Willdenowia18 - 1989 409

JOSEFBOGNER & PETERBOYCE

A remarkablenew Biarum() from Turkey

Abstract Bogner,J.& Boyce, P.: A remarkablenew Biarum(Araceae) from Turkey. - Willdenowia18: 409-417. 1989.- ISSN 0511-9618. Biarumditschianum from SWTurkey is describedas a speciesnew to scienceand illustrated.Its possible relationshipswithin the are discussed.

Discovery of the new species

Duringan excursionto SWTurkeyorganised by the BotanicalInstitute of the Universityof Bonn in March/April1987 Friedrich Ditsch, a studentat Bonn, collectedmaterial of a speciesof Biarum.He col- lectedone sterile,living specimen which was grown on in Bonn.Whenthe plantflowered in May 1987it was obvious that it did not matchany describedspecies and Mr.Ditsch took colourphotographs of the , but made no herbariumspecimen. Unfortunately the plant died soon afterflowering. In October 1987Mr. Ditsch, accompaniedby ManfredKoenen from the BotanicalGarden of the Universityof Bonn,visited Turkey in orderto tryto recollectthe plant.They succeeded in findingonly seedlingsand immatureplants and were unable to locateany flowering specimens. It thereforeappeared thatthe springflowering, so unusualin Biarum,exhibited by the cultivatedplant was typical and not sim- ply a responseto disturbance.A trip by Mr.Koenen in April 1988was, however, successful, resulting in good vegetative,flowering and fruitingspecimens. Some of the tuberscollected had developing inflores- cenceswhich subsequently opened in cultivationat Bonn at the beginningofMay. From these spirit and dried materialwas prepared. Despitethe unusualappearance of the new species,it is clearlya Biarumon the basisof the unilocu- lar,uniovulate ovaries with the ovule attachedto a basal placenta. It gives us great.pleasure to name this new species in honour of Mr.Friedrich Ditsch.

Biarumditschianum Bogner & Boyce, sp. nova - Fig. 1-8. Typus:Bonn BotanicalGarden Accession no. 22592 (holotypus:K; isotypi: B, BONN (Fig.2), K, M).

Laminafoliorum angustissimeelliptica vel lanceolata;petiolus a lateribuscompressus. Spatha 4 usque5 cm longa,marginibus basi per ca. 2.5 cm connatis,tubum formans, lamina reducta, breviter sub- triangulari.Spadicis pars flores femineos masculinis discedens nuda, pistillodia carehtia, appendix stipi- tata basi incrassataprocessibus piliformibus retroversis instructa. A perennial,tuberous herb. Tuber light brown,depressed-globular to sub-globular,2.5-3 cm in dia-

Botanischer Garten und Botanisches Museum, Berlin-Dahlem is collaborating with JSTOR to digitize, preserve, and extend access to Willdenowia ® www.jstor.org 410 Bogner& Boyce: A new Biarumfrom Turkey

Fig.1. Biarumditschianum Bogner & Boyce (left)in naturalhabitat growing In limestone holes. - Photo:M. Koenen.

meter,1. 5-2 cm thick,with a centralgrowth point andperipheral adventitious buds on the uppersufface. Roots 1.4-2 mm thick,contractile roots ringed.Leaves annual, 2-3, rarelyup to 5, leaf basesencased by several5-10 cm long, membranaceous,whitish cataphylls; petioles 8-20 cm long, 1.5-3 mm wide,chan- neled, laterallycompressed, whitish basally below soil level,mid-green, sometimes reddish tinged distal- ly; lamina linear, narrowly-elliptic, lanceolate or oblanceolate, 6-15(-20) cm long, 0.6-3(-3.5) cm wide, basecuneate, decurrent, apex acute to obtuse,lamina mid-green, veins mostly lighter coloured; venation reticulate,mid-vein pronounced, especially in the basalpart of the lamina,4-7 primarylateral veins on each side. Inflorescenceappearing when plant leafless,base enclosed by 5-6 cataphylls,these at first membranaceous,whitish, soon witheringto become paperyand brown, 1.5-6(-7) cm long and up to 1.5 cm wide, the longest equallingthe spathe;peduncle subterranean,2-5 cm long, 3.5-5 mm thick, whitish;spathe 4-5 cm long, spathetube ca. 3 cm long, 1.8-2 cm in diameter,margins basally connate for ca.2.5 cm, freeand overlappingdistally for 5 mm, exteriorwhitish,sometimes slightly reddish tinged, interiorreddish-purple, spathe lamina much reduced,1.8-2 cm long, subtriangular,terminating in a ca. 2 mm long mucro,exterior greenish to lightgreen, sometimes with a reddishtinge, interior purplish-red; spadix 7-8 cm long, appendixelongate-conoid to somewhatsub-cylindric, short-stipitate, 4-4.5 cm long, 7-11 mm diam.,apex obtuse, base rounded, the basal7-10 mm furnishedwith reflexed,filiform, acuminate,transparent white processes,1-2.5 mm long, 0. 1-0.25 mm thick,appendix dark yellow, ex- cept for the reddish-purplebasal 7-1 0 mm; stipe 5-6 mm long, 3 -4 mm thick,cream. Flowers unisexual, naked;pistillate and staminateflowers separated by a naked2.3-3.5 cm long,2.5-3.5(-4) mm thicklight Willdenowia18 - 1989 411

