm, white or pink, weakly lobed, midpetaline band pubescent; ovary and style glabrous; style divided 2–3 mm above base, stigmas c. 1.5 mm. Capsule glabrous, 4–6 mm in diameter; seeds winged, raised-tuberculate. [
Generally distributed in Australia but most common in central-western areas (
This species is extremly variable in leaf form and flowering specimens are indistinguishable from
AUSTRALIA, New South Wales,
A local endemic close to
Australia: New South Wales (
AUSTRALIA, South Australia,
Perennial herb with trailing stems to at least 50 cm, vegetative parts sparsely to roughly pubescent with adpressed or spreading hairs. Leaves dimorphic, petiolate; petioles 5–20 mm; leaves of non-flowering shoots 1–2.5 × 0.5–0.9 cm, oblong-subrectangular, truncate at apex and base, sometimes cordate and weakly auriculate, margins coarsely dentate; leaves of flowering shoots 1.2–3 × 0.5–1.6 cm, ovate or lanceolate in outline but deeply incised lobed, characteristically with the terminal lobe prominent, linear or narrowly oblong, acute to emarginate, the margin undulate to coarsely dentate, basal part deeply bi-quadrilobed. Flowers solitary (rarely paired), pedunculate, axillary; peduncles 0.8–2.2 cm, becoming recurved in fruit; bracteoles c. 1 mm, linear; pedicels 2–6 mm; sepals 3–5 × 3–3.5 mm, obovate or elliptic, acute or rounded with a small recurved mucro; corolla 5–9 mm, white or pink, weakly lobed, midpetaline band pubescent; ovary and style glabrous; style divided 2 mm above base, stigmas 1–2 mm. Capsule glabrous, 4–4.5 mm in diameter; seeds obscurely winged and with irregular ridging. [
We recognise two subspecies:
Larger in all its parts, the sepals 4–5 mm and corolla 7–9 mm.
Australia: eastern South Australia, New South Wales and Victoria (
AUSTRALIA, South Australia,
A relatively distinctive subspecies because of the small corolla 5–7 mm in length and the very small calyx, the sepals 3–4 mm long. The leaves on the flowering stems are usually sparsely hairy and the central lobe is narrowly oblong with a distinctive emarginate apex.
Australia: South Australia and adjacent parts of Western Australia centred on the Nullabar plains (
AUSTRALIA, Victoria,
Perennial herb with densely adpressed pubescent, trailing or twining stems, similar in facies to
Australia: Victoria and New South Wales (
This species occupies a situation somewhat intermediate both geographically and morphologically between
Plate in Bot. Mag. t.1007 (1807), lectotype, designated here; AUSTRALIA, plant from Hawkesbury River, New South Wales, collected by R. Brown on left side of sheet
Perennial herb with trailing or twining stems reaching at least 50 cm, stems crisped-pubescent, stouter than in other Australian species, commonly exceeding 2 mm in width. Leaves petiolate, variable in size but not markedly dimorphic, 2.5–8 × 1–3.5 cm, deltoid, apex obtuse and mucronate, margin crenate or repand, base broadly cordate and cuneate onto the petiole with prominent auricles, these variable, simple, toothed or laciniately lobed; petioles 1.2–2.5 cm, diminishing in size upwards. Flowers 1–4, usually clearly cymosely arranged, axillary, pedunculate, not recurved in fruit; peduncles 1–2 per axil, 2–6 cm, usually straight; bracteoles 1–3 mm long, filiform; pedicels 8–25 mm, very variable in length and strikingly unequal in individual inflorescences, often sinuate; sepals 5.5–7 × 3.5–5 cm. narrowly elliptic, terminating in a recurved mucro; corolla 1.2–1.5 cm, pinkish, lobed with triangular lobes, midpetaline bans pubescent near apex; ovary glabrous; style glabrous, divided 3–7 mm above the base, stigma 2 mm. Capsule glabrous, seeds tuberculate, unwinged. [
Australia: eastern coast of New South Wales and Queensland (
The most robust Australian species, the peduncles usually bearing several flowers and unusually sometimes with two peduncles per leaf axil. Most similar to
All Australian and New Zealand native species were once treated under this name following
AUSTRALIA, Tasmania, “Van Dieman’s Land near Risdon Cove”
Perennial herb with trailing or twining stems, pubescent to subglabrous to at least 40 cm but commonly short. Leaves extremely variable and often di/trimorphic on the same plant, petiolate, petioles 0.5–7 cm, diminishing in length upwards; lowermost leaves (if present) 1–2 × 0.3–1.5 cm, ovate-deltoid, obtuse, margin entire, undulate or sinuate-lobed especially towards the base, base cordate or truncate, the auricles poorly developed, entire to bi(tri-)fid; lower stem leaves 2–3 (-6) × 1.3–5 cm, broadly or narrowly ovate-deltoid in outline, entire undulate or deeply sinuate lobed, the basal auricle prominent, lobed with a short ascending lobe; upper stem leaves usually finely lobed, the central lobe linear-oblong, mostly 2.5–4.5 × 0.1–0.3 cm, acute or apiculate; auricles usually with a prominent ascending lobe resembling the terminal lobe but half its length together with short bifid reflexed lobes. Flowers pedunculate, axillary, usually solitary; peduncles 0.5–5 cm long, becoming recurved in fruit; bracteoles filiform, 1–2 mm long; pedicels 3–20 mm; sepals 4–7 × 2–2.5 mm, elliptic, rounded or acute, minutely mucronate, inner sepals slightly smaller; corolla 1.2–2 cm long, usually pink, weakly lobed, midpetaline bands pubescent only near apex; ovary glabrous; style glabrous, divided 3–10 mm above the base, stigmas 1–2 mm. Capsule glabrous; seeds 3–4 mm long, covered in low reticulate ridges. [
Principally southeastern Australia: Tasmania, Victoria, New South Wales, South Australia, Queensland, Western Australia (
NEW ZEALAND, South Island,
Greyish-pubescent creeping herb arising from underground rhizome; stems to 30 cm. Leaves petiolate, 1.5–3.6 × 0.2–0.8 cm, dimorphic and very variable, mostly deltoid or ovate, sometimes hastate and always with some leaves with an oblong or linear terminal lobe 1–5 cm long, combined with small basal auricles arising at right angles to the terminal lobe; petioles 1–5 cm. Flowers solitary, axillary, pedunculate; peduncles 2–6 cm long, 1-flowered, slender, pubescent; bracteoles 2–3 mm long, linear; pedicels 2–6 mm, pubescent; outer sepals 6–8 × 5–6 mm, broadly ovate, pubescent, larger than inner sepals; corolla 1.7–2 cm long, white, midpetaline bands pink; ovary glabrous; style glabrous, divided 6–7 mm above base. Capsule glabrous; seeds finely tuberculate. [
New Zealand: South Island (
Similar to forms of the Australian
NEW ZEALAND, South Island,
Perennial herb from an underground rhizome, stems decumbent and trailing to ascending up to 20 cm long, thinly pubescent on vegetative parts. Leaves petiolate, 6.5–11.5 × 5–12.5 mm, deltoid, ovate or broadly oblong, always lacking basal auricles, retuse or obtuse at apex, margin undulate, base
New Zealand: South Island (
Based on
Similar in overall morphology to
New Zealand: North and South Islands (
This is a morphologically and geographically heterogeneous group formed from Clade B (Figure
IRAN, “In rupestris apricis. inter Abuschir et Schiras”,
Intricately branched spiny shrub reaching 1 m in height; branches woody, finely appressed sericeous, small stem spines present. Leaves shortly petiolate, 1–2.4 × 0.5–0.9 cm, variable in shape, oblong, lanceolate, ovate or suborbicular, acute, entire, base cuneate, truncate or auriculate, glabrous, pubescent or sericeous; petioles 2–4 mm. Flowers axillary, pedunculate, solitary; peduncle 1–10 mm, stout, woody; bracteoles minute, c. 1 mm, squamose, caducous or absent; pedicels 2–7 mm, compressed, finely pubescent, often not differentiated from peduncle; sepals lax, somewhat scarious, 4–7 × 2.5–5 mm, ovate or broadly elliptic to obovate, obtuse or acute, obviously veined or not, somewhat scarious, becoming more or less erect and adpressed to capsule in fruit or spreading or reflexed; corolla 2–3 cm long, white or pinkish, unlobed, midpetaline bands pilose near the tips only; filaments glandular below; ovary pubescent or glabrous; style glabrous, divided c. 10 mm above the base, stigmas c. 2 mm. Capsule glabrous; seeds glabrous, smooth. [
Characterised by the lanceolate to ovate or obovate outer sepals 4–7 × 2.5–5 mm wide, these patent to erect in fruit. Leaves and ovary glabrous or hirsute.
Tajikistan (
AFGHANISTAN,
This differs from
Restricted to Afghanistan, Farah Province (
Intermediates with
ALGERIA, Djebel Babor,
Perennial herb from a rhizomatous rootstock with branched trailing stems to at least 40 cm; stem and vegetative plants adpressed pubescent or (rarely) glabrous. Leaves petiolate, 2–5 × 1.5–3 cm, ovate-deltoid, apex obtuse, margin undulate to sinuate, base truncate to very shallowly cordate; petioles 0.3–0.8(-1.5) cm. Flowers solitary, axillary, pedunculate; peduncles 4–9 cm, often bent at apex; bracteoles 1–2 mm, filiform; pedicels 5–7 mm; outer sepals 9–10 × 4–5 mm, broadly elliptic, mucronate; inner sepals slightly larger c. 11 × 6 mm, somewhat scarious; corolla 2.5–3.1 (-4.5) cm long, white with dark centre, midpetaline bands pink, pubescent near the apex, unlobed but margin slightly undulate; filaments glandular; ovary glabrous, conical; style glabrous, divided c. 12–14 mm above base; stigmas 2–3 mm, stout. Capsule and seed not known. [
Djebel Tazekka and Rif Mountains of Morocco (
Distinctive because of the long pedunculate, solitary flowers, triangular, basally more or less truncate leaves and very long style.
A distinctive variety was described from Djebel Tazekka, an isolated massif in Morocco, by
TURKEY, Constantinople,
Perennial branched undershrub with woody rootstock and lower branches, the whole plant shortly tomentose; stems very stout, 3–5 mm thick. Leaves petiolate, coriaceous, 2–5 × 1–3.5 cm, elliptic, oblong-elliptic or obovate, apex obtuse to rounded, margin entire, base broadly cuneate to rounded, slightly asymmetric; petioles 0–6 cm. Flowers solitary, axillary, pedunculate, usually arising from the middle part of the stem; peduncles 0.8–3.3 cm; bracteoles 3–5 × 1–3 mm, ovate to oblong-elliptic, acute; pedicels 5–7 mm; sepals 12–15 × 7–9 mm, ovate, obtuse or acute; corolla 3–4.5 cm, white, unlobed, undulate, midpetaline bands pilose; filaments glandular below; ovary very narrow, thinly pilose at apex; style divided c. 18 mm above base; stigmas 2 mm, narrowly elliptic. Capsule glabrous, seeds tuberculate. [
Coasts of the Black Sea: Georgia (
A very distinct, apparently isolated species.
LIBYA, Cyrenaica, “Messe a ovesti di Cirene Sfonta–Ruheina, 8 Mag 1934,”
Perennial trailing, scrambling or twining herb with puberulent, sharply 4-angled to weakly winged stems of “a considerable height”, presumably 1–2 m. Leaves petiolate, somewhat dimorphic, 2.5–7 × 2–5.5 cm, younger leaves ovate, apex acute to obtuse, margin undulate to irregularly dentate with some large teeth, base cordate and briefly cuneate with a broad right-angled sinus, minutely adpressed pubescent; mature leaves more deltoid, deeply lobed with acute segments in the lower part, strongly auriculate; petioles 0.5–2 cm, puberulent. Flowers in a terminal cluster of up to 7 at the apex of a long axillary peduncle; peduncles 8–22 cm, pubescent; bracteoles filiform to linear, 4–8 mm, pubescent; pedicels 3–8 mm, pubescent; outer sepals 8–9 × 5–6 cm, broadly obovate-elliptic, mucronate, pubescent; inner sepals 4–5 cm wide,less hirsute and with scarious margins; corolla 2.8–4.5 cm, white with purplish throat, unlobed, basal tube narrow, midpetaline bands thinly adpressed pubescent; filaments glandular; ovary pilose; style divided c. 9 mm above base; stigmas 2 or 3, 2 mm long, rather stout. Capsule and seeds not seen. [
Libya: Cyrenaica (
A remarkable species with flowers clustered at the apex of a very long peduncle. The stigmas are short and stout and commonly 3 in number.
AFGHANISTAN,
Trailing or scrambling herb, probably branched at base with stems to at least 40 cm, all vegetative parts densely pubescent to tomentose. Leaves petiolate, 2.2–3.5 × 0.7–1.5 cm, ovate-deltoid, apex acute, margin entire, base truncate; petioles 4–5 mm. Flowers up to 3 in pedunculate, axillary cymes, but often reduced to 1–2; peduncles 2.5–5 cm; bracteoles 2–3 mm, linear-lanceolate; pedicels 3–6 mm; outer sepals 9–10 × 4–4.5 mm, broadly oblong-rectangular, obtuse to truncate and mucronate, inner sepals scarious-margined and subglabrous; corolla 2.5–2.7 cm, colour unknown, unlobed, midpetaline bands thin, pilose; filaments glandular near base; ovary glabrous; style glabrous, divided c. 6 mm above base, stigmas 2 mm, relatively stout. Capsule glabrous, seeds minutely hirsute.
Afghanistan–only known from the type collected at “Seh Baba”.
This species and
AFGHANISTAN,
Perennial herb with woody rootstock from which arise many wiry, probably scrambling stems to at least 25 cm; stems striate, glabrescent. Leaves very shortly petiolate, 1–3.5 × 0.4–0.6 cm, lanceolate-deltoid, acute, margin entire to sinuate-lobed, base truncate to auriculate, shortly crisped pubescent when young, glabrescent; petioles 1–3 mm. Flowers 1–3, borne on axillary peduncles; peduncles 1–1.5 cm; bracteoles 2.5 mm, filiform; pedicels 4–10 mm; outer sepals 9–10 × 6 mm, broadly oblong-rectangular, truncate and minutely mucronate, margin scarious, purplish, pubescent and with ciliate margins, inner sepals oblong, c. 4 mm wide; corolla 2.3–3.2 cm long, pinkish, midpetaline bands pubescent; filaments glandular below; ovary glabrous, style glabrous, divided c. 7 mm above base; stigmas not seen. Capsule glabrous; seeds wrinkled. [
Afghanistan (
Although placed as distantly related in different sections by Sa’ad,
Based on
Perennial undershrub with trailing or twining stems to 4 m, becoming woody below with age, the whole plant glabrous to thinly adpressed pilose on younger parts. Leaves petiolate, 4–11 × 1.5–6 cm, ovate, acute to shortly acuminate, entire, truncate to cordate at the base, more or less reticulate veined on abaxial surface, glabrous or nearly so; petioles 1.5–4 cm. Flowers 1–6, in very lax, pedunculate, axillary cymes; peduncles 3–6 cm; bracteoles 14 × 0.5–1 mm, filiform to linear; pedicels 15–30 mm; sepals 9–15 × 4–7 mm, obovate or oblanceolate, with a triangular apiculate apex; corolla 2–2.5 cm, white, unlobed, midpetaline bands pink, pilose; filaments glandular; ovary pubescent at apex, style pubescent, divided 4 mm above the base, stigmas 4 mm. Capsule large, acute c. 6 mm long, glabrous; seeds (immature), smooth, with pale reticulation. [
Endemic to Madeira (
Very similar to
CANARY ISLANDS (lectotype LINN 218.17!, designated by
A liana or scrambling shrub to 10 m, old stems woody with brown bark, young stems villous. Leaves petiolate, 4–10 × 2–5 cm, ovate to oblong-ovate, acute, entire, cordate at base, densely villous, the veins prominent on the lower surface; petioles 1–1.5 cm. Flowers 3–7 in axillary, pedunculate cymes; peduncles c. 2.5–3.5 cm, bracteoles linear, acuminate; pedicels 5–18 mm; sepals 8 × 4 mm, elliptic-rhomboid, apiculate, pilose, the inner sepals with glabrous, membranous margins; corolla 1.8–2.2 cm long, pale blue with a white centre, unlobed, midpetaline bands pilose; filaments glandular; ovary sparsely pilose at apex; style glabrous, divided 3–4 mm above base; stigmas c. 4 mm. Capsule glabrous; seeds smooth, glabrous. [
Endemic to the Canary Islands, but common in and around laurel forest, 400–1000 m. Gran Canaria, Tenerife, La Palma, La Gomera, El Hierro (
CANARY ISLANDS, Tenerife,
A liana or scrambling shrub, the stems woody below, vegative parts glabrous to thinly pilose. Leaves petiolate, 5–7 × 0.5–2.2 cm, linear-lanceolate, acute to obtuse, entire, base rounded, glabrescent to thinly pilose, veins prominent beneath; petioles 4–12 mm. Flowers 1–3 in pedunculate axillary cymes; peduncles 1–2.5 cm, slender; bracteoles 1–3 mm, filiform; pedicels 3–7 mm; outer sepals 6 × 3 mm, broadly oblong, slightly constricted below triangular, acute, with a greenish apical portion, inner sepals ovate, slightly smaller; corolla 2 cm, whitish with pink, pilose midpetaline bands, deeply lobed with lanceolate lobes; ovary glabrous; style glabrous, divided c. 3 mm above base, stigmas 6 mm. Capsule not seen.
