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	<front>
		<journal-meta>
			<journal-id journal-id-type="publisher-id">AJBM</journal-id>
			<journal-title-group>
				<journal-title>Anales del Jard&#xed;n Bot&#xe1;nico de Madrid</journal-title>
				<abbrev-journal-title abbrev-type="publisher">Anal. Jard. Bot. Madr.</abbrev-journal-title>
			</journal-title-group>
			<issn publication-format="print">0211-1322</issn>
			<issn publication-format="electronic">1988-3196</issn>
			<issn-l>0211-1322</issn-l>
			<publisher>
				<publisher-name>Consejo Superior de Investigaciones Cient&#xed;ficas</publisher-name>
			</publisher>
		</journal-meta>
		<article-meta>
			<article-id pub-id-type="publisher-id">ajbm.2588</article-id>
			<article-id pub-id-type="doi">10.3989/ajbm.2588</article-id>
			<article-categories>
				<subj-group subj-group-type="heading">
					<subject>Short note</subject>
				</subj-group>
			</article-categories>
			<title-group>
				<article-title>Morphology, anatomy and micromorphology of the genus <italic>Oliveria</italic> (Apiaceae)</article-title>
				<trans-title-group xml:lang="es">
					<trans-title>Morfolog&#xed;a, anatom&#xed;a y micromorfolog&#xed;a del g&#xe9;nero <italic>Oliveria</italic> (Apiaceae)</trans-title>
				</trans-title-group>
			</title-group>
			<contrib-group>
				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0002-2708-9944</contrib-id>
					<name>
						<surname>&#xc7;elik</surname>
						<given-names>Mustafa</given-names>
					</name>
					<aff id="aff1"><institution content-type="research-centre">Advanced Technology Research and Application Center</institution>, <institution>Sel&#xe7;uk University</institution>, <addr-line>Konya</addr-line>, <country>Turkey</country></aff>
				</contrib>
				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0001-9043-0394</contrib-id>
					<name>
						<surname>Alatli</surname>
						<given-names>Hamdiye</given-names>
					</name>
					<aff id="aff2"><institution content-type="school">Graduate School of Natural and Applied Sciences</institution>, <institution>Sel&#xe7;uk University</institution>, <addr-line>Konya</addr-line>, <country>Turkey</country></aff>
				</contrib>
				<contrib contrib-type="author">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-4591-1126</contrib-id>
					<name>
						<surname>G&#xfc;rl&#xfc;k</surname>
						<given-names>Leyla</given-names>
					</name>
					<aff id="aff3"><institution content-type="school">Graduate School of Natural and Applied Sciences</institution>, <institution>Sel&#xe7;uk University</institution>, <addr-line>Konya</addr-line>, <country>Turkey</country></aff>
				</contrib>
				<contrib contrib-type="author" corresp="yes">
					<contrib-id contrib-id-type="orcid">https://orcid.org/0000-0003-2886-3409</contrib-id>
					<name>
						<surname>&#xc7;etin</surname>
						<given-names>&#xd6;zlem</given-names>
					</name>
					<email xlink:href="ozlemcetin8419@gmail.com">ozlemcetin8419@gmail.com</email>
					<aff id="aff4"><institution content-type="department">Department of Biotechnology</institution>, <institution content-type="faculty">Faculty of Science</institution>, <institution>Sel&#xe7;uk University</institution>, <addr-line>Konya</addr-line>, <country>Turkey</country></aff>
				</contrib>
				<contrib contrib-type="editor">
					<name>
						<surname>Aedo</surname>
						<given-names>Carlos</given-names>
					</name>
				</contrib>
			</contrib-group>
			<pub-date pub-type="epub">
				<day>17</day>
				<month>12</month>
				<year>2021</year>
			</pub-date>
			<pub-date pub-type="collection">
				<month>12</month>
