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ITS phylogeny of Middle Asian geophilic Umbelliferae-Apioideae genera with comments on their morphology and utility of psbA-trnH sequences

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Abstract

Generic delimitation and relationships within the taxonomically controversial group of geophilic Umbelliferae from Middle Asia are problematic. Geophilic plants in arid regions have a short period of development during spring and early summer and survive unfavorable seasons due to the presence of tuber-like underground storage organs. To investigate the circumscription of recently recognized Middle Asian geophilic genera and their relationships, we conducted phylogenetic analyses using DNA sequence data from rapidly evolving nuclear (nrITS) and chloroplast (psbA-trnH intergenic spacer) regions as well as multivariate analyses of the morphological and ITS data. Our results show that Galagania and Mogoltavia are monophyletic, whereas Bunium, Hyalolaena and Oedibasis are polyphyletic. Elaeosticta, one of the larger genera of geophilic Middle Asian umbellifers, is rendered paraphyletic by the inclusion of the enigmatic Hyalolaena melanorrhiza. The genus Scaligeria, in which most species of Elaeosticta were initially described, is not grouped with these genera and has other affinities. Our analyses also show that the morphological characters used for the delimitation of genera within this group of Umbelliferae (such as life form, cotyledon number, presence/absence of petiolules of basal leaflets, shape of mericarp ribs, width of mericarp commissure, number of secretory ducts in mericarp furrows) are rather homoplastic. The psbA-trnH sequences were not informative enough to reveal relationships among the genera studied, although some interesting features in the spacer evolution were revealed (maximum reduction in length among Umbelliferae, homoplasious directed substitutions to restore secondary structure).

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Acknowledgments

This work is supported by grants from the Russian Foundation for Basic Research (## 10-04-00675, 12-04-31543). We are grateful to the curators of AA, E, LD, LE, MW, TAD TARI, and TASH herbaria for use of their facilities. Anthony R. Magee kindly checked the manuscript linguistically.

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Authors and Affiliations

  1. Botanical Garden, Biological Faculty, Moscow State University, Moscow, 119991, Russia

    Galina V. Degtjareva, Eugene V. Kljuykov & Michael G. Pimenov

  2. A.N. Belozersky Institute of Physico-Chemical Biology, Moscow State University, Moscow, 119991, Russia

    Tahir H. Samigullin & Carmen M. Valiejo-Roman

Authors
  1. Galina V. Degtjareva
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  2. Eugene V. Kljuykov
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  3. Tahir H. Samigullin
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  4. Carmen M. Valiejo-Roman
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  5. Michael G. Pimenov
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Correspondence to Galina V. Degtjareva.

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Appendices

Appendix 1

Voucher information and GeneBank accession numbers for plants used in the present study

Aegopodium podagraria L.: ITS U305371, U305361, psbA-trnH EU4457142. Bunium alpinum Waldst. & Kit.: ITS DQ4352042, DQ4352432. Bunium avromanum (Boiss. et Hausskn. ex Boiss.) Drude: ITS DQ4352062, DQ4352452, psbA-trnH EU4457182. Bunium bulbocastanum L.: ITS DQ4437222, DQ4437242, psbA-trnH DQ4571622. Bunium cornigerum (Boiss. et Hausskn.) Drude: ITS DQ4352142, DQ4352532, psbA-trnH EU4457252. Bunium elegans (Fenzl) Freyn: ITS AF0735433, AF0735443, psbA-trnH DQ4571632. Bunium fallax Freyn: ITS DQ4352172, DQ4352562, psbA-trnH EU4457282. Bunium ferulaceum Sm.: ITS DQ4352192, DQ4352582, psbA-trnH EU4457302. Bunium mauritanicum Batt.: ITS DQ4352262, DQ4352652, psbA-trnH EU4457382. Bunium microcarpum (Boiss.) Freyn et Sint. ex Freyn: ITS DQ4352272, DQ4352662, psbA-trnH DQ4571642. Bunium pachypodum P.W.Ball: ITS DQ4352282, DQ4352672, psbA-trnH EU4457392. Bunium paucifolium DC.: ITS DQ4352292, DQ4352682, psbA-trnH EU4457402. Bunium pinnatifolium Kljuykov: ITS DQ4352312, DQ4352702, psbA-trnH EU4457422. Bunium rectangulum (Boiss. & Hausskn.) H.Wolff: ITS DQ4352322, DQ4352712, psbA-trnH DQ4571652. Bunium scabrellum Korovin: ITS DQ4352342, DQ4352732, psbA-trnH EU4457442. Bunium simplex (K. Koch) Kljuykov: ITS DQ4352372, DQ4352762, psbA-trnH DQ4571692. Carum appuanum (Viv.) Grande [as Carum apuanum Grande]: ITS AY8409854, AY8409844, psbA-trnH EU4457472. Carum carvi L.: ITS AF0778785, psbA-trnH DQ4571712. Carum heldreichii Boiss.: ITS AY8409894, AY8409884. Crithmum maritimum L.: ITS U305411, U305401, psbA-trnH EU4457512. Elaeosticta aitchisonii (H. Wolff) Kljuykov et al.: Afghanistan, Badakhshan prov., valley of Warduj river, 50 km above from Barak settlement, 09.vii.1974, Gubanov et al. 444a (MW), ITS HM229350, psbA-trnH HM474795. Elaeosticta alaica (Lipsky) Kljuykov et al.: (1) Uzbekistan, valley of Angren river, near Angren, 26.05.1978, Pimenov et al. 71 (MW), ITS HM229351, psbA-trnH HM474796; (2) Kirghizia, Djalalabad prov., Toguztoroo distr., Kokirimtoo Mts., Koksha river, S slope of Atdjailau Mt., 10.viii.1995, Pimenov et al. K95–310 (MW), ITS HM229352, psbA-trnH HM474797; (3) Tadzhikistan, Karategin ridge, S slope of Kabud-krym ridge, 03.vii.1988, Pimenov et al. 243 (MW), ITS HM229353, psbA-trnH HM474798. Elaeosticta allioides (Regel & Schmalh.) Kljuykov et al.: (1) Tadzhikistan, valley of Obihingou river, Peter I ridge, 25.vii.1974, Pimenov & Kljuykov 622 (MW), ITS HM229354, psbA-trnH HM474799; (2) Uzbekistan, valley of Aksakatau-saj river, 01.vii.1976, Kljuykov et al. 896 (MW), ITS HM229355, psbA-trnH HM474800. Elaeosticta bucharica (Korovin) Kljuykov et al.: Tadzhikistan, gorge of Varzob, mouth of Guskharf river, 04.vi.1976, Pimenov & Kljuykov 409 (MW), ITS HM229356, psbA-trnH HM474801. Elaeosticta conica Korovin: (1) Uzbekistan, near Bajsun, v.1979, Pimenov et al. 426 (MW), ITS HM229357, psbA-trnH HM474802; (2) Uzbekistan, Kashka-darja prov., mountains SW of Dehkanabad, Sakyrtma ridge, 01.vi.1976, Kljuykov & Pimenov 314 (MW), ITS HM229358, psbA-trnH HM474803. Elaeosticta meifolia Fenzl: Iran, Chaharmahal-e Bakhtiari, 29.vi.1986, 54987 (TARI), ITS HM229359, psbA-trnH HM474804. Elaeosticta ferganensis (Lipsky) Kljuykov et al.: (1) Kirghizia, Talass Alatau ridge, 18.vii.1976, Kljuykov & Baranova 1179 (MW), ITS HM229360, psbA-trnH HM474805; (2) Kirghizia, Kichik-Alaj ridge, 17.vii.1981, Kljuykov et al. 404 (MW), ITS HM229361, psbA-trnH HM474806, HM474807. Elaeosticta glaucescens (DC.) Boiss.: Armenia, Vedi prov., Urcskij ridge, near Surenavan settlement, 08.vi.1977, Pimenov et al. 348 (MW), ITS HM229362, psbA-trnH HM474808. Elaeosticta hirtula (Regel & Schmalh.) Kljuykov et al.: (1) Tadzhikistan, Anzob pass, 23.vii.1974, Pimenov et al. 587 (MW), ITS HM229364, psbA-trnH HM474810; (2) Afghanistan, Badakhshan, Yavarsan, c.30 km S of Qeshm, 01.vii.1969, Hedge & Wendelbo W9358 (MW), ITS HM229363, psbA-trnH HM474809. Elaeosticta knorringiana (Korovin) Korovin: (1) Kirghizia, pass between Osh and Naukat, 14.vii.1987, Pimenov et al. 378 (MW), ITS HM229365, psbA-trnH HM474811; (2) Kirghizia, Osh prov., near Suzak, 15.vii.1987, Pimenov et al. 392 (MW), ITS HM229366, psbA-trnH HM474812; (3) ITS DQ4228162, DQ4228352, psbA-trnH EU4457522. Elaeosticta korovinii (Bobr. ex Korovin) Kljuykov et al.: Turkmenistan, Great Balkhan ridge, 29.vii.1978, Pimenov & Kljuykov 129 (MW), ITS HM229367, psbA-trnH HM474813. Elaeosticta lutea (M.Bieb. ex Hoffm.) Kljuykov et al.: (1) Russia, Rostov prov, near Kamensk-Shahtinskij, 23.vii.1978, Pimenov & Kljuykov 27 (MW), ITS HM229369, psbA-trnH HM474815; (2) Russia, Kalmykia, Iki-Burul’sk area, 16.vi.2000, Novikova s.n. (MW), ITS HM229370, psbA-trnH HM474816; (3) Ukraine, Lugansk prov., 17.vi.1966, Pavlov s.n. (MW), ITS HM229371, psbA-trnH HM474817; (4) Russia, Volgograd prov., 07.vi.1950, Smirnov et al. s.n. (MW), ITS HM229372, psbA-trnH HM474818; (5) ITS DQ4228152, DQ4228342, psbA-trnH DQ4571722. Elaeosticta nodosa (Boiss.) Boiss.: ITS AY9412716, AY9412996, psbA-trnH EU4457532. Elaeosticta paniculata (Korovin) Kljuykov & Pimenov: (1) Uzbekistan, Urgut, 30.v.1978, Pimenov et al. 175 (MW), ITS HM229379, psbA-trnH HM474819; (2) ITS DQ4228172, DQ4228362, psbA-trnH DQ457173. Elaeosticta platyphylla (Korovin) Kljuykov et al.: Turkmenistan, Badhys, Kushka, 07.vi.1978, Pimenov et al. 285 (MW), ITS HM229373, psbA-trnH HM474820. Elaeosticta polycarpa (Korovin) Kljuykov et al.: (1) Tadzhikistan, southern spurs of Darvaz ridge, Khozretishakh ridge, Turkoni settlement, 06.viii.1975, Pimenov & Kljuykov 1082 (MW), ITS HM229374, psbA-trnH HM474821, (2) Kirghizia, Osh prov., N slope of Turkestan ridge, valley of Isfara river, between Proletarsk and Kol’co station, 12.06.1976, Kljuykov & Pimenov 561 (MW), ITS HM229375, psbA-trnH HM474822. Elaeosticta ramosissima Kljuykov: Iran, Chaharmahal-e Bakhtiari, Darreh Bazott, Mavraz, 13.viii.1986, Mozaffarian 58046 (TARI), ITS HM229376, psbA-trnH HM474823. Elaeosticta samarkandica (Korovin) Kljuykov et al.: Tadzhikistan, Turkestan ridge, Kul’-Keryz breakage, 30.v.1976, Pimenov & Kljuykov 252 (MW), ITS HM229377, psbA-trnH HM474824. Elaeosticta scariosibracteata (Rech.f. & Riedl) Kljuykov et al.: Afghanistan, Parvan prov., Panjshir valley, Shanez, 21.vii.1962, Hedge & Wendelbo W5338 (MW), ITS HM229378, psbA-trnH HM474825. Elaeosticta transcaspica (Korovin) Kljuykov et al.: (1) Afghanistan, Herat prov., Cheshmeh Obeh, 12.v.1969, Hedge et al. W7927 (MW), ITS HM229380, psbA-trnH HM474827; (2) Turkmenistan, Karabil’ between Tahta-Bazar and Kerki, 21.v.1975, Kljuykov & Baranova 595 (MW), ITS HM229381, psbA-trnH HM474826. Elaeosticta transitoria (Korovin) Kljuykov et al.: (1) Uzbekistan, Kazykurt pass, between Tashkent and Chimkent, 07.vi.1975, Kljuykov & Baranova 707 (MW), ITS HM229382, psbA-trnH HM474828; (2) Tadzhikistan, Mogoltau, Spa Mt., 02.vii.1976, Kljuykov & Vassilieva 904 (MW), ITS HM229383, psbA-trnH HM474829; (3) Kirghizia, Kirgiz ridge, Shamsi gorge, 14.vii.1985, Kljuykov 120 (MW), ITS HM229384, psbA-trnH HM474830; (4) Kazakhstan, Chimkent prov., Syrdarya Karatau ridge, valley of Bir-Isek river, 07.vii.1959, Parfent’eva & Pavlov s.n. (MW), ITS HM229385, psbA-trnH HM474831; (5) Kazakhstan, W Tien Shan, Talas Alatau ridge, Taldy-Bulak gorge, 05.viii.1981, Baranova & Lavrova 1037 (MW), ITS HM229368, psbA-trnH HM474814. Elaeosticta tschimganica (Korovin) Kljuykov et al.: Kirghizia, Osh prov., S slope of Susamyr ridge, 07.vii.1985, Kljuykov 29 (MW), ITS HM229386, psbA-trnH HM474832; ITS DQ4228182, DQ4228372, psbA-trnH EU4457542. Elaeosticta ugamica (Korovin) Korovin: Kirghizia, Osh prov., valley of Chatkal river, above mouth of Sandalash river, 16.vi.1976, Kljuykov et al. 723 (MW), ITS HM229387, psbA-trnH HM474833. Elaeosticta vvedenskyi (Kamelin) Kljuykov et al.: Uzbekistan, Nuratau ridge, Aktau Mts., 19.vii.1975, Pimenov et al. 749 (MW), ITS HM229388, psbA-trnH HM474834. Elwendia afghanica (Beauverd) Pimenov & Kljuykov (= Bunium afghanicum Beauverd): ITS DQ4352032, DQ4352422, psbA-trnH EU4457162. Elwendia angreni (Korovin) Pimenov & Kljuykov (= Bunium angreni Korovin): ITS DQ4352052, DQ4352442, psbA-trnH EU4457172. Elwendia badachschanica (Kamelin) Pimenov & Kljuykov (= Bunium badachschanicum Kamelin): ITS DQ4352072, DQ4352462, psbA-trnH EU4457192. Elwendia capusii (Franch.) Pimenov & Kljuykov (= Bunium capusii (Franch.) Korovin): ITS DQ4352112, DQ4352502, psbA-trnH EU4457222. Elwendia caroides Boiss. (= Bunium caroides (Boiss.) Hausskn. ex Bornm.): ITS DQ4352122, DQ4352512, psbA-trnH EU4457232. Elwendia chaerophylloides (Regel & Schmalh.) Pimenov & Kljuykov (= Bunium chaerophylloides (Regel & Schmalh.) Drude): ITS DQ4352132, DQ4352522, psbA-trnH EU4457242. Elwendia cylindrica (Boiss. & Hausskn.) Pimenov & Kljuykov (= Bunium cylindricum (Boiss. & Hohen.) Freyn): ITS DQ4352162, DQ4352552, psbA-trnH EU4457272. Elwendia fedtschenkoana (Korovin ex Kamelin) Pimenov & Kljuykov (= Bunium fedtschenkoanum Korovin ex Kamelin): ITS DQ4352182, DQ4352572, psbA-trnH EU4457292. Elwendia hissarica (Korovin) Pimenov & Kljuykov (= Bunium hissaricum Korovin): ITS DQ4352212, DQ4352602, psbA-trnH EU4457322. Elwendia intermedia (Korovin) Pimenov & Kljuykov (= Bunium intermedium Korovin): ITS DQ4352222, DQ4352612, psbA-trnH EU4457332. Elwendia latiloba (Korovin) Pimenov & Kljuykov (= Bunium latilobum Korovin): ITS DQ4352242, DQ4352632, psbA-trnH EU4457362. Elwendia longipes (Freyn) Pimenov & Kljuykov (= Bunium longipes Freyn): ITS DQ4352252, DQ4352642, psbA-trnH EU4457372. Elwendia persica (Boiss.) Pimenov & Kljuykov (= Bunium persicum (Boiss.) B.Fedtsch.): ITS DQ4352302, DQ4352692, psbA-trnH EU4457412. Elwendia salsa (Korovin) Pimenov & Kljuykov (= Bunium salsum Korovin): ITS DQ4352332, DQ4352722, psbA-trnH EU4457432. Elwendia seravschanica (Korovin) Pimenov & Kljuykov (= Bunium seravschanicum Korovin): ITS DQ4352352, DQ4352742, psbA-trnH DQ4571672. Elwendia setacea (Schrenk) Pimenov & Kljuykov (= Bunium setaceum (Schrenk) H.Wolff): ITS DQ4352362, DQ4352752, psbA-trnH DQ4571682. Elwendia stewartiama (Nasir) Pimenov & Kljuykov (= Bunium stewartianum Nasir): ITS DQ4352382, DQ4352772, psbA-trnH EU4457342. Elwendia vaginata (Korovin) Pimenov & Kljuykov (= Bunium vaginatum Korovin): ITS DQ4352392, DQ4352782, psbA-trnH DQ4571702. Elwendia kuhitangi (Nevski) Pimenov & Kljuykov (= Bunium kuhitangi Nevski): ITS DQ4352232, DQ4352622, psbA-trnH EU4457352. Elwendia wolffii (Kljuykov) Pimenov & Kljuykov (= Bunium wolffii Kljuykov): ITS DQ4352412, DQ4352802, psbA-trnH EU4457462. Falcaria vulgaris Bernh.: ITS AF0778885, psbA-trnH DQ4571742. Fuernrohria setifolia K. Koch: ITS AF0091128, AF0086338, psbA-trnH EU4457552. Galagania ferganensis (Korovin) M. Vassiljeva & Pimenov: ITS DQ4228192, DQ4228382, psbA-trnH EU4457562. Galagania fragrantissima Lipsky: ITS DQ4228202, DQ4228392, psbA-trnH DQ4571772. Galagania gracilis (Kamelin & Pimenov) Kamelin & Pimenov: Tadzhikistan, S slope of Peter I ridge, valley of Obihingou river, Dashti-Khasan, 23.vi.1990, Pimenov & Kljuykov 190 (MW), ITS HM229389, psbA-trnH HM474835. Galagania margiana Pimenov & M. Vassiljeva: Turkmenistan, N slopes of hills near Kushka, 04.vi.1978, Pimenov & Kljuykov 267 (MW), ITS HM229390, psbA-trnH HM474836. Galagania neglecta M. Vassiljeva & Pimenov: Uzbekistan, Kashkadarja prov., Sakartma ridge, 01.vi.1976, Pimenov et al. 320 (MW), ITS HM229391, psbA-trnH HM474837. Galagania platypoda (Aitch. & Hemsl.) M. Vassiljeva & Pimenov: Turkmenistan, Kopet Dagh, between Nukhur and Alljam Mt., 11.vi.1978, Pimenov et al. 440 (MW), ITS HM229392, psbA-trnH HM474838. Galagania tenuisecta (Regel & Schmalh.) M. Vassiljeva & Pimenov: ITS DQ4228212, DQ4228402, psbA-trnH DQ4571762. Gongylotaxis rechingeri Pimenov & Kljuykov: Afghanistan, Bamyan prov., 23.vii.1962, Rechinger 18664 (E), ITS DQ5163587, psbA-trnH HM474839. Hellenocarum multiflorum (Sm.) H.Wolff (= Carum multiflorum (Sm.) Boiss.): ITS AY8409874, AY8409864. Hyalolaena bupleuroides (Schrenk ex Fisch. & C.A. Mey.) Pimenov & Kljuykov: ITS DQ4228222, DQ4228412, psbA-trnH EU4457582. Hyalolaena depauperata Korovin: Uzbekistan, Kashka-darja prov., to N from Karshi, 31.v.1978, Pimenov & Kljuykov 232 (MW), ITS HM229393, psbA-trnH HM474840. Hyalolaena intermedia Pimenov & Kljuykov: (1) Kirghizia, Dzhalalabad prov., N slope of Fergana Mts., valley of Bekachal river between Karakul and Tashkumyr, 11.vi.1996, Pimenov et al. K96–123 (MW), ITS HM229394, psbA-trnH HM474841; (2) Kirghizia, valley of Naryn river, between KaraKul’ and Tashkumyr, 27.vi.1969, Pimenov 828 (MW), ITS HM229395, psbA-trnH HM474842. Hyalolaena issykkulensis Pimenov & Kljuykov: Kirghizia, Terskei Alatau, valley of Chon-Kysylsu river, 14.vii.1974, Pimenov & Kljuykov 907 (MW), ITS HM229396, psbA-trnH HM474843. Hyalolaena jaxartica Bunge: ITS DQ4228232, DQ4228422, psbA-trnH EU4457592. Hyalolaena lipskyi (Korovin) Pimenov & Kljuykov: Turkmenistan, Badhys, Erojlandus lake, 07.vi.1978, Pimenov et al. 316 (MW), ITS HM229397, psbA-trnH HM474844. Hyalolaena melanorrhiza Pimenov & Kljuykov: Tadzhikistan, Badakhshan, valley of Pjandsh river, between Shipad and mouth of Shisgu river, 30.vii.1981, Pimenov et al. 1004 (MW), ITS HM229398, psbA-trnH HM474845. Hyalolaena transcaspica (Korovin) Pimenov & Kljuykov: Turkmenistan, Central Kopet Dagh, near Ashhabad, road to Bekrova, 02.v.1990, Pimenov & Kljuykov 64 (MW), ITS HM229399, psbA-trnH HM474846. Hyalolaena trichophylla (Schrenk) Pimenov & Kljuykov: ITS DQ4228242, DQ4228432, psbA-trnH EU4457602. Hyalolaena tschuiliensis (Pavlov ex Korovin) Pimenov & Kljuykov: ITS DQ4228252, DQ4228442, psbA-trnH EU4457612. Hyalolaena viridiflora Kljuykov: Kirghizia, Central Tien Shan, northern slope of At-Bashy ridge, valley of Bash-Kaindi river, 19.vii.1985, Kljuykov 207 (MW), ITS HM229400, psbA-trnH HM474847. Mogoltavia narynensis Pimenov & Kljuykov: Kirghizia, Osh prov., Toktogul distr., basin of Naryn river, at confluence of Kokomeren river, Ak-Tatyr tract, 13.06.1977, Aidarova et al. (MW), ITS HM229401, psbA-trnH HM474848. Mogoltavia sewerzowii (Regel) Korovin: ITS DQ4228272, DQ4228462, psbA-trnH DQ4571782. Oedibasis apiculata (Kar. & Kir.) Koso-Pol.: (1) Kazakhstan, Chimkent prov., Bel’tau Mts., 26.v.1976, Pimenov et al. 166 (MW), ITS HM229402, psbA-trnH HM474849; (2) ITS DQ4228282, DQ4228472, psbA-trnH DQ4571792. Oedibasis platycarpa (Lipsky) Koso-Pol.: ITS AF0091118, AF0086328, psbA-trnH EU4457632. Oedibasis tamerlanii (Lipsky) Korovin ex Nevski: Uzbekistan, Nuratau Mts., between Nawoi and Nurata, Karatau ridge, Kara-Karga pass, 24.iv.1987, Pimenov & Kljuykov 48 (MW), ITS HM229403, psbA-trnH HM474850. Physospermum cornubiense (L.) DC.: ITS AF0779045, psbA-trnH EU4457652. Postiella capillifolia (Post ex Boiss.) Kljuykov: ITS DQ4228292, DQ4228482, psbA-trnH EU4457662. Pyramidoptera cabulica Boiss.: ITS AF0086318, AF0091108, psbA-trnH DQ4571802. Rhabdosciadium aucheri Boiss.: ITS AH0088953, psbA-trnH EU4457672. Scaligeria napiformis (Willd. ex Spreng.) Grande: ITS DQ4228302, DQ4228492, psbA-trnH EU4457682. Scaligeria halophila (Rech. f.) Rech. f.: Greece, Safora, the island of Makri Safora, 06.v.1967, Bentzer & Runemark 28236 (LD), ITS HM229404, psbA-trnH HM474851. Scaligeria moreana Engstr.: Greece, Lakonia, 2 km N of Areopolis. 22.v.1964, Runemark & Snogerup 20727 (LD), ITS HM229405, psbA-trnH HM474852. Schulzia albiflora (Kar. & Kir.) Popov: ITS DQ4228312, DQ4228502, psbA-trnH DQ4571812. Schulzia crinita (Pall.) Spreng.: ITS AY3289499, AY3305159, psbA-trnH EU4457702. Schulzia prostrata Pimenov & Kljuykov: Kirghizia, Central Tien Shan, At-Bashy distr., basin of Aksaj river, valley of Terek river, 08.viii.1987, Pimenov & Kljuykov 790 (MW), ITS HM229406, psbA-trnH HM474854, HM474853. Tamamschjanella rubella (E.Busch) Pimenov & Kljuykov: ITS DQ4228332, DQ4228522, psbA-trnH EU4457752. Trachyspermum ammi (L.) Sprague: ITS U7838010, U7844010, psbA-trnH EU4457762.

1 Downie and Katz-Downie (1996), 2 Degtjareva et al. (2009), 3 Downie et al. (2000a), 4 Papini (2006), 5 Valiejo-Roman et al. (1998), 6 Valiejo-Roman et al. (2006b), 7 Spalik and Downie (2007), 8 Katz-Downie et al. (1999), 9 Valiejo-Roman et al. (2006a), 10 Downie et al. (1998).

Appendix 2

Morphological characters and character states used in the analysis

  1. 1.

    Life form (0, perennial polycarpic herb; 1, perennial monocarpic herb).

  2. 2.

    Thickened part of root (0, not branched; 1, branched).

  3. 3.

    Tubers (0, superficial; 1, deeply submerged in soil).

  4. 4.

    Leaf (0, soft; 1, rigid).

  5. 5.

    Basal leaflets (0, sessile; 1, petiolulate).

  6. 6.

    Shape of terminal leaflets (0, linear to filiform; 1, lanceolate; 2, ovate, dentate).

  7. 7.

    Rachis and petiolules of leaf (0, not refracted; 1, refracted).

  8. 8.

    Petioles (0, glabrous; 1, covered with short hairs).

  9. 9.

    Terminal leaflets (0, not dense; 1, dense).

  10. 10.

    Sheats of stem leaves (0, not inflated; 1, inflated).

  11. 11.

    Central (terminal) umbel (0, developed; 1, not developed).

  12. 12.

    Bracts (0, absent; 1, herbaceous; 2, broadly white-margined).

  13. 13.

    Pedicels (0, very short (up to 3 mm; 1, middle-sized (4–10 mm); 2, long (11–20 mm)).

  14. 14.

    Pedicels during fructification (0, thin (<0.7 mm in diam.); 1, thick (>0.7 mm in diam.)).

  15. 15.

    Calyx teeth (0, absent; 1, developed).

  16. 16.

    Petal shape and orientation (0, slightly emarginate, deflexed under 45°–90°; 1, slightly emarginate, deflexed downwards; 2, deeply emarginate, strict).

  17. 17.

    Petal color (0, white; 1, yellow or greenish-yellow; 2, purple).

  18. 18.

    Fruit separation at maturity (0, from below; 1, from above).

  19. 19.

    Mericarp shape (0, orbicular; 1, elliptic to ovate; 2, lanceolate or oblong; 3, linear).

  20. 20.

    Beak (the seed does not fill the top of the fruit) (0, present; 1, absent).

  21. 21.

    Mericarp in transverse section [0, strongly compressed dorsally (mericarp width more than three times the thickness); 1, dorsally convex; 2, slightly compressed dorsally (mericarp width less than three times the thickness)].

  22. 22.

    Dorsal mericarp ribs (0, almost obsolete; 1, keeled; 2, short-winged).

  23. 23.

    Marginal mericarp ribs (0, not winged; 1, winged).

  24. 24.

    Mericarps (0, not pubescent; 1, pubescent).

  25. 25.

    Stylopodium shape (0, flat; 1, short conical; 2, conical; 3, cylindrical).

  26. 26.

    Styles (0, recurved; 1, erect).

  27. 27.

    Exocarp cell size (0, large (>25 mcm); 1, small (<20 mcm)).

  28. 28.

    Width of mericarp commissure (0, narrow (exocarp almost reaches carpophore); 1, broad (exocarp interrupts near edges of marginal ribs)).

  29. 29.

    Secretory ducts (vittae) in mericarp furrows (0, solitary; 1, 2–3 per furrow; 2, numerous or cyclical).

  30. 30.

    Endosperm on commissural side (0, flat; 1, with a groove).

  31. 31.

    Cotyledon number (0, two; 1, one).

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Degtjareva, G.V., Kljuykov, E.V., Samigullin, T.H. et al. ITS phylogeny of Middle Asian geophilic Umbelliferae-Apioideae genera with comments on their morphology and utility of psbA-trnH sequences. Plant Syst Evol 299, 985–1010 (2013). https://doi.org/10.1007/s00606-013-0779-9

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