Endemic to the Canary Islands: Tenerife and La Gomera; 300–800 m (
CANARY ISLANDS,
Similar to
Endemic to Lanzarote in the Canary Islands at 400–600 m (
Twining perennial of unknown size, stems pubescent. Leaves petiolate, 3.5–7 × 1.2–2 cm, narrowly ovate, apiculate, entire, base cordate, weakly auriculate, veins prominent beneath, both surfaces minutely tomentellous. Flowers 2–5 in axillary, pedunculate cymes; peduncles 1.5–2.5 cm; bracteoles 7–10 × 0.5–2 mm, shortly petiolate, oblong, acuminate; pedicels 6–18 mm; outer sepals 10–13 × 3–4.5, narrowly ovate, acuminate, tomentose, inner sepals 8–10 mm long, acute; corolla 2.2–2.4 cm long, lobed, midpetaline bands pilose; ovary c. 4 mm long, densely hirsute; style pilose, divided 3–4 mm above base. Capsule and seeds not seen.
Only known from La Palma in the Canary Islands (
This appears to be a distinct species but we hesitate to describe it given the complexity of the Canary species and the fact that it is only known from a single collection. Its sepals are distinctly longer than those of both
CANARY ISLANDS, a plant grown at Paris from seed sent by Collignon from the Canary Islands (holotype P [Herb. Lam.]!).
Woody-based scrambling plant with long trailing stems to 1.5 m. Leaves shortly petiolate, 1–5 × 0.5–1.4(-2.5) cm, oblong, apiculate, entire, base subcordate, weakly auriculate, thinly to densely tomentellous on both surfaces; petioles 2–4 mm. Flowers 1–3 in shortly pedunculate, axillary cymes the flowers and leaves crowded together; peduncles 0–10 mm; bracteoles, 5–6 mm, filiform, caducous; pedicels 5–12 mm; sepals 6–7 (-9) × 2.5–3 mm, pubescent, outer sepals broadly oblong-elliptic, obtuse to subacute, inner sepals oblong, acute to mucronate; corolla 1.5–1.7 cm long, pale blue, weakly lobed, midpetaline bands pilose; filaments glandular, ovary 3.5 mm long, pilose at apex or glabrous; style glabrous except sometimes near the base, divided 3 mm above base; stigmas 3 mm. Capsule globose-conical, apiculate, c. 4–8 mm long, pilose apically or glabrous; seeds tuberculate, glabrous. [
Characterised by the obtuse to subacute outer sepals. Plants are almost always densely tomentellous and leaves usually subcordate.
Endemic to the Canary Islands: Tenerife (
Plants referred to
Based on
Best distinguished from
Endemic to the Canary Islands: Gran Canaria, La Palma, 500–1000 m.
Indumentum varies from nearly completely glabrous (
PORTUGAL,
A scrambling or twining shrub, old stems woody, glabrescent, young stems pubescent. Leaves petiolate, 1–6 × 2.5–3.5 cm, elliptic or oblong-elliptic, retuse, entire, base rounded to subcordate, puberulent on the nerves beneath but soon glabrescent; petiole 1–2.5 cm. Flowers 3–6 in axillary, pedunculate cymes; peduncles 1–2 cm; bracteoles 5–9 × 1–2.5 mm, oblanceolate; pedicels 5–10 mm; sepals 6–9 × 3–3.5 mm, obovate to elliptic, mucronate, puberulent, inner sepals scarious, puberulent near base only; corolla 1.5–1.7 cm, white, sinuate, midpetaline bands tomentose; filaments glandular near base; ovary glabrous; style glabrous, divided c. 4 mm above base, stigmas c. 2 mm. Capsule and seeds unknown. [
Portugal: Cabo Espichel. Rare, narrow endemic of dolomitic rock at 125 m.
Although seeds are clearly visible on the image of the type specimen no description has been provided.
This species is related to
SPAIN, Valencia, Alicante,
Perennial herb from a rhizomatous rootstock with decumbent stems to 40 cm long; vegetative parts appressed hairy to pilose, often with both types of hair on the same plant. Leaves shortly petiolate, 1.5–4 × 0.2–1(-2.3) cm, lanceolate, oblong or oblong-elliptic, often falcate, acute, entire, base truncate; petioles 1–6 mm. Flowers 1–2 in pedunculate axillary cymes; peduncles 1–4 cm; bracteoles 8–20 × 0.5–2 mm, linear or linear-lanceolate, pedicels 0–3 mm, very short; outer sepals 7–9 × 2.5–4 mm, oblong-ovate to ovate, acuminate, inner sepals oblong-elliptic, cuspidate with broad membranous margins; corolla 2–2.5 cm, blue, pale violet, weakly lobed, midpetaline bands darker on the exterior, adpressed-pilose; filaments glandular below; ovary conical, glabrous; style divided 4–6 mm above base, glabrous, stigmas 3.5–5 mm. Capsule glabrous; seeds glabrous, tubercled. [
SE Spain (
The unusually short pedicels and often falcate leaves serve to make this species distinct.
ITALY, Liguria,
Rather variable perennial herb from a woody rootstock, the stems sometimes short and straight and sometimes with trailing, flexuose stems at least 40 cm long; vegetative parts varying from appressed puberulent to villous. Leaves petiolate, 0.5–3 × 0.3–2.2 cm, ovate to suborbicular, rounded to obtuse, entire, base truncate to subcordate and shortly cuneate onto the petiole; petioles 1–5 mm. Flowers 1–3 in shortly pedunculate axillary dichasial cymes; peduncles 0.5–3.5 cm, commonly flexuose and sometimes recurving in fruit; bracteoles 3–13 × 0.5–2.5 mm, linear to oblong-lanceolate; pedicels 3–12 mm; outer sepals 5–7 × 2–2.5 mm, oblong-lanceolate, acute; inner sepals broader (c. 3 mm wide), thinly pubescent except ciliate margins, membranous; corolla 1.6–2 cm long, blue or violet, unlobed, midpetaline bands pilose; filaments glandular below; ovary glabrous; style glabrous, divided c. 4 mm above ovary, stigmas 3 mm. Capsule glabrous; seeds glabrous, tuberculate. [
Sepals pubescent with appressed hairs, the margins glabrous.
Italy: Liguria (
The type of
Based on
Distinguished by the prominent spreading hairs on the calyx and often also on the stem and leaves.
Morocco (
The flexuose peduncles, prominent bracteoles, essentially broadly oblong leaves and violet flowers make
ALGERIA, Oran,
Perennial herb from a deep, somewhat woody rootstock, usually branched at base with trailing herbaceous stems to c. 40 cm long, vegetative parts all (thinly to) densely lanate. Leaves shortly petiolate, 0.6–1.6 × 0.3–0.8 cm, oblong to oblong-elliptic or oblong-ovate, obtuse, acute or shortly mucronate, entire, abruptly narrowed to a truncate or cordate base; petioles 0–1 mm. Flowers 1–3 in shortly pedunculate axillary dichasial cymes, the inflorescence as a whole appearing racemose; peduncles 0.5–3 cm; bracteoles 5–9 × 0.5–1 mm, linear; pedicels 0–3(-8) mm, outer sepals 8–11 × 3 mm. oblong-elliptic, acute; inner sepals ovate, c. 4 mm wide, membranous; corolla 1.8–2.8 cm long, yellow or creamy-yellow, unlobed, midpetaline bands pilose; filaments glandular below; ovary glabrous or with a few hairs; style glabrous, divided 6–10 mm above ovary, stigmas 3–4 mm. Capsule glabrous or with a few hairs; seeds glabrous, tuberculate. [
There is much variation in indumentum and two varieties can be recognised:
Leaves thinly to densely lanate.
North African Maghreb: Libya (
Based on
Leaves glabrous or glabrescent on the upper surface.
Principally in northeastern Morocco with scattered populations in Algeria: Morocco (
Distinctive because of its lanate indumentum, small, very shortly petiolate leaves and usually elongate, raceme-like inflorescence with yellow corollas and short pedicels, the bracteoles immediately below the sepals.
A relatively well-defined group of annual species essentially Mediterranean in their distribution except
MOROCCO,
Annual herb, commonly much branched at the base, reaching c. 35 cm, stems thinly pubescent. Leaves 2–8 × 1–2.5 cm, obovate-spathulate, obtuse to acute, entire, base attenuate or, above, abruptly cuneate, clearly sessile, glabrous or with a few marginal cilia. Flowers several in a sessile terminal head; bracts 1.5–2.3 × 0.8–1.5 cm, ovate, acute, rounded at the base, thinly pilose with long white hairs, green with a palid area near the base; peduncles absent; bracteoles 12–15 × 1–3 mm, oblong, very variable in size, pilose with long white hairs; pedicels absent; outer sepals 9–10 × 2 mm, oblong, acute, densely pilose with white hairs; inner sepals similar but narrowly lanceolate; corolla 1.5–2.5 cm long, blue, unlobed, midpetaline bands pilose, terminating in a small tooth; filaments with sessile glands; ovary glabrous; style glabrous, divided c. 7 mm above base, stigmas 5 mm. Capsule glabrous, much exceeding calyx; seeds tuberculate.
Endemic to Morocco (
ITALY, Sicily (lectotype LINN 218.40!, designated by
Annual herb, commonly branched at base with prostrate to erect stems to 40 cm long, thinly pilose with brownish hairs on vegetative parts. Leaves petiolate, 1–5 × 0.4–2 cm, ovate-deltoid, acute, base cuneate to abruptly truncate, margin entire; petioles 2–5 (-11) mm. Flowers 1–3 (-4); peduncles 0.6–2.5 cm, bracteoles 2–11 × 1 mm, filiform, linear or linear-lanceolate; pedicels 0–10 mm, becoming recurved in fruit; outer sepals 5–6 × 1.5–3 mm, lanceolate, ovate to subrhomboid, acute; corolla 5–7 mm long, white or lilac, deeply lobed for c. 2 mm, midpetaline bands glabrous; filaments glandular below; ovary glabrous; style glabrous, divided c. 2 mm above base, stigmas 2 mm. Capsule glabrous; seeds glabrous, tuberculate. [
We recognise two subspecies, which intergrade occasionally (e.g.
Bracteoles narrowly lanceolate, but variable in size: 15 × 3.5 mm in the Linnean type but usually much smaller to 4 × 1 mm; pedicels very short (0–1 mm) so bracteoles adjacent to sepals.
European Mediterranean, main Mediterranean islands and Western Middle East from Turkey south to Egypt, Jordan and northern Saudi Arabia: Portugal; Spain (
Based on
Bracteoles filiform, pedicels 5–10 mm long so bracteoles distant from sepals. [
Principally in NW Africa and the southern Red Sea area: Canary Islands (
As a species
Some publications (e.g.
LIBYA,
This hybrid differs from
Found once in Libya (Cyrenaica).
This taxon represents
Without locality,
Annual herb with slender rootstock, often branched at base; stems adpressed pubescent. Lower leaves c. 3–6 × 0.7–1.2 cm; spathulate with an attenuate, pseudopetiolar base, apex obtuse, margin entire, nearly glabrous but more or less ciliate on the margins; upper stem leaves and bracts clasping, 2–4 (-6) × 0.4–0.8 cm, lanceolate (to oblanceolate). Flowers solitary, pedunculate, axillary; peduncles 5–18 mm, pubescent, becoming recurved in fruit; bracteoles 1–3 mm, filiform to narrowly lanceolate; pedicels 3–8 mm, pubescent; sepals somewhat scarious, 5 × 2.5 mm, ovate, acute and mucronate, glabrous apart from long basal trichomes; corolla 0.7–0.9 mm long, blue, shallowly lobed, midpetaline bands pubescent with brown hairs; filaments sparsely glandular below; ovary glabrous or with a few very long trichomes; style glabrous, divided 3–4 mm above the base, stigmas c. 2 mm. Capsule glabrous, strongly exserted from the sepals; seeds covered in pointed tubercles. [
Circum-mediterranean, east to Iraq: Portugal (
Similar morphologically to
PORTUGAL, Coimbra,
Annual herb, commonly branched at the base, reaching c. 40 cm, stems and leaves with long, stiff spreading hairs. Basal leaves 2.5–4.5 × 0.7–1 cm, obovate-spathulate, obtuse, entire, base attenuate into a pseudopetiole, stem leaves 1.5–5.5 × 0.2–1(-1.8) cm, oblong or lanceolate, sessile. Flowers solitary, axillary, pedunculate; bracts lanceolate, acute; peduncles 1.5–5 cm, slender, flexuose, becoming recurved in fruit; bracteoles 0.5 mm, triangular, acute; pedicels 2–4 mm, not well differentiated from the peduncles; sepals 5–6 × 2–2.5 mm, ovate, acute to apiculate, margin, scarious, ciliate or glabrous, sepals otherwise more or less glabrous; corolla 1.5–2.4 cm long, tricoloured blue, white and yellow, weakly 5-angled with apices of lobes pointed, midpetaline bands adpressed pilose; ovary glabrous; style glabrous, divided c. 5 mm above base, stigmas 2.5–3 mm. Capsule glabrous, much exceeding calyx; seeds tuberculate. [
West Mediterranean region, perhaps only in Europe: Algeria (?); Morocco (?); Portugal (
Somewhat resembling
Herb. Clifford 68,
Annual herb, commonly branched at the base, reaching c. 40 cm, stems and leaves with long, stiff spreading hairs mixed with short, appressed hairs. Lower leaves 2.5–4.5 × 0.7–1.4 cm, obovate-spathulate, obtuse or emarginate, entire, base tapering, stem leaves 1.5–4 (-5) × 0.2–1(-1.8) cm, oblong, obovate or oblanceolate, sessile. Flowers solitary, axillary, pedunculate; bracts lanceolate, acute, resembling upper leaves; peduncles 1–4 cm, slender, flexuose, becoming recurved in fruit; bracteoles 2–3 mm, filiform, acute; pedicels 3–7 mm, not well differentiated from the peduncles; sepals 5.5–7 × 2.5–3 mm, broadly oblong to pandurate, scarious-margined, pilose, clearly differentiated into two parts, the upper green, acute to apiculate, the lower part colourless; corolla 2–3 (-3.5) cm long, tricoloured blue, white and yellow, weakly 5-angled with apices of lobes pointed, midpetaline bands adpressed pilose; filaments glandular below, ovary pilose; style glabrous, divided c. 5–6 mm above base, stigmas 5–6 mm. Capsule pilose, much exceeding calyx; seeds tuberculate. [
We recognise two subspecies whose ranges overlap in North Africa:
The upper, green part of the sepals is acute and shorter than the lower colourless part.
Mostly western Mediterranean: Spain (
ITALY, Sicily, “in campis argillosis–Palermo,”
The upper green part of the sepals is acuminate and longer than the lower colourless part.
Central Mediteranean: Malta; Sicily; Morocco (
Todaro never provided a description for
Plant of unknown origin, cultivated in Vienna by Jacquin (not found at W).
Annual herb, commonly branched at the base with decumbent or ascending stems 5–25 cm long, vegetative parts glabrescent or shortly pubescent. Leaves 0.5–5 × 0.3–1.5 cm, oblong-oblanceolate, apex obtuse to rounded, margin entire, basal leaves gradually narrowed into a long petiole-like base up to 2 cm in length (so appearing spathulate), stem leaves abruptly narrowed at base, sessile and sometimes clasping and auriculate. Flowers solitary, subsessile in the axils of the upper leaves, becoming crowded towards the apex; peduncles and pedicels not clearly differentiated, 0–1 mm long; bracteoles filiform, minute; sepals 2.5–3.5 × 1.5–3 mm, narrowly elliptic to obovate, acute to rounded, thinly pilose; corolla 1–1.1 cm long, blue with a pale tube, distinctly lobed with triangular lobes, midpetaline bands pubescent, darkish; ovary long-pilose, style divided 1–1.5 mm above base, glabrous, stigmas 2 mm. Capsule pilose with stiff coarse hairs; seeds strongly tuberculate. [
Nearly circum-mediterranean but apparently absent from France, Turkey and the Balkans: Spain (
Similar morphologically to
Based on
Annual with adventious root; stems 5–25 cm long from central rootstock, decumbent, indumentum of scattered long hairs mixed with some shorter pubescence. Leaves 1–5 × 0.2–0.6 cm, oblong-oblanceolate; apex rounded; base attenuate, petiole-like, margin entire. Flowers solitary, axilary; peduncles 0.8–1.6 cm, often becoming recurved in fruit; bracteoles 3–6 (-9) × 1–2 mm, oblanceolate; pedicels 1–3 mm; outer sepals 5 × 1 mm, lanceolate-oblong, acute to apiculate, differing from the oblong-obovate inner sepals; corolla 5–6 mm long, deeply lobed, pale blue with white midpetaline bands radiating from the centre; midpetaline bands glabrous, terminating in a mucro; filaments eglandular; ovary glabrous; style glabrous, divided 1–1.5 mm above base, stigmas 1 mm. Capsule glabrous; seeds tuberculate.