				<year>2021</year>
			</pub-date>
			<volume>78</volume>
			<issue>2</issue>
			<elocation-id>e117</elocation-id>
			<history>
				<date date-type="received">
					<day>14</day>
					<month>02</month>
					<year>2021</year>
				</date>
				<date date-type="accepted">
					<day>18</day>
					<month>06</month>
					<year>2021</year>
				</date>
				<date date-type="pub">
					<day>29</day>
					<month>11</month>
					<year>2021</year>
				</date>
			</history>
			<permissions>
				<copyright-statement>&#xa9; 2021 CSIC</copyright-statement>
				<copyright-year>2021</copyright-year>
				<license license-type="open-access" xlink:href="https://creativecommons.org/licenses/by/4.0/">
					<license-p>This is an open-access article distributed under the terms of the Creative Commons Attribution 4.0 International (CC BY 4.0) License.</license-p>
				</license>
			</permissions>
			<self-uri xlink:href="http://asclepio.revistas.csic.es/index.php/asclepio/article/view/XXXX/XXXX"/>
			<abstract>
				<title>Abstract</title>
				<p>
					<italic>Oliveria</italic> is a little-known, monotypic genus of the family Apiaceae. The only accepted species, <italic>O. decumbens</italic>, is distributed throughout Mesopotamia. It is an annual, aromatic plant that has been used in traditional medicine since ancient times. The aim of this study was to examine morphological, micromorphological, and anatomical characters of <italic>O. decumbens</italic> in order to increase the current botanical knowledge of this genus. </p>
			</abstract>
			<trans-abstract xml:lang="es">
				<title>Resumen</title>
				<p>
					<italic>Oliveria</italic> es un g&#xe9;nero monot&#xed;pico y poco conocido de la familia Apiaceae. La &#xfa;nica especie aceptada, <italic>O. decumbens</italic>, se distribuye en toda Mesopotamia. Es una planta anual y arom&#xe1;tica que ha sido usada en la medicina tradicional desde tiempos antiguos. El objetivo de este estudio fue examinar los caracteres morfol&#xf3;gicos, micromorfol&#xf3;gicos y anat&#xf3;micos de <italic>O. decumbens</italic> para incrementar el conocimiento bot&#xe1;nico actual del g&#xe9;nero.</p>
			</trans-abstract>
			<kwd-group>
				<kwd>Anatolia</kwd>
				<kwd>monotypic genus</kwd>
				<kwd>Iran</kwd>
				<kwd>Iraq</kwd>
				<kwd>pollen</kwd>
				<kwd>Turkey</kwd>
			</kwd-group>
			<kwd-group xml:lang="es">
				<kwd>Anatolia</kwd>
				<kwd>g&#xe9;nero monot&#xed;pico</kwd>
				<kwd>Iran</kwd>
				<kwd>Iraq</kwd>
				<kwd>polen</kwd>
				<kwd>Turqu&#xed;a</kwd>
			</kwd-group>
			<counts>
				<fig-count count="6"/>
				<table-count count="0"/>
				<equation-count count="0"/>
				<ref-count count="19"/>
				<page-count count="5"/>
			</counts>
		</article-meta>
	</front>
	<body>
		<p>The Apiaceae Lindl. family contains around 460 genera and 3700 species distributed worldwide, with Turkey being one of the richest countries in southwest Asia in terms of number of species. The country has 105 genera and approximately 495 species of Apiaceae followed in richness by Iran, Georgia, Syria, Azerbaijan, and Iraq. There are 60 genera endemic to Asia, of which 45 are monotypic (<xref ref-type="bibr" rid="B18">Sales &amp; Hedge 2013</xref>). Genera <italic>Chamaesciadium</italic> C.A.Mey.<italic>, Cymbocarpum</italic> DC. ex C.A.Mey<italic>., Diplotaenia</italic> Boiss.<italic>, Exoacantha</italic> Labill.<italic>, Fuernrohria</italic> K.Koch<italic>, Froriepia</italic> K.Koch<italic>, Gongylosciadium</italic> Rech.f.<italic>, Lisaea</italic> Boiss.<italic>, Oliveria</italic> Vent.<italic>, Ormosciadium</italic> Boiss.<italic>, Petroedmondia</italic> Tamamsch.<italic>, Rhabdosciadium</italic> Boiss.<italic>, Szovitsia</italic> Fisch. &amp; C.A.Mey<italic>, Thecocarpus</italic> Boiss. and <italic>Trigonosciadium</italic> Boiss. have small geographic ranges and are endemic to Turkey, Iran, and Iraq. Genera <italic>Aegokeras</italic> Raf.<italic>, Crenosciadium</italic> Boiss. &amp; Heldr.<italic>, Postiella</italic> Kljuykov<italic>, Ekimia</italic> H.Duman &amp; M.F.Watson are only endemic to Turkey.</p>