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Fig.2. Biarumditschianum Bogner & Boyce. - Herbariumspecimens, in vegetativestage showing the leafvaria- tion (BonnBotanical Garden Accession no. 22592, BONN). - Photo: K. Liedi. 412 Bogner& Boyce: A new Biarumfrom Turkey

Fig3 Biarum dztschianum o

- W. Barthlott. Fig. 3. Biarum ditschianum Bogner & Boyce. - One flowering and two fruiting specimens. Photo: Willdenowia18 - 1989 413

Biarum Bogner & Boyce. - ; note the filiform processes at the base of the appendix, spathe partly removed: (left) female stage, note the still closed thecae; (right) male stage with pollen being extruded in strands, stigmas and styles shrivelled. - Photos: W. Barthlott.

purplish-redaxis; pistillodes absent; pistillate flowers in ca. 4 spiralrows arranged in a 2-2.5 mm high ditschianum hemisphericalFig. 4. cluster at the base (Fig.5); pistils lageniform,styles and stigmascurved outwards; style 1.2-1.3 mm long, ca. 0.4 mm thick basally,thickening slightly and purplishintensifying towards the ovary;stigma sub-capitate, 0.5-0.6 mm in diameter,slightly papillose,yellow; ovary ca. 1 mm diam.,pur- plish-red;ovule orthotropouson a basalplacenta, ca. 0.5 mm long, funiclevery short; staminate flowers arrangedin an oblong zone (Fig.6), each consistingmostly of 2 stamens,rarely 1, sub-sessile;stamen 1 mm long, ca. 0.6 mm wide,yellow basally, purplish-red apically, thecae ellipsoid,dehiscing by an api- cal slit; pollen extrudedin strands,pollen grainsinaperturate, ellipsoid, 42-45 pm x 30-35 pm, exine smooth (Fig. 8). Infructescenceborne at soil level, peduncle 3-5 cm long, ca. 5 mm thick,whitish, in- fructescence3-3.5 cm in diameter,ca. 2 cm thick,consisting of ca.50 berriesin a depressed-globularclus- ter(Fig. 7); berriesobovoid, 6-10 mm long,4-7 mm wide, pericarpwhitish, sometimes with a veryslight reddishtinge basally,0.4-0.7 mm thick.Seed obovoid, 5.5-7.5 mm long and 4-5.5 mm wide;testa with upperpart very slightly irregular-reticulate, lower part smooth; embryoelongate, ca. 3.5 mm long, endo- sperm abundant. Chromosomenumber: 2n = 26. 414 Bogner & Boyce: A new Biarum from Turkey

Fig. 5. Biarumditschianum Bogner & Boyce. - Zone of female flowersat base of the spadix.- Photo: W. Barthlott.

Distribution

Turkey,Anatolia, Province of Lycia, in the Esen River valley, 30-60 m (BonnBotanical Garden Acces- si'onno. 22592). We are not disclosing the exact locality since this species has a restricted local distribution and is also quite rare. Another collection from the same locality is B. Mathew11058 (K); it was made in late April 1988 and the inflorescence was in bud. Vernacular name: Yilan Bica'gi.

Ecology

Bi' ditschianumgrows in the low- to middle-high garigue usually in association with Quercus aucheri'andDaphne gnidioides but occasionally in transition vegetation of Asphodelusmicrocarpus plains with Urgineamaritima, Pistacia lentiscus, Olea sylvestris, Euphorbia characias subsp. wu/fenii, Selaginella denti- culata,Phiomis fruticosa and Cyclamengraecum. Biarumditschianum is confined to deep, partly loam-filled chimney-like holes, or more rarely cre- vices, in limestone rocks (Fig. 1). This type of habitat is unusual, but not unique for the genus; B. bovei Blume frequently grows in, but is not restricted to, similar situations in NW Syria (Boyce, pers. obs.). Willdenowia18 - 1989 415