United States of America: California (
Although separated geographically from related species,
Vegetatively extremely variable including annual and perennial herbs, spiny and unarmed undershrubs and some species with fastigiate branching. The outstanding feature of many (but not all) species lies in the structure of the stigmas. In many species in this clade (Clade A in Figure
SUDAN, Kordofan [Kurdufan],
Annual herb, often branched at base, pubescent in all vegetative parts, stems prostrate to ascending, up to 60 cm long. Leaves shortly petiolate, 0.5–4.5 × 0.1–1.3 cm. ovate, lanceolate, elliptic or oblanceolate, obtuse or acute, margin entire, base cuneate, rarely truncate; petioles 2 mm long. Flowers 3–6 in subsessile, axillary, bracteate heads; peduncles 0–0.5(-3) cm long; bracteoles 4–8 × 2–3 mm, lanceolate or ovate; outer sepals 4–7 × 0.5–1.5 mm, lanceolate, acute, densely villous; corolla 4–5 mm long, white or pale pink, deeply lobed, the lobes longer than broad, 2–2.5 mm long, more or less hidden by calyx hairs, midpetaline bands glabrous, ovary glabrous; style glabrous, divided 2–2.5 mm above base, stigmas c. 1.5 mm long; capsule glabrous, seeds tuberculate or (
A Saharo-Sindian species, unexpectedly absent from southern Arabia and the Gulf region: India (
The glabrous midpetaline bands of this, the previous and the following species are unusual.
SOMALIA,
Annual or briefly perennial herb with a small tap root, similar to
This species is quite variable, approaching
The type variety has small leaves and bracts 0.5–1.5 cm long, equalling or only slightly exceeding the flower heads. The corollas are only 7–8 mm long.
Apparently restricted to Somalia and Djibouti, where it is the common variety (
Based on
Leaves and bracts 3–6 cm long giving the inflorescence a leafy appearance. Corolla 9–12 mm long.
This seems to be the only subspecies in Kenya (
Distinguished from
KENYA,
Perennial herb with prostrate or twining stems to 50 cm, vegetative parts adpressed pilose, when young with silvery hairs. Leaves shortly petiolate, 1.5–4 × 0.35–1(-2) cm, oblong, ovate or elliptic, apex rounded to acute, base cuneate to subhastate, margin entire; petioles 1–4 mm long. Flowers 2–5 in pedunculate, bracteate heads; peduncles 1–4.5 cm; bracteoles 6–14 × 2–8 mm, lanceolate or oblong; outer sepals 5–9 × 1–4 mm, elliptic to obovate, villous; corolla 8–10 mm long, blue, unlobed, midpetaline bands pubescent; ovary glabrous; style glabrous, divided 5 mm above base, stigmas 3–3.5 mm long, linear. Capsule glabrous; seeds glabrous, tuberculate. [
East Africa, coastal sand, coral outcrops and grassland near the sea: Tanzania (
This species is not so distinct from
SOMALIA,
Perennial herb with prostrate stems to 50 cm, vegetative parts appressed pilose. Leaves shortly petiolate, 0.7–3 × 0.1–0.3 cm, linear to narrowly oblong, acute, base cuneate, margin entire; petioles 0.5–1.5 mm long. Flowers 1 (– 2) in pedunculate, axillary, bracteolate heads; peduncles 12–30 mm long, often flexuose or reflexed; bracteoles 4–7 × 1.25–1.5 mm, linear to lanceolate, appressed to sepals; pedicels absent; outer sepals 5–8 × 2–3 mm, lanceolate to ovate; corolla 12 mm long, blue, unlobed, midpetaline bands pubescent; ovary glabrous; style glabrous, divided c. 2 mm above base, stigmas 2.5 mm, linear. Capsule glabrous; seeds glabrous, sinuate-ridged. [
Somalia (
Easily recognised by its flexuose peduncles, one-flowered heads and bracteoles appressed to the sepals.
Two specimens (
ETHIOPIA,
Perennial herb with numerous prostrate stems to 50 cm from a central woody rootstock, vegetative parts finely adpressed pubescent. Leaves shortly petiolate, 1–3.7 × 0.1–0.5 cm, linear to narrowly oblong, apex apiculate, margin entire, base narrowly cuneate; petioles 1–2 mm. Flowers 1–3, subsessile in shortly pedunculate, axillary, bracteate heads; peduncles 2–5 mm long; bracteoles 7–11 × 1–2 mm, narrowly oblong, glabrous; pedicels 0–1.5 mm; outer sepals 8–10 × 1.5–2 mm, narrowly elliptic, acute; corolla 8–10 mm long, blue with a whitish centre, unlobed, weakly crenate, midpetaline bands thinly pilose; ovary and style glabrous, style divided 4–5 mm above base, stigmas 2 mm, linear. Capsule glabrous; seeds glabrous, tuberculate. [
Endemic to Ethiopia (
Perhaps most similar to
ETHIOPIA,
Perennial herb with prostrate stems to 40 cm, vegetative parts covered in adpressed silvery hairs. Leaves shortly petiolate, 1–2.5 × 0.3–0.6 cm, linear-oblong, acute, margin entire, base more or less truncate; petioles 1–1.5 mm. Flowers 1–2, subsessile in shortly pedunculate, axillary bracteolate heads; peduncles 0–2.5(-5) mm, bracteoles 6–9 × 1.5–2 mm, lanceolate; outer sepals 8–9 × 2.5–4 mm, ovate, shortly acuminate; corolla 9–12 mm, blue, very shallowly lobed, midpetaline bands pilose; ovary glabrous, style glabrous, divided c. 3 mm above base; stigmas c. 2 mm, linear. Capsule glabrous; seeds glabrous, tuberculate. [
Ethiopia (
Similar to
SOMALIA,
Perennial herb with prostrate or twining stems to 50 cm long, vegetative parts densely sericeous becoming golden-brown when old. Leaves shortly petiolate, 0.6–2 × 0.6–2 cm, suborbicular, apex rounded or, rarely, emarginate or obtuse, base cordate, margin entire; petioles 1–6 mm long. Flowers 2–5 in shortly pedunculate, bracteate heads; peduncles 0.4–1.5 cm; bracteoles 7–10 × 5–7 mm, oblong-elliptic to ovate; outer sepals 5–7 × 2–3 mm, oblong to obovate, villous; corolla 9–11 mm long, blue, unlobed, midpetaline bands pilose; ovary glabrous; style glabrous, divided 3 mm above base; stigmas 1.75–2 mm, narrowly cylindrical and thicker than style. Capsule glabrous; seeds glabrous, tuberculate. [
Endemic to Somalia (
IRAN,
Undershrub forming a small bush up to 40 cm high and 60 cm wide; stems from a deep woody taproot, many, ascending, rigid, green, glabrous, sometimes spinescent at the tips, weakly divaricate. Leaves sessile, 1.7–2.5 × 0.2–0.4 cm, lanceolate or linear-lanceolate, glabrous, acute or acuminate, margin entire, base truncate to obscurely auriculate. Inflorescence of few-flowered, axillary, pedunculate, hirsute heads; peduncles 1.5–4.5(-8) cm, rigid, woody; bracteoles 5–12 × 1–2 mm, very variable in size, linear or lanceolate, acute; pedicels 1–2 mm, often bent at a sharp angle to the peduncle; sepals 8–9 × 2–2.5 mm, ovate, acuminate, villous; corolla 1.3–2.1 cm, usually white, sometimes pinkish, shallowly lobed, the midpetaline bands ending in teeth, pubescent, sometimes darker pink; ovary glabrous; style glabrous, divided c. 5.5 mm above the base, stigmas 1–1.25 mm, elliptic. Capsule glabrous; 1–2-seeded; seeds tuberculate, glabrous. [
We recognise two varieties:
Plant lax in habit, the branches neither very numerous nor markedly spinescent; leaves all or mostly > 2 cm long; corolla > 1.5 cm long.
Locally frequent in desert in southern Gulf region: U.A.E. (
Based on
Differs from the type in being very compact with short, intricately branched. spinescent shoots, relatively short, broad leaves, small heads and short corollas. Possibly an adaptation to extreme arid conditions.
Oman (
SAUDI ARABIA, Jiddah,
Perennial herb with prostrate, ascending or twining stems to 1 m, rootstock thick and somewhat woody, pubescent on vegetative parts; stems sometimes slightly woody. Leaves shortly petiolate, 1–4.5 × 0.5–1.5 cm, lanceolate or ovate, acute, base truncate to cordate, margin entire; petioles 0–5 mm long. Flowers 4–10 in axillary, pedunculate, villous, bracteate heads formed of compact scorpioid cymes; peduncles 1–4 (-7) cm long, straight or recuvedbracteoles 8–12 (-28) × 2.5–5 (-9) mm, ovate, acuminate, villous; outer sepals 8–12 × 3.5–4.5 mm, ovate, long-acuminate, broader than the inner sepals; corolla 8–12 mm long, white to pale blue, undulate, midpetaline bands pilose; ovary glabrous; style glabrous, divided c. 4 mm above base, stigmas ellipsoid, 1.5–2 mm. Capsule glabrous, seeds glabrous, tuberculate. [
We recognise two varieties:
Corolla 8–12 mm long; sepals usually < 10 m long; pedunces usually straight.
A characteristic Saharo-Sindian species: Niger (
A subsp. typo corolla 15–20 mm longa (non < 12 mm longa) et usque 2.5 cm diametro dignoscenda.
OMAN, Dhofar, Cliff near Dalkut,
The key feature of this variety is the very large corolla 1.5–2 cm long when dry and up to 2.5 cm in diameter when living. The leaves are always relatively large, up to 4 × 2.5 cm, the peduncles long and commonly gently curved, reaching 7 cm in length, and the sepals may reach 12 mm. However, all characters apart from the corolla size can be matched in other populations of
Oman (Dhofar):
There has been some uncertainty about the correct name for this species.
OMAN,
Low undershrub from a woody base reaching c. 30 cm; stems densely tomentellous with white, spreading hairs, becoming stiff and rather woody when old and said to form a candelabra-shaped plant. Leaves alternate below but mostly opposite upwards, subsessile, 5–13 × 2–5 mm, elliptic or oblong-elliptic, apex acute to rounded, margin entire, base cuneate to truncate, glandular and hirsute with short spreading hairs on both surfaces; petioles 0–1 mm. Flowers 1–2 in small sessile, axillary clusters near the branch tips; peduncles and pedicels absent or nearly so; bracteoles 6 × 2 mm, oblong, acute; outer sepals 7–8 × 1.8–2 mm, lanceolate, acute, green near apex but membranous below, sericeous; inner sepals narrower (1.5–1.8 mm wide) with membranous margin; corolla 11–13 mm long, pinkish or “pale blue”, midpetaline bands villous; ovary glabrous; style glabrous, divided c. 6 mm above base, stigmas short, clavate. Capsule glabrous, seeds minutely rugulose.
Endemic to Oman (
Very distinct because of the short clavate stigmas and small, tomentose leaves arranged oppositely towards the branch tips. Some plants have very reduced leaves giving the plant a somewhat different facies (e.g.,
SOMALIA,
Slender virgate unarmed shrub of fastigiate habit to 1 m high; young stems glabrous to thinly adpressed pubescent, bluish-grey, rigid and woody when mature. Leaves sessile, usually erect, 1.5–4 × 0.5 mm, linear to narrowly elliptic, acute, glabrous or minutely pubescent. Flowers 1–5 (- 8) in subsessile, bracteate, silky villous heads; peduncles 0.5–3 mm long, bracteoles 3–7 × 1–2 mm, linear-lanceolate, acute, villous; outer sepals 7–9 × 1–1.5 mm, lanceolate to oblong -oblanceolate, acuminate, villous, inner sepals 5–7 mm; corolla 10–12 mm long, pale blue, unlobed, midpetaline bands pilose; ovary glabrous; stigma narrowly elliptic. Capsule glabrous; seeds 1, glabrous, smooth. [
Northern Somalia (
The type collection virtually lacks leaves. The Somalia plants and
Based on
Subshrub, much branched with entangled spinescent branchlets to 1 (- 2) m, vegetative parts glabrous, sericeous, pubescent or densely pilose. Leaves subsessile, 4–10 (-15) × 1–3 (-5) mm, narrowly oblong, elliptic or ovate, acute, base broadly cuneate, truncate to auriculate, margin entire. Flowers 1–6 in subsessile, elliptic to suborbicular axillary clusters; peduncles absent; bracteoles 3–7 × 2–5 mm, narrowly elliptic to obovate; pedicels 0–1 mm; outer sepals 7–9 × 6–8 mm, broadly ovate or elliptic, acuminate, wider than inner ones; corolla 0.8–1.4 cm long, pale to dark violet, weakly lobed, midpetaline bands pubescent, brown; ovary glabrous; style glabrous, divided 6–7 mm above base, arms unequal c. 1–1.5 mm, stigmas short and narrowly elliptic, c. 1 mm. Capsule glabrous; seeds glabrous.
Within Arabia and along the Red Sea this species is reasonably constant although there is variation in whether hairs are appressed or spreading and in the number of flowers in each head, but much of this variation seems random geographically. In Somalia variation is much greater and three subspecies are here recognised:
Plant spiny with numerous short spine-like lateral branchlets arising on the main spinescent branches.Stems and leaves pubescent to pilose. Flowers in clusters of up to 6 forming a suborbicular head; bracteoles obovate, about as broad as long, corolla 1.2–1.4 cm long.
Red Sea coastal regions of Egypt (
The type of
Based on
Similar to
Ogaden region of Ethiopia (
Some specimens from Somalia (
Based on
Plant virtually unarmed, the side branches long, relatively slender, spinescent when old, short lateral branchlets absent or, when present, neither rigid nor spine-like. Indumentum and flower heads similar to
Bari and Nogol regions of NE Somalia (
Occasional specimens intermediate with subspecies
Two other specimens from NE Somalia (
Based on
Fastigiate undershrub reaching 60 cm in height but more in width, silvery-sericeous when young with most hairs appressed but some spreading, glabrescent and becoming brown when old; branches rigid but not spinescent. Leaves sessile, 2–6 × 0.5–1 (-2) mm, linear-lanceolate, acute, entire, cuneate at the base, sericeous. Flowers solitary, axillary, sessile; bracts resembling small leaves; bracteoles 1.5 × 0.5 mm, ovate, acute; sepals 5–7 × 2–3 mm, ovate to elliptic, concave, the apex acute, bent outwards, pubescent; corolla 5.5–7 mm long, white, midpetaline bands pilose; filaments glandular below; ovary glabrous; style glabrous, divided 3–4 mm above base, arms somewhat unequal 0.5–1 mm long, stigmas 0.5 mm, obovoid. Capsule 1-seeded, glabrous, seeds smooth, glabrous.
Endemic to Socotra (
Based on
Undershrub to 60 cm, branches fastigiate, sharply spinescent, sericeous when young but later glabrescent. Leaves subsessile, 4–12 × 1–1.5 mm, acute, entire, cuneate at the base, silvery pubescent. Flowers solitary, axillary, subsessile; peduncles absent; bracteoles 2–3 × 0.5 mm, oblong, acute; pedicels c. 1 mm; outer sepals 6 × 3 mm, ovate to elliptic, acute, pubescent on the margins; inner sepals c. 4 mm wide, subrhomboid with hyaline margins; corolla 10–11 mm long, white, midpetaline bands pubescent; filaments glandular basally; ovary glabrous, style glabrous, divided c. 8 mm above base; style arms 0 and 0.5 mm, unequal, stigmas subglobose. Capsule and seeds not known.
Endemic to Abd al Kuri Island in the Socotra group (
Based on
Grey undershrub to 50 cm, branches straight and rigid, densely grey canescent when young, becoming more sparsely hirsute with age and reddish-brown in colour. Leaves shortly petiolate, 2.5–8 × 0.5–1.5 mm, narrowly lanceolate, subacute, cuneate at base, densely grey-sericeous; petioles 0.5–1 mm. Flowers solitary, axillary, subsessile; peduncle absent; bracteoles 4–5 × 1.5 mm, narrowly oblong-elliptic, densely pilose; pedicels 0.5 mm; outer sepals 6–6.5 × 1.5 mm, oblong, acuminate; inner sepals shorter, scarious; corolla 7–8.5 mm long, white faintly flushed pink, midpetaline bands long-sericeous to base; filaments glandular at base; ovary pilose; style divided 4 mm above base, stigmas 0.75 mm, obovoid. Capsule pilose at apex; seeds glabrous.
Endemic to Semha Island in the Socotra islands of Yemen.
All species in this group (Clade I of Figure
SOMALIA,
Spiny undershrub, appressed pubescent/strigose in its vegetative parts, branches spinescent; sterile spines also present. Leaves sessile, 10–35 × 1–2.5 mm, linear to narrowly lanceolate, obtuse, cuneate at the base. Flowers borne on usually paired spinescent side branches 1.5–3 cm long, the individual flowers solitary or paired arising in the axils of minute bracteoles; bracteoles 0.5 × 0.3 mm, oblong, caducous; pedicels 1–2 mm long; outer sepals 3–4.5 × 1.5–2 mm, oblong to lanceolate, apex obtuse and apiculate; corolla 8 –11 mm long, white, sometimes flushed pink or pale blue, very shallowly lobed, midpetaline bands pubescent; ovary glabrous; style glabrous, divided 1.5–3.5 mm above base; stigmas 3.5–4.5 mm, linear. Capsule glabrous; seeds shortly pubescent, smooth.
