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  <front>
    <article-meta>
      <title-group>
        <article-title>Isoëtes gymnocarpa and Utricularia × neglecta – new taxa for Montenegro</article-title>
      </title-group>
      <contrib-group content-type="author">
        <contrib contrib-type="person">
          <name>
            <surname>Romanov</surname>
            <given-names>Roman E.</given-names>
          </name>
          <email>streptophytes@gmail.com</email>
          <xref ref-type="aff" rid="aff-1"/>
        </contrib>
        <contrib contrib-type="person">
          <name>
            <surname>Dragićević</surname>
            <given-names>Snežana</given-names>
          </name>
          <email>sneza.dragicevic72@gmail.com</email>
          <xref ref-type="aff" rid="aff-2"/>
        </contrib>
        <contrib contrib-type="person">
          <name>
            <surname>Troia</surname>
            <given-names>Angelo</given-names>
          </name>
          <email>angelo.troia@unipa.it</email>
          <xref ref-type="aff" rid="aff-3"/>
        </contrib>
      </contrib-group>
      <aff id="aff-1">
        <institution>Komarov Botanical Institute of the Russian Academy of Sciences, Professora Popova 2, 197022 St. Petersburg, Russia</institution>
        <country>Montenegro</country>
      </aff>
      <aff id="aff-2">
        <institution>Montenegrin Academy of Sciences and Arts, Rista Stijovića 5, 81000 Podgorica, Montenegro</institution>
        <country>Montenegro</country>
      </aff>
      <aff id="aff-3">
        <institution>Dipartimento STEBICEF, Università degli Studi di Palermo, via Archirafi 38, 90123 Palermo, Italy</institution>
        <country>Italy</country>
      </aff>
      <history>
        <date date-type="received" iso-8601-date="2023-07-14">
          <day>14</day>
          <month>07</month>
          <year>2023</year>
        </date>
        <date data-type="published" iso-8601-date="2024-09-27">
          <day>27</day>
          <month>09</month>
          <year>2024</year>
        </date>
      </history>
    <pub-date pub-type="epub"><day>27</day><month>09</month><year>2024</year><volume>83</volume></pub-date></article-meta>
  </front>
  
  
<body id="body">
    <sec id="heading-49e01b4231cf761c091ee47cd99b1e63">
      <title>Introduction</title>
      <p id="_paragraph-3">The flora of Montenegro (SE Europe) is easily recognized as one of the richest in Europe by the ratio of the ­number of species per square km (Pulević 2022). This is a reflection of evident habitat heterogeneity and richness ­resulting in large amounts of different ecological niches available for plants. A long tradition of botanical research has resulted in the more than 3,600 species and subspecies of vascular plants known from the Montenegrin flora (Rohlena 1942, Pulević 2005, 2022, Stešević et al. 2008, Stešević &amp; Caković 2013, 2021). However, some habitat types, e.g., small temporary and permanent water bodies and wetlands in coastal region, have received less attention (cf. Bubanja 2016, Bubanja et al. 2016). A lot of them can be referred to “Mediterranean temporary ponds”, a priority habitat according to the European Union 92/43 Habitats ­Directive, showing a clearly regressive trend. Distributional patterns of plants associated with these habitats in the ­Balkan Peninsula are still poorly known. </p>
    </sec>
    <sec id="heading-c39c48600503ce5646237bf181193b5d">
      <title>Materials and methods</title>
      <p id="_paragraph-5">Some localities in the coastal region and Zeta plain of Montenegro were surveyed during 2022 and 2023 as potential habitats for charophytes (Characeae), focussing on ­species of plants associated with them. All types of water bodies and wetlands encountered were checked, including Mediterranean temporary ponds, inundated sand pits, quarries, and artificial depressions, excavated ponds, drainage channels, puddles, concrete ponds, temporary and ­permanent streams and springs and the water bodies associated with them. The strong seasonality in the traits of ­water bodies and the appearance of aquatic plants as well as habitat heterogeneity are remarkable for the area surveyed. The specimens were collected by hand and studied in a ­living state. Voucher specimens were deposited in the Herba­rium of the Natural History Museum of Montenegro (NHMM) and the Herbarium Mediterraneum Panormi­tanum (PAL). The nomenclature for lycophytes follows Troia &amp; Greuter (2014), for <italic id="_italic-1">Utricularia</italic>, Bobrov et al. (2022). </p>