The soft leavesare partly grazed by goats,but it seemsthat the deep holes favouredby B. ditschianum help to reducedamage since the tuberis often situatedat the base of the hole with most of the foliage containedin the empty chimneyand thus inaccessible.Furthermore, the narrow,deep holes arepracti- callyfree of otherperennial plants, the Biarumapparently growing with a minimumof competition.Pre- ciselyhow the seed is spreadfrom one hole to anotheris unclear.However, the infructescencemorpholo- gy and seed type suggestseed dispersalby ants.This is knownto occurin manyspecies of Biaruminclud- ing B. bovei. The strongsmell of carrionand excrementproduced by the open inflorescenceattracts carrion flies which are presumablythe primarypollinators. The spadixis a strongUV light absorber,giving a good contrastto the surroundinglimestone. Carrionflies are known to perceivemainly UV light.

Fig.6. Biarumditschianum Bogner& Boyce. -Upper part of spadixshowing the malezone and baseof appendixwith fi- liform processes. - Photo: W. Barthlott. 416 Bogner & Boyce: A new Biarum from Turkey

Fig. 7. Biarum ditschianum Bogner & Boyce. - Infructescence. - Photo: W. Barthlott.

Fig. 8. Biarum ditschianumBogner &Boyce. Pollen grain (longitudinal depressions resulting from dehydration). - Photo: W. Barthlott. Willdenowia18 - 1989 417

Growth cycle

Biarumditschianum begins growth in earlywinter after the firstrains. Leaf shape is veryvariable.The firstleaves produced are broadly spathulate but as the seasonprogresses subsequent leaves are narrower. This may be in responseto wateravailability in the confines of the limestone holes. Floweringoccurs towards the end of the growthperiod from early May to lateJune. At this time the previousyear's infructescences ripen. By the time the inflorescencesopen most of the plantsare leafless. Eachinflorescence lasts for two days.During the firstday, the pistillateflowers are at anthesisand the spa- dix appendixproduces the characteristicfoetid smell.By the secondday the staminateflowers reach an- thesis, the odour declines and the stigmasare no longerreceptive.

Relationships

Biarumditschianum is perhapsthe most remarkablespecies of Biarumdescribed to date.The much reducedspathe limb togetherwiththe filiformprocesses at the baseof the appendix(Fig. 3, 4) clearlyse- parateB. ditschianumfrom all otherspecies. The morphologyof the spatheand appendixand the oblong staminateflower zone suggesta linkwith Biarumsect. Ischarum Blume (1837), but the lackof pistillodes togetherwith the filiformprocesses preclude the inclusion of B. ditschianumin this section.A reduced spathelimb and absenceof pistillodesis foundin B. davisiiTurrill,but the vegetativefeatures ofB. davisii aredifferent, as also is the shapeof the staminateflowercluster. B. davisiihas been tentativelyincluded in Biarumsect. Ischarum (Riedl 1980) but this is probablyincorrect because of the elongatestaminate flower zone, lackof pistillodesand sweet,not foetid,odour. It would appearthat the delimitationof sectionsin Biarumis in need of revisionand althoughit would be bestto accommodateB. ditschianumin a section of its own this would necessitatea completereappraisal of the genus.Since a revisionis presentlybeing undertakenby the second authorit is felt thatthe publicationof a new sectionat this time is premature.

Adknowledgements

Wewould like to expressour thanks to the staffat the BotanicalInstitute and Gardenof the Univer- sityof Bonn who helpedus by providingthe necessarymaterial and information.Particular thanks go to Mr.Friedrich Ditsch for his valuablehabitat data, Mr. Manfred Koenen for recollectingfertile material and Prof.Wilhelm Barthlott for providingthe photographsand SEMmicrographs. Thanks also to Miss GittePetersen of the BotanicalInstitute of the Veterinaryand Agriculture University of Copenhagenfor countingthe chromosomesand to Dr.AlarichKress of the BotanicalGarden Munchen,forpreparing the Latindiagnosis. The secondauthorwould like to expresshis thanksto Dr.D. A. Simpsonand Dr.J. Drans- field of the Royal Botanic Gardens,Kew, for their helpful suggestions.

References

Blume, C. L. von 1837: Rumphia1. - Lugduni-Batavorum. Riedl,H. 1980:Tentative keys for the identificationof speciesin Biarumand ,with notes on some taxa included in Biarum.- Aroideana3: 24-31.

Addressesof the authors: Josef Bogner,Botanischer Garten Miunchen, Menzinger Strage 63, D-8000 Miunchen19. PeterBoyce, The Herbarium,Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE,England, U.K.