		<p>
			<italic>Oliveria decumbens</italic> Vent. is scatteredly distributed in small areas in Iran, Iraq, Syria, and Turkey. The genus <italic>Oliveria</italic> was established by <xref ref-type="bibr" rid="B19">Ventenat (1801)</xref>. <italic>Oliveria aucheri</italic> Jaub. &amp; Spach, <italic>O. bruguieri</italic> Jaub. &amp; Spach, and <italic>O. orientalis</italic> DC. were then published as new species and are currently regarded as synonyms of <italic>O. decumbens</italic>. <italic>Oliveria</italic> was placed into tribe <italic>Smyrneae</italic> by de <xref ref-type="bibr" rid="B7">Candolle (1830)</xref> and then transferred into tribe <italic>Ammineae</italic> by <xref ref-type="bibr" rid="B5">Bentham (1867)</xref> followed by <xref ref-type="bibr" rid="B6">Boissier (1872)</xref>. Afterwards, <italic>Oliveria</italic> was placed in tribe <italic>Apieae</italic> by <xref ref-type="bibr" rid="B16">Rechinger &amp; Hedge (1987)</xref>. These traditional <italic>Apiaceae</italic> classification systems were largely based on fruit anatomy and morphology. <italic>Apiaceae</italic> is undergoing a major taxonomic rearrangement within the angiosperms given that current systematic knowledge has changed in light of molecular phylogenetics (e.g., <xref ref-type="bibr" rid="B11">Jim&#xe9;nez-Mej&#xed;as &amp; Vargas 2015</xref>). A comprehensive molecular phylogenetic analysis of tribal classification of Apiaceae, based on DNA sequences, showed that <italic>Oliveria</italic> is sister to <italic>Trachyspermum</italic> within <italic>Pyramidoptereae</italic> (<xref ref-type="bibr" rid="B14">Mousavi et al. 2020</xref>).</p>
		<p>Plants from the <italic>Apiaceae</italic> family are commonly used for food, flavoring, fragrance, and medicines. They are also known for their uses as household remedies for centuries. Many recent experimental and biological investigations have been conducted to validate the ethnomedicinal claims of the plants belonging to this family (<xref ref-type="bibr" rid="B1">Ahmad &amp; al. 2018</xref>). In traditional medicine, <italic>Oliveria</italic> has been used for indigestion, diarrhea, abdominal pain, fever, and skin wounds since ancient times (<xref ref-type="bibr" rid="B3">Amin &amp; al. 2005</xref>; <xref ref-type="bibr" rid="B12">Mahboubi &amp; al. 2016</xref>; <xref ref-type="bibr" rid="B2">Alizadeh-Behbahani &amp; al. 2018</xref>). There are many studies on the antioxidant, antihemolytic, anticancer, and anti-inflammatory properties of <italic>Oliveria</italic>. However, there are no detailed morphological, anatomical, and palynological studies about this genus (<xref ref-type="bibr" rid="B17">Sajjadi &amp; Hoseini 2002</xref>; <xref ref-type="bibr" rid="B3">Amin &amp; al. 2005</xref>; <xref ref-type="bibr" rid="B12">Mahboubi &amp; al. 2016</xref>; <xref ref-type="bibr" rid="B9">Eftekhari &amp; al. 2019</xref>). For this reason, the aim of this study was to investigate the morphological, carpological, and palynological characteristics of <italic>O. decumbens</italic> and to evaluate their significance in Apiaceae taxonomy.</p>
		<sec id="sec1">
			<title>Updated morphological description</title>