Endemic to northern Somalia (
Superficially rather similar to
ALGERIA, Oran,
Intricate, nearly leafless, spiny undershrub to 50 cm, stems and branches spinescent, branches arising at right angles, stems and vegetative parts glabrous to strongly adpressed pubescent. Leaves alternate on younger shoots but commonly clustered on very short thick brachyblasts on older shoots, sessile, 0.4–2.2 × 0.1–0.4 cm, oblanceolate, obtuse, entire, attenuate at the base. Flowers solitary (very rarely paired), axillary but sometimes appearing to be in clusters on brachyblasts from which the leaves have fallen; peduncles 0–6 mm, woody, persistent and spinescent; bracteoles c. 1 mm long, scale-like; pedicels 1–3.5 mm, commonly recurved; calyx somewhat globose, sepals similar, 4–5 × 2.5–3 mm, elliptic, rounded and minutely mucronulate, pubescent; corolla 1.4–1.6 cm, white, very shallowly lobed, midpetaline bands pilose, pink; ovary very sparsely pilose; style sparsely pilose below, divided 5 mm above base, stigmas 4–6 mm, somewhat unequal. Capsule and seeds not seen. [
Restricted to the Maghreb of northwestern Africa: Mauritania (
A rather variable plant, sometimes glabrous, sometimes densely adpressed pubescent so approaching
INDIA,
Annual herb from a thin tap root, commonly branched at base with erect or ascending stems, 10–45 cm high, vegetative parts adpressed pilose, occasionally with some stiff spreading hairs. Leaves mostly on lower part of stem, sessile, 1.5–3.5 (-7.5) × 0.2–0.6 (-1) cm, lanceolate, oblong or narrowly oblong-lanceolate, mucronate, entire, gradually narrowed to a pseudopetiolate base. Flowers 1–3 in axillary pedunculate cymes; peduncles 1–5 cm, ascending; bracteoles 2–5 × 0.5 mm, fiiform to linear-lanceolate; pedicels 2–5 mm; outer sepals 4–6 × 2–2.5 mm, ovate to elliptic, acuminate and mucronate, glabrous or adpressed pilose; inner sepals slightly smaller, glabrous; corolla 0.7–1.2 cm long, pink, distinctly lobed, midpetaline bands pilose; ovary glabrous; style glabrous, divided 2–4 mm above base, stigmas 3 mm long. Capsule glabrous; seeds pubescent with patches of adpressed hairs.
We recognise two subspecies:
Outer sepals adpressed pilose. [
India, (
Based on
Outer sepals glabrous.
Endemic to Pakistan: Balochistan and Sind (
YEMEN, Mor,
Very variable perennial herb with ascending or prostrate stems up to 70 cm long, vegetative parts thinly to densely pubescent or villous, stems often somewhat rigid and woody when old. Leaves subsessile, 0.8–3 × 0.2–0.6 cm, oblong, linear, lanceolate, narrowly lanceolate or oblanceolate, acute or obtuse, margin entire, cuneate, the lowermost tapered at the base. Flowers 1–3 (-11) in sessile or pedunculate bracteate clusters, these occasionally rather lax with individual flowers clearly separate; bracts leaf-like, exceeding, equalling or shorter than the flowers; peduncles 0–5 (-10) cm long; bracteoles 3–7 × 1–2 mm, filiform, linear to lanceolate or narrowly oblong, sometimes caducous; pedicels 0–3 (-6) mm long; outer sepals 4–8 × 1.5–6 mm, lanceolate to ovate, acuminate, green in the upper half, paler in the lower half; inner sepals caudate; corolla (6-)10–15 mm long, white or pale pink, unlobed, midpetaline bands pilose; ovary glabrous; style glabrous, divided 3–4 mm above the base, stigmas c. 4 mm. Capsule glabrous, seeds smooth, pubescent. [
A characteristic Saharo-Sindian species, apparently absent from Palestine/Israel, Syria, most of Iraq and countries north of Iran: Cape Verde Islands (
In the types of
In India, Pakistan and Afghanistan plants similar to (i) occur commonly but a second form corresponding to the type of
In Arabia and rather rarely elsewhere there occur plants which accord with the types of
Occasional plants with spinescent branches occur. These appear to be restricted to Arabia and include
Plants with very woody stems which could be treated as shrubs occur occasionally, particularly in Oman. Good examples are
Dwarf forms with very small leaves occur on Bahrain (
There is much variation in the number of flowers in each cluster but occasional plants occur where the flowers are solitary. Examples include
No attempt has been made here to give formal names to any of this variation but it is very clear that the
“Levant”,
Perennial herb with a woody taproot, stems decumbent or ascending to 70 cm, plant usually thinly adpressed pubescent but sometimes villous. Lower leaves with petioles to 5 cm, narrowly oblanceolate, obtuse, stem leaves sessile, 1.5–9 × 2–1.5 cm, linear, lanceolate or oblong, acute, margin entire or wavy, attenuate at base. Flowers pedunculate, sometimes solitary but mostly clearly separated and arranged in somewhat congested, pedunculate monchasial or diachasial cymes; peduncles (1-) 3–5 (-10) cm. much exceeding the subtending bracts; bracteoles 3–6 × 1–2 mm, linear or narrowly oblong; pedicels 1.5–6 mm; sepals with green herbaceous acute, upper part, outer sepals 5–8 × 3–5 mm, oblong-oblanceolate, acute, inner sepals ovate, caudate, corolla 10–15 mm, pink or white, unlobed (or, rarely, divided to base), midpetaline bands pubescent; ovary and style glabrous; style divided 3–4 mm above base, stigmas c. 4 mm; capsule glabrous, seeds smooth, shortly pubescent. [
We recognise two varieties of this species:
Leaves lanceolate or oblong, at least 2 cm wide. Inflorescence usually of several flowers.
Middle East and Central Asia but apparently absent from the Indian subcontinent and Africa, apart from Egypt and Libya; in Arabia rare and absent from the south: Libya (
IRAN, Bushehr,
This is a delicate form with linear-lanceolate leaves to 2 cm wide and mostly solitary flowers.
The occasional specimen of this variety occurs throughout most of the range of
IRAN,
Erect perennial undershrub with a woody base and glabrous, herbaceous branches to at least 1 m, nearly leafless above. Leaves sessile, somewhat dimorphic, basal leaves 2–3.5 × 0.5–1 cm, spathulate to oblanceolate with a long petiole-like base; stem leaves 2.5–3 × 0.2–2 cm, linear to lanceolate, acute, entire, base cuneate. Flowers in a large, nearly leafless, branched terminal inflorescence composed of axillary cymes of 1–6-flowered cymes, the flowers irregularly grouped; ultimate branches rather slender, not rigid; bracts 7 × 1 mm, linear, bracteoles minute; pedicels 0–10 mm, densely pubescent; sepals 2–4 × 1.5–4 mm, ovate to suborbicular, mucronulate, pubescent, the inner sepals broader, scarious; corolla 1.3 cm long, white, midpetaline bands pilose; ovary pilose; style glabrous, divided c. 3 mm above base; stigmas 3 mm. Capsule pubescent at apex, 1-seeded; seeds glabrous, black, smooth.
We recognise two varieties of which only
Sepals pubescent.
Turkey (
IRAN, Tehran,
Distinguished by the glabrous sepals
Apparently very rare and reported only from Iran.
YEMEN, Aden,
Slender, erect branched shrub 0.5–3 m high, vegetative parts characteristically sericeous to strigose. Leaves subsessile, 4–13(-40) × 1–5 (-10) mm, filiform, oblong to obovate, acute. Inflorescence with the appearance of long narrow branched racemes; flowers in dichasial cymes commonly reduced to clusters of 1 –2(-3), arising from the axils of linear bracts; peduncles 0–3.5 cm long; bracteoles linear 1–5 mm long; pedicels 2–3 mm long; outer sepals 2.5–3.5 × 1.5–2 mm, ovate, acute, glabrous to sericeous; corolla 6–10 mm long, white, weakly lobed, midpetaline bands strigose, red; ovary glabrous, style glabrous, divided 3 mm above base, stigmas 1 mm. Capsule glabrous; seeds pubescent. [
Yemen: south (
Specimens collected after good rainfall have a rather different morphology with well-developed, obovate lower leaves,
SOCOTRA, Abd al Kuri Island,
Herb with thickened woody taproot and a basal rosette; stems ascending to c. 30 cm, vegetative parts sericeous and with some spreading hairs. Leaves petiolate, dimorphic; rosette leaves 3.5–4.5 × 0.7–1.4 cm. obovate to oblanceolate-spathulate, obtuse to acute, margin entire to undulate, base attenuate onto the petiole; stem leaves 0.3–1.5(-3) × 0.2–1 cm, acute, oblanceolate, oblong or elliptic, margin deeply sinuate, base attenuate; petioles 2–5 mm long. Flowers solitary, pedunculate; peduncles c. 3 mm long; bracteoles 2 × 0.75 mm, linear; pedicels c. 3.5 mm long; outer sepals 4–4.5 × 2–2.5 mm, ovate, acute or shortly acuminate; corolla 10–12 mm long, white or pinkish, lobed with broadly triangular lobes, midpetaline bands pilose; ovary glabrous; style glabrous, divided c. 2.5–3 mm above base; stigmas 4 mm long. Capsule glabrous; seeds glabrous. [
Endemic to Abd al Kuri Island in the Socotra archipelago (
Very distinct because of the rosette habit and small, sinuately-lobed stem leaves. The corolla is slightly larger than in
SOCOTRA,
Perennial herb with numerous decumbent stems to 30 cm long, vegetative parts densely greyish strigose-pubescent when young but indumentum much sparser on older parts. Leaves forming a somewhat persistent basal rosette, sessile, 0.8–2.5 × 0.2–0.8 cm, obovate to oblanceolate, apex obtuse, margin entire, at base attenuate into a pseudopetiole, densely silvery strigose; stem leaves narrower upwards and merging with bracts, thinly strigose. Inflorescence a raceme-like thyrse formed of single-flowered cymes; peduncles 2–10(-16) mm long; bracteoles 1–3 mm long, filiform to narrowly elliptic; pedicels 3–5 mm long; outer sepals 2.5–4 × 1.5–2 mm, ovate, acute, green with scarious margin; corolla (7-)9–11 mm long, white, unlobed, midpetaline bands strigose; ovary glabrous; style white, glabrous, divided c. 2 mm above base, stigmas 2.5 mm, red. [
Yemen: Socotra (
The sericeous basal leaves separate
SOMALIA, Wodderi,
Perennial herb, base woody, stems erect, to 70 cm, more or less herbaceous to somewhat woody and fastiagiate, pilose below with hairs up to 3 mm long, becoming strigose upwards, glabrescent when old and woody. Leaves crowded towards the base of the stem, diminishing in size upwards, 1–4 × 0.2–0.4 cm, oblanceolate, acute, gradually narrowed into a long petiole-like base, long-pilose, often absent; stem leaves 1–2.8 × 0.1–0.2 cm, diminishing in size upwards, all linear to very narrowly linear-oblanceolate, apparently caducous. Inflorescence a large, lax terminal thyrse up to 30 cm long, formed of dichasial cymes bearing up to 3 flowers each but commonly reduced to single flowers; bracts on main axis 5–7 mm, linear; peduncles 0.6–6 cm long, becoming shorter on the secondary and tertiary flowers, often single-flowered; bracteoles 1 mm, linear; pedicels 2–4 mm long; outer sepals 2.5–3.5 × 1.5 mm, ovate, acuminate, somehat scarious but with green midrib, inner sepals narrower, strigose; corolla 8–9 mm long, white, weakly lobed, midpetaline bands strigose; ovary 2 mm long, glabrous (
Somalia (type only); Socotra (
The capsule is only 2.5 mm wide, not 5 mm as described by
We agree with
Sepalis ovatis acuminatis 2.5–4 mm longis
OMAN, Breik Qotait, Mussandan Peninsula, 30 March 1980,
Small unarmed undershrub c. 15 cm high, stems rigid, minutely adpressed pubscent. Leaves almost completely absent, probably lanceolate, certainly acute, entire, thinly pubescent. Inflorescence elongate, formed of pedunculate dichasial cymes arising in the axils of bracts at right angles to the stem; bracts 2–10 × 0.5–2 mm, linear to narrowly oblong, acute, entire, cuneate at base, thinly appressed pubescent; peduncles 1.5–3 cm long, straight, rigid, appressed pubescent; bracteoles 1–2 × 0.5 mm, linear, pubescent; pedicels c. 5–15 mm; flowers almost always solitary; sepals 3.5–4 × 1.5 mm, ovate, acuminate, pubescent with spreading hairs, inner sepals similar but somewhat scarious-margined; corolla c. 10 mm long, of unknown colour and shape, midpetaline bands pubescent; filaments c. 1 mm; anthers c. 1.5 mm; ovary c. 1.25 mm, acuminate, glabrous; style glabrous, divided c. 2 mm above base, stigmas c. 2 mm.
Known only from the type collection made on the Musandam Peninsula in Oman.
This species is very similar morphogically to
This species is only known from a single collection. It may be rare and threatened or simply overlooked but for the time being should be classified as Data Deficient (DD) within
IRAN,
Much-branched fastigiate undershrub to c. 40 cm, the branches usually arising at right angles to the main stem, adpressed pubescent, hairs resembling cystoliths. Leaves commonly absent, sessile, 1. 5–2.5 cm, linear to linear oblanceolate, acute, entire, attenuate at base, subglabrous or with few adpressed hairs. Flowers 2–3 in very lax axillary dichasial cymes borne on rigid branches, somewhat spinescent after flowers have fallen; peduncles 1.5–2 cm; bracteoles 1 mm, oblong-elliptic, minute; pedicels 2–8 mm, very fine; sepals 3–4 × 2 mm, obovate, shortly mucronate, uniformly pale green, adpressed pubescent, inner sepals slightly longer and broader; corolla 11–14 mm long, white, weakly lobed, midpetaline bands pilose; ovary glabrous; style glabrous, divided c. 3 mm above base, stigmas 4 mm. Capsule glabrous; seeds brown, glabrous, smooth. [
Endemic to southern Iran (
The very short pubescent sepals are a good character as are the very slender peduncles arising at or near 90°. There seems to be some variation in inflorescence branching, sometimes clearly dichasial (
The following species (species 120–126) form a complex of mostly ill-defined taxa. Although frequently misidentified,
AZERBAIJAN (?), “Caspian Sea”,
Much-branched undershrub with somewhat zigzag, rigid branches forming an intricate ball-like plant to 1(-3) m in height, the lower part sometimes with a distinct trunk up to 1.5 cm in diameter; young stems sericeous, branching at 80–90°. Basal leaves 10–30 × 1 mm, stem leaves usually few, sessile, 6–7 (-20) × 0.5–1 mm, linear, pubescent. Flowers usually solitary (rarely paired) borne on thin, rigid peduncles 4–15 (-20 )mm long; bracteoles 1 mm, triangular, scale-like, puberulent; pedicels 1–5 mm, puberulent, often recurved; outer sepals 3–4 × 2.5 mm, elliptic-obovate, rounded to emarginate, sericeous-pubescent; inner sepals narrower, 1–1.5 mm wide; corolla 0.6–0.9 cm long, deeply lobed to about quarter of its length, white or pink, midpetaline bands narrowly triangular, pubescent, terminating in a point; ovary sericeous; style glabrous, very short, divided c. 2 mm above base, stigmas c. 1.5 mm; capsule pubescent or glabrous, 1-seeded; seeds pubescent. [
Russia: Dagestan (
Distinguished by the sericeous stem, short obtuse sepals, small, lobed corolla and tendency to have short rigid peduncles giving the plant a characteristic intricate habit.
The recently described
Based on
Erect undershrub, intricately branched from the base to at least 40 cm; stems, thinly sericeous. Leaves 1–5 × 0.2 cm, linear-oblong, acute, base tapering into a long pseudopetiole, thinly adpressed pubescent and appearing greenish. Flowers 1 (-2) at the apex of rigid, straight, relatively slender woody peduncles 0.8–1.5 cm long; bracteoles minute, linear, c. 1 mm, pedicels 0.5–1 mm, bent at 90° to peduncle; sepals (3-) 4 (-5) × 1.5 mm, narrowly oblong, acute, densely pubescent; corolla (8-)10 mm, white, unlobed, broadly infundibuliform, midpetaline bands pilose, terminating in a tooth; ovary pilose. Capsule pilose, seeds pubescent.
Kazakhstan (
The indumentum and corolla size suggests this lies between
Sa’ad treated this species as a synonym of
KAZAKHSTAN, Balkash,
Branched (often divaricately) undershrub to at least 50 cm, stems sericeous to adpressed puberulent, somewhat zigzag, branches arising at an acute angle. Basal leaves unknown, stem leaves sessile, 1–3 × 0.1–0.3 cm, linear-lanceolate, acute, entire, base cuneate, sericeous, Flowers up to 3 in pedunculate, cymes but commonly reduced to solitary flowers; peduncles 2.5–6 cm, characteristically long and slender, often bent at transition to pedicel; bracteoles 2 × 0.5 cm, deltoid-lanceolate, caducous; pedicels very short, 1–2 mm; sepals 5–7 × 2 mm, narrowly ovate, shortly acuminate, pubescent; corolla 1.1–1.5 cm, white, shallowly lobed, the midpetaline bands, pilose, terminating in teeth; ovary pilose; style glabrous, divided c. 5 mm above base, stigmas 2 mm. capsule pilose at apex; seeds pubescent.
