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An undescribed species ofPentanemaCass. (Asteraceae–Inuleae) from Turkey; with notes on the phylogenetic status of the genus
Botanical Journal of the Linnean Society (1999), 129: 333–338. With 1 figure Article ID: bojl.1998.0228, available online at http://www.idealibrary.com on
An undescribed species of Pentanema Cass. (Asteraceae–Inuleae) from Turkey, with notes on the phylogenetic status of the genus HAYRI DUMAN Gazi University, Faculty of Science and Literature, Department of Biology, Teknikokullar, Ankara, Turkey ARNE A. ANDERBERG∗ Department of Phanerogamic Botany, Swedish Museum of Natural History, P.O. Box 50007, S-104 05 Stockholm, Sweden
Received July 1997; accepted for publication September 1998
A new species of Pentanema Cass. (Asteraceae–Inuleae), Pentanema alanyense H. Duman & Anderb., sp. nov. is described. This is the first record of a species of this genus from Turkey. An illustration of the new species and a brief description of its habitat is provided along with a discussion of the phylogenetic status of the genus Pentanema with reference to generic delimitations in the Inula complex. 1999 The Linnean Society of London
ADDITIONAL KEY WORDS:—Systematics.
CONTENTS
Introduction . . . . . . . . . . The Inula complex . . . . . . . . Pentanema Cass. . . . . . . . . . Pentanema alanyense H. Duman & Discussion . . . . . . . . . . Acknowledgements . . . . . . . References . . . . . . . . . .
. . . . . . . . . Anderb. . . . . . . . . .
. . . . . . . . . . . . sp. nov. . . . . . . . . . . . .
. . . . . . .
. . . . . . .
. . . . . . .
. . . . . . .
. . . . . . .
. . . . . . .
333 334 334 335 337 338 338
INTRODUCTION
A few years ago, the senior author (H.D.) collected material of a daisy in the mountains of Antalya in southern Anatolia in Turkey. The plant was a perennial ∗ Corresponding author. Email: [email protected] 0024–4074/99/040333+06 $30.00/0
333
1999 The Linnean Society of London
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H. DUMAN AND A. A. ANDERBERG
herb with delicately branched stems and small radiate capitula. It was identified as belonging to the Inula complex of the tribe Inuleae s.s., as outlined by Anderberg (1989, 1991), but its generic position could not be ascertained. The plant has a unique set of characters, involving a distinctly conical epaleate receptacle, and large bulb-like resting buds basally, formed by thick silky-hairy leaves. The cypselas have a number of sclerenchymatous ribs, as in the genus Inula L., but only very few capillary pappus bristles as in the genus Pentanema Cass. It soon became clear that the material represented an undescribed species, but less clear as to in which genus it should be placed.
THE INULA COMPLEX
The only cladistic analysis of morphological data so far undertaken in the Inuleae s.s. (Anderberg, 1991) identified an informal, loosely composed group of genera, the paraphyletic or possibly polyphyletic Inula complex. The assemblage comprised those genera of the tribe that combined epaleate receptacles with a pappus of scabrid or barbellate capillary bristles, but lacking scales or other specializations characterizing other genera, or groups of genera. The complex included the genera Inula L. (incl. Codonocephalum Fenzl), Limbarda Adans., Varthemia DC., and Pentanema, with close but unresolved relationships to genera such as Duhaldea DC., Dittrichia Greuter, Chrysophthalmum Sch.Bip., Amblyocarpum Fisch. & Mey., and even Carpesium L. Also the relationships to the Pulicaria complex, i.e. taxa with double pappus composed of both capillary bristles and short scales, was pointed out as interesting in relation to the phylogeny of the Inula complex. A recent study of a few taxa from the Inuleae s.s., based on nucleotide sequences from the Internal Transcribed Spacer (ITS) region of the 18S–26S nuclear ribosomal DNA (Eldena¨s, Anderberg & Ka¨llersjo¨, 1998), revealed that the phylogenetic interrelationships in the tribe may be even more complex, with paleate genera with scaly pappus such as Asteriscus Tourn. ex Mill., and Buphtalmum L. being more closely related to the Pulicaria complex, and the likewise paleate but epappose genus Telekia Baumg. being closely related to Inula. A detailed investigation of both morphological and molecular data is thus needed before a well corroborated hypothesis of generic relationships in the tribe can be gained. Most of the genera of the Inula complex are monospecific, very small and homogeneous, or easily diagnosed due to morphological specialization, e.g. Dittrichia Greuter. The central problem is the heterogeneity and possible polyphyly of Inula. It is likely that Inula should be split into smaller, more homogeneous and monophyletic groups, each related to various other genera of the complex. Apart from the genera of the Inula complex that are diagnosed by particular apomorphic features, only two genera remained: the large residual genus Inula, diagnosed by the mere absence of the characters diagnosing the other genera, and the medium sized genus Pentanema.