    </sec>
    <sec id="heading-35d599aca6f0bb4bce55de14213d215f">
      <title>Results and discussion</title>
      <p id="_paragraph-7">Fifteen species of charophytes, belonging to the genera <italic id="_italic-2">Chara</italic> L., <italic id="_italic-3">Lamprothamnium</italic> J.Groves, <italic id="_italic-4">Nitella</italic> C. Agardh, <italic id="_italic-5">Tolypella</italic> (A.Braun) A. Braun, and <italic id="_italic-6">Sphaerochara</italic> Mädler, were found in the research area. Our efforts aimed at the listing of all associated vascular plant species resulted in the finding of one genus and two species, new for the flora of Montenegro.</p>
      <p id="_paragraph-8"><italic id="_italic-7">Isoëtes gymnocarpa</italic> (Gennari) A. Braun (Fig. 1A): North of the Nature Park Ulcinjska Solana (US), unshaded drainage channel within farmland moderately damaged by livestock and probably briefly inundated during winter and ­early spring, 41.92966 N, 19.27361 E, 3 m a.s.l., 17 IV 2023, leg: Roman Romanov (RR) and Snežana Dragićević, det: Angelo Troia (NHMM, PAL). </p>
      <fig id="fig1">
        <label>Figure 1</label>
        <caption>
          <p id="_paragraph-9"/>
        </caption>
        <graphic id="_graphic-1" mimetype="image" mime-subtype="jpeg" xlink:href="image1.jpeg"/>
      </fig>
      <p id="_paragraph-10">
        <bold id="_bold-1">Fig. 1. New genus and species records for Montenegro: A – <italic id="_italic-8">Isoëtes gymnocarpa</italic>, general appearance of unearthed plant, B, C – <italic id="_italic-9">Utricularia × neglecta</italic>, B – isolated corolla showing large flat lower lip, view from above, C – isolated corolla showing short straight spur and nearly right angle between upper and lower lips, lateral view. Scale: B, C – 1 cm. All photos were taken from the voucher specimens by R. E. Romanov.</bold>
      </p>
      <p id="_paragraph-11">This habitat is characterized by the specific water ­regime, which implies an alternation of wet (winter, spring) and dry phases (summer and autumn), just like that in the meadows in the immediate vicinity with mosaic habitats and communities of wet grasslands and pastures which host ­Mediterranean therophytes and geophytes. These agricultural lands are pastures grazed by livestock (sheep, cows) or partly arable land, with other negative impacts in the ­immediate environment (like cutting of surrounding trees and shrubs and the associated physical disturbance). The cover abundance of <italic id="_italic-10">Isoëtes</italic> was low, not exceeding 20%. The plants were spotted within an area of less than 10 square meters. </p>
      <p id="_paragraph-12">The locality is currently outside the Nature Park Ulcinjska Salina, but because of the presence of this rare species of <italic id="_italic-11">Isoëtes</italic>, it should be protected by the Park area being extended to include it.</p>
      <p id="_paragraph-13"><italic id="_italic-12">Isoëtes gymnocarpa</italic> is close to <italic id="_italic-13">I. histrix</italic> Bory (for its terrestrial habitat, and for the presence of phyllopodia and tuberculate megaspores), the main diagnostic character being the length of phyllopodia lateral teeth (not longer and thinner than the central tooth as in <italic id="_italic-14">I. histrix</italic>, but as long and as thick as the central tooth, cf. Troia &amp; Greuter 2014, 2015a, 2015b). The group needs further investigation, but we follow here the taxonomic view of Troia &amp; Greuter (2015a) with <italic id="_italic-15">I. gymnocarpa</italic> = <italic id="_italic-16">I. sicula</italic> Tod. = <italic id="_italic-17">I. subinermis</italic> (Gennari) Cesca &amp; Peruzzi. The taxon so circumscribed is spread around the Mediterranean area, from the Iberian Peninsula to Anatolia: the population here reported is the northernmost along the Adriatic coasts, both in the Balkan Peninsula and in Italy. A previous report of <italic id="_italic-18">I. histrix</italic> for southern Albania (Barina et al. 2013) should be checked to assess if it belongs to <italic id="_italic-19">I. histrix</italic> or to <italic id="_italic-20">I. gymnocarpa</italic>. </p>