Kazakhstan (
UZBEKISTAN, Chiwa (Khiva),
Perennial undershrub from a woody rootstock forming a small bush; stems numerous to 50 cm, rigid and woody below, herbaceous above, slightly divaricate, densely grey-pilose. Basal leaves not seen; stem leaves sessile, 0.3–1.5 × 0.3–1.5 cm, very variable in size from plant to plant, oblanceolate, lanceolate, ovate or suborbicular, acute, entire to somewhat undulate, basally cuneate to rounded, pilose. Flowers 1–3 (- 5) in axillary dichasial cymes but often solitary; peduncle 0.5–6 cm, somewhat terete to rigid; bracteoles 3 mm, filiform; pedicels 2–10 mm; outer sepals 5–6 × 2.5 mm, lanceolate, ovate or elliptic, acuminate or caudate, densely lanate; inner sepals broader (c. 3 mm), glabrous; corolla 1.1–1.3 (-1.5) cm long, cream or pinkish, unlobed, midpetaline bands darker, pilose, terminating in a tooth; ovary glabrous or with a few apical hairs; style glabrous, divided c. 2 mm above base, stigmas c. 2 mm; capsule glabrous, 1-seeded (?always); seeds hirsute. [
Uzbekistan/Kazakhstan: Kzyl-Kum (
Distinctive because of the pilose indumentum with spreading hairs combined with the lanceolate to ovate stem leaves but very close to
Specimens intermediate between
TAJIKISTAN,
Small grey-pilose undershrub similar in general facies to
Tajikistan, Tujun-tau, apparently only known from the type.
This species needs revision. It may or may not prove to be distinct from
IRAN, Kashan,
Undershrub from a thickened woody rootstock to 60 cm with rigid (but not spiny) branches spreading widely at almost a right angle and forming an entwined mass; stems and leaves pubescent with spreading hairs. Leaves sessile, 0.5–5.5 × 0.1–0.9 cm, linear to oblong, acute, margin entire to undulate, pubescent, merging upwards into the bracts. Flowers pedunculate, solitary or in 2–3-flowered cymes; bracts 3–20 × 0.5–2.5 mm, suborbicular to broadly ovate; peduncles 0.8–5.5 cm, straight, rigid, usually arising at 90° to the stem; bracteoles 2–3 mm, filiform; pedicels 0–1 mm, often at an angle from the peduncle; outer sepals 4–5 × 2 mm, ovate, acute with mucronate apex, pilose, pubescent or, rarely, glabrous on the abaxial surface, sometimes glabrescent in fruit; inner sepals similar but less hairy and commonly convex; corolla 1.1–1.8 cm, white or purplish-flushed, somewhat undulate but not lobed, midpetaline bands pubescent; ovary glabrous to hirsute, style glabrous; style divided 5–6 mm above base; capsule glabrous or with a few apical hairs, seeds hirsute. [
Turkmenistan (
We are proposing a broad concept of this ill-defined species. It is characterised by the presence of spreading hairs at least on the lower part of the stem, acute to mucronate sepals and straight rigid peduncles arising at 80–90° to the branch. The types of
AFGHANISTAN,
Much-branched undershrub to 30 cm, stem and all vegetative parts grey-sericeous, branches rigid and with few leaves, ascending at about 60°. Leaves sessile, 6–20 × 1–5 mm, linear to narrowly oblong, obtuse, entire, narrowed at both ends. Flowers 1–3, arising in the axils of small bracts on a rigid peduncle; bracts 2 × 1 mm, triangular; peduncles 0.8–2.5 cm; bracteoles c. 1 mm; pedicels 0–1.5 mm, sepals 3–3.5 × 2 mm, oblong-elliptic, densely sericeous, outer rounded, inner mucronulate; corolla 0.9–1 cm long, white, midpetaline bands sericeous, weakly lobed; ovary globose, pubescent (or glabrous in
Afghanistan (
This poorly known species is characterised by the grey sericeous stems and leaves, small sericeous mucronulate calyx and short corolla. It may be only a form of
IRAQ, between Kirkuk and Sulaymaniyah,
Undershrub with fastigiate branches to 40 cm, stems appressed pubescent. Leaves sessile, the basal leaves 2.5–6 × 0.5–3 cm, oblanceolate, acute to obtuse, attenuate at the base, appressed pubescent; stem leaves linear 1.5–3.5 × 0.2–0.4 cm. Flowers axillary, solitary or paired forming a lax, sparsely branched inflorescence, which is commonly dichasial in structure apically; peduncles 0.5–2 cm long, slender, bracteoles minute; pedicels 2–6 mm; sepals 6–7 × 3–4 mm, elliptic to obovate, mucronate, the margins scarious, the inner sepals narrower; corolla 12 mm long, white, midpetaline bands pubescent; ovary comose; style pilose, divided 4 mm above base, stigmas 1–2 mm. Capsule not seen. [
Iraq (
This species is distinguished by the relatively large, glabrous sepals and narrow, sparingly branched inflorescence which is clearly cymose in structure apically. It seems close to
IRAN, Kermanshah,
Perennial undershrub with woody taproot, from which arise many sericeous stems to 50 cm. Basal leaves unknown; stem leaves sessile, 20–40 cm long, linear-filiform. Flowers 3–5 in axillary and terminal dichasial cymes; bracteoles linear, acute; sepals 8 mm long (fide description) or 4–4.5 mm (fide illustration), ovate, mucronate, glabrous; corolla c. 1.8 cm long, white, midpetaline bands pubescent; ovary glabrous; style glabrous, much longer than stigmas (fide illustration). Capsule glabrous, 1-seeded.
Iran; Iraq (
We have not seen the type material.
IRAN, Luristan, Chah-Bazan,
Undershrub with woody taproot, from which arise many rigid branched stems to c 40 cm, stems minutely adpressed pilose, the hairs resembling cystoliths. Leaves sessile, 10–30 × 0.5 mm, filiform, acute, adpressed pubescent. Flowers 1–3 in long-pedunculate, axillary and terminal dichasial cymes; peduncles up to 20 cm long, straight, slender and not very rigid; bracteoles 1–2 × 0.25 mm, filiform; sepals 5–6 × 4 mm, obovate-elliptic, apiculate, glabrous; corolla1.4–1.5, colour unknown, midpetaline bands pubescent; ovary comose; style divided 2 mm above base; stigmas 3 mm. Capsule and seeds not known. [
Iran (
Appears to be close to
IRAN, Tehran,
Slender, sericeous, rigid undershrub of grey appearance to c. 30 cm with numerous erect stems and very rigid branches arising at c. 60°. Leaves sessile, 0.8–2.8 × 0.2–0.3 cm, linear to oblanceolate, acute, entire, decurrent at the base. Inflorescence of solitary pedunculate flowers, becoming somewhat dichasial cymose apically; bracts oblong, caducous; peduncles 5–20 mm, slender; bracteoles minute, scale-like; pedicels 3–7 mm, curved; sepals 2 × 2 mm, suborbicular to broadly elliptic, obtuse, the outer ones pubescent, the inner glabrous, membranous, mucronate; corolla 0.6 cm long, white, midpetaline bands pilose; ovary pilose, style pilose, divided 1 mm above base, stigmas 2 mm. Capsule and seeds not seen. [
Iran. Only known from the type collection.
The tiny sepals and corolla and the very rigid grey stems are characteristic.
KAZAKHSTAN, Koksu River,
Perennial undershrub with tough woody rootstock from which arise several erect, woody, branched fastigiate stems to 50 cm; stems adpressed pubescent. Basal leaves sessile, 2–4 × 0.2–0.5 cm. oblong, acute, entire, attenuate to somewhat abruptly narrowed at the base, glabrous to thinly adpressed pubescent on the veins and lower surface; stem leaves similar but linear, diminishing in size upwards. Inflorescence terminal and panicle-like, formed of terminal dichasial cymes and solitary flowers at the apex of lateral branches; bracteoles 2–5 mm, filiform; pedicels 1–10 mm; sepals 5–7 × 2 mm, oblong to obovate, acute to mucronate, glabrous, the inner sepals larger, c. 3 mm wide, obovate, with a clearly demarcated wing-like margin; corolla 1.8–2.2 cm, pink, undulate, midpetaline bands pilose, terminating in a broad tooth; ovary glabrous, style glabrous, divided c. 4 mm above base, stigmas 3 mm. Capsule glabrous, 1-seeded, seeds pubescent (fide Sa’ad).
Two well-defined subspecies can be recognised:
Characterised by the shorter sepals, c. 5 mm long, which are obovate and abruptly mucronate.
China (fide
Based on
Characterised by having oblong, acuminate sepals, c. 7 mm in length.
Apparently restricted to Iran and Turkmenistan: Northern Iran (
A relatively distinct group of shrubs or undershrubs or, if herbaceous, woody at base, never twining or trailing, the leaves always lacking a distinct petiole. All species are hirsute and usually sericeous, often cushion-forming and/or with spinescent branches. The ovary is hirsute except in
RUSSIA, Siberia,
Grey-sericeous perennial herb with thick woody taproot; stems short, decumbent or weakly ascending, more or less herbaceous, to 12 (-22) cm long. Leaves sessile, 0.7–3 × 0.15–0.5 mm, linear or linear-oblanceolate, obtuse, tapering at the base. Flowers pedunculate from the axils of leaf-like bracts or, rarely, terminal only; peduncles 0.4–2.2 cm, 1-flowered; bracteoles, 4–13 × 0.5 mm, linear; pedicels 1–6 mm, often bent or curved; outer sepals 4.5–6 × 2–2.5 mm, ovate, abruptly narrowed and drawn out to an obtuse apex, the inner sepals much broader, c. 7 × 4 mm; corolla 0.8–1.3 cm long, white or very pale pink, very obscurely lobed, midpetaline bands pilose, terminating in a blunt tooth; ovary and style pilose; style divided c. 3 mm above base, stigmas c. 3 mm long. Capsule pilose at apex only; seeds smooth, pubescent. Figure
Northern China and Manchuria (
Maxime affine
CHINA, Shensi, Hancheng Sian, 1914,
Much branched perennial undershrub with erect stems to c. 20 cm; old stems woody and rigid, weakly divaricate, up to 10 mm thick, new growth herbaceous grey-sericeous. Leaves sessile, 4–40 × 0.5 mm, linear, acute, grey-sericeous. Flowers terminal, solitary or in small dichasial cymes, the branches appressed pilose, up to 6 mm long; bracts and bracteoles not clearly differentiated, 1–10 × 0.5 mm, linear; peduncles not differentiated from the stem; pedicels 1–3 mm; outer sepals 6–7 × 2.5–3.5 mm, elliptic to obovate, caudate, densely pilose; inner sepals 5 × 3–4 mm, ovate, caudate, convex, somewhat scarious, thinly pilose; corolla 1.5–2 cm, unlobed, midpetaline bands pilose; filaments 4 mm long, glabrous, anthers 2 mm, sagittate; ovary 2 mm long, conical, comose, style pilose below, divided 3–7 mm above base, persistent; stigmas 1.75 mm; capsule 2 × 3.5 mm, pubescent, capped by the persistent style; seeds 1.5 × 1 mm, conical with a rounded base, pubescent, blackish-brown.
China: Shanxi, Hanchengsian; Henan, Loyan, Shan Xian, Sanmenxia Gorge, on left bank of Yellow River (
Similar to
This species is only known from two collections from the Hwang Ho (Yellow River) region of north China. The Petrov collection at LE has entirely solitary terminal flowers, while in the Purdom collection (K) some branches terminate in solitary flowers while in others the flowers are arranged in terminal dichasial cymes.
This species may be rare and threatened or simply overlooked but for the time being should be classified as Data Deficient (DD) within
KYRGYZSTAN, Jalalabad,
Low undershrub with thick woody rootstock from which arise various short stiff stems to c. 20 cm, stems densely pilose, dead branches persistent and spinescent. Leaves sessile, 12–20 × 1–1.5 mm, linear-oblanceolate, obtuse, flat, silvery sericeous. Flowers subsessile from the uppermost leaf axils, 1–4 together forming a subterminal cluster; peduncles absent; bracteoles 1–3 mm, linear; pedicels 0–2 mm, pilose; sepals 6–7 × 2.5, elliptic or obovate, narrowed to an apiculate apex, densely pilose; corolla 1.7–2 cm, pink, obscurely lobed, the lobes pubescent, midpetaline bands densely sericeous; ovary and style pilose. Capsule and seeds not seen.
Kyrgyzstan (
“near Tashkent”,
Densely grey canescent undershrub with thick woody base, up to 1.5 cm in diameter, from which arise numerous, erect, straight, rigid but not spinescent woody stems to 40 cm, these near leafless upwards and somewhat scape-like. Leaves mostly on lower part of stem, numerous, sessile, 5–25 × 0.5 mm, linear to very narrowly linear-oblanceolate, obtuse. Flowers subsessile, 1–5 grouped at the apex of the stem, sometimes with a single flower on the stem below the apical cluster; bracts and peduncles absent; bracteoles 1–2 mm, linear-lanceolate, obtuse; pedicels 1–2 mm; sepals 4–6 × 2.5–3 mm, broadly ovate to suborbiculate, shortly apiculate, margins scarious, densely silky-pubescent; corolla 1.3–1.5 cm long, white, unlobed but with pilose border, midpetaline bands densely pilose, terminating in teeth; ovary densely pilose; style pilose, divided c. 8 mm above base; stigmas 3 mm. Capsule pilose apically. [
Endemic to Kyrgyzstan (
Very distinct because of the dense grey-canescent indumentum, the plentiful linear leaves and the long, scape-like stem with subsessile flowers at or near the apex
The cited type locality is thought to be an error as this species has never subsequently been found near Tashkent (Uzbekistan).
MONGOLIA, Tzagan-Balgassu,
Low grey-sericeous undershrub with woody rootstock, often more or less cushion-forming; branches short, woody, spinescent when old. Leaves sessile, 1–2.8 × 0.1–0.3 cm, linear to narrowly linear-oblanceolate, obtuse, entire, tapering at the base. Flowers in clusters of 1–4 in the axils of the leaves towards the tips of the branches; peduncles absent; bracteoles c. 2 mm long, filiform or linear; pedicels 1–2 mm; outer sepals 4–6 × 3.5 mm, obovate, rounded to truncate and slightly fimbriate, pilose abaxially; inner sepals broader, 4–4.5 mm wide, glabrous to thinly pilose; corolla 1.5–2.5 cm long, pink, unlobed, midpetaline bands pubescent; ovary pilose; style thinly pilose, divided c. 4.5 mm above base, stigmas c. 3 mm. Capsule pubescent; seeds glabrous. [
Widespread in central Asia: China (
KAZAKHSTAN,
Sometimes treated as a synonym of
China : Kashgar (
RUSSIA, Irtysh,
Intricately branched spiny shrub to 50 cm, stems adpressed pubescent. Leaves sessile, 1–3.5 × 0.3–0.6 cm, oblong to narrowly oblong-elliptic, acute, margin entire, base cuneate, adpressed-pubescent to sericeous; cauline spines abundant on young shoots. Flowers borne on short spinescent axillary peduncles, usually solitary, sometimes up to 3 in a diachasial cyme; peduncles 12 mm, spine-like; bracteoles 9–11 × 1–2 mm, linear to linear-elliptic; pedicels 2 mm, recurved; sepals very lax, outer sepals 10–11 × 3–6 mm, elliptic or rhomboid, acute to mucronate, densely pubescent; inner sepals much shorter, 7–8 mm long; corolla 2.3–2.5 cm long, pink, somewhat undulate, midpetaline bands darker pink, adpressed pilose; ovary and style pilose; style divided 7 mm above the base, stigmas 3 mm long. Capsule pilose; seeds puberulous. [
China (fide
KYRGYZSTAN,
Low spiny undershrub, often cushion-like, 10–20(-30) cm high, stems divaricate, branches arising at right angles, grey-sericeous when young, persistent and woody-spinescent when old; cauline spines also present. Leaves sessile, 1–2.5 × 0.2–0.5 cm, oblanceolate, acute, entire, tapering to a long petiole-like base. Flowers solitary, axillary, pedunculate; peduncles 5–7 mm, woody, straight, spinescent; bracteoles 6–12 × 1–2 mm, foliose, linear to linear-oblong; pedicels 1–2 mm; outer sepals 7–11 × 7–11 mm, orbicular, apex rounded and apiculate, base cordate, glabrous, papery in texture; inner sepals 5–7 × 4–5 mm, ovate, acuminate; corolla 1.8–2.2 cm long, pink, undulate but not lobed, midpetaline bands brownish-pilose terminating in a small tooth; ovary pubescent; style divided 7 mm above base, pubescent below; stigmas 3 mm. Capsule apically pubescent; seeds not seen.
Similar in inflorescence, habit and overall morphology to
Kyrgyzstan (
There are a number of specimens in which the sepals are pilose but similar in shape to those of
IRAN,
Intricately branched spiny undershrub to 60 cm, stems finely adpressed pubescent, branches spinescent and with some stem spines near branch tips formed from old peduncles. Leaves sessile, 0.5–1.2 × 0.3–0.4 cm, oblong to oblanceolate, obtuse, margin entire, base attenuate, adpressed-pubescent. Flowers axillary, usually solitary, sometimes up to 3 in a diachasial cyme; peduncles 4–8 mm, woody, spinescent, bent at apex; bracteoles 1–1.5 mm, scale-like; pedicels c. 1 mm; outer sepals 3 × 2 mm, oblong to elliptic, obtuse and mucronate, convex, pubescent with spreading hairs; inner sepals 4.5–5 × 3 mm, pubescent with spreading hairs; corolla 1.3–1.6 cm long, white, midpetaline bands adpressed pilose; ovary pubescent; style pubescent, divided c. 4 mm above base, stigmas 3 mm; capsule comose; seeds smooth, glabrous. [
Iran (
The inner sepals are noticeably longer than the outer sepals.