			<p>
				<bold>Oliveria decumbens</bold> Vent., Descr. Pl. Nouv., pl. 21 (1801). Type: cultivated in Hort. Cels, s.d., E.P. Ventenat s.n. (lectotype designated by <xref ref-type="bibr" rid="B8">Callmander &amp; al. 2017: G 00341532, image!</xref>). <xref ref-type="fig" rid="f1">Fig. 1</xref>-<xref ref-type="fig" rid="f2">2</xref>. </p>
			<fig id="f1">
				<label>Fig. 1</label>
				<caption>
					<title>
						<italic>Oliveria decumbens Vent.</italic></title>
						<p><bold>a,</bold> plant in the field; <bold>b,</bold> umbel [based on &#xc7;elik 1103 (GAZI)].</p>
				</caption>
				<graphic id="gra-1" xlink:href="AJBM-78-02-e117-gf1.png"/>
			</fig>
			<fig id="f2">
				<label>Fig. 2</label>
				<caption>
					<title>
						<italic>Oliveria decumbens</italic> Vent.</title>
						<p><bold>a,</bold> flowers; <bold>b,</bold> bracts and bracteoles [based on &#xc7;elik 1103 (GAZI)].</p>
				</caption>
				<graphic id="gra-2" xlink:href="AJBM-78-02-e117-gf2.png"/>
			</fig>
			<p>Ascending to erect or rarely decumbent, aromatic, annual herbs. Stem (13-)20-45(-75) cm tall, rigid, whitish, &#xb1; much branched, glabrous or rarely hirsute, solid, terete, 2-4 mm in diameter at base. Basal leaves soon withering; 3-10 cm long (inc. petiole), lamina oblong in outline, &#xb1; glabrous, 1-pinnate with 4-5 pairs of segments; each segment divided into smaller, decurrent lobes; ultimate segments lanceolate, &#xb1; toothed; petioles 0.6-2.5 cm long with sheathing base. Lower cauline leaves similar to basal leaves; middle and upper cauline leaves ovate, subsessile or sessile, pilose, pinnatisect. Umbels 1.5-2.5 cm in diameter, &#xb1; condensed, (2-)3-5(-7) rayed; rays (2-)3-10 mm long, slightly thickened, hairy, &#xb1; equal. Bracts 3-5, 3-8 mm long, obovate, dissected, densely pilose. Flowers 15-35(45), hermaphrodite, condensed, pedicellate. Pedicels 0.5-2 mm long, slightly thickened, rounded, hairy. Bracteoles 6-8, 2.5-5 mm long, dissected, obovate, densely pilose. Sepals 5, 0.5-0.75 mm, distinct, persistent, ovate-triangular, acute, hairy. Petals 5, 1-1.6 &#xd7; (0.6-)0.8-1.1 mm, white or pink, &#xb1; oblong-obovate, emarginate, with narrow tip bent inward, hairy on abaxial surface. Ovary hairy. Stamens enrolled; filaments whitish, c. 1.5 mm long; anthers 0.4-0.9 mm, sub-globular, dorsally inserted. Fruits 2.0-3.6 &#xd7; 1-1.6 mm, oblong; mericarps densely hirsute, laterally compressed; ribs filiform, obscured by hair; stylopodium &#xb1; conical; style 1.0-1.8 mm, slender, erect; vittae large, oblong, vallecular vitta 1; commissural vittae 2. </p>
			<p>
				<italic>Distribution and phenology.-Oliveira decumbens</italic> is distributed along the northwest-southeast transect of a narrow area in the western and southern region of Iran and the northern regions of Iraq (<xref ref-type="fig" rid="f3">Fig. 3</xref>; <xref ref-type="app" rid="app1">Appendix 1</xref>) at elevations between 150 and 1700 m. In Turkey, the species is only known from the Ceylanp&#x131;nar region near the border with Syria. It grows in steppes and flowers in May and July. </p>
			<fig id="f3">
				<label>Fig. 3</label>
				<caption>
					<title>Map showing the distribution and locations of the studied specimens of <italic>Oliveria decumbens</italic> Vent.</title>
					<p>(numbers correspond to those given in <xref ref-type="app" rid="app1">Appendix 1</xref>)</p>
				</caption>
				<graphic id="gra-3" xlink:href="AJBM-78-02-e117-gf3.png"/>
			</fig>
			<p>
