Abstract
The monospecific genus Aitchisonia (Rubiaceae), a suffrutescent plant endemic to Afghanistan, Iran, and Pakistan, was merged in the genus Plocama of the monogeneric tribe Putorieae sensu Backlund et al. (Spermacoceae alliance, subfamily Rubioideae). This decision was based on the phylogenetic placement of the specimen Rafei and Zangooei 25651 (FUMH) erroneously identified as Aitchisonia rosea in Plocama. This specimen was later correctly re-identified as Plocama dubia, rending the position of Plocama rosea (= Aitchisonia rosea) in Plocama (thus Putorieae) doubtful. We reconstructed a new phylogeny of Spermacoceae alliance based on trnL-F and nrITS sequence data and using the Bayesian method to: 1) assess the phylogenetic placement of P. rosea within Rubioideae, using new material from Iran; 2) pinpoint the phylogenetic positions of Plocama asperuliformis, P. crucianelloides, P. dezfulensis, P. ehrendorferi, P. kandaharensis, P. szowitsii, and P. vassilczenkoi; and 3) re-assess the monophyly of Plocama bruguieri and P. eriantha. The trnL-F and nrITS trees further support the monophyly of the Rubieae complex, but were not fully congruent. The trnL-F tree resolved Plocama rosea as sister to tribe Rubieae (including Kelloggia), while the nrITS tree was inconclusive. We described a new monogeneric tribe Aitchisonieae to accommodate P. rosea. Our nrITS analysis resolved Plocama crucianelloides in the Plocama Middle-Eastern clade and P. asperuliformis, P. dezfulensis, P. ehrendorferi, P. kandaharensis, P. szowitsii, and P. vassilczenkoi in the Plocama Central-Asian clade, both sensu Rincón-Barrado et al. The non-monophyly of P. bruguieri and P. eriantha is confirmed.
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Data availability
All sequence data are available in GenBank, and our DNA sequences are available as Online Resource 1 and 2 associated with this manuscript.
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Acknowledgements
The authors would like to thank the directors and curators of TUH, G, LE, K, B herbaria and the herbarium of Agricultural and Natural Resources Research Center of Khuzestan for providing plant materials. No funding was received to assist with the preparation of this manuscript.
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Appendices
Appendix
List of taxa investigated in molecular analyses, GenBank accession numbers for the sequences used, and voucher information for the new sequences in the following order: Taxon, voucher information (if applicable), nrITS, trnL–F (‒ indicates missing data). * new sequences for this study.
Aitchisonia rosea Hemsl. ex Aitch. Iran, S, Fars, Fasa, Mianjangal, Tang-e Ahru, 29°05′14"N, 53°26′05"E, 2044 m a. s. l., 20 Aug 2018, Bordbar 3115 (MIR) [MZ412534*, MZ388054*]; Amphidasya ambigua (Standl.) Standl. [‒, EU145576]; Anthospermum herbaceum L.f. [MK141272, MK141631]; Argostemma hookeri King [FJ695432, EU145545]; Asperula purpurea (L.) Ehrend.[‒, GU357130]; Asperula pinifolia (Boiss.) Ehrend. & Schönb.-Tem. [LM997574, ‒]; Asperula pulvinaris Heldr. ex Boiss. [LM997575, ‒]; Carphalea glaucescens (Hiern) Verdc.[AM266993, AM266906]; Coccocypselum condalia Pers. [EU145547]; Colletoecema dewevrei (De Wild.) E.M.A.Petit [‒, EU145532]; Coussarea pilosula C.M.Taylor [‒, MH95686]; Craterispermum longipedunculatum Verdc. [‒, LN680875]; Danais xanthorrhoea (K.Schum.) Bremek. [EU145364, AM409329]; Dunnia sinensis Tutcher [EU145394, EU145587]; Didymaea alsinoides (Cham. & Schltdl.) Standl. [‒, EU145570]; Galium album Forssk. [‒, GU357146]; Galium elongatum C.Presl [‒, GU357122]; Galium spurium L. [MG906007, ‒]; Galium. trifidum L. [MG905977, ‒]; Kelloggia chinensis Franch. 1 [MG906046, MF780799]; Kelloggia chinensis 2 [‒, MF780800]; Kelloggia galioides Torr. [FJ695443, DQ662146]; Leptodermis crassifolia Collett & Hemsl. [MH275603, MH323901]; Mitchella repens L. [‒, JK412389]; Mycetia tonkinensis (Pit.) Razafim. & B.Bremer [KP212781, KP212890]; Neurocalyx zeylanicus Hook. [‒, EU145560]; Palicourea guianensis Aubl. [AF152615]; Paederia praetermissa Puff [‒, KC306164]; Paederia foetida L. [HG004853, ‒]; Pauridiantha paucinervis (Hiern) Bremek. [‒, EU145578]; Pentas lanceolata (Forssk.) Deflers [AB247275, AM266963]; Plocama afghanica (Ehrend.) M.Backlund & Thulin, Afghanistan, Kandahar, 80 km W Kandahar, toward Girishk, 31°36'N, 65°47'E, 1100 m a. s. l., 21 May 1973, Rechinger 34814 (G) [MZ412538*, ‒]; Plocama asperuliformis (Lincz.) M.Backlund & Thulin, Pamir, Badakhshan, Tajikistan, the valley of the Viskharvi river, 2 km from the mouth, in the rocks, 2000 m a. s. l., K. Stanjukovich, L. Sidorov, M. Krivonogova, G. Ladygina and S. Ikonnikov 18884 (LE) [MZ412537*, ‒]; Plocama aucheri (Guill.) M.Backlund & Thulin, Iran, SE, Kerman, Kahnuj, 55 km to Manujan, 27°49′05.0"N, 57°40′41.3"E, 562 m a. s. l., 29 Mar 2011, Mirtadzadini 2351 (MIR) [MZ388057*, ‒]; Plocama botschantzevii (Lincz.) M.Backlund & Thulin, Uzbakistan, E, Prov.Termez, Mt. Pamir-Alay, Aktash, ca. 1000 m a. s. l., 15 Jun 1971, Botschantzev 50445 (LE) [MZ388110*, ‒]; Plocama brevifolia (Coss. & Durieu ex Pomel) M.Backlund & Thulin, Morocco, Berkane Prov., 850 m a. s. l., s.d., Faure 88387 (K) [MZ388127*, ‒]; Plocama bruguieri (A.Rich. ex DC.) M.Backlund &Thulin 1, Iran, NE, Khorasan, N Nehbandan, E Kalata Seied Ali village, 31°37′15.5"N, 60°01′13.8"E, 1435 m a. s. l., 2 May 2017, Mirtadzadini and Jafari 2373 (MIR) [MZ389910*, ‒]; Plocama bruguieri 2, Iran, SE, Kerman to Kuhpayeh, 26 May 2002, Mirtadzadini 2372 (MIR)[MZ407637*, ‒]; Plocama bruguieri 3, Iran, C, Yazd, W Ardekan, 1000 m a. s. l., May 1994, s.col., 28168 (TUH) [MZ389906*, ‒]; Plocama bruguieri 4, Iran, C, 80 km Isfahan to Kashan, 29 May 2007, Mirtadzadini 2368 (MIR) [MZ412545*, ‒]; Plocama bruguieri 5, Iran, S, Hormozgan, E Qeshm Island, 21 Apr 1995, Ghahreman, Attar and Hamzeh 19212 (TUH) [MZ409596*, ‒]; Plocama bruguieri 6, Iran, C, Tehran, Khojir Park, 28 Jul 1984, s.col., 28038 (TUH) [MZ389976*, ‒]; Plocama calabrica (L.f.) M.Backlund & Thulin [FJ695457, DQ662166]; Plocama calycoptera (Decne.) M.Backlund & Thulin, Iran, SE, NE of Giroft, between Saghder and Jiroft, 11 km before the bridge of Shur river, 28°49.204 N, 57°49.809E, 1431 m a. s. l., 9 May 2013, Mirtadzadini 2355 (MIR) [MZ389930*, ‒]; Plocama crucianelloides (Jaub. & Spach) M.Backlund &Thulin, Iran, SE, Kerman, Jiroft, Qale Gabrha, 2 May 2000, Mirtadzadini 2354 (MIR) [MZ412544*, ‒]; Plocama dezfulensis (Naanaie & Assadi) Bordbar & Mirtadz., Iran, Khuzestan Prov., Dezful, Shahion, 10 km after road Miankuh to Salan mountain, 1045 m a. s. l., 29 May 2001, Naanaie and Mohammadi 5173 (isotype, herbarium of Agricultral and Natural Resources Research Center of Khuzestan) [MZ412541*, ‒]; Plocama dubia (Aitch. & Hemsl.) M.Backlund & Thulin, Iran, NE: Khorasan, between Qaen and Khuaf, on slopes SW of Estend village, 33°50′42.03"N, 59°49′18.52"E, 17 Jun 2013, Mirtadzadini 2357 (MIR) [MZ412547*, DQ662134]; Plocama eriantha (Jaub. & Spach) M.Backlund & Thulin 1, Iran, SE, Balutshistan, between Zahedan and Khash, near Putshenki village, 28°50′12.0"N, 60°45′50.0"E, 2039 m a. s. l., 1 May 2017, Mirtadzadini 2362 (MIR) [MZ412539*, ‒]; Plocama eriantha 2, Iran, S, Fars, Fasa, Mianjangal, 29°12′39.21"N, 53°20′18.18"E, 1779 m a. s. l., 9 Jun 2017, Bordbar 2365 (MIR) [MZ389909*, ‒]; Plocama eriantha 3, Iran, SE, Kerman, Bardsir, 2 km to Bidkhun village, 19 Jun 2015, Bordbar 2366 (MIR) [MZ389975*, ‒]; Plocama ehrendorferi Mirtadz. & Bordbar, Iran, SE., Kerman Prov., 35 km from Jiroft to Esfandaqeh, 28°45′07.09"N, 57°29′35.78"E, 1175 m a. s. l., 8 May 2014, Mirtadzadini 2358 (MIR) [MZ412540*, ‒]; Plocama hymenostephana (Jaub. & Spach) M.Backlund & Thulin, Iran, SE, Balutshistan, Nikshahr to Sarhe, 11 Apr 1998, Mirtadzadini 2352 (MIR) [MZ412542*, DQ662163]; Plocama jolana (Thulin) M.Backlund & Thulin, Yemen, Hadheralmout, Jol Berka, 1200–1400 m a. s. l., 14°35–36'N, 48°40'E, 30 Nov 1999, Kilian et al. NK6465 (B) [MZ389931*, ‒]; Plocama kandaharensis (Ehrend. & Qarar ex Ehrend. & Schönb.-Tem.) M.Backlund & Thulin, Afghanistan, 52 km W Kandahar, 12 Jun 1958, Pabot A283 (G) [MZ409597*, ‒]; Plocama macrantha (Blatt. & Hallb.) M.Backlund &Thulin, Pakistan, W, Quetta, above Harnai, in the lower part of Torkhan, 900 m a. s. l., 14–15 May 1973, Rechinger 29510 (G) [MZ389878*, ‒]; Plocama olivieri [‒, DQ662142]; Plocama pendula Aiton [FJ695459, DQ662162]; Plocama reboudiana (Coss. & Durieu) M.Backlund & Thulin [‒, DQ662137]; Plocama szowitsii (DC.) M.Backlund & Thulin, Armenia, Ararat Prov., Ararat valley, 12 km NNW of Ararat town, 7 km NW of Vedi, Erah range, 1030 m a. s. l., s.m., 39°56'N, 44°42'E, 10 Apr 2002, Fayvush et al. OPTIMA Inter XI/0964 (G) [MZ412546*, ‒]; Plocama thymoides (Balf.f.) M.Backlund & Thulin, Yemen, Socotra, above village Qaysoh, N flank of Ma'ala massif, 12°38'N, 53°27'E, 150–300 m a. s. l., 30 Mar 2002, Kilian et al. YP2342 (B) [MZ389876*, ‒]; Plocama tinctoria (Balf.f.) M.Backlund & Thulin, s.loc., s.d., s.col., 88,386 (K) [MZ389887*, ‒]; Plocama vassilczenkoi (Lincz.) M.Backlund & Thulin, Kyrgyzstan, Sokh river basin, near the village Ak-Tupak, 880 m a. s. l., 40°16', 71°14', 4 Jun 2012, G.A. Lazkov 50444 (LE) [MZ408919*, ‒]; Plocama yemenensis (Thulin) M.Backlund & Thulin, Yemen: Lawdar, 26 Mar 1997, Hein 3626 (B) [MZ398140*, ‒]; Psychotria schliebenii E.M.A.Petit[‒, AM945361]; Rubia chinensis Regel & Maack [‒, MF780801]; Rubia tinctorum L. [KP098121, FJ695421]; Rubia hangii L.E Yang & Z.L.Nie [KX827253; ‒]; Saprosma foetens (Wight) K.Schum.[‒, DQ662168]; Saprosma fruticosa Aiton[FJ695461, ‒]; Saprosma ternatum (Wall.) Hook.f. [MH275998, ‒]; Schismatoclada sp. [JQ729845, EU145553]; Schizocolea linderi (Hutch. & Dalziel) Bremek. [‒, AM945338]; Schradera subandina K.Krause [‒, AM945343]; Serissa foetida (L.f.) Lam. [‒, AF152618]; Serissa serissoides (DC.) Druce [FJ980385, ‒]; Seychellea sechellarum (Baker) Razafim., Kainul. & Rydin [‒, MN437540]; Sherardia arvensis L. [‒, EU145571]; Spermadictyon suaveolens Roxb. [‒, DQ662171]; Theligonum cynocrambe L. 1 [‒, AF152621]; Theligonum cynocrambe 2 [FJ695469, FJ695426]; Theligonum cynocrambe 3 [FJ695470, FJ695427]; Theligonum formosanum (Ohwi) Ohwi & T.S.Liu [‒, MF780802]; Theligonum japonicum Ôkubo & Makino [‒, MF780803]; Theligonum macranthum Franch. [‒, MF780805].
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Online resource 1. Nexus file of our DNA sequence matrix of trnL-F dataset.
Online resource 2. Nexus file of our DNA sequence matrix of nrITS dataset.
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Bordbar, F., Mirtadzadini, M. & Razafimandimbison, S.G. Phylogenetic re-assessment of the delimitation of Plocama and its species relationships and limits (Rubiaceae, Putorieae): resurrection of the monospecific genus Aitchisonia and a description of trib. nov. Aitchisonieae. Plant Syst Evol 308, 7 (2022). https://doi.org/10.1007/s00606-021-01799-4
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DOI: https://doi.org/10.1007/s00606-021-01799-4