Iran,
Grey spiny undershrub to 60 cm, stems grey-sericeous, spinescent, lateral spines formed from dead peduncles present near branch tips. Leaves sessile, 1–1.7 × 0.3–0.6 cm, obtuse, margin entire, base attenuate, grey-sericeous. Flowers solitary (or, fide
Iran: Kerman, Fars (
Resembles
IRAN, Shiraz,
Intricately branched spiny undershrub, 10–60 cm high, stems woody, weakly divaricate and spreading at a wide angle, grey-sericeous, spine-tipped and with numerous short lateral spines on older shoots formed from old peduncles. Leaves sessile, 0.5–1.5 × 0.2–0.4 cm, oblong-elliptic, oblanceolate or oblong-lanceolate, obtuse, margin entire, attenuate at the base, grey-sericeous. Inflorescence of few-flowered terminal clusters, sometimes also with solitary flowers in the leaf axils below; peduncles 3–8 mm, rigid, spinescent, sericeous; bracteoles 4–6 × 0.5–1.5 mm, linear, acute; pedicels 1–3 mm, villous, sometimes recurved; outer sepals 7–10 × 2–3 mm, ovate, abruptly narrowed into an acuminate and mucronate apex, densely pilose with long, pinkish hairs, inner sepals narrower and with scarious margins; corolla 1.5–2.6 cm, pink or white, very shallowly lobed, the midpetaline bands with long hairs; ovary and style pilose; style divided 6–7 mm above base, the stigmas relatively short, c. 2 mm long; capsule pilose at apex, one-seeded; seeds puberulous. [
Pakistan (fide
The spiny sericeous habit, spinescent peduncles and sepals with very long hairs make this distinct from all but
IRAN, Fars province, 20 km from Shiraz along road to Bushehr, 26 Jun 1972,
Much branched, spiny woody undershrub forming a compact intricate bushlet c. 20–25 cm high, young stems pale brown with appressed hairs. Branch tips spinescent, lateral spines present, formed from old fertile and (towards the branch tips) sterile peduncles. Leaves sessile, 7–10 × 3–4 mm, oblanceolate to obovate, obtuse, entire, attenuate at the base, appressed pubescent with whitish hairs. Flowers solitary, axillary; bracts resembling reduced leaves, 2–3 × 1.5–2 mm, oblong-elliptic, obtuse; peduncles 12–14 mm, straight, rigid, spinescent, arising at 90° from the branch, appressed pubescent; bracteoles c. 0.25 mm, minute, filiform; pedicels 1–1.5 mm, often bent at 90° to the peduncle, distinctly more densely pubescent with spreading hairs that the peduncle; outer sepals 4–4.5 × 3 mm, elliptic-rhomboid, acute to shortly acuminate, thinly pilose, often (?always) purplish, inner sepals c. 4 × 1.5 mm, colourless, very thinly pilose; corolla 1.5–1.7 cm, base somewhat inflated, colour unknown, midpetaline bands densely pilose; filaments glabrous, anthers 1.25 mm, ovary subglobose, c. 1 mm, thinly pilose; style 5 mm long, pilose, stigmas 2 mm, slightly widened apically. Capsule and seeds unknown.
Only known from the type.
Most similar to
This species is only known from a single collection. It may be rare and threatened or simply overlooked but for the time being should be classified as Data Deficient (DD) within
IRAN,
Similar to
Endemic to Iran (
IRAN, Karevandar,
Small undershrub to c. 25 cm, stems and branches sericeous, spinescent, sterile cauline spines present, few. Leaves sessile, 1.5–3 × 0.5 cm, obovate or elliptic, obtuse, entire, the base attenuate to a petiole-like base, densely sericeous. Flowers in compact capitate cymes, terminal on naked peduncle-like branches; bracts ovate, acuminate, densely pilose; bracteoles c. 8 × 2 mm, linear-lanceolate, finely acuminate, densely pilose; pedicels absent; sepals 8–9 × 1.5–2.5 mm, lanceolate, acuminate, pilose; corolla c. 1.8 mm long, cream, midpetaline bands pilose; ovary pilose; style pubescent, divided c. 5 mm above base, stigmas c. 3 mm. Capsule not seen. [
Endemic to southeastern Iran (
Superficially similar to
“In Italia, Sicilia, Narbona, Verona” (lectotype LINN No. 218.48!, designated by
Undershrub with stout woody base and erect or ascending stems to 50 cm; stems commonly branched in the lower half; all vegetative parts pilose to pubescent. Leaves sessile, the basal and lowermost leaves 4–8 × 0.5–2 cm, obovate-spathulate with a long attenuate petiole-like base; cauline leaves 2–5 × 0.2–0.8(-1.5) cm, linear, linear-oblong, lanceolate or oblanceolate, acute, entire, cuneate at the base. Flowers in long-pedunculate axillary and terminal diachasial cymes, these sometimes congested and subcapitate, often forming a large lax, panicle-like inflorescence or, sometimes, appearing in the form of a lax raceme; bracts leaf-like, linear to oblong, diminishing in size upwards; peduncles 1–9 cm long; bracteoles 5–7 mm, linear, pedicels 0–10 mm; outer sepals 7–8 × 4–5 mm, lanceolate, acuminate to caudate, densely pilose, the lower half colourless, the apical portion green, the inner sepals slightly broader, ovate; corolla 1.7–2.5 cm long, pink, unlobed, midpetaline bands pilose, darker; ovary pilose, style pilose, divided c. 4 mm above base, stigmas 3 mm. Capsule pilose, seeds puberulent. [
Mediterranean region east to Afghanistan: Morocco (
This species can look very similar to forms of
TURKEY, (Gazi)antap,
Similar to
Apparently endemic to Turkey, being found principally around Gaziantap (
IRAN,
Woody based perennial with deep taproot, usually branched at base with several ascending stems in a loose tuft to c. 35 cm in height, stems and all vegetative parts obscurely to densely appressed pubescent. Leaves sessile, 3 × 0.2–0.3 cm, linear, acute, entire, attenuate at base. Inflorescence of terminal heads, occasionally with a few flowers in the uppermost leaf axil; bracts 6–9 × 1.5–2 mm, lanceolate; peduncles and pedicels absent; bracteoles 7 × 1 mm, linear, acute, often overtopping the head; outer sepals 8–9 × 2.5 mm, ovate, abruptly narrowed around the middle with a long acuminate apex, villous; inner sepals similar but with glabrous margins; corolla 1.7–2.5 cm long, cream, midpetaline bands pilose; ovary pilose; style pilose, divided 7 mm above the base, stigmas 4 mm. Capsule pilose. [
Endemic to Iran (
Distinguished from other species with a capitate inflorescence by its narrow, linear leaves and adpressed stem indumentum. The sepals are shorter than in
IRAQ, Mosul,
Woody based perennial with thick woody taproot, usually branched at base, sometimes more or less cushion-forming, with several erect stems in a loose tuft to c. 40 cm in height, stems and all vegetative parts densely appressed pilose. Basal leaves 3–5 × 0.4–0.8 cm, oblanceolate to spathulate, acute, entire, attenuate at base; stem leaves sessile, 1.5–4.5 × 0.2–0.8 cm, oblong-lanceolate, acute, entire, base cuneate to attenuate. Inflorescence mostly of sessile terminal heads but sometimes with 1–2 flowers in the axils of the upper leaves; heads with 1–5 flowers; bracts 10–22 × 1.5–3 mm; peduncle absent; bracteoles 4–6 × 1 mm, linear, pedicels 0–3(-9) mm; outer sepals 11–13 × 3–4 mm, broadly lanceolate, abruptly narrowed to a long attenuate apex, pilose; inner sepals ovate but slightly shorter, c. 10 mm long; corolla 2–2.8 cm long, pale pink or white, unlobed, midpetaline bands adpressed pilose, pink; ovary pilose, style pilose, divided 6–8 mm above base, stigmas 3 mm; capsule pilose, 1-seeded (fide
Iran (
The short-pedicellate flowers which give the inflorescence a looser appearance than in related species are noteworthy. The appressed indumentum especially on the stem serves to separate this species from
IRAN, Ilam,
Very similar to
Endemic to Iran and only known from the type collection.
TURKEY,
Variable woody based perennial with thick woody taproot, sometimes cushion-forming, with several ascending stems in a loose tuft to c. 20 cm in height, stems and all vegetative parts with long spreading hairs. Basal leaves 4–8 × 0.3–0.8 cm, oblong or oblanceolate, acute, entire, narrowed to a long petiole-like base; stem leaves sessile, 3–5 × 0.6–0.9 cm, oblong-elliptic, acute, entire, cuneate at the base. Flowers in many-flowered terminal heads, occasionally also 1–2 flowers in the axil of the uppermost leaf; bracts 1.3–3 × 0.1–0.6 cm, linear-oblong or lanceolate, acute, pilose; bracteoles 11–16 × 0.5–1 mm, linear, pilose; pedicels 0–3 mm; outer sepals 6–10 × 1.5–2 mm, lanceolate, long acuminate, pilose; inner sepals abruptly narrowed around the middle with a caudate apex; corolla 1.5–2 cm, white or pink, not lobed, midpetaline bands pilose, darker; ovary pilose; style pilose, divided c. 3 mm above base, stigmas 3–5 mm. Capsule pilose; seeds pubescent. [
We recognise two subspecies:
Stems and leaves with prominent spreading hairs as well as appressed sericeous hairs. Leaves oblong or oblanceolate, 5–10 times longer than broad.
Turkey (
Based on
Leaves and stem silvery with appressed sericeous hairs; spreading hairs absent or almost so. Leaves broadly oblong to obovate, up to three times as long as broad.
Crimea (
CRIMEA,
Perennial herb with leaves arranged in a basal rosette, from which arise erect stems 30–40 cm high; stems adpressed pilose. Leaves sessile, mostly basal, 2–5 × 0.3–0. 5 cm, linear to oblanceolate, acute, tapered at the base to a pseudopetiole, adpressed pilose to subsericeous. Flowers congested at the top of the stem forming a headlike inflorescence with a single head arising from the uppermost leaf axil; bracts resembling linear-lanceolate reduced leaves, erect, slightly exceeding the inflorescence; peduncles of lateral capitula 2–22 mm, bracteoles filiform, pedicels 0–3 mm, sepals 14–15 × 5–6 mm, obovate, strongly cuspidate, adpressed pubescent and with a few spreading hairs, the inner sepals smaller; corolla 1.8–2 cm long, pink, midpetaline bands pilose; ovary hirsute; style pilose, divided 5 mm above the base; stigmas 4 mm. Capsule not seen.
Crimea (
This species appears to be almost certainly the hybrid between
CRIMEA,
Undershrub with a thick woody taproot from which arise numerous short prostrate to weakly ascending stems to 30 cm forming a mat, vegetative parts all grey-sericeous. Leaves sessile, 2–4.5 × 0.2–0.6 cm, linear, oblong or oblanceolate, acute, entire, base long-attenuate.. Flowers in terminal cymose clusters, usually with single flowers or few-flowered, diachasia, subsessile or borne on peduncles up to 1.7 cm long from the axils of the uppermost leaves; bracts resembling the leaves but smaller, linear- oblong; bracteoles 3–6 × 0.5 mm, linear; pedicels 1–4 mm; outer sepals 10–15 × 5–12 mm, ovate to suborbicular, acuminate, gibbous with broad membranous wing-like margins; inner sepals slightly smaller; corolla 2–2.7 cm long, pale yellow or cream; unlobed but slightly undulate, midpetaline bands adpressed pilose; ovary pubescent, style pubescent, divided c. 7–8 mm above ovary, stigmas 5 mm. Capsule pubescent, seeds puberulent. [
We recognise two subspecies based on sepal size, although intermediates are not uncommon.
Sepals relatively small and narrow, 7–10 × 5–10 mm.
Croatia (?); Greece; Bulgaria; Macedonia (
TURKEY, Harput,
Sepals large, suborbicular 11–17 × 11–13 mm, the type being an extreme example of the subspecies.
Turkey (
The gibbous sepals are very distinctive in this species.
Based on
Cushion-forming undershrub, stems short, spreading, the bases covered in leaf remains; vegetative parts silvery-sericeous. Leaves sessile, 0.3–2 × 0.2–0.6 cm, obovate to oblanceolate, obtuse or acute, entire, cuneate to a broad base, the venation very prominent, especially below. Flowers usually solitary, terminal or axillary; peduncle 1–3 mm long; bracteoles 4–8 × 0.5–1 mm, linear; pedicels 1–3 mm; sepals 7–10 × 2–3 mm, oblong-lanceolate with caudate apex, base membranous, apex pilose, inner sepals oblong-ovate with membranous margins; corolla 1.7–2.1 cm long, pink, weakly lobed, midpetaline bands darker, pilose; ovary sericeous; style sericeous, divided 3–4 mm above the ovary, stigmas c. 4 mm. Capsule hirsute; seeds glabrous, smooth. [
Divisable into two, geographical disjunct but morphologically similar subspecies:
Leaves obovate or oblanceolate, the hairs weakly appressed to somewhat spreading and rather similar to the indumentum of the calyx.
Spain (
Based on
Leaves obovate to narrowly linear-oblanceolate, hairs strongly appressed, sometimes sericeous and clearly differentiated from the spreading hairs of the calyx. [
Balkan peninsular and Turkey: Croatia (
TURKEY,
This apparently sterile hybrid differs from
Reported from a single locality in central Anatolia.
This taxon represents
TURKEY,
It resembles
This hybrid is reported from a single locality in south central Anatolia.
This taxon represents
TURKEY,
Resembles
Reported from a single locality in SW Anatolia.
This taxon represents
Plant from Bulgaria, Ali Botush Mountain, cultivated in Berlin,
Intermediate between
Endemic to the area of Ali Botush Mountain, Bulgaria
Without locality,
Perennial herb, often mat-forming, from a thick, branched underground rhizome; stems ascending to 30 cm, vegetative parts densely but minutely puberulent to shiny white-sericeous. Basal leaves 4.5–8 × 0.8–1.8 cm, oblong to oblanceolate, acute, entire, attenuate at base into a long pseudopetiole; stem leaves distinctly sessile, 1.5–5 × 0.2–0.7 cm, narrowly linear-oblanceolate, acute. Flowers in terminal diachasial cymes and axillary cymes of 1–5 flowers; bracts and bracteoles not clearly differentiated, 3–25 × 1–3 mm, linear to oblanceolate, acute, very variable in size; pedicels 3–4 mm; sepals of two parts–pale base and acute, commonly reflexed, green apex; outer sepals 6–10 × 2 mm, oblong-lanceolate; inner sepals 6–8 × 3 mm, broadly ovate, margins scarious; corolla 1.8–2.5 cm long, pink with white centre and paler midpetaline bands, shallowly lobed, midpetaline bands appressed pilose; ovary conical, sericeous, style pilose, divided 7–10 mm above base, stigmas 5–7 mm; capsule pubescent; seeds shortly pubescent. [
Around the Mediterranean and Black Seas and through Iran and central Asia to western China (Xinjiang) and Siberia (Altai): Portugal; Spain (
Although we have not seen the type of
Apart from
CYPRUS, Lania,
Similar to
Endemic to Cyprus: Cape Gata (
This taxon represents
“Levant”, plant cultivated in Paris (holotype P [Herb. Lam.]).
60 cm, the old growth woody, young shoots herbaceous, flower-bearing; vegetative parts grey-sericeous. Leaves sessile, (1-)2.5–6 × (0.1-) 0.4–0.8 cm, linear, oblong or oblanceolate, obtuse to acute, entire, the base long-attenuate. Flowers in terminal diachasial clusters, sometimes with single flowers or few-flowered, diachasia borne on peduncles up to 3 cm long from the axils of the uppermost leaves; bracts resembling the leaves but smaller, always linear-oblong; bracteoles 8–10 × 1–1.5 mm, linear; pedicels 0–10 mm; outer sepals 6–9 × 2.5–5 mm, ovate, shortly acuminate to an obtuse apex, villous; inner sepals broader with scarious margins; corolla 2–2.5 cm long, pink, very shallowly lobed, the midpetaline bands appressed pilose, brown, terminating in a tooth; ovary pilose, style pilose, divided c. 3 mm above base, the stigmas 5 mm. Capsule pilose; seeds, densely pubescent. [
Very variable in habit but commonly ascending with stems > 10 cm in length and branches herbaceous. Leaves very variable but usually 2.5–6 × 0.4–0.8 cm.
Eastern Mediterranean: Malta (
LIBYA, Cyrenaica,
Distinguished by its prostrate, pulvinate habit – it does not normally exceed 10 cm in height. The leaves are oblanceolate, small, 1–2 × 0.4–0.6 cm.
Appears to be a maritime ecotype and is reported from Libya and Cyprus (
LIBYA, Cyrenaica,
Distinguished by its rigid, more or less fastigiate habit, the branches all woody. Leaves are linear, 1–1.5 × 0.2 cm and mostly absent below. The hairs on the bracteoles and calyx are shorter than in the type.
Reported from Libya, Cyprus and Turkey (
GREECE, Crete, Ierápetra.
Undershrub clearly related to
Endemic to Crete.