				<italic>Fruit anatomy and micromorphology.-</italic>The fruit of <italic>Oliveira decumbens</italic> is oblong-elliptic in the transverse section. Mericarps are homomorphic, nearly circular, and have five projecting ribs. The exocarp is a single layer and consists of rectangular-like cells near the distal ends of the marginal ribs. Their walls are slightly thickened and the outer walls are covered with a smooth cuticle layer and simple unicellular trichomes, which originate from the epidermal cells. The mesocarp is multilayered with thin-walled parenchymatous cells. They are irregularly shaped and polygonal. Vascular bundles are located under the ribs and are surrounded by mesocarp cells. The vittae are large and oblong-elliptic or nearly orbicular. They are lined with thin-walled epithelial cells. There is one vitta per vallecula and two commissural vittae. The endocarp is single layer and consists of thin-walled cells. Endosperm cells are irregular, polygonal in shape, and are filled with granular contents (<xref ref-type="fig" rid="f4">Fig. 4</xref>). The fruit of <italic>O. decumbens</italic> is covered with dense, long, hirsute hairs that have a papillose surface (<xref ref-type="fig" rid="f6">Fig. 6</xref>).</p>
			<fig id="f4">
				<label>Fig. 4</label>
				<caption>
					<title>Cross-section of a fruit of <italic>Oliveria decumbens</italic> Vent.</title>
					<p>[based on &#xc7;elik 1103 (GAZI); scale: 100 &#x3bc;m]</p>
				</caption>
				<graphic id="gra-4" xlink:href="AJBM-78-02-e117-gf4.png"/>
			</fig>
			<p>
				<italic>Palynology.-</italic>Pollen materials were taken from herbarium specimen. Light microscopy (LM) observations were carried out on acetolyzed pollen and prepared according to <xref ref-type="bibr" rid="B10">Erdtman (1952)</xref>. Measurements, based on 50 samples from one population, were taken with a Leica microscope. For SEM analyses, the acetolyzed pollen grains were mounted on stub and coated with gold, using a Polaron SC7620 sputter coater. Microphotographs were taken using an incorporated Zeiss LS-10 camera. The terminologies of <xref ref-type="bibr" rid="B10">Erdtman (1952)</xref> and Faegri &amp; Iversen (1975) were used for the micromorphological features. The pollen grains of <italic>Oliveria decumbens</italic> are monad, isopolar, radially symmetric, and tricolporate (<xref ref-type="fig" rid="f5">Fig. 5</xref>). The polar axis (P) ranges from 28.30 to 35.55 &#xb5;m and the equatorial axis (E) ranges from 11.39 to 16.58 &#xb5;m. The mean length of the polar axis (P) is 30.05 &#xb1; 2.30 &#x3bc;m, and the mean of the equatorial diameter (E) is 12.68 &#xb1; 1.74 &#x3bc;m. P/E averages 2.29 &#xb1; 0.16, making the pollen grain shape perprolate. In polar view, the pollen grains are nearly triangular with obtuse angles. In an equatorial view, the grains are narrowly oblong with slightly equatorial constriction and obtuse polar caps. The colpus extends nearly to the poles. The pore area is slightly protruding and located in the middle of colpus. The sculpturing pattern is striate or psilate at the polar area and irregularly rugulate at the equatorial area, but sometimes striate at the polar area and psilate-foveolate toward the polar area. The fruit of <italic>O. decumbens</italic> is covered with dense, long, hirsute hairs that have a papillose surface (<xref ref-type="fig" rid="f6">Fig. 6</xref>).</p>
			<fig id="f5">
				<label>Fig. 5</label>
				<caption>
					<title>SEM images of pollen grains of <italic>Oliveria decumbens</italic> Vent.</title>
					<p><bold>a,</bold> general view; <bold>b,</bold> detail of pollen grain surface [based on &#xc7;elik 1103 (GAZI)].</p>
				</caption>
				<graphic id="gra-5" xlink:href="AJBM-78-02-e117-gf5.png"/>
			</fig>
