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<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:ali="http://www.niso.org/schemas/ali/1.0/" article-type="research-article" dtd-version="1.2" xml:lang="en"><front><journal-meta><journal-id journal-id-type="publisher-id">Paleontological Journal</journal-id><journal-title-group><journal-title xml:lang="en">Paleontological Journal</journal-title><trans-title-group xml:lang="ru"><trans-title>Палеонтологический журнал</trans-title></trans-title-group></journal-title-group><issn publication-format="print">0031-031X</issn><issn publication-format="electronic">3034-5871</issn><publisher><publisher-name xml:lang="en">The Russian Academy of Sciences</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="publisher-id">677965</article-id><article-id pub-id-type="doi">10.31857/S0031031X24060053</article-id><article-id pub-id-type="edn">QILTVE</article-id><article-categories><subj-group subj-group-type="toc-heading" xml:lang="en"><subject>Articles</subject></subj-group><subj-group subj-group-type="toc-heading" xml:lang="ru"><subject>Статьи</subject></subj-group><subj-group subj-group-type="article-type"><subject>Research Article</subject></subj-group></article-categories><title-group><article-title xml:lang="en">Effect of posterior gut reduction on the evolution of rhynchonelliform brachiopods</article-title><trans-title-group xml:lang="ru"><trans-title>Влияние редукции заднего отдела кишечника на эволюцию ринхонеллиформных брахиопод</trans-title></trans-title-group></title-group><contrib-group><contrib contrib-type="author"><name-alternatives><name xml:lang="en"><surname>Selischeva</surname><given-names>А. А.</given-names></name><name xml:lang="ru"><surname>Селищева</surname><given-names>А. А.</given-names></name></name-alternatives><address><country country="RU">Russian Federation</country></address><email>selav21@mail.ru</email><xref ref-type="aff" rid="aff1"/></contrib></contrib-group><aff-alternatives id="aff1"><aff><institution xml:lang="en">Lomonosov Moscow State University</institution></aff><aff><institution xml:lang="ru">Московский государственный университет имени М.В. Ломоносова</institution></aff></aff-alternatives><pub-date date-type="pub" iso-8601-date="2024-12-19" publication-format="electronic"><day>19</day><month>12</month><year>2024</year></pub-date><issue>6</issue><fpage>37</fpage><lpage>55</lpage><history><date date-type="received" iso-8601-date="2025-04-01"><day>01</day><month>04</month><year>2025</year></date></history><permissions><copyright-statement xml:lang="en">Copyright ©; 2024, Russian Academy of Sciences</copyright-statement><copyright-statement xml:lang="ru">Copyright ©; 2024, Российская академия наук</copyright-statement><copyright-year>2024</copyright-year><copyright-holder xml:lang="en">Russian Academy of Sciences</copyright-holder><copyright-holder xml:lang="ru">Российская академия наук</copyright-holder></permissions><self-uri xlink:href="https://transsyst.ru/0031-031X/article/view/677965">https://transsyst.ru/0031-031X/article/view/677965</self-uri><abstract xml:lang="en"><p>Brachiopods are a group of animals known since the Early Cambrian and thrived in the Paleozoic. After the Permian–Triassic extinction, there was a significant reduction in the taxonomic diversity of brachiopods. According to one hypothesis, in the Mesozoic, brachiopods with blind gut could not reinstate their numbers due to the predominance of shelly plankton. We assume that the terebratulids, the most widespread group of modern brachiopods, were able to adapt to the changed composition of food due to a more efficient filtration mechanism of the plectolophous lophophore. Extant rhynchonellids have a blind–closed bulbus end of digestive tract, which is probably used for crushing and digesting shelly plankton.