It is impossible to confirm whether this is a distinct relict species or some peculiar adaptation of
MOROCCO, Middle Atlas, J. Guebb-er-Rahal,
A small cushion plant, 7–13 cm in diameter with a stout woody taproot, stem sericeous. Leaves sessile, 2–3 × 0.2–0.4 cm, linear to linear-oblanceolate, acute, entire, base attenuate, grey sericeous-pubescent on the lower surface and margins of upper surface, glabrous on central area. Flowers solitary or paired, sessile or on very short terminal peduncles up to 10 mm long; bracteoles linear, 3 × 0.5 mm, pedicels 2 mm; outer sepals 5–6 × 2 mm, lanceolate, obtuse, pilose; inner sepals ovate, c. 4 mm wide, membranous, abruptly narrowed to a green, pilose mucro; corolla 1.6–1.7 cm long, white, lobed with red, pubescent, midpetaline bands terminating in a broadly triangular lobe; ovary conical, pilose at apex; style pilose, divided c. 4 mm above base, stigmas 3 mm. Capsule and seeds not seen. [
Morocco (
Dwarf plants of
TURKEY, Eski-Scheher,
Compact cushion-forming undershrub with thick woody taproot, branched at base with short woody prostrate to ascending branches to c. 15 cm, young stems sericeous. Leaves sessile, 1.2–2 × 0.3–0.4 cm, linear-oblanceolate, acute, attenuate at the base, sericeous and with some spreading hairs. Flowers solitary, axillary, shortly pedunculate but becoming crowded, sessile and subcapitate towards the branch tips; peduncles 0–5 mm, pilose; bracts leaf-like; bracteoles similar but shorter; sepals 8–11 × 3 mm, oblong, acute to apiculate, concave, adpressed pilose, not bicoloured; corolla 1.7–2.3 cm long, white with distinct pink, pubescent midpetaline bands, unlobed; ovary pilose; style glabrous, divided c. 4 mm above base; stigmas 4 mm. Capsule and seeds unknown.
Endemic to Turkey (
Could be confused with
Although
LEBANON,
Cushion plant with branched woody rootstock, herbaceous flowering stems to 5 cm, vegetative parts adpressed-pilose to subglabrous. Leaves sessile, 1–2.5(-6) × 0.2–0.4 cm, linear to oblanceolate, obtuse to acute, entire, base attenuate; glabrous or very sparsely hairy on the upper surface. Flowers in small terminal diachasial cymes of up to 5 flowers with solitary axillary flowers or axillary cymes; bracts leaf-like; peduncles 0–3 cm; bracteoles 3 × 0.5 mm; sepals 5–8 × 1.5–2 mm, oblong-obovate, scarious with distinct, acute, green triangular apex, inner sepals scarious; corolla 1.2–1.5 cm, white or pink, unlobed, midpetaline bands pilose, terminating in a tooth; ovary pilose; style pubescent, divided c. 2 mm above the base; stigmas c. 4 mm. Capsule pubescent, seeds puberulent. [
Lebanon (
TURKEY,
Perennial cushion-forming plant with woody taproot and short spreading woody branches, the herbaceous parts densely covered with weakly appressed to spreading, stiff hairs. Leaves sessile, dimorphic, small obovate-spathulate leaves 3–5 mm long with a broad base mixed with linear to oblanceolate, acute entire leaves, 1–2 × 0.2–0.5 cm, with an attenuate base. Flowers terminal, solitary (?always), sessile; bracteoles 5–7 mm, linear; outer sepals 3 × 1.5 mm, obovate, cuspidate, whitish, pilose; inner sepals 3 × 2 mm, broadly obovate, cuspidate, corolla pink, 2–2.2 cm, unlobed, midpetaline bands pilose; stamens very unequal; ovary pilose, style thinly pilose, divided c. 3 mm above base; stigmas 6–10 mm long; capsule pilose; seeds puberulent. [
Endemic to Turkey (
A very distinctive species on account of its indumentum, dimorphic leaves, tiny calyx, pink corolla and unusually long stigmas.
TURKEY, Beryt Dagh,
Woody based perennial, branched at base with a thick, woody tap root, somewhat cushion-forming with short spreading woody branches and ascending herbaceous stems to c. 20 cm, adpressed pilose with some spreading hairs. Leaves mostly basal, sometimes folded, sessile, 1–6 × 0.2–0.3 cm, linear or linear-oblanceolate, acute, entire with long tapering base; lower surface and margins pilose, upper surface glabrous except at margins. Flowers in few-flowered terminal heads, occasionally also with solitary sessile or very shortly pedunculate heads from the axils of the uppermost leaves; bracts 10–20 × 1–2 cm, linear, acute; peduncles 0–2 cm; bracteoles 3–6 mm, linear; pedicels 0–1 mm; sepals 6–10 × 2 mm, ovate, the scarious base abruptly narrowed to a green caudate apex 2–3 mm long, shortly pilose; inner sepals with broad, scarious, glabrous margins; corolla 1.5–2 cm long, white, unlobed, midpetaline bands pilose, terminating in a tooth; ovary pubescent; style divided c. 4 mm above base, stigmas 3 mm. Capsule pilose; seeds not seen. [
A rare species of central and eastern Turkey (
Similar to
TURKEY,
Woody based perennial similar to
Endemic to Turkey and apparently rare.
This species may prove only to be a variety or subspecies of
Without locality (lectotype P [Herb. Lam.], designated by
Woody based perennial, branched at base with erect stems to 40 cm; vegetative parts appressed pilose to villous with spreading hairs. Basal leaves 1.5–6 × 1–4 cm. oblanceolate with a long attenuate base; stem leaves sessile, 2.5–5 × 0.1–0.3 cm. linear or oblong, acute or obtuse, entire, somewhat narrowed at the base. Flowers in many-flowered terminal heads; bracts 7–22 × 2–6 mm, oblong-elliptic or ovate, acute; bracteoles 7–8 × 0.5 mm, linear; sepals 8–9 × 2.5–3 mm, narrowly ovate and abruptly narrowed into a long fine point, margin somewhat scarious; corolla 1.5–2. 8 cm long, white or pink with darker midpetaline bands, weakly lobed with broadly triangular lobes; midpetaline bands dark, pilose; ovary glabrous; style glabrous, divided c. 4 mm above base; stigmas c. 4 mm. Capsule glabrous, seeds smooth, pubescent. [
France (
A somewhat variable plant, the type adpressed pubescent, but villous plants with spreading indumentum, e.g.
Plate “
Perennial undershrub to c. 30 cm, the flowering shoots herbaceous with all vegetative parts densely grey-sericeous. Leaves sessile, 2–3.5(-5) × 0.3–0.8(-1.2) cm, oblong to oblanceolate, acute or obtuse, entire, attenuate at base. Flowers in a dense terminal cymose cluster, sometimes with one or two flowers in the axils of bracts immediately below the cluster, borne on peduncles 1–2(-4) cm long; bracts as for leaves but smaller; bracteoles 9–14 × 1 mm, linear, acuminate and apiculate; pedicels 0–3 mm; sepals 7–9 × 2–3 mm, oblong-oblanceolate, acute, densely pilose, the inner sepals broader (c. 3.5 mm) with scarious margins; corolla 2–7 cm, white, unlobed, the midpetaline bands densely pilose; ovary pilose; style glabrous or pilose at base, divided c. 3 mm above the base; stigmas 5 mm. Capsule pilose; seeds pubescent. [
We recognise two varieties:
Plants representative of the type subspecies have linear-oblong leaves and are apparently restricted to Sicily.
Endemic to Sicily (
CROATIA,
Distinguished by the oblanceolate leaves.
Much the most common variety: Croatia (
“Oriente,”
Erect, perennial, much-branched undershrub from a woody rootstock to 1 m, the branches rigid and woody, adpressed pubescent. Leaves sessile, 1.5–6 × 0.2–2 cm, narrowly oblong to narrowly oblanceolate, acute, entire, attenuate at base, villous, the stem leaves smaller than those at the base. Flowers in a large, nearly leafless, branched terminal inflorescence composed of axillary cymes of 1–3(-7)-flowered diachasial cymes, the flowers appearing solitary; branches stout and woody; bracteoles 2–3 mm, linear; pedicels 0–4 mm; sepals 2.5–5 × 2–5 mm, very variable in form, oblong-elliptic to obovate, acuminate to emarginate and mucronate, adpressed pubescent, the inner sepals broader than the outer sepals; corolla 1.2–1.7 cm, pink (very rarely pure white), the midpetaline bands pilose; ovary glabrous, style glabrous, divided c. 5 mm above base, often persistent in fruit, stigmas c. 3 mm; capsule glabrous, seeds subglobose, puberulent. [
We recognise three subspecies but intermediates are quite frequently found:
Branches rigid, woody; stem leaves few; sepals broadly obovate, truncate and mucronate.
Tunisia (
Based on
Branches rigid, woody; sepals oblong-elliptic to lanceolate, acuminate. [
Syria (
Based on
Stems leafy and more or less herbaceous; sepals narrowly oblong-obovate, more or less abruptly narrowed to a mucronate apex.
Apparently abundant in Central Asia: Northeastern Iran (
Western Tian Shan,
As rightly noted by
Endemic to Uzbekistan (
The status of this species is unclear and it may be of hybrid origin.
CANARY ISLANDS, Fuerteventura,
Hummock-forming undershrub 10 to 30 cm high, spreading horizontally to c. 60 cm; stems white-sericeous, stiff, spinescent when old. Leaves mostly alternate but sometimes clustered in short brachyblasts, sessile, 0.4–1.5 × 1.5–2.5 cm, oblanceolate to spathulate, acute, entire, cuneate at base, grey-sericeous with short, appressed scaly hairs. Flowers more or less sessile, axillary, 1 (-2) together; bracts similar to leaves but smaller, 5–6 × 1.5 mm; pedicels c. 1 mm, bracteoles 1.5 × 0.5 mm, oblong, acute; sepals all similar, 3–4 × 1.5–2 mm, broadly oblong-obovate, acute to mucronate, pubescent; corolla 8–10 mm long, white or pale pink, unlobed, midpetaline bands adpressed pilose, reddish; ovary with a bright orange disc, pilose; style pilose, divided c. 2.5 mm above base; stigmas 2.5 mm. Capsule reddish, densely pilose with the style persistent; seeds not seen. [
Canary Islands: Gran Canaria, Fuerteventura (
CANARY ISLANDS, Barrancas,
A branched unarmed undershrub to 2 m, vegetative parts glabrous or sparsely adpressed pilose. Leaves sessile, caducous, 0.5–4.5 × 0. 1 mm, filiform, acute, entire. Flowers (1-) 5–6 in pedunculate, terminal and axillary cymes; peduncles 2–7 (-11) mm, pubescent; bracteoles 2–3 × 1 mm, lanceolate, acuminate, base clasping, appressed to the calyx, pubescent; pedicels 3–7 mm, stout, pubescent; outer sepals 4–6 × 2–2.5 mm, broadly oblong, mucronate; inner sepals obovate, abruptly narrowed to a mucronate apex; corolla 1–1.2 cm long, white or pinkish, deeply lobed, midpetaline bands pilose; ovary pilose; style pilose, divided 3–4 mm above base, stigmas c. 3 mm. Capsule not seen but presumably pilose. [
Endemic to the Canary Islands: Tenerife, Gran Canaria, La Gomera, La Palma (?), El Hierro (
Based on
The hybrid
La Gomera and Tenerife in the Canary Islands (
This taxon represents
CANARY ISLANDS,
A branched unarmed shrub up to 4 m in height, vegetative parts shortly adpressed pubescent, somewhat glabresent on older parts. Leaves sessile, 2–14 × 0.5–2.6 cm, narrowly to broadly oblong, dark green, acute to obtuse, entire, base attenuate. Inflorescemce branches from the upper leaf axils forming a terminal, panicle-like inflorescence of branched cymes with primary branches to 7 cm long, the axes densely pubescent; bracteoles 1 × 0.5 mm, scale-like, caducous, pedicels 2–15 mm; outer sepals 4 × 2 mm, broadly lanceolate, apiculate, the margin ciliolate; inner sepals 4 × 4 mm, mucronate, elliptic or suborbicular, membranous; corolla 1.1–1.5 cm long, white, unlobed, midpetaline bands pilose; anthers exserted; ovary pilose; style pilose, divided c. 3 mm above base; stigmas c. 3 mm. Capsule acute, pilose, 1-seeded; seeds minutely hirsute. [
Endemic to the Canary Islands: Gran Canaria, Tenerife, La Palma, La Gomera, Lanzarote, Fuerteventura (
A very distinctive species because of its shrubby habit and white flowers in a terminal paniculate inflorescence.
All species are perennial undershrubs, usually with leaves densely villous and lacking distinct petioles. Most are unarmed. The flowers are arranged in sessile or pedunculate capitulae. They are plants mostly of the Middle East from Egypt and Saudi Arabia east to Pakistan, but not in the republics of former Soviet Central Asia.
The first two species,
IRAN,
A very variable undershrub 25–50 cm high from a woody rootstock, with woody, branches which are usually spinescent, vegetative parts usually densely white-tomentellous to villous but indumentum sometimes very short; side branches long and slender, reaching 30cm or short, stout and spinescent. Basal leaves sessile 0.7–2.5 (-4.5) × 0.3–0.7, oblong-lanceolate, apex acute and mucronate, margin entire, narrowed to a petiole-like base, both surfaces tomentose to tomentellous, but the adaxial surface often greener; stem leaves and bracts smaller than the lower leaves, commonly ovate, acute to apiculate. Flowers 1–several in numerous subsessile villous axillary capitula (very rarely helicoid cymose in form), forming an elongate spicate inflorescence; peduncles absent, bracteoles 3.5–8 × 1.5–3 mm, lanceolate to oblong, acute; outer sepals 5–8 × 2–3.5, narrowly elliptic to oblong-obovate, acute, villous; inner sepals slightly narrower; corolla 1–1.4 cm, white or pink, weakly lobed, midpetaline bands pilose; ovary and style pilose; style divided c 5 mm above base, the stigmas 2.5 mm long. Capsule not known. [
We recognise two subspecies:
The type subspecies is characterised by the sessile, sometimes clasping, very acute, sometimes apiculate stem leaves and bracts, long slender inflorescence branches with flowers usually 2 -3 together, the sepals acute and the corolla 8–13 mm long. Spinescent side branches are few or absent.
Iraq (
IRAQ, between Ramadi and Rutbah,
Lateral branches numerous, short (2–4 cm), rigid, stout, strongly spinescent. Leaves somewhat caducous. Flowers several together, the inflorescence often elongating at maturity, reaching 3 cm in length
Iraq (
The type of this subspecies is immature and almost flowerless. It could easily be interpreted as a form of
IRAQ,
Undershrub with spinescent branches forming a low bush to 60 cm high and up to 1 m wide; young branchlets white-tomentellous, side branches rather short, c. 5 cm long, sometimes spine-like. Basal leaves 0.5–3 × 0.3–0.6, oblanceolate, obtuse, margin undulate, base attenuate into a long petiole-like base, densely puberulent on both surfaces, paler beneath; stem leaves and bracts 3 × 2 mm, oblong to oblanceolate, obtuse to subacute, cuneate at base. Flowers 1 (-2), sessile in the axils of bracts; bracteoles 2 × 0.5 mm, linear-oblong, pedicels 0–1 mm; sepals 5–6 × 2–3 mm, oblong to oblanceolate, obtuse, densely pilose; corolla 0.6–1cm long, white, slightly lobed, midpetaline bands pilose; ovary hirsute, style glabrous, divided c. 4 mm above base, stigmas 4 mm. Capsule not seen.
Locally abundant principally in sandy desert: Syria (?); Iraq (
Arabian specimens of this species have been misidentified as
This species possibly intergrades with
IRAN, Shiraz,
Weakly cushion-forming plant with a woody rootstock, from which arise ascending stems to 45 cm, vegetative parts pilose with somewhat stiff spreading hairs. Basal leaves 3–8 × 0.5–0.9 cm, oblong-oblanceolate, acute, entire, base long-attenuate and petiole-like up to 4 cm in length; stem leaves 2–5 × 0.5–1 cm, oblanceolate to oblong, acute, entire, base cuneate to attenuate. Flowers in sessile (above) or shortly pedunculate (below) 2–3-flowered axillary heads; bracts usually < 3 cm long, lanceolate (if longer, than narrowly lanceolate); peduncles 0–1.5 cm; bracteoles 9–11 × 1 mm, linear-oblong, acute, densely pilose; pedicels absent; outer sepals 9–11 × 2–2.5 mm, lanceolate, acuminate, densely pilose; inner sepals 8–9 × 1 mm, linear-lanceolate; corolla 2–2.5 cm long, pink, shallowly lobed, midpetaline bands darker, adpressed pilose; ovary glabrous; style glabrous, divided 3 mm above base, stigmas 4.5–5 mm; capsule glabrous; seeds glabrous. [
Pakistan; southern Iran (
This species has been confused with
AFGHANISTAN,
Vigorous perennial with stout, woody rootstock and stem base, to 50 cm high, the whole plant densely covered in long, soft white hairs. Basal leaves 3–13 × 0.6–2.5 cm, oblong-lanceolate, acute at both ends and with a long winged petiolate base; stem leaves sessile, 3–12 × 0.3–3.8 cm, elliptic or oblong-elliptic, acute, entire, broadly to narrowly cuneate at base, diminishing in size upwards, veins prominent. Inflorescence of sessile (above) or shortly pedunculate (below) capitula or scorpioid cymes arising from the upper part of the stem; bracts mostly 3–4 cm × 1.2–2 cm, ovate; peduncles 0–3 cm, bracteoles 12–23 × 1.5–3 mm, linear-lanceolate, acuminate; pedicels 0–1 mm; outer sepals 10–14 × 3.5 mm, ovate, acuminate, covered with an indumentum of short dense hairs mixed with longer spreading hairs; inner sepals convex, glabrous; corolla 2–2.2 cm. white, weakly lobed, midpetaline bands pilose; ovary and style glabrous, style divided c. 2.5 mm above base, stigmas c.2.5 mm; capsule glabrous, seeds pubescent. [
Afghanistan (
Based on
Undershrub from a woody base to 35 cm, the lower branches woody and spinescent, younger brances herbaceous; vegetative parts softly white-tomentose with spreading hairs. Leaves sessile, 1–3 × 0.3–0.5 cm, oblanceolate below, oblong above, acute, margin entire, base cuneate. Flowers in subsessile heads along the upper part of the stem; bracts oblong-elliptic to ovate; peduncles 0–5 mm; bracteoles 10 × 5 mm, ovate; pedicels absent; outer sepals 10 × 4 mm ovate, acute; inner sepals 7 × 2 mm, lanceolate, much narrower than outer sepals; corolla 1.8–2.3 cm, pale pink or white, somewhat undulate, midpetaline bands pilose; ovary and style glabrous; style divided 5 mm above base, the stigmas 4 mm. Capsule glabrous; seeds smooth, glabrous (fide
Lower Egypt and Sinai (
Replaces
“Levant”,
Perennial herb from a woody base, stems prostrate, reaching 50 cm, the whole plant densely villous. Leaves sessile, 2–4 × 0.4–1.3 cm, oblanceolate or oblong, becoming elliptic upwards, obtuse, margin entire, lower leaves attenuate at base, upper leaves and bracts cuneate, veins prominent, internodes short so leaves and flower heads more crowded than in related species. Inflorescence characteristically narrow, flowers 1–5 in subsessile, congested head-like scorpioid cymes; peduncle 0–1.5 cm; bracteoles 8–11 × 3–5 mm, lanceolate to ovate, acute; pedicels absent; outer sepals 12–15 × 3–4 mm, oblong, acuminate, flat; middle sepal asymmetric; inner sepals slightly shorter and narrower; corolla 2–2.8 cm long, usually creamy with a yellowish centre, weakly lobed, midpetaline bands long-sericeous; ovary and style glabrous, style divided 6 mm above base, stigmas 3mm; capsule glabrous, 1-seeded (fide
Turkey (
Unlike
Species 178–181 are extremely similar and may prove to be conspecific.