			<fig id="f6">
				<label>Fig. 6</label>
				<caption>
					<title>SEM images of fruits of <italic>Oliveria decumbens</italic> Vent.</title>
					<p><bold>a,</bold> general shape of fruit; <bold>b, c, d,</bold> detail of fruit surface [based on &#xc7;elik 1103 (GAZI)]</p>
				</caption>
				<graphic id="gra-6" xlink:href="AJBM-78-02-e117-gf6.png"/>
			</fig>
			<p>
				<italic>Comments.-Oliveria</italic> is very isolated from other members of Apiaceae and its morphological characteristic is rather distinctive and apparently does not closely resemble other genera in tribe <italic>Apieae</italic>. <italic>Oliveria</italic> is characterized by a much-branched and bright whitish stem; conspicuous and persistent sepals; densely hairy and divided bracts and bracteoles; dorsally hairy and white or pink petals; and slightly laterally compressed, densely hairy, and ovoid-oblong fruits.</p>
			<p>Although our description corresponds to that in Flora of Turkey (<xref ref-type="bibr" rid="B13">Matthews 1972</xref>), there are several differences as follows: the stem is (13)20-45(60) cm (not 20-40 cm), basal leaves 3-10 cm long (not determined), the rays are (2-)3-5(-7) (not 3-5) in number and 3-5(-10) mm long (not 5-10 mm long), bracts are 3-5 (not determined) in number, 3-8 mm long (not determined), sepals are 0.5-0.75 mm long (not determined), petals are 1-1.6 x (0.6-) 0.8-1.1 mm in size (not determined). </p>
			<p>One of main characteristics of <italic>Oliveria</italic> is its shiny white and glabrous stem. Similar stem structure is also seen in <italic>Ergocarpon</italic> C.C.Towns., <italic>Eryngium</italic> L., and <italic>Schumannia</italic> Kuntze. <xref ref-type="bibr" rid="B4">Amiri &amp; al. (2011)</xref> have studied the anatomical characteristics of the leaf and stem of <italic>Oliveria</italic>. Their studies have shown that the surface of leaves is covered by non-glandular hairs, while the cross section of the stems show secretory canals in the cortex region. In the present study, we examined the anatomy and morphology of the fruit. According to our findings, the fruit of <italic>O. decumbens</italic> is oblong, laterally compressed, and densely hirsute. The bracts of <italic>Oliveria</italic> are divided and resemble those of <italic>Ammi</italic> L., <italic>Artedia</italic> L., <italic>Cuminum</italic> L., <italic>Daucus</italic> L., <italic>Eremodaucus</italic> Bunge, <italic>Kalakia</italic> Alava, <italic>Lagoecia</italic> L., <italic>Ormosciadium</italic> Boiss., <italic>Scandix</italic> L., and <italic>Trachyspermum</italic> Link. </p>
			<p>According to <xref ref-type="bibr" rid="B10">Erdtman (1952)</xref>, Umbelliferae species are stenopalynous. Observations by <xref ref-type="bibr" rid="B15">Perveen &amp; Qaiser (2006)</xref> have indicated that the pollen grains of Apiaceae are usually radially symmetrical, isopolar, and prolate to perprolate and tricolporate (rarely 4-colporate). The tectal surface is commonly striate-rugulate, rugulate-striate, or simple striate. Our result confirms the stenopalynous characteristic of the family Apiaceae, and the palynological observations reveal that the pollen grains are perprolate in shape and irregularly rugulate in the equatorial area.</p>
			<p>Our study provides a detailed account of the morphology, palynology, and micromorphology of <italic>Oliveria</italic>, increasing our botanical knowledge of a species that has been traditionally used since ancient times.</p>
		</sec>
	</body>
	<back>
		<ref-list>
			<title>References</title>
			<ref id="B1">
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