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Pericarp Structure in Some Species in the Tribe Sileneae DC

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Pericarp Structure in Some Species in the Tribe Sileneae DC Botanica Pacifica. A journal of plant science and conservation. 2019. 8(2): 25–34 DOI: 10.17581/bp.2019.08211 Pericarp structure in some species in the tribe Sileneae DC. (Caryophyllaceae, Viscaria group) Tatyana I. Kravtsova1* & Yana V. Bolotova2 Tatyana I. Kravtsova1* ABSTRACT e-mail: [email protected] The pericarp structure was studied in the representatives of genera Atocion, Min­ Yana V. Bolotova2 jaevia, and Viscaria, forming a separate Viscaria clade on the phylogenetic tree of e-mail: [email protected] the tribe Sileneae, and in Ixoca, previously a member of this group. It was found that, despite of differences in the capsule morphology and dehiscence in A. ar­ meria and M. rupestris on the one hand, and V. vulgaris on the other, there are many 1 Komarov Botanical Institute RAS, Saint similar features in their pericarp anatomy and micromorphology that supports Petersburg, Russia their close relationships. The highest similarity in the pericarp structure, as well as 2 Amur Branch of the Botanical Garden- in the capsule morphology, is between A. armeria and M. rupestris. Pericarp struc- Institute FEB RAS, Blagoveshchensk, ture in V. vulgaris is less derived compared to them and differs by the absence of Russia silicified palisade layer in the exocarp and indistinct differentiation of pericarp sclerenchyma into different (morphologically and in orientation) cell layers. Peri- carp anatomy confirms the separation of genusIxoca from the Viscaria group. Keywords: Caryophyllaceae, Sileneae, Atocion, Ixoca, Minjaevia, Viscaria, fruit, molecular phylogeny, pericarp micromorphology and anatomy, taxonomy * corresponding author РЕЗЮМЕ Manuscript received: 11.10.2018 Кравцова Т.И., Болотова Я.В. Строение перикарпия у некоторых Review completed: 10.10.2019 ви дов трибы Sileneae DC. (Caryophyllaceae, Viscaria group). Строение Accepted for publication: 29.10.2019 пери карпия изучено у представителей родов Atocion, Minjaevia и Viscaria, со­ Published online: 02.11.2019 ставляющих на молекулярно­филогенетическом дереве трибы Sileneae кладу Viscaria, а также у Ixoca, ранее входившего в эту кладу. Выяснено, что, не- смотря на различия в морфологии и способе вскрывания коробочек у A. ar­ meria и M. rupestris с одной стороны, и V. vulgaris с другой, в анатомическом строении и микроморфологии их перикарпия имеется много общих черт, что подтверждает близкое родство этих таксонов. Наибольшее сходство в стро ении перикарпия, как и в морфологии коробочек, обнаружено между A. armeria и M. rupestris. Менее продвинутая по сравнению с ними структура перикарпия у V. vulgaris отличается отсутствием палисадного окремневшего экзокарпия и неотчетливой дифференциацией склеренхимы на слои, раз- личающиеся по морфологии и ориентации клеток. Анатомия перикарпия подтверждает обособленность рода Ixoca от родов группы Viscaria. Ключевые слова: Caryophyllaceae, Sileneae, Atocion, Ixoca, Minjaevia, Viscaria, микро­ мор фология и анатомия перикарпия, молекулярная филогения, плод, таксономия The systematics of the tribe Sileneae (Caryophyllaceae) is Vis caria group, the Lychnis group, and the terminal one, the cur rently insufficiently developed; the taxonomic struc ture largest Silene group, including two subclades. The Viscaria and the phylogenetic relationships of the genus Silene L., the group initially included four mor pho logically distinct small largest in the Caryophyllaceae, numbering about 700 species, ge ne ra: Atocion Adans., Ixoca Raf. (=Heliosperma (Rchb.) are debatable issues. It is considered some times to be very Rchb.), Minjaevia Tzvel. and Viscaria Bernh., the close rela- wide and includes almost all of other ge ne ra of the tribe, tion ships of which is very doubtful in the opinion of Tzvelev except Agrostemma L. (Greuter 1995), then divides into large (2001). The close relationships between Atocion and Viscaria num bers of small genera, the lar gest number in Tzvelev is one of the unexpected re sults obtained by Oxelman & (2001) classification. The taxonomic re vi sions of the tribe Li dén (1995), which was con firmed by all subsequent mo­ al so vary significantly in the opinion of authors from our leu c lar studies (Fraiman, Heidari & Oxelman 2009). In the country (Tzvelev 2001, Lazkov 2003, 2006). Molecular phy- previous classifications of the tribe, on the base of tra di­ loe g netic studies of the Sileneae with dif ferent molecular mar- tio nal methods, the Atocion was related to Silene s.str., and kers of nuclear and chloroplast ori gin (Oxelman & Lidén Vis caria to Lychnis (for example, Chowdhuri 1957). The 1995, Oxelman et al. 1997), allowed elucidation of genera con tradictions with the traditional systematics of the tribe re la tion ships in the tribe. These investigations revealed that Sileneae are associated with the possible reticular character of se pa rate clades on the mo le cular phylogenetic trees are as their evolution (Erixon & Oxelman 2008, Fraiman, Heidari follows: basal clades Agrostemma, Eudianthe, Petrocoptis, the & Oxelman 2009, Mikhailova 2016b). ©Botanical Garden-Institute FEB RAS. 2019 25 Kravtsova & Bolotova According to recent data (Fraiman & Oxelman 2007, in Ixoca arcana (Heliosperma alpestre) at the site of cracking Frai man, Eggens & Oxelman 2009), the position of Ixoca cells of the exocarp are smaller (lower) than neighboring as a sister group, in relation to Viscaria vulgaris, is erroneous; cells, in Viscaria vulgaris, the cells of subepidermal layer are its real position within Sileneae remains unclear. A study of smaller. This character, however, varies in the second spe- this gen us with the involvement of low-copy nuclear genes cies. In a number of works, there are data on exocarp mic­ in di cates its hyb rid origin. At least two lineages have been ro mor pho logy for several species (Melikian & Devyatov in vol ved in the origin of Ixoca: an ancestor of Viscaria and 2000, Kravtsova & Romanova 2015, Kravtsova 2017): it Ato cion and the other of Eudianthe Rchb. and/or Petrocoptis was found to be colliculate in Atocion, Minjaevia, Viscaria, A. Braun ex Endl. (Fraiman, Eggens & Oxelman 2009). Re­ and a different type in Ixoca. cent results (Mikhailova 2016а, b) have shown that gen us The aim of this study was to characterize the peri- Ixo ca forms a se parate basal clade on the molecular phy lo- carp, its micromorphology and anatomical structure, in genetic tree. spe cies combined in the smolkova clade (Viscaria group) The data of molecular phylogenetic analysis resulted in and to consider the importance of these characteristics for the revision of the tribe Sileneae classification, in which 8 systematics of the group. A comparative analysis also in- ge nera were recognized (Oxelman et al. 2001). Monotypic cludes two species of the genus Ixoca, previously attributed ge nus Minjaevia (Minjaevia rupestris (L.) Tzvel.) was included in to this group. Atocion, as Atocion rupestre (L.) Oxelman (Oxelman et al. 2001, Frai man, Heidari & Oxelman 2009, Fraiman et al. 2013). MATERIAL AND METHODS The close relationships between Atocion and Viscaria The study is based on mature fruits (capsules) of five can be considered as an example of glaring contradictions spe cies (15 specimens) from Herbaria LE and KW, car po- bet w een morphology and molecular phylogeny, which are logical collection (Lec) of Botanical Museum of Ko ma- not uncommon in modern botany. There are significant rov Botanical Institute, and those collected in nature. The va riations within the Viscaria group in several carpological accepted taxonomy of the group corresponds to the clas si­ cha rac ters used in Sileneae taxonomy to distinguish genera: fication of Tzvelev (2001). The following specimens were car pel and capsule teeth number; septa presence or ab sence; investigated: septicidal or loculicidal dehiscence. Capsules of A. armeria Atocion armeria (L.) Raf.: Park of Komarov Botanial In- and M. rupestris have some characteristic features of Silene sti tute , 4 IX 2004, N.N. Tzvelev 26 (LE); Gomelskaya Pro­ s.str.: three styles, a loculicidal-septicidal capsule with in- vince, Dobrush station, 25 VII 1926, Ju. Krechetovich 24 com plete septa, dehiscent via 6 teeth. These genera are in- (LE); St.­Petersburg, lawn in S. Kowalewskaya street, IX clu ded sometimes into Silene s.l., in the sections Compactae 2014, G.Ju. Konechnaya; Far East, Blagoveshchensk, Amur Branch of the Botanical Garden-Institute FEB RAS, IX (Boiss.) Schischk. and Nanosilene Otth (or Rupifraga Otth), 2016, Ya.V. Bolotova. res pecti vely (Schishkin 1936, Chowdhuri 1957). Ixoca arcana (Zapał.) Ikonn.: West Ukraine, Stanislavskaya and capsules have another structure. In Viscaria Ixoca (Ivanovo-Frankowskaya) oblast, 21 VII 1940, A.I. Pojarkova V. vul garis, there are five styles, capsules are five­locular with et al. (LE); Botanical Museum of Komarov Botanical in comp lete septa, with unique loculicidal dehiscence via five Institute, LEc № 9833. teeth. In Ixoca species, there are three styles, capsules are Ixoca carpatica (Zapał.) Ikonn.: Herb. Mus. Univ. Leopol­ one-locular, without septa, loculicidal-septicidal, dehiscent di nensis N 104699, Carpathians, Charnohora, VIII. 1927, via six teeth. J. Mą dalski (LE); Ivanovo­Frankowskaya oblast, 28 VI The anatomical structure of fruits in the species, consti tu- 1963, I.V. Artemchuk (LE); ting the Viscaria group, as well as in the whole tribe Sileneae, is Minjaevia rupestris (L.) Tzvelev. Karelia, the bank of Lake insufficiently studied. The pericarp structure was in vestigated Onega, cape Cherest, 26 VII 2013, G.Ju. Konechnaya
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