</p></abstract><trans-abstract xml:lang="ru"><p>Брахиоподы – группа животных, известная с раннего кембрия и достигшая расцвета в палеозое. После пермско-триасового вымирания произошло значительное сокращение таксономического разнообразия брахиопод. По одной из гипотез, в мезозое брахиоподы со слепо замкнутым кишечником не смогли восстановить свою численность из-за преобладания в планктоне панцирных одноклеточных водорослей. Мы предполагаем, что теребратулиды, которые являются самой распространенной группой среди современных брахиопод, смогли приспособиться к изменившемуся составу питания за счет более эффективного механизма фильтрации, характерного для плектолофного лофофора. Современные ринхонеллиды на конце кишечника имеют слепое расширение, которое, по-видимому, используется для измельчения и переваривания панцирного планктона.</p></trans-abstract><kwd-group xml:lang="en"><kwd>brachiopods</kwd><kwd>lophophore</kwd><kwd>digestive tract</kwd><kwd>evolution</kwd><kwd>phytoplankton</kwd><kwd>extinction</kwd></kwd-group><kwd-group xml:lang="ru"><kwd>брахиоподы</kwd><kwd>лофофор</kwd><kwd>кишечник</kwd><kwd>эволюция</kwd><kwd>фитопланктон</kwd><kwd>вымирание</kwd></kwd-group><funding-group><award-group><funding-source><institution-wrap><institution xml:lang="ru">Российский научный фонд</institution></institution-wrap><institution-wrap><institution xml:lang="en">Russian Science Foundation</institution></institution-wrap></funding-source><award-id>23-14-00020</award-id></award-group></funding-group></article-meta></front><body></body><back><ref-list><ref id="B1"><label>1.</label><mixed-citation>Афанасьева Г.А., Невесская Л.А. Анализ причин различных последствий кризисных ситуаций на примере замковых брахиопод и бивальвий // Экосистемные перестройки и эволюция биосферы. Вып. 1. М.: Недра, 1994. С. 101–108.</mixed-citation></ref><ref id="B2"><label>2.</label><mixed-citation>Кузьмина Т.В., Георгиев А.А. Особенности питания брахиоподы Hemithiris psittacea (Rhynchonelliformea: Rhynchonellida) // Сб. материалов всеросс. науч. конф. с междунар. участием, посвященной 85-летию Беломорской биостанции им. Н.А. Перцова Биол. фак-та МГУ им. М.В. Ломоносова. М.: Тов-во науч. изданий КМК, 2023. С. 106–107.</mixed-citation></ref><ref id="B3"><label>3.</label><mixed-citation>Невесская Л.А. Этапы развития бентоса фанерозойских морей. Палеозой. М.: Наука, 1998. 503 с.</mixed-citation></ref><ref id="B4"><label>4.</label><mixed-citation>Пунин М.Ю. Исследование организации эпителия пищеварительного тракта замковой брахиоподы Hemithyris psittacea. II. Электронно–микроскопический анализ // Цитология. 1981. Т. 23. № 10. С. 1109–1115.</mixed-citation></ref><ref id="B5"><label>5.</label><mixed-citation>Пунин М.Ю. Гистологическая организация кишечных эпителиев приапулид, брахиопод, двустворчатых моллюсков и полихет. СПб.: Наука, 1991. 248 с.</mixed-citation></ref><ref id="B6"><label>6.</label><mixed-citation>Пунин М.Ю., Филатов М.В. Организация железы замковой брахиоподы Hemithyris psittacea // Цитология. 1980. Т. 22. № 3. С. 277–286.</mixed-citation></ref><ref id="B7"><label>7.</label><mixed-citation>Atkins D. A new species and genus of Kraussinidae (Brachiopoda) with a note on feeding // Proc. Zool. Soc. London. 1958. V. 131. № 4. P. 559–581.</mixed-citation></ref><ref id="B8"><label>8.</label><mixed-citation>Atkins D., Rudwick M.J.S. The lophophore and ciliary feeding mechanisms of the brachiopod Crania anomala (Müller) // J. Mar. Biol. Assoc. U.K. 1962. V. 42. № 3. P. 469–480.</mixed-citation></ref><ref id="B9"><label>9.</label><mixed-citation>Bramlette M.N. Mass extinctions of Mesozoic biota // Science. 1965. V. 150. № 3701. P. 1240–1240.</mixed-citation></ref><ref id="B10"><label>10.</label><mixed-citation>Carlson S.J. The evolution of Brachiopoda // Ann. Rev. Earth and Planetary Sci. 2016. V. 44. P. 409–438.</mixed-citation></ref><ref id="B11"><label>11.</label><mixed-citation>Chuang S.H. The ciliary feeding mechanisms of Lingula unguis (L.) (Brachiopoda) // Proc. Zool. Soc. London. 1956. V. 127. № 2. P. 167–189.</mixed-citation></ref><ref id="B12"><label>12.</label><mixed-citation>Chuang S.H. The structure and function of the alimentary canal in Lingula unguis (L.) (Brachiopoda) // Proc. Zool. Soc. London. 1959. V. 132. № 2. P. 283–311.</mixed-citation></ref><ref id="B13"><label>13.</label><mixed-citation>Chuang S.H. An anatomical, histological, and histochemical study of the gut of the brachiopod, Crania anomala // J. of Cell Science. 1960. V. 3. № 53. P. 9–18.</mixed-citation></ref><ref id="B14"><label>14.</label><mixed-citation>Clarkson E.N.K. Invertebrate Paleontology and Evolution. L.: George Allen &amp; Unwin, 1979. 323 p.</mixed-citation></ref><ref id="B15"><label>15.</label><mixed-citation>Curry G.B., Brunton C.H.C. Stratigraphic distribution of brachiopods // Treatise on Invertebrate Paleontology. Part H. Brachiopoda (revised). Vol. 6: Suppl. / Ed. Kaesler R.L. Boulder, Lawrence: Geol. Soc. America; Univ. Kansas Press, 2007. P. 2901–2965.</mixed-citation></ref><ref id="B16"><label>16.</label><mixed-citation>Dhar S.R., Logan A., Macdonald B.A., Ward J.A. Endoscopic investigations of feeding structures and mechanisms in two plectolophous brachiopods // Invertebr. Biol. 1997. V. 116. № 2. P. 142–150.</mixed-citation></ref><ref id="B17"><label>17.</label><mixed-citation>D’Hondt J.L. Etude de l’intestin et de la glande digestive de Terebratulina retusa (L.)(Brachiopode). IV. Comparaison avec les activités enzymatiques d’autres brachiopodes du même biotope // Les Brachiopodes Fossiles et Actuels / Eds. Racheboeuf P.R., Emig C. Brest, 1986. P. 301–306. (Actes du 1er Congrès intern. sur les Brachiopodes, Biostratigr. Paléozoïque. V. 4).</mixed-citation></ref><ref id="B18"><label>18.</label><mixed-citation>D’Hondt J.L., Boucaud–Camou E. Étude l’intestin et de la glande digestive de la Terebratulina retusa (L.) (Brachiopode). Ultrastructure et recherche d’activités amylasiques et protéasiques // Bull. Soc. Zool. France. 1982. V. 107. № 2. P. 207–216.</mixed-citation></ref><ref id="B19"><label>19.</label><mixed-citation>Elyakova L.A. Distribution of cellulases and chitinases in marine invertebrates // Compar. Biochemistry and Physiology. Pt B: Compar. Biochemistry. 1972. V. 43. № 1. P. 67–70.</mixed-citation></ref><ref id="B20"><label>20.</label><mixed-citation>Raven J., Falkowski P.G., Katz M.E. et al. The evolution of modern eukaryotic phytoplankton // Science. 2004. V. 305. № 5682. P. 354–360.</mixed-citation></ref><ref id="B21"><label>21.</label><mixed-citation>Gilmour T.H.J. Ciliation and function of the food-collecting and waste-rejecting organs of lophophorates // Canad. J. Zool. 1978. V. 56. № 10. P. 2142–2155.</mixed-citation></ref><ref id="B22"><label>22.</label><mixed-citation>Gilmour T.H.J. Food-collecting and waste-rejecting mechanisms in Glottidia pyramidata and the persistence of lingulacean inarticulate brachiopods in the fossil record // Canad. J. Zool. 1981. V. 59. № 8. P. 1539–1547.</mixed-citation></ref><ref id="B23"><label>23.</label><mixed-citation>Gould S.J., Calloway C.B. Clams and brachiopods – ships that pass in the night // Paleobiology. 1980. V. 6. № 4. P. 383–396.</mixed-citation></ref><ref id="B24"><label>24.</label><mixed-citation>Guo Z., Flannery-Sutherland J.T., Benton M.J. et al. Bayesian analyses indicate bivalves did not drive the downfall of brachiopods following the Permian–Triassic mass extinction // Nature Commun. 2023. V. 14. № 1. P. 5566.</mixed-citation></ref><ref id="B25"><label>25.