      <p id="_paragraph-14"><italic id="_italic-21">Isoëtes gymnocarpa</italic> should be recognized as indigenous to Montenegro because no cases of non-native species are known for this genus. The inconspicuous general habit of terrestrial <italic id="_italic-22">Isoëtes</italic>, having the appearance of sterile monocots, can be a possible explanation why species of this genus were not spotted in Montenegro before. </p>
      <p id="_paragraph-15"><italic id="_italic-23">Utricularia × neglecta </italic>Lehm. (Fig. 1B, C): Morača River, the lower reaches, the island of Vranjina, the wetland of ­Bakine Tigle (BT), 19 VI 2023, 42.28217 N, 19.15145 E, 4 m a.s.l., 19 VI 2023, leg: RR, det: RR (NHMM); BT, small floodplain pool, 42.28746 N, 19.15261 E, 5 m a.s.l., 19 VI 2023, leg: RR, det: RR (NHMM). The cover abundance was low, not exceeding 20 % in both cases.</p>
      <p id="_paragraph-16">This hybrid is widely distributed in temperate and ­tropical regions of the Old World (Taylor 1989, Uotila 2013, Bobrov et al. 2022, <ext-link id="_external-link-1" xlink:href="http://www.plantsoftheworldonline.org/">http://www.plantsoftheworldonline.org/</ext-link>). For a long time, it was reported as <italic id="_italic-24">U. australis</italic> R.Br. It was mentioned from Montenegro, from Lake Skadar, based on images of sterile plants, not showing the leaf-teeth bristle character (<ext-link id="_external-link-2" xlink:href="https://www.inaturalist.org/observations/31338617">https://www.inaturalist.org/observations/31338617</ext-link>), a trait only visible with microscopy. ­Moreover, the use of solely this character can be unreliable (­Taylor 1989). Therefore, our records are either the first ­verified ones for Montenegro, or else can be recognized as a confirmation of an earlier uncheckable record. The infrequent and short period of flowering, usually associated with habitats difficult of access, hampers reliable recording of <italic id="_italic-25">Utricularia</italic> species. This hybrid seems to be indigenous to Montenegro; no ­cases of introduction are known for it in the Balkan Peninsula.</p>
    </sec>
    <sec id="heading-bfc3116adff3ad97d005ac7db9fe11b5">
      <title>Acknowledgments</title>
      <p id="_paragraph-18">The anonymous reviewers are gratefully acknowledged for their encouragement and kind suggestions for the improvement of the manuscript.</p>
      <p id="_paragraph-19">This research is supported by the project “BioMon – ­Establishing biodiversity monitoring in Ulcinj Salina”, ­supported by the IKI Small Grants programme, which is part of the International Climate Initiative (IKI). IKI Small Grants is funded by the Federal Ministry of Economic ­Affairs and Climate Action (BMWK) and the Federal ­Foreign Office (AA), Germany, and carried out by Deutsche Gesellschaft für Internationale Zusammenarbeit (GIZ) GmbH. The BioMon project is implemented by the Center for Protection and Research of Birds (CZIP), a BirdLife ­partner in Montenegro. This research is partly supported by the project of the Komarov Botanical Institute RAS (No. 121021600184-6). </p>
    </sec>
    <sec id="heading-7772b5ec883269242d09c2b501fdfe52">
      <title>Author contribution statement</title>
      <p id="_paragraph-21">Roman E. Romanov – field studies, preparation and identification of specimens, manuscript writing; Snežana Dragićević – field studies, preparation of specimens, manuscript writing; Angelo Troia – preparation and identification of specimens, manuscript writing.</p>
    </sec>
    <sec id="heading-9889794762fb415c9dbb20a04d8e13a9">
      <title>References</title>
      <p id="_paragraph-23">Barina, Z., Rakaj, M., Pifkó, D., 2013: Contributions to the flora of Albania, 4. Willdenowia 43(1), 165–184. <ext-link id="external-link-f69c19140ba6c45c30b1bdc2baec7dc2" xlink:href="http://dx.doi.org/10.3372/wi.43.43119">http://dx.doi.org/10.3372/wi.43.43119</ext-link></p>
      <p id="_paragraph-24">Bobrov, A. A., Volkova, P. A., Kopylov-Guskov, Y. O., Mochalova, O. A., Kravchuk, A. E., Nekrasova, D. M., 2022: Unknown sides of <italic id="_italic-26">Utricularia</italic> (Lentibulariaceae) diversity in East Europe and North Asia or how hybridization explained old taxonomical puzzles. Perspectives in Plant Ecology, Evolution and Systematics 54, 125649. <ext-link id="external-link-38dc83c4b98b303cc3787edf63462a00" xlink:href="https://doi.org/10.1016/j.ppees.2021.125649">https://doi.org/10.1016/j.ppees.2021.125649</ext-link></p>
      <p id="_paragraph-25">Bubanja, N., 2016: Geobotanical characterization of vascular flora of long Ulcinj beach and its hinterland in Montenegro. Doctoral dissertation. University of Belgrade, Faculty of biology, Belgrade.</p>
      <p id="_paragraph-26">Bubanja, N, Šinžar-Sekulić, J., Stevanović,V., 2016: Assessing the influence of environmental parameters on aquatic plants of ponds in the hinterland of Long Beach in Montenegro. Limnetica 35(2), 385–396. <ext-link id="external-link-ec8887598edd65bda095e7466ed776f4" xlink:href="https://doi.org/10.23818/limn.35.31">https://doi.org/10.23818/limn.35.31</ext-link></p>
      <p id="_paragraph-27">Pulević, V., 2005: Material for the vascular flora of Montenegro. Posebno izdanje Republičkog zavoda za zaštitu prirode Crne Gore, Podgorica. </p>
      <p id="_paragraph-28">Pulević, V. (ed.), 2022: Botanical lexicon of Montenegro. Montenegrin Academy of Sciences and Arts, Lexicographic center, Lexicons, Book 3, Podgorica.</p>
      <p id="_paragraph-29">Rohlena, J., 1942: Conspectus florae Montenegrinae [An overview of the Montenegrin flora]. Preslia 20–21, 1–506.</p>
      <p id="_paragraph-30">Stešević, D., Caković, D., 2013: Katalog vaskularne flore Crne Gore [The catalog of the vascular flora of Montenegro]. T. 1. Podgorica.</p>
      <p id="_paragraph-31">Stešević, D., Caković, D., 2021: Katalog vaskularne flore Crne Gore [The catalog of the vascular flora of Montenegro]. T. 2. Podgorica.</p>
      <p id="_paragraph-32">Stešević, D., Petrović, D., Bubanja, N., Vuksanović, S., Biberdžić, V., 2008: Contribution to the flora of Montenegro (Supplementum to the material for vascular flora of Montenegro). Natura Montenegrina, Podgorica 7(3), 463–480. </p>
      <p id="_paragraph-33">Taylor, P., 1989: The genus <italic id="_italic-27">Utricularia</italic>: a taxonomic monograph. London.</p>
      <p id="_paragraph-34">Troia, A., Greuter, W., 2014: A critical conspectus of Italian <italic id="_italic-28">Isoetes</italic> (Isoetaceae). Plant Biosystems 148(1), 13–20. <ext-link id="external-link-235517708d8093664efb59df06683155" xlink:href="https://doi.org/10.1080/11263504.2013.878409">https://doi.org/10.1080/11263504.2013.878409</ext-link></p>
      <p id="_paragraph-35">Troia, A., Greuter, W., 2015a: Isoetaceae (vers. 1.0). In: Peruzzi, L., Cecchi, L., Cristofolini, G., Domina, G., Greuter, W., Nardi, E., Raimondo, F. M., Selvi, F., Troia, A. (eds.), Flora critica d’Italia. Fondazione per la Flora Italiana, Firenze. Published online on 25 February 2015. Available at: <ext-link id="_external-link-3" xlink:href="https://floraitaliana.it/contenuti/flora-critica/6">https://floraitaliana.it/contenuti/flora-critica/6</ext-link></p>
      <p id="_paragraph-36">Troia, A., Greuter W., 2015b: A conspectus of and key to Greek <italic id="_italic-29">Isoetes</italic> (Isoetaceae), based on a reassessment of Haussknecht’s gatherings of 1885. Willdenowia 45(3), 391–403. <ext-link id="external-link-b0add79f54b2406e26be9cd2a1a904bd" xlink:href="https://doi.org/10.3372/wi.45.45303">https://doi.org/10.3372/wi.45.45303</ext-link></p>
      <p id="_paragraph-37">Uotila, P., 2013: Lentibulariaceae. Euro+Med Plantbase – the ­information resource for Euro-Mediterranean plant di­versity.</p>
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