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Micromorphology of an Involucral Bract and Cypselae of the Endemic and Relict Species Amphoricarpos Elegans Albov (Asteraceae, Cardueae) from the Caucasus

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Micromorphology of an Involucral Bract and Cypselae of the Endemic and Relict Species Amphoricarpos Elegans Albov (Asteraceae, Cardueae) from the Caucasus (2020) 44 (1): 37-46 DOI: https://doi.org/10.2298/BOTSERB2001037G journal homepage: botanicaserbica.bio.bg.ac.rs Original Scientific Paper Micromorphology of an involucral bract and cypselae of the endemic and relict species Amphoricarpos elegans Albov (Asteraceae, Cardueae) from the Caucasus Milan Gavrilović1✳, Núria Garcia-Jacas2, Alfonso Susanna2, Arsena Bakhia3, Petar D. Marin1 and Pedja Janaćković1 1 University of Belgrade, Faculty of Biology, Department of Morphology and Systematics of Plants, Studentski trg 16, Belgrade 11000, Serbia 2 Botanic Institute of Barcelona (IBB, CSIC-ICUB), Pg. del Migdia s. n., 08038 Barcelona, Spain 3 Ilia State University - School of Natural Sciences and Engineering, Cholokashvili Avenue 3/5, 0160 Tbilisi, Georgia. ✳ correspondence: [email protected] Keywords: ABSTRACT: dimorphic cypselae, SEM, In this study, micromorphology of an involucral bract and inner and outer dimor- Xerantheminae phic cypselae of the relict Amphoricarpos elegans, endemic from western Caucasus, is examined using a scanning electron microscope (SEM) and presented here for the first time. The middle involucral bract is nearly glabrous, with a reticulate-rugu- lose surface and an oblong-obtusate mucro. Both cypselae exhibit similar features, viz., narrowly oblong-cylindrical or obovate shape with more or less conspicuous longitudinal ribs, an adaxial detachment area, an asymmetrical carpopodium, ab- sence of a pericarp crown and a barbate-aristate pappus made up of narrow, subu- late, basally smooth and apically barbate bristles. However, some differences are evident. The outer cypsela is glabrescent and flattened, with an obscure outline of epidermal cells lacking an end wall and with a sunken periclinal surface and striate- reticulate structure. Also, the outer cypsela has two entire, narrow, lateral wings, rounded above; and less numerous pappus bristles in one row. In contrast to this, the inner cypsela has a distinct outline of the epidermal cells, which are rod-shaped with a short acute end wall, a swollen periclinal surface and reticulate structure. The pappus of the inner cypsela possesses many more bristles organised in two UDC: 581.47:582.99(234.9) rows (a biseriate pappus). Also, slightly pentagonal or round nectary remnants are found in the centre of the upper part of the inner cypsela. The taxonomic value of Received: 26 August 2019 the analysed characters is briefly discussed. Revision accepted: 04 December 2019 INTRODUCTION Amphoricarpos includes heterocarpic perennial plants. The Amphoricarpos taxa are limestone chasmophytic The genus Amphoricarpos Vis. belongs to the subtribe Xer- mountain plants with a disjunct distribution. Two spe- antheminae (Cardueae-Asteraceae) (Herrando-Morai- cies grow on the Balkan Peninsula [A. autariatus Blečić ra et al. 2019). Xerantheminae includes unarmed annual & E. Mayer and A. neumayerianus (Vis.) Greuter], two in (Chardinia Desf., Xeranthemum L. and Siebera J.Gay) and Anatolia (A. exsul O. Schwarz and A. praedictus Ayasligil perennial (Shangwua Yu J. Wang, Raab-Straube, Susanna & Grierson) and one in the Caucasus (A. elegans Albov). & J. Quan Liu and Amphoricarpos) plants with extremely Such a disjunct distribution indicates that Amphoricarpos diverse and complicated geographic distribution (Wang taxa belong to an orophytic flora of Tertiary age and rep- et al. 2013; Herrando-Moraira et al. 2019). The genus resent a connection between the Balkan Peninsula, Asia © 2020 Institute of Botany and Botanical Garden Jevremovac, Belgrade 38 | vol. 44 (1) Minor and the Caucasus (Turril 1929). Taxonomy of the graphs. Thus, A. elegans is almost unexplored from the mi- genus is rather complicated, especially in the case of the cromorphological point of view. Therefore, the aims of our Balkan taxa (Blečić & Mayer 1967; Caković et al. 2015). present study were to: (1) examine micromorphology of Amphoricarpos elegans inhabits limestone rocks and the involucral bract and both (inner and outer) cypselae; stony slopes in the alpine zone (1900 – 2300 m a.s.l.) of and (2) briefly evaluate their taxonomic value. the Caucasus. Compared to the Mediterranean region, the Caucasus is distinctive by virtue of its unique habitats and MATERIAL AND METHODS genetic species diversity, which are a result of its edaphic and climatic conditions, well expressed geographical isola- Plant Material. Plant material (one middle involucral tion and high hypsometric levels (Shetekauri & Kutate- bract and two cypselae) of A. elegans was collected by ladze 2017). The Caucasus region is characterised by high Arsena Bakhia