EGYPT, Sinai, Wadi Feiran,
Villous perennial to 40 cm. Leaves sessile, 1.3–7 × 0.4–1.2 cm, oblong to oblanceolate, acute, margin entire, attenuate at the base, softly white-villous. Flowers in dense axillary heads; peduncles 1.5–3.5 cm, exceeding bracts; bracteoles 6–15 × 1–2 mm, linear-lanceolate, acuminate, densely villous; outer sepals 7–10 × 2–3 mm, lanceolate, acuminate, inner sepals narrower, 1–2 mm wide; corolla 2–2.5 cm, unlobed, midpetaline bands pubescent, terminating in a tooth; ovary and style glabrous; style divided c. 3 mm above base, the stigmas 5 mm, longer than the united part of the style. [
Egypt: Sinai (
Differs from the two previous species in the pedunculate heads but superficially identical to
The lectotype is a meagre specimen and all the other cited specimens represent more complete material.
JORDAN,
Similar to the previous species. Perennial to 40 cm; stems slightly zigzag. Leaves sessile, 1.3–4 × 0.3–0.4 cm, oblong, acute (not spinescent as stated by Sa’ad), margin entire, shortly tomentose. Flowers in dense axillary heads, becoming scorpioid at maturity; peduncles 1.5–3 cm, exceeding bracts; bracteoles 6–7 × 3.5–4 mm, ovate, caudate (not spinescent), densely villous; outer sepals 7 × 3 mm, elliptic, shortly acuminate or caudate, inner sepals oblong, c. 1 mm wide; corolla 2–2.5 cm, unlobed, midpetaline bands pubescent, terminating in a tooth; ovary and style glabrous; style divided c. 2 mm above base, the stigmas 5–6 mm long.
Endemic to Jordan where it is known from one unlocalised record.
The most distinct species in this complex because of the narrowly oblong, shortly tomentose leaves combined with the pointed bracteoles and sepals but only known from the type.
EGYPT, Sinai,
Perennial herb from a stout woody rootstock to 50 cm in height, all parts covered in long spreading hairs, which are rust-coloured when dry. Basal leaves petiolate, 1.5–6.5 × 0.5–1 cm, obovate or oblanceolate, obtuse, margin strongly undulate, with long petiole-like base; stem leaves sessile, shorter (< 2.5 cm long), linear-oblanceolate. Flowers in pedunculate heads with up to 5 flowers; bracts 1–1.5 × 0.5–0.7 cm, oblong-elliptic; peduncles 0–11 mm long, about equalling the subtending bract; bracteoles 12 × 2 mm, linear-oblanceolate; pedicels absent; outer sepals 10–11 × 2–3 mm, lanceolate, acute, villous; inner sepals narrower and shorter, c. 8–9 × 1 mm; corolla 2–2.2 cm, colour unknown, wavy at the apex, scarcely lobed, midpetaline bands pilose; ovary and style glabrous divided c. 5 mm above base, stigmas c. 5 mm. Capsule not seen. [
Endemic to Sinai in Egypt (
Distinguished from
IRAN, Makran,
Perennial herb from a woody taproot, from which arise ascending stems 30–35 cm high. Branches stout; whole plant densely pubescent with grey hairs. Leaves sessile, 1.5–4 × 0.4–1 cm, oblong-oblanceolate, obtuse, margin crisped-undulate, attenuate to a long petiole-like base, sometimes the abaxial surface notably paler. Inflorescence of axillary, pedunculate heads, which are occasionally elongated into more or less scorpioid cymes; peduncles 1.2–1.5 cm; bracteoles 8 × 5 mm, ovate-elliptic, acute or shortly acuminate; outer sepals 10–12 × 3 mm, broadly lanceolate to ovate, acuminate; inner sepals shorter and narrower, c. 8 × 2 mm; corolla 1.5–2 cm, pinkish, unlobed, midpetaline bands pilose; ovary comose; style glabrous or with a few hairs, divided c. 2.5 mm above base, stigmas c. 2 mm long; capsule comose, seeds pubescent. [
We recognise two, not very well-defined subspecies, which are bridged by intermediates:
Leaves relatively short, usually less than 5 cm long, indumentum asperous. Style glabrous or with a few hairs.
Pakistan (
One collection from Iranian Baluchestan (
Fruiting heads often become distinctly scorpioid in structure as in
Based on
Differs from
Principally around the head of the Persian/Arabian Gulf: Iran (
IRAQ [probably], inter Arrah et Deïr,
Perennial herb from a woody taproot and base with stems to 40 cm, plant roughly tomentose with longish white hairs. Basal leaves 5–11 × 0.6–1.7 cm, oblanceolate, obtuse or rounded, margin entire to slightly undulate, base narrowed into a pseudopetiole c. 2–4 cm long; stem leaves and bracts sessile, 2–5 × 1.5–2 cm, ovate, acute, base broadly cuneate. Flowers in many-flowered, axillary, pedunculate heads, mostly 2–2.5 cm in diameter, occasionally somewhat elongate; peduncles 1.5–5 cm; bracteoles 10–15 × 1–3 mm, linear to lanceolate, long acuminate, pilose; pedicels absent; sepals 8–9 × 2.5–3 mm, lanceolate, acuminate, long-pilose, inner sepals slightly narrower, c. 2 mm wide; corolla 2–2.3 cm long, pink, midpetaline bands pilose, very shallowly lobed with midpetaline bands terminating in a comose point; ovary glabrous; style glabrous, divided 5 mm above base, stigmas 4 mm. Capsule glabrous; seeds glabrous. [
Iran, Iraq (
Resembles
SAUDI ARABIA, Asir,
Cushion-like perennial from a woody rootstock with decumbent to ascending stems up to 75 cm long, plant densely white-tomentellous. Basal leaves petiolate, 0.3–2 × (0.1-)0.4–0.8 cm, broadly oblanceolate, acute, base cuneate, petiole up to 2 cm long; stem leaves sessile, 2–5 × 0.5–0.8 cm, similar in shape, diminishing in size upwards and merging into bracts. Inflorescence of sessile axillary clusters towards the branch tips, raceme-like; bracteoles 3 × 2 mm, elliptic, acute; outer sepals 7 × 2 mm, oblong-lanceolate, acute, pilose; inner sepals narrower (c 1 mm wide) and slightly shorter, the margins membranous; corolla 1.6–2 cm, pink or white, undulate, midpetaline bands thinly pilose; ovary comose; style glabrous; capsule comose, seeds glabrous (or puberulent fide R.R.Mill). [
Endemic to Saudi Arabia: Asir region (
Distinguished by the cushion habit, sessile heads and usually small leaves.
AFGHANISTAN/PAKISTAN, Kurram Valley,
Cushion/hummock forming plant with woody taproot, branched just below the ground surface to produce many very short shoots from which arise flowering stems 1–9 cm high. Leaves aggregated at the shoot tips, sessile, 2–2.5 × 0.2–0.3(-0.4) cm, oblanceolate, acute, margin entire, attenuate at the base, densely villous. Flowers 1–4 in subsessile helicoid cymes, usually reduced to heads which arise in the axils of the upper bracts; bracts 1.5 cm long, oblong, villous; peduncles 0–2 mm; bracteoles 10 × 0.5 mm, linear, acuminate, villous; pedicels absent; outer sepals villous; outer sepals 11–13 × 2–3 mm, ovate acuminate; inner sepals narrower, lanceolate c. 2 mm wide; corolla 1.4–1.5 cm long, cream (?), weakly lobed, midpetaline bands pilose; ovary and style glabrous; style divided c. 3 mm above base, stigmas 4–4.5 mm long; Capsule and seeds not known. [
Pakistan (
Distinguished by its cushion-like habit and subsessile heads borne on a short stem.
IRAQ,
Prostrate perennial to 50 cm from a woody taproot, the whole plant densely velvety-tomentose of characteristic dark brown colour in herbarium specimens; stems 2–3 mm thick. Leaves bullate above, reticulate and greener beneath, becoming leathery when old; lower leaves up to 9 × 2.2 cm, oblong-elliptic, obtuse, entire, attenuate at base; stem leaves and bracts ovate, acute, abruptly rounded at base. Flowers in pedunculate (below) or subsessile (above) or all subsessile, axillary several-flowered heads; bracts c. 1.5–3 × 1–2.2 cm, ovate, nearly as broad as long; peduncles 0–5 cm, tomentose; bracteoles 6–10 × 2–5 mm, lanceolate to elliptic, acute; sepals 8–11 × 1.5–2.5 mm, lanceolate, acuminate, pilose; inner sepals narrower, c. 0.5–1 mm wide; corolla 1.2–1.5 cm long, white, unlobed but slightly undulate, midpetaline bands adpressed brown-pilose; ovary glabrous; style glabrous, divided c. 3 mm above base; stigmas 4 mm; capsule glabrous; seeds pubescent (fide Sa’ad). [
We recognise two subspecies:
Stems relatively slender 2–3 mm wide, bracteoles lanceolate 2–3 mm wide, sepals lanceolate, 1.5 to 2.5 cm wide.
Western Iran (
Based on
Distinguished by its stout stems, c. 5 mm thick, elliptic bracteoles 4–5 mm wide and obovate sepals c. 4 mm wide. [
Iran (
Appears to be mostly sympatric with
IRAN, W. Kazerun,
Perennial herb from a woody rootstock stems to c. 30 cm, the whole plant densely velvety-tomentose of characteristic dark brown colour in herbarium specimens. Basal leaves unknown; stem leaves sessile, 4–7 × 1.2–2 cm, oblong-lanceolate, acute, margin entire, base cuneate, leathery in texture. Flowers in sessile clusters in the axils of bracts along the upper part of the stem; bracts triangular-ovate, 3.5 cm long below but diminishing in size upwards, bracteoles 10 × 1 mm, linear, villous; sepals 10–11 × 3 mm, ovate, finely acuminate to a long aristate point, densely villous; corolla 2–2.2 cm long, colour unknown, unlobed, midpetaline bands adpressed pilose; ovary apically pilose; style glabrous, divided c. 12 mm above base; stigmas (fide
Endemic to Iran (
Shares with
IRAN,
Perennial herb from a woody rootstock with stems to at least 40 cm, the whole plant densely but very shortly tomentose leaving leaf veins visible; branches somewhat woody. Lower leaves c. 8 × 1.5 cm, oblanceolate, narrowed to a long petiole-like base; stem leaves 2.5–4.5 × 1.5–2.5 cm, ovate, obtuse, margin entire, base cordate and subsessile. Flowers in dense axillary heads, sessile above, shortly pedunculate below; bracts 1.3–2.5 × 1–2.2 cm, ovate, acuminate, basally cordate; peduncles 0–1.5 cm; bracteoles 10–15 × 1–2 mm, linear, long-pilose; pedicels absent; outer sepals 14–16 × 3.5–4 mm, ovate, abruptly narrowed, rounded and with a long, aristate point, long pilose; inner sepals somewhat smaller, c. 12 × 1.5–3 mm; corolla 1.5–1.8 cm, pink, midpetaline bands long-pilose; ovary comose; style glabrous, divided c. 6 mm above base, stigma 3 mm; capsule not seen. [
Endemic to Iran (
Sa’ad described the ovary as glabrous but it is comose as in her illustration.
IRAN,
Perennial undershrub to about 25 cm with stems and branches rigid and somewhat spinescent, tomentellous with matted hairs. Leaves sessile, 1–2.5 × 0.2–0.8 cm, linear-oblanceolate, obovate-elliptic to subspathulate, acute, margin entire, base attenuate, adpressed pubescent with matted hairs. Inflorescence of more or less globose terminal heads; bracts 9–12 × 4–6 mm, broadly ovate, acute; pedicels 0; bracteoles 14 × 1–1.5 mm, lanceolate, acuminate; outer sepals 10–12 × 2.5–3.5mm, oblong-elliptic, acute, densely pilose; inner sepals similar but lanceolate; corolla 1.5–1.8 cm long, colour unknown, midpeline bands pilose; ovary and style glabrous; style divided 3 mm above base, stigmas c. 6 mm long; capsule glabrous. [
Endemic to southern Iran (
This species has an inflorescence of terminal heads like
OMAN, Muscat,
Intricately branched spiny undershrub to c. 30 cm, both the main and side branches spinescent, young stems sericeous. Leaves sessile, (6-) 8–11 × 4–6 mm, obovate or spathulate, apex rounded, margin entire, base attenuate, sericeous especially abaxially. Flowers in dense sessile, axillary clusters of up to 4; bracts 2–4 × 1–3 mm, oblong-elliptic; peduncles absent; bracteoles 1–3 × 1–2 mm, elliptic, apiculate or retuse, sericeous and with spreading hairs; outer sepals 5–6 × 2 mm, narrowly elliptic, acute, densely covered in long woolly hairs; inner sepals similar but smaller, c. 4 × 1 mm; corolla 11 mm long, white, midpetaline bands woolly; ovary apically pilose; style glabrous, divided 2–3 mm above base; stigmas 6 mm. Capsule not seen. [
Almost restricted to Oman (
A little-known species similar to
PAKISTAN, Sind,
Perennial undershrub with rigid but not spinescent branches, 20–60 cm high, branches softly grey-tomentellous. Leaves alternate or more or less clustered, sessile, coriaceous, 5–15 × 4–6 mm, obovate or elliptic, rounded to obtuse, margin undulate, gradually narrowed at base, grey or rufous-tomentose, veins strongly impressed. Inflorescence of sessile terminal heads covered in reddish-brown hairs; bracteoles 6–7 × 4 mm, obovate-elliptic, lacking veins, villous; sepals 6–7 × 1.5 mm, lanceolate, densely reddish-brown villous; corolla 10–12 mm long, white, shallowly lobed, midpetaline bands pubescent; ovary and style glabrous; capsule glabrous, apparently always 1-seeded; seeds glabrous, smooth. [
Probably endemic to Pakistan (
Very distinctive because of the reddish-brown-haired terminal heads combined with the woody, but not spinescent, branches and the small, often clustered leaves with strongly impressed veins.
The following is a list of species placed in
We have had the help of numerous people and institutions during the preparation of this monograph and wish to thank them all, Many institutions have loaned specimens or sent images to clarify specific issues or provide material for artwork or identification. These are AD, B, BM, BRI, C, E, FI, FT, G, GOET, HBG, K, KW, LISE, M, MA, MICH, MO, NY, P, PRE, TEX, UPS, URT and W. We are also grateful to curators and staff at institutions visited by John Wood during the preparation of this monograph, Jonathan Gregson, John Hunnex and Jacek Wajer at BM, Antonio Krapovickas and Maria Mercedes Arbo at CTES, Adele Smith and Suzanne Cubey at E, the former for processing loans and the latter DNA samples, David Goyder and Tim Harris at K, Alexei Oskolski at LE, Stephan Beck at LPB, Thierry Deroin at P and Ernst Vitek at W. We have had useful discussions and correspondence with many people in different parts of the world and would like to thank the following for their interest and comments: George Staples, R.W. (Bob) Johnson (BRI), now sadly deceased, Chiara Nepi and Piero Cuccuini (FT), Rosangela Bianchini (SP), Priscilla Ferreira (ICN), Tom Wendt (Texas) and Alfredo Reyes Betancort (Canary Islands), all of whom have contributed in different ways to the improvement of this monograph. Dick Brummitt (deceased), Nicholas Hind and Nick Turland provided helpful advice on nomenclatural matters. We would also like to thank Shahina Ghazanfur (
All references cited in the text are provided in full below. However the user’s attention is drawn to the increasing availability of images and particularly photographs on the internet. The following website, not specifically cited in the text, for example, provides photographs and distribution maps of all Italian species: