diff --git a/data/03/F8/48/03F84846FF959D08FF1FF892FA27FCC1.xml b/data/03/F8/48/03F84846FF959D08FF1FF892FA27FCC1.xml new file mode 100644 index 00000000000..08cfe72df6e --- /dev/null +++ b/data/03/F8/48/03F84846FF959D08FF1FF892FA27FCC1.xml @@ -0,0 +1,715 @@ + + + +Molecular phylogeny, morphology and distribution of Amphitetras antediluviana Ehrenberg from the Black Sea and the Sea of Azov + + + +Author + +Nevrova, Elena +0000-0001-9963-4967 +A. O. Kovalevsky Institute of Biology of the Southern Seas RAS, IBSS RAS, 2 Nakhimov Ave., Sevastopol, Crimea, 299011, Russia +el_nevrova@mail.ru + + + +Author + +Maltsev, Yevhen +0000-0003-4710-319X +К. А. Timiryazev Institute of Plant Physiology RAS, IPP RAS, 35 Botanicheskaya St., Moscow, 127276, Russia +ye.maltsev@gmail.com + + + +Author + +Kezlya, Elena +0000-0002-5263-9338 +К. А. Timiryazev Institute of Plant Physiology RAS, IPP RAS, 35 Botanicheskaya St., Moscow, 127276, Russia +melosira@mail.ru + + + +Author + +Mironov, Andrei +К. А. Timiryazev Institute of Plant Physiology RAS, IPP RAS, 35 Botanicheskaya St., Moscow, 127276, Russia & Faculty of Biology, M. V. Lomonosov Moscow State University, 1 / 12 Leninskie Gory, Moscow, 119234, Russia + + + +Author + +Kulikovskiy, Maxim +0000-0003-0999-9669 +К. А. Timiryazev Institute of Plant Physiology RAS, IPP RAS, 35 Botanicheskaya St., Moscow, 127276, Russia +max-kulikovsky@yandex.ru + +text + + +Phytotaxa + + +2024 + +2024-11-19 + + +672 + + +1 + + +49 +63 + + + + +https://doi.org/10.11646/phytotaxa.672.1.3 + +journal article +10.11646/phytotaxa.672.1.3 +1179-3163 +14520563 + + + + + +Amphitetras antediluviana +Ehrenberg (1840: 62) + + + + + +Homotypic synonyms: + +Triceratium antediluvianum +(Ehrenberg) +Grunow (1867: 24) + +; + +Biddulphia antediluviana +(Ehrenberg) +Van Heurck (1885: 207) + +; + +Odontella antediluviana +(Ehrenberg) +Peragallo (1903: 686) + +. + + + + +Morphological description: +Living cells of + +A. antediluviana + +form chain-shaped, tangled colonies of dark brown color ( +Fig. 2 +). Each cell with numerous small chloroplasts of discoid shape ( +Fig. 3 +). The valve is quadrangular with broadly rounded apices and slightly concave margins ( +Figs 4–7 +). The Black Sea specimens differs from +type +material by a more smoother frustules shape; the outgrowths with ocelli protrudes less clearly. Frustules are heavily silicified and have significant wall thickness (2.3–2.5 µm on average). Numerous girdle bands of valve have small elevations on corners and slightly depressions on the sides of mantle. In girdle view, frustule length significantly (by 2–3 times) exceeds its width ( +Figs 8–10 +). Valve diameter in the Black Sea population on the Southern coast of +Crimea +(Laspi Bay and Cape Fiolent) varies from 23.4–43.6 μm, height 70.0–82.1 μm in girdle view. Parameters of specimens from the Eastern coast of +Crimea +(Feodosiya Bay) vary from 20.3 to 35,1 μm in valve diameter, and from 62 to 65 μm in valve height. Off the coast of the Azov Sea, cell dimensions were 26,2–28,5 μm in valve diameter, 68–70 μm in valve height ( +Figs 7, 10 +). + + + +FIGURES 2–3. + +Amphitetras antediluviana +, LM. Strain + +BL-5_12092019 (Laspi Bay, Black Sea). 2. Cell macrocolony. 3. Alive cell, girdle view. Scale bars = 100 μm (Fig. 2), 10 μm (Fig. 3). + + + +Each valve is equipped with four ocelli located at the corners ( +Figs 11–12 +, +17 +, +23, 26 +, +29 +, black arrows), with a diameter of 8–9.5 μm. Ocelli are riddled with small, densely arranged ( +50–55 in +10 μm) poroids ( +Figs 21 +, +26 +). The diameter of poroids is around 0.17 μm. In SEM, it is visible that poroids are arranged in more or less distinguishable rows, divided by thin hyaline strips ( +Fig. 32 +, black arrowheads). The polar region of each valve is moderately depressed, surrounded by a circular elevation, which is resolvable in angled valve view ( +Fig. 29 +, white arrowhead). + + +The central part of valve is constructed by thick vimines separating the areolae of +two types +. Areolae of the first +type +( +Figs 15 +, +20 +, +33–34 +, white arrows) are more common, round to angular, 1.2–2.9 μm in diameter, arranged by +4–5 in +10 μm. They are covered by cribrate occlusions. The latter are formed by simple poroids (0.08–0.15 μm in diameter) separated by delicate crossbars. Externally, openings are round to hooked. In close view, areolae of the first +type +possess large spathulate formations with C-shaped slits at the perimeter ( +Fig. 27 +, white arrowheads). In addition, there are numerous particles of silica situated at the crossbars separating the poroids ( +Fig. 27 +, black arrowheads). Internally, cribrate occlusions are supported by several (5–12) relatively wide props ( +Figs 16 +, +22 +, +34 +, white arrows). Second-type areolae are smaller (less than 1 μm in diameter) and less common, equipped with volate occlusions ( +Figs 15 +, +20 +, +33–34 +, black arrows). In this case, outgrowths of occlusions are spathulate, moderately branched, resulting in slits of irregular shape ( +Fig. 28 +, black arrows). Notably, vimines throughout the valve are equipped with angular, mostly tetrahedral particles of biosilica ( +Figs 15 +, +20 +, +26 +, +32, 33 +, white arrowheads) with a diameter of 0.5–0.8 μm. + + +The girdle is connected to the epitheca and hypotheca by one valvocopula on each side ( +Fig. 25 +, white arrowheads). There are large, round-squarish areolae at the girdle, arranged in rows by +4–6 in +10 μm ( +Figs 13 +, +19 +, +25 +, +29 +, white arrows). + + + +FIGURES 4–10. + +Amphitetras antediluviana +, LM. + +4–6, 8–9. Strain BL-5_12092019 (Laspi Bay, Black Sea). 7, 10. Strain АZ-12_24122019 (Sea of Azov). 4–7. Valve view. 8–10. Girdle view. Scale bars = 10 μm. + + + +In this study, the reconstruction of phylogenetic relationships was carried out using the ML and BI methods. Consistently, with previous study on + +Amphitetras + +phylogeny ( + +Ashworth +et al. +2013 + +), the analysis identified united clade for + +Odontella rostrata +(Hustedt) +Simonsen (1987: 250) + +, + +Amphipentas pentacrinus +Ehrenberg (1841: 205) + +, and + +Amphitetras antediluviana + +( +Fig. 1 +). The ML and BI phylogenetic analysis of V4 region of the 18S rRNA gene and the plastid +rbc +L gene associates novel + +Amphitetras antediluviana + +strains (BL-5_ +12092019 +and AZ-12_ +24122019 +) with strain + +Amphitetras antediluviana +ETC + +3627 with maximum statistical support (BS 100, PP 1.0; +Fig. 1 +). It is worth mentioning that during the construction of the phylogram, we analyzed two strains from GenBank library with the same index as the strain from the Sea of Azov investigated herein: “Azo-12 Amphi-A11 OP297425 OP304753” and “Azo-12 Amphi-B4 OP297426 OP304754”. Strain “Azo-12 Amphi-A11…” was originally identified as + +Amphipentas pentacrinus + +, while strain “Azo-12 Amphi-B4…” belongs to + +Amphitetras antediluviana + +according to the GenBank. This discrepancy can indicate that the mentioned strains were misidentified. However, taking into account the detached position of “Azo-12 Amphi-B4…” at the presented tree, + +A. antediluviana + +may be understood as a cryptic taxon from now on. + + +Type +locality of the studied species is +Isle Tjörn +, situated at the +Southwestern +coast of +Sweden +( +Jahn & Kusber 2006 +). +Around +the +World Ocean +, this species has been noted by different authors: as + +Amphitetras antediluviana + +from +Europe +: +Greece +( +Jahn & Kusber 2006 +), +Albania +( +Miho & Witkowski 2005 +); +North America +, +Mexico +( +López-Fuerte & Siqueiros-Beltrones 2016 +); +South America +, +Argentina +( + +Garibotti +et al. +2011 + +) and +Uruguay +( +Metzeltin & García-Rodríguez 2003 +). As + +Triceratium antediluvianum + +the species was recorded from +Canary Islands +( + +Gil-Rodríguez +et al. +2003 + +; + +Moro +et al. +2011 + +), Adriatic Sea ( + +Viličić +et al. +2002 + +), Britain ( + +Hartley +et al. +1986 + +; +Sims 1996 +), +France +( + +Méléder +et al. +2007 + +), +North Carolina +( +Hustedt 1955 +), +Colombia +( + +Lozano-Duque +et al +. 2010 + +), +Gambia +( +Foged 1986 +), +Egypt +( +Zalat 2002 +), +China +( +Liu 2008 +). At last, it was mentioned as + +Biddulphia antediluviana + +in the samples from Britain ( +Hendey 1954 +, +1974 +), +Ireland +( +Adams 1908 +), +Portugal +( +Moita & Vilarinho 1999 +), +Romania +( +Cărăus 2012 +), +Brazil +( + +Eskinazi-Leça +et al. +2010 + +), +Ghana +( + +Smith +et al. +2015 + +) and +New Zealand +( + +Harper +et al. +2012 + +). However, the species is not mentioned in the sinopsis of marine benthic diatoms ( + +Witkowski +et al. +2000 + +). + + + +FIGURES 11–16. + +Amphitetras antediluviana +, SEM. Strain + +BF-36_18122019 (Feodosiya Bay, Black Sea). 11–12. External valve view. Black arrows point to ocelli. 13. External girdle view. White arrows point to rows of poroids at the girdle. 14. Internal valve view. 15. Details of areolae structure, externally. Black arrow points to volate occlusion, white arrow points to cribrate occlusion. 16. Details of areolae structure, internally. White arrows point to supporting props. Scale bars = 10 μm (Figs 11–12, 14), 15 μm (Fig. 13), 1 μm (Figs 15–16). + + + + +FIGURES 17–22. + +Amphitetras antediluviana +, SEM. Strain + +CF-4_23072020 (Cape Fiolent, Black Sea). 17. External valve view. Black arrows point to ocelli. 18. Internal valve view. 19. External girdle view. White arrows point to rows of poroids at the girdle. 20. Details of areolae structure, externally. Black arrow points to volate occlusion, white arrow points to cribrate occlusion, white arrowhead points to siliceous particle. 21. Details of ocellus structure, externally. 22. Details of ocellus structure, internally. White arrows point to supporting props. Scale bars = 5 μm (Figs 17–18), 10 μm (Fig. 19), 1 μm (Fig. 20), 2 μm (Figs 21–22). + + + + +FIGURES 23–28. + +Amphitetras antediluviana +, SEM. Strain + +AZ-12_24122019 (Sea of Azov). 23. External valve view. Black arrows point to ocelli. 24. Internal valve view. 25. External girdle view. White arrows point to rows of poroids at the girdle, white arrowheads point to valvocopulae. 26. Details of ocellus structure, externally. White arrowheads point to siliceous particles. 27. Details of cribrate areolae structure, externally. Black arrowheads point to siliceous particles at the crossbars of occlusions, white arrowheads point to C-shaped slits. 28. Details of volate areolae structure, externally. Black arrows point to spathulate formations of volae. Scale bars = 5 μm (Fig. 23), 10 μm (Figs 24–25), 2 μm (Fig. 26), 0.5 μm (Figs 27–28). + + + + +FIGURES 29–34. + +Amphitetras antediluviana +, SEM. Strain + +BL-5_12092019 (Laspi Bay, Black Sea). 29. External valve view. Black arrows point to ocelli, white arrows point to the rows of poroids at the girdle, white arrowhead points to polar depression. 30–32. Internal valve view. 32. Details of ocellus structure, externally. Black arrowheads point to pores of ocelli. 33. Details of polar area structure, externally. Black arrow points to volate areolae, white arrow points to cribrate areolae, white arrowheads point to siliceous particles. 34. Details of polar area structure, externally. Black arrows point to volate areolae, white arrows point to cribrate areolae. Scale bars = 20 μm (Fig. 29), 10 μm (Fig. 30), 5 μm (Fig. 31), 2 μm (Fig. 32), 4 μm (Figs 33–34). + + + +In the Black Sea, + +A. antediluviana + +was rarely recorded, found as single specimens: in the Northwestern part of the Black Sea on silty-sandy soil off the coast of +Romania +( +Bodeanu 1987 +); in +Ukraine +, particularly in macrophyte algae fouling and on silty-sandy soil in the Tendrovsky and Jarylgach Bays, in Tuzlov and Shabolatsky estuaries ( + +Guslyakov +et al. +1992 + +); in +Russia +: in the area of the Zernov’s Phyllophora field ( +Nevrova 2014a +), in the Sevastopol Bay ( +Kucherova 1973 +), in the bays of the southern coast of Crimea and the Caucasus ( +Proshkina-Lavrenko 1963 +; +Nevrova 2014b +, +2015 +, +2016 +; +Nevrova & Petrov 2019 +; +Nevrova & Revkov 2003 +). In all the listed sources, the species is mentioned as + +Triceratium antediluvianum + +. According to the results of our recent research ( +Nevrova 2022 +), in the last decade, the finds of + +Amphitetras antediluviana + +have become frequent, its macrocolonies in the form of chains have been found in many habitats off the Western, Southern and Eastern Black Sea coasts of Crimea, off the Caucasian coast, as well as off the Sea of Azov coast (see +Table 2 +). + + + +TABLE 2. +Records of + +Amphitetras antediluviana + +in the Black Sea and the Sea of Azov. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
+Sampling Location + +Date of sampling + +Type of substrate + +Depth, m + +Cell condition +
+Black Sea +
Zernov’s Phyllophorae field 45°50’38”N, 30°44’32”E2010silty36, 48scarce empty cells
Coast near Marjino village 45°22’67”N, 33°12’40”E2010sand6scarce empty cells
Omega Bay 44°35’55”N, 33°26’56”E2004sand3, 14scarce empty cells
Golubaya Bay 44°33’54”N, 33°23’57”E2010sand6scarce empty cells
Cape Fiolent2009sand3scarce empty cells
44°30’53”N, 33°28’18”E2020sand4alive colonies
Cape Ajya 44°28’25”N, 33°37’58”E2006sand12scarce empty cells
Laspi Bay1996silty-sandy5, 31, 52scarce empty cells
44°25’10”N, 33°42’27”E2019sand5alive colonies
Karadag Natural Reserve 44°54’53”N,35°13’51”E2010silty-sandy8scarce empty cells
Lisya Bay 44°53’45”N, 35°09’30”E2008silty-sandy5scarce empty cells
Dvuyakornaya Bay 44°59’42”N, E 35°22’34”E2022silty-sandy5alive colonies
Feodosiya Bay 45°04’49”N, 35°34’11”E2019silty36alive colonies
Caucasian coast, Sukhaya Shchel’ 44°40’45”N, 37°31’44”E1999coarse sand, shell debris50scarce empty cells
Caucasian coast, Cape Bol’shoj Utrish 44°45’33”N, 37°22’2”E1999silty-sandy75scarce empty cells
+Sea of Azov +
Semisotskoye village, Leninskiy district 45°41’8”N, 35°00’22”E2019silty-sandy12alive colonies
+ + +It can be concluded that despite significant morphometric (see +Table 3 +) and morphological differences, clones of + +Amphitetras antediluviana + +from the Black and Azov Seas belong to the same species. Multiple detections of + +Amphitetras antediluviana + +colonies at various depths in the coastal habitats of the Black Sea and the Sea of Azov in the last decade indicate a change in environmental factors.As we suggest, the reason for that can be the recent climatic fluctuation and confounding of optimal factors for the survival and development of this species in the marine sublittoral ecosystems. + + + + \ No newline at end of file