in 2015 during the flowering period from endemism and represents one of the world’s biodiversity plants growing in a natural habitat: on Mt. Migaria, Same- hotspots (Akhalkatsi et al. 2012). Out of 4130 species of grelo, Georgia (N 42.6479800; E 42.63992537). Species the flora of Georgia, 1304 (32.3%) are endemics of the Cau- identification and classification were done according to casus, while 261 (6.6%) are endemics of Georgia (Schatz Webb (1976) and Linczevsky (1962). Voucher specimens et al. 2009). Among calciphytes of the botanical-geograph- were deposited in the herbarium of the Institute of Botany ical province of Cochis, one of the most distinctive species and Botanical Garden “Jevremovac”, Faculty of Biology, is A. elegans (Shetekauri & Kutateladze 2017), which University of Belgrade (accession number: BEOU 17420). belongs to the group of “limestone endemics” of the west- ern Caucasus Mountains (Linczevsky 1962). Micromorphological analysis. The micromorphological Micromorphological characters, especially those of the analysis was carried out using a scanning electron micro- cypsela, have been widely used in detecting taxonomic scope (SEM). Dry involucral bracts and both cypselae (in- and phylogenetic relationships within the family Asterace- ner and outer) were sputter-coated with gold for 180 s at ae (Abid & Qaiser 2009; Gavrilović et al. 2019; Ozcan 30 mA (BAL-TEC SCD 005) and observed using a JEOL & Acinci 2019). Petit (1997) performed a cladistic analy- JSM-6460LV electron microscope at an acceleration volt- sis on 45 Cardueae genera, including Amphoricarpos, and age of 20 kV. The description of cypsela wall sculpturing presented 75 characters, including characters of involucral follows Barthlott (1981), that of trichome morphology bracts and cypselae. Regarding Amphoricarpos, some of his follows Wurdack (1986), the pappus description follows notes were as follows: the keeled portion of true involucral Small (1919) and the detachment area of cypselae is de- bract is narrow in A. elegans, whereas it is wider, often (ir- scribed according to Häffner (2000). regularly) split lengthwise in A. neumayerianus. Moreover, Petit (1997) mentioned that the ancestral type of true RESULTS bracts is certainly a simple tongue. Also, most trichomes on the cypsela surface in A. elegans are twin trichomes, Involucral bract micromorphology but a few trichomes were found to consist of four to five Micrographs of the middle involucral bract are shown in cells (Petit 1997). Additionally, the indicated author sug- Fig. 1. The involucral bract is paleaceous, glabrous or near- gested that multicellular trichomes could be derived from ly so, with vermiform (lanate) trichomes sparsely distrib- twin trichomes. In writing about the taxonomic position uted near the margins and on the tip (Fig. 1a, b). The sur- of A. elegans, Linczevsky (1962) discussed the characters face of the involucral bract is reticulate-rugulose covered used by Albov in describing the section Chodatella Alb., with epicuticular wax (Fig. 1a, b). Sunken stomata are also where he classified A. elegans. Some of the characters were: observed (Fig. 1c). The mucro is oblong-obtusate (Fig. 1b). cypselae of pistillate florets have wings, rounded above and without corniculate projections; pappus scales are narrow, Cypsela micromorphology almost setaceous, numerous (30 – 40); and stems are leafed Amphoricarpos elegans possesses dimorphic cypselae. Mi- almost to the tip. However, Linczevsky (1962) mentioned crographs of the inner cypsela are shown in Fig. 2, while that receptacular scales in A. elegans are not always entire, micrographs of the outer cypsela are shown in Fig. 3. In- and that slightly lacerated scales are very common. ner cypselae, derived from a bisexual floret, are yellow- It was shown that cypsela features are very infor- ish-brownish, sericeous, narrowly oblong-cylindrical or mative at the generic level within the Xerantheminae obovate, with more or less conspicuous longitudinal ribs (Gavrilović et al. 2019). In a previous investigation, we (costae) (Fig. 2a). Outer cypselae, derived from a female examined leaf micromorphology of A. elegans and its re- floret, are yellowish-brownish, glabrescent, flattened, nar- lationship with production of sesquiterpenes lactones and rowly oblong-cylindrical or obovate, with more or less antioxidative activity (Gavrilović et al. 2018a). The mor- conspicuous longitudinal ribs (costae) and with two nar- pho-anatomical study of A. elegans done by Petit (1977) row entire lateral wings, rounded above (Fig. 3a, b). The represents a valuable contribution to knowledge about the detachment area of both cypselae is adaxial (Figs. 2b and micromorphology of this species, but it lacks SEM micro- 3b). The abscission zone is surrounded by a special struc- M. Gavrilović et al.: Micromorphology of Amphoricarpos elegans | 39 the carpopodium (Figs. 2b and 3b). The cells of the cypse- lae surface are parallel
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