PENTANEMA CASS.
The genus Pentanema Cass. (syn. Vicoa Cass.) is a medium sized genus of about 18 species, mainly distibuted in Central Asia, Iran, Afghanistan, Iraq, Pamir-Altai, and
A NEW SPECIES OF PENTANEMA
335
Tien-Shan. Most species are confined to rather restricted areas of distribution. The annual, weedy species have a wider distribution within East Africa, India, Indonesia and China (Gorschkova, 1959; Ling, 1979; Rechinger, 1980). The genus was originally diagnosed from its pappus composed of a few capillary bristles only, but several species have up to 10–15 bristles (Rechinger, 1980). The genus has no synapomorphy found in all its species, but rather characters that are nested within each other. The perennial species have lanate resting buds on the woody stems, a feature that Rechinger (1980) referred to as ‘gemmae quiescentes’, and which was interpreted as a synapomorphy of a large group of Pentanema species by Anderberg (1991). Within that group of species the number of pappus bristles varies from 5 to 15. Many of these species have disciform capitula with an outer row of almost tubular ray-florets, but there is a continuous variation from species with very reduced ray-floret lamina to species with well developed rays. The annual species in Pentanema have well developed ray-florets, and like most perennials they have ovate or elliptic leaves and almost unsclerified cypselas with few bristles. The lack of apparent synapomorphies diagnosing the genus indicate that it does not constitute a monophyletic group as presently circumscribed, and that some species may have to be removed. We conclude that the undescribed species is best placed in the genus Pentanema, sharing specialized characters with several of the perennial species near the type of the genus, few (4–5) pappus bristles, silky-hairy resting buds (‘gemmae quiescentes’), divaricately branched stems with delicate branches, are characters that are found in species of Pentanema but hardly in Inula or any other genus of the tribe.
Pentanema alanyense H. Duman & Anderb. sp. nov. (Fig. 1) Type: Turkey C4, Antalya, Alanya, Cayarasi-Gevne, 1240 m, clearings of Pinus nigra forest. 2.ix.1993, H. Duman 5504 & F. A. Karaveliogullar. Holotype: GAZI; Isotypes: E, S). Diagnosis. Species nova ab omnibus aliis speciebus generis Pentanematis receptaculo distincte conico, habitu omnino herbaceo et gemmis quiescentibus forma ad instar bulborum sericeopilosis differt et praesentia typica etiam radicum multarum secundariarum absentiaque radicis primariae dominantis insignis. Speciei annuae P. divaricato aspectu (praecipue propter ramificationem divaricatam) maxime simile P. alanyense est sed haec species a P. alanyensi forma folii, receptaculo fere plano et in involucro indumento denso pilorum longorum patentium obtecto recedit. Description. A small delicate perennial herb, basally with several bulb-like resting buds covered with long silky hairs. numerous but dominating primary root absent. terete, up to c. 15 cm high and about 1 mm wide, divaricately branched in the upper part with almost thread-like branches, sparingly branched below, reddish-brown, glandular-hairy and with scattered bristly hairs. linear to lanceolate, lower up to 8 cm long and 7 mm wide, gradually attenuating towards the base; lamina with minute glandular hairs; margin almost entire or with minute and inconspicuous teeth; upper leaves linear, sessile; leaves of resting buds involute, 3–12 mm, glaucous to purple, broad. c. 5 mm wide, heterogamous, solitary
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Figure 1. Pentanema alanyense H. Duman & Anderb sp. nov. A, habit of plant; B, bulb-shaped resting-bud; C, capitulum; D, capitulum (L. S.) showing conical receptacle; E, ray-floret; F, disc-floret; G, cypsela with pappus bristles; H, anther; I, anther-tails; J, endothecial tissue; K, style-branches. A–K, Duman 5504 (S).
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T 1. List of taxa growing in the type locality of Pentanema alanyense. Taxa marked with ∗ are local endemics in southern Anatolia Ostrya carpinifolia Scop. ∗Gypsophila curvifolia Fenzl ∗Potentilla isaurica (P. H. Davis) Pawł ∗Cicer isauricum P. H. Davis ∗Bupleurum subuniflorum Boiss. & Heldr. ∗Eryngium palmito Boiss. & Heldr. ∗Ferulago isaurica Pesmen Pimpinella tragium Vill. ssp. lithophila (Schischkin) Tutin Ampelopsis orientale (Lam.) Planch. ∗Digitalis davisiana Heywood ∗Origanum bilgeri P. H. Davis Origanum saccatum P. H. Davis ∗Phlomis monocephala P. H. Davis
Sideritis arguta Boiss. & Heldr. ∗Sideritis argyrea P. H. Davis ∗Sideritis erythrantha Boiss & Heldr. var. cedretorum P. H. Davis ∗Thymbra sintenisii Bornm. & Aznav. ssp. isaurica P. H. Davis ∗Cephalaria gazipashensis Su¨mbu¨l ∗Cephalaria isaurica V. A. Matthews ∗Campanula myrtifolia Boiss. & Heldr. Campanula trachyphylla Schott & Kotschy Chrysophthalmum dichotomum Boiss. & Heldr. ∗Leucocyclus formosus Boiss. ssp. amanicus (Rech. f.) Hub.Mor. & Grierson
in the end of the branches. Involucral bracts 2–4 mm, glandular-hairy, in several rows, imbricate; outer bracts green, obtuse; inner bracts scarious, acute, marginally fimbriate. Receptacle conical, epaleate, glabrous. Ray-florets in one row, female, radiate, yellow, with scattered glandular hairs; lamina c. 5 mm long, with elongated epidermis cells, apically 3-dentate. as in disc-florets, but pappus of 3(−4) capillary bristles. - yellow, perfect, 5-lobed, c. 3 mm long; corollalobes short, glandular-hairy; anthers ecalcarate, with short slightly branched tails; endothecial tissue radial, i.e. with thickenings confined to the vertical cell walls; filament collar with square cells; style bifid with slightly enlarged base; style-branches slightly flattened, obtuse, dorsally with acute sweeping hairs not reaching the furcation; stigmatic surface forming two apically confluent bands. c. 1 mm, elliptic, sparsely hairy with 8–10 sclerenchymatous ribs; epidermis with elongated crystals; pappus of 4(−5) barbellate, c. 2 mm long capillary bristles; carpopodium oblique, white; cells thick-walled, in several rows. The new species differs from all other species of Pentanema by its distinctly conical receptacle, entirely herbaceous growth habit combined with silky-hairy, bulb-like basal resting buds. Also, the presence of a large number of secondary roots and absence of a dominating primary root is typical. In general appearance, Pentanema alanyense is most similar to the annual P. divaricatum Cass., mainly because of its divaricate branching, but the latter species differs, e.g. in leaf-shape, in having an almost flat receptacle and a dense indumentum of long spreading hairs on the involucre. Habitat. The new species was found growing on calcareous rock in clearings in Pinus nigra forests. A number of species with restricted distribution, several being local endemics of southern Anatolia, were found growing in the same locality (Table 1.). Other examined material Turkey C4, Antalya, Alanya, Hadim-Alanya, 1250–1300 m., on calcareous rocks, 9.viii.1994, H. Duman, Z. Aytac & A. Duran 5597 (GAZI).
DISCUSSION
From a biogeographical point of view, the presence of Pentanema alanyense in southern Turkey is also interesting. The genus has its highest species diversity in
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H. DUMAN AND A. A. ANDERBERG
Iran, Afghanistan and Pamir-Altai. The annual species P. divaricatum is found in Arabia, the Middle East and Central Asia. The closest occurrence of a perennial species is P. flexuosum in northern Iraq. The discovery of a Pentanema in Antalya widens the total distribution range of the genus to include the Eastern Mediterranean region, and adds to the number of endemic taxa that show a disjunct distribution between Antalya and Konya in Turkey with Iran and Afghanistan. Other examples with the same distribution pattern include Thermopsis turcica Kit Tan, Vural & Ku¨cu¨ko¨du¨k with T. alpina (Pall.) Ledeb. from Turkestan (Tan, Vural & Ku¨cu¨ko¨du¨k, 1983), Arnebia purpurea Erik & Su¨mbu¨l with A. euchroma (Royle) I. M. Johnst. from Afghanistan and Iran (Erik & Su¨mbu¨l, 1986), Pyrus serikensis Gu¨ner & H. Duman with P. boissieriana from Iran (Gu¨ner & Duman, 1994), and Aethionema alanyae H. Duman with the genus Moriera Boiss. from Iran (Duman, 1994).
ACKNOWLEDGEMENTS
H.D. wishes to thank the Edinburgh Botanic Garden (Sibbald) Trust for financial support during his visit to Edinburgh, and the Curator of the Royal Botanic Garden Edinburgh for study facilities. The new species was collected during the [Turkish] Endemic Plants Project, which is funded by TUBITAK (Project No.: TBAG-DPT/ C.SEK4). The authors thank Dr Robert Mill (Royal Botanic Garden Edinburgh) for the Latin diagnosis and Ms Pollyanna Lidmark (Swedish Museum of Natural History) for the illustration.
REFERENCES
Anderberg AA. 1989. Phylogeny and reclassification of the tribe Inuleae (Asteraceae). Canadian Journal of Botany 67: 2277–2296. Anderberg AA. 1991. Taxonomy and phylogeny of the tribe Inuleae (Asteraceae). Plant Systematics and Evolution 176: 75–123. Duman H. 1994. A new species from South Anatolia – Aethionema alanyae (Brassicaceae). The Karaca Arboretum Magazine 2: 171–176. Eldena¨s P, Anderberg AA, Ka¨llersjo¨ M. 1998. Molecular phylogenetics of the tribe Inuleae s. str. (Asteraceae), based on ITS sequences of nuclear ribosomal DNA. Plant Systematics and Evolution 210: 159–173. Erik S, Su¨mbu¨l H. 1986. Three new taxa from Turkey. Notes from Royal Botanic Garden Edinburgh 44: 151–156. Gorschkova SG. 1959. Pentanema Cass. In: Schischkin BK, ed. Flora SSSR 25. Izdatelstvo Akademij Nauk SSSR. Moscow and Leningrad, 477–486. Gu¨ner A, Duman H. 1994. Critics on Pyrus boissieriana Buhse subsp. crenulata Browicz. The Karaca Arboretum Magazine 2: 165–170. Ling Y. 1979. Pentanema Cass. In: Ling Y, ed. Flora reipublicae popularis sinicae 75. Academia Sinica, 281–286. Rechinger KH. 1980. Pentanema Cass. In: Rechinger KH, ed. Flora Iranica 145. Compositae IV – Inuleae. Graz: Akademische Druck- und Verlagsanstalt, 96–105. Tan K, Vural M, Ku¨cu¨ko¨du¨k M. 1983. An unusual new Thermopsis from Turkey. Notes from Royal Botanic Garden Edinburgh 40: 515–518.
An undescribed species of Pentanema Cass. (Asteraceae–Inuleae) from Turkey, with notes on the phylogenetic status of the genus HAYRI DUMAN Gazi University, Faculty of Science and Literature, Department of Biology, Teknikokullar, Ankara, Turkey ARNE A. ANDERBERG∗ Department of Phanerogamic Botany, Swedish Museum of Natural History, P.O. Box 50007, S-104 05 Stockholm, Sweden
Received July 1997; accepted for publication September 1998
A new species of Pentanema Cass. (Asteraceae–Inuleae), Pentanema alanyense H. Duman & Anderb., sp. nov. is described. This is the first record of a species of this genus from Turkey. An illustration of the new species and a brief description of its habitat is provided along with a discussion of the phylogenetic status of the genus Pentanema with reference to generic delimitations in the Inula complex. 1999 The Linnean Society of London
ADDITIONAL KEY WORDS:—Systematics.
CONTENTS
Introduction . . . . . . . . . . The Inula complex . . . . . . . . Pentanema Cass. . . . . . . . . . Pentanema alanyense H. Duman & Discussion . . . . . . . . . . Acknowledgements . . . . . . . References . . . . . . . . . .
. . . . . . . . . Anderb. . . . . . . . . .
. . . . . . . . . . . . sp. nov. . . . . . . . . . . . .
. . . . . . .
. . . . . . .
. . . . . . .
. . . . . . .
. . . . . . .
. . . . . . .
333 334 334 335 337 338 338
INTRODUCTION
A few years ago, the senior author (H.D.) collected material of a daisy in the mountains of Antalya in southern Anatolia in Turkey. The plant was a perennial ∗ Corresponding author. Email: [email protected] 0024–4074/99/040333+06 $30.00/0
333
1999 The Linnean Society of London
334
H. DUMAN AND A. A. ANDERBERG
herb with delicately branched stems and small radiate capitula. It was identified as belonging to the Inula complex of the tribe Inuleae s.s., as outlined by Anderberg (1989, 1991), but its generic position could not be ascertained. The plant has a unique set of characters, involving a distinctly conical epaleate receptacle, and large bulb-like resting buds basally, formed by thick silky-hairy leaves. The cypselas have a number of sclerenchymatous ribs, as in the genus Inula L., but only very few capillary pappus bristles as in the genus Pentanema Cass. It soon became clear that the material represented an undescribed species, but less clear as to in which genus it should be placed.
THE INULA COMPLEX
The only cladistic analysis of morphological data so far undertaken in the Inuleae s.s. (Anderberg, 1991) identified an informal, loosely composed group of genera, the paraphyletic or possibly polyphyletic Inula complex. The assemblage comprised those genera of the tribe that combined epaleate receptacles with a pappus of scabrid or barbellate capillary bristles, but lacking scales or other specializations characterizing other genera, or groups of genera. The complex included the genera Inula L. (incl. Codonocephalum Fenzl), Limbarda Adans., Varthemia DC., and Pentanema, with close but unresolved relationships to genera such as Duhaldea DC., Dittrichia Greuter, Chrysophthalmum Sch.Bip., Amblyocarpum Fisch. & Mey., and even Carpesium L. Also the relationships to the Pulicaria complex, i.e. taxa with double pappus composed of both capillary bristles and short scales, was pointed out as interesting in relation to the phylogeny of the Inula complex. A recent study of a few taxa from the Inuleae s.s., based on nucleotide sequences from the Internal Transcribed Spacer (ITS) region of the 18S–26S nuclear ribosomal DNA (Eldena¨s, Anderberg & Ka¨llersjo¨, 1998), revealed that the phylogenetic interrelationships in the tribe may be even more complex, with paleate genera with scaly pappus such as Asteriscus Tourn. ex Mill., and Buphtalmum L. being more closely related to the Pulicaria complex, and the likewise paleate but epappose genus Telekia Baumg. being closely related to Inula. A detailed investigation of both morphological and molecular data is thus needed before a well corroborated hypothesis of generic relationships in the tribe can be gained. Most of the genera of the Inula complex are monospecific, very small and homogeneous, or easily diagnosed due to morphological specialization, e.g. Dittrichia Greuter. The central problem is the heterogeneity and possible polyphyly of Inula. It is likely that Inula should be split into smaller, more homogeneous and monophyletic groups, each related to various other genera of the complex. Apart from the genera of the Inula complex that are diagnosed by particular apomorphic features, only two genera remained: the large residual genus Inula, diagnosed by the mere absence of the characters diagnosing the other genera, and the medium sized genus Pentanema.
PENTANEMA CASS.
The genus Pentanema Cass. (syn. Vicoa Cass.) is a medium sized genus of about 18 species, mainly distibuted in Central Asia, Iran, Afghanistan, Iraq, Pamir-Altai, and
A NEW SPECIES OF PENTANEMA
335
Tien-Shan. Most species are confined to rather restricted areas of distribution. The annual, weedy species have a wider distribution within East Africa, India, Indonesia and China (Gorschkova, 1959; Ling, 1979; Rechinger, 1980). The genus was originally diagnosed from its pappus composed of a few capillary bristles only, but several species have up to 10–15 bristles (Rechinger, 1980). The genus has no synapomorphy found in all its species, but rather characters that are nested within each other. The perennial species have lanate resting buds on the woody stems, a feature that Rechinger (1980) referred to as ‘gemmae quiescentes’, and which was interpreted as a synapomorphy of a large group of Pentanema species by Anderberg (1991). Within that group of species the number of pappus bristles varies from 5 to 15. Many of these species have disciform capitula with an outer row of almost tubular ray-florets, but there is a continuous variation from species with very reduced ray-floret lamina to species with well developed rays. The annual species in Pentanema have well developed ray-florets, and like most perennials they have ovate or elliptic leaves and almost unsclerified cypselas with few bristles. The lack of apparent synapomorphies diagnosing the genus indicate that it does not constitute a monophyletic group as presently circumscribed, and that some species may have to be removed. We conclude that the undescribed species is best placed in the genus Pentanema, sharing specialized characters with several of the perennial species near the type of the genus, few (4–5) pappus bristles, silky-hairy resting buds (‘gemmae quiescentes’), divaricately branched stems with delicate branches, are characters that are found in species of Pentanema but hardly in Inula or any other genus of the tribe.
Pentanema alanyense H. Duman & Anderb. sp. nov. (Fig. 1) Type: Turkey C4, Antalya, Alanya, Cayarasi-Gevne, 1240 m, clearings of Pinus nigra forest. 2.ix.1993, H. Duman 5504 & F. A. Karaveliogullar. Holotype: GAZI; Isotypes: E, S). Diagnosis. Species nova ab omnibus aliis speciebus generis Pentanematis receptaculo distincte conico, habitu omnino herbaceo et gemmis quiescentibus forma ad instar bulborum sericeopilosis differt et praesentia typica etiam radicum multarum secundariarum absentiaque radicis primariae dominantis insignis. Speciei annuae P. divaricato aspectu (praecipue propter ramificationem divaricatam) maxime simile P. alanyense est sed haec species a P. alanyensi forma folii, receptaculo fere plano et in involucro indumento denso pilorum longorum patentium obtecto recedit. Description. A small delicate perennial herb, basally with several bulb-like resting buds covered with long silky hairs. numerous but dominating primary root absent. terete, up to c. 15 cm high and about 1 mm wide, divaricately branched in the upper part with almost thread-like branches, sparingly branched below, reddish-brown, glandular-hairy and with scattered bristly hairs. linear to lanceolate, lower up to 8 cm long and 7 mm wide, gradually attenuating towards the base; lamina with minute glandular hairs; margin almost entire or with minute and inconspicuous teeth; upper leaves linear, sessile; leaves of resting buds involute, 3–12 mm, glaucous to purple, broad. c. 5 mm wide, heterogamous, solitary
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Figure 1. Pentanema alanyense H. Duman & Anderb sp. nov. A, habit of plant; B, bulb-shaped resting-bud; C, capitulum; D, capitulum (L. S.) showing conical receptacle; E, ray-floret; F, disc-floret; G, cypsela with pappus bristles; H, anther; I, anther-tails; J, endothecial tissue; K, style-branches. A–K, Duman 5504 (S).
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337
T 1. List of taxa growing in the type locality of Pentanema alanyense. Taxa marked with ∗ are local endemics in southern Anatolia Ostrya carpinifolia Scop. ∗Gypsophila curvifolia Fenzl ∗Potentilla isaurica (P. H. Davis) Pawł ∗Cicer isauricum P. H. Davis ∗Bupleurum subuniflorum Boiss. & Heldr. ∗Eryngium palmito Boiss. & Heldr. ∗Ferulago isaurica Pesmen Pimpinella tragium Vill. ssp. lithophila (Schischkin) Tutin Ampelopsis orientale (Lam.) Planch. ∗Digitalis davisiana Heywood ∗Origanum bilgeri P. H. Davis Origanum saccatum P. H. Davis ∗Phlomis monocephala P. H. Davis
Sideritis arguta Boiss. & Heldr. ∗Sideritis argyrea P. H. Davis ∗Sideritis erythrantha Boiss & Heldr. var. cedretorum P. H. Davis ∗Thymbra sintenisii Bornm. & Aznav. ssp. isaurica P. H. Davis ∗Cephalaria gazipashensis Su¨mbu¨l ∗Cephalaria isaurica V. A. Matthews ∗Campanula myrtifolia Boiss. & Heldr. Campanula trachyphylla Schott & Kotschy Chrysophthalmum dichotomum Boiss. & Heldr. ∗Leucocyclus formosus Boiss. ssp. amanicus (Rech. f.) Hub.Mor. & Grierson
in the end of the branches. Involucral bracts 2–4 mm, glandular-hairy, in several rows, imbricate; outer bracts green, obtuse; inner bracts scarious, acute, marginally fimbriate. Receptacle conical, epaleate, glabrous. Ray-florets in one row, female, radiate, yellow, with scattered glandular hairs; lamina c. 5 mm long, with elongated epidermis cells, apically 3-dentate. as in disc-florets, but pappus of 3(−4) capillary bristles. - yellow, perfect, 5-lobed, c. 3 mm long; corollalobes short, glandular-hairy; anthers ecalcarate, with short slightly branched tails; endothecial tissue radial, i.e. with thickenings confined to the vertical cell walls; filament collar with square cells; style bifid with slightly enlarged base; style-branches slightly flattened, obtuse, dorsally with acute sweeping hairs not reaching the furcation; stigmatic surface forming two apically confluent bands. c. 1 mm, elliptic, sparsely hairy with 8–10 sclerenchymatous ribs; epidermis with elongated crystals; pappus of 4(−5) barbellate, c. 2 mm long capillary bristles; carpopodium oblique, white; cells thick-walled, in several rows. The new species differs from all other species of Pentanema by its distinctly conical receptacle, entirely herbaceous growth habit combined with silky-hairy, bulb-like basal resting buds. Also, the presence of a large number of secondary roots and absence of a dominating primary root is typical. In general appearance, Pentanema alanyense is most similar to the annual P. divaricatum Cass., mainly because of its divaricate branching, but the latter species differs, e.g. in leaf-shape, in having an almost flat receptacle and a dense indumentum of long spreading hairs on the involucre. Habitat. The new species was found growing on calcareous rock in clearings in Pinus nigra forests. A number of species with restricted distribution, several being local endemics of southern Anatolia, were found growing in the same locality (Table 1.). Other examined material Turkey C4, Antalya, Alanya, Hadim-Alanya, 1250–1300 m., on calcareous rocks, 9.viii.1994, H. Duman, Z. Aytac & A. Duran 5597 (GAZI).
DISCUSSION
From a biogeographical point of view, the presence of Pentanema alanyense in southern Turkey is also interesting. The genus has its highest species diversity in
338
H. DUMAN AND A. A. ANDERBERG
Iran, Afghanistan and Pamir-Altai. The annual species P. divaricatum is found in Arabia, the Middle East and Central Asia. The closest occurrence of a perennial species is P. flexuosum in northern Iraq. The discovery of a Pentanema in Antalya widens the total distribution range of the genus to include the Eastern Mediterranean region, and adds to the number of endemic taxa that show a disjunct distribution between Antalya and Konya in Turkey with Iran and Afghanistan. Other examples with the same distribution pattern include Thermopsis turcica Kit Tan, Vural & Ku¨cu¨ko¨du¨k with T. alpina (Pall.) Ledeb. from Turkestan (Tan, Vural & Ku¨cu¨ko¨du¨k, 1983), Arnebia purpurea Erik & Su¨mbu¨l with A. euchroma (Royle) I. M. Johnst. from Afghanistan and Iran (Erik & Su¨mbu¨l, 1986), Pyrus serikensis Gu¨ner & H. Duman with P. boissieriana from Iran (Gu¨ner & Duman, 1994), and Aethionema alanyae H. Duman with the genus Moriera Boiss. from Iran (Duman, 1994).
ACKNOWLEDGEMENTS
H.D. wishes to thank the Edinburgh Botanic Garden (Sibbald) Trust for financial support during his visit to Edinburgh, and the Curator of the Royal Botanic Garden Edinburgh for study facilities. The new species was collected during the [Turkish] Endemic Plants Project, which is funded by TUBITAK (Project No.: TBAG-DPT/ C.SEK4). The authors thank Dr Robert Mill (Royal Botanic Garden Edinburgh) for the Latin diagnosis and Ms Pollyanna Lidmark (Swedish Museum of Natural History) for the illustration.
REFERENCES
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