</label><mixed-citation>Hammen C.S. Brachiopod metabolism and enzymes // Amer. Zool. 1977. V. 17. № 1. P. 141–147.</mixed-citation></ref><ref id="B26"><label>26.</label><mixed-citation>Hancock A. XXXIV. On the organization of the Brachiopoda // Phil. Trans. Roy. Soc. London. 1858. V. 148. P. 791–869.</mixed-citation></ref><ref id="B27"><label>27.</label><mixed-citation>Harper D.A.T., Popov L.E., Holmer L.E. Brachiopods: origin and early history // Palaeontology. 2017. V. 60. № 5. P. 609–631.</mixed-citation></ref><ref id="B28"><label>28.</label><mixed-citation>James M.A., Ansell A.D., Collins M.J. et al. Biology of living brachiopods // Adv. in Mar. Biol. 1992. V. 28. P. 175–387.</mixed-citation></ref><ref id="B29"><label>29.</label><mixed-citation>Kuzmina T.V., Malakhov V.V. Structure of the brachiopod lophophore // Paleontol. J. 2007. V. 41. № 5. P. 520–536.</mixed-citation></ref><ref id="B30"><label>30.</label><mixed-citation>Kuzmina T.V., Ratnovskaya A.A., Madison A.A. Lophophore evolution from the Cambrian to the present // Paleontol. J. 2021. V. 55. № 10. P. 1109–1140.</mixed-citation></ref><ref id="B31"><label>31.</label><mixed-citation>Kuzmina T.V., Temereva E.N. Rejection mechanism of plectolophous lophophore of brachiopod Coptothyris grayi (Terebratulida, Rhynchonelliformea) // Moscow Univ. Biol. Sci. Bull. 2018. V. 73. P. 136–141.</mixed-citation></ref><ref id="B32"><label>32.</label><mixed-citation>Kuzmina T.V., Temereva E.N. Structure of the oral tentacles of early ontogeny stage in brachiopod Hemithiris psittacea (Rhynchonelliformea, Rhynchonellida) // J. Morphol. 2024. V. 285. № 4. P. e21686.</mixed-citation></ref><ref id="B33"><label>33.</label><mixed-citation>LaBarbera M. Water flow patterns in and around three species of articulate brachiopods // J. Experim. Mar. Biol. Ecol. 1981. V. 55. № 2–3. P. 185–206.</mixed-citation></ref><ref id="B34"><label>34.</label><mixed-citation>Lee D.E. The terebratulides: the supreme brachiopod survivors // Brachiopoda: Fossil and Recent / Eds. Harper D.A.T., Long S.L., Nielsen C. Wiley, 2008. P. 241–249 (Fossils and Strata. V. 54).</mixed-citation></ref><ref id="B35"><label>35.</label><mixed-citation>Liow L.H., Reitan T., Harnik P.G. Ecological interactions on macroevolutionary time scales: clams and brachiopods are more than ships that pass in the night // Ecology Letters. 2015. V. 18. № 10. P. 1030–1039.</mixed-citation></ref><ref id="B36"><label>36.</label><mixed-citation>McCammon H.M. The food of articulate brachiopods // J. Paleontol. 1969. V. 43. P. 976–985.</mixed-citation></ref><ref id="B37"><label>37.</label><mixed-citation>McCammon H.M. Physiology of the brachiopod digestive system // Ser. in Geol., Notes for Short Course. 1981. V. 5. P. 170–189.</mixed-citation></ref><ref id="B38"><label>38.</label><mixed-citation>Morton J.E. The functions of the gut in ciliary feeders // Biol. Rev. 1960. V. 35. № 1. P. 92–139.</mixed-citation></ref><ref id="B39"><label>39.</label><mixed-citation>Nielsen C. The development of the brachiopod Crania (Neocrania) anomala (OF Müller) and its phylogenetic significance // Acta Zool. 1991. V. 72. № 1. P. 7–28.</mixed-citation></ref><ref id="B40"><label>40.</label><mixed-citation>Popov L.Ye., Holmer L.E. Trimerellida and Chileata // Treatise on Invertebrate Paleontology. Part H, Brachiopoda (Revised). Vol. 2: Linguliformea, Craniiformea, and Rhynchonelliformea (part) / Ed. Kaesler R.L. Boulder, Lawrence: Geol. Soc. America; Univ. Kansas Press, 2000a. P. 184–200.</mixed-citation></ref><ref id="B41"><label>41.</label><mixed-citation>Popov L.Ye., Holmer L.E. Class Obolellata // Treatise on Invertebrate Paleontology. Part H, Brachiopoda (Revised). Vol. 2: Linguliformea, Craniiformea, and Rhynchonelliformea (part) / Ed. Kaesler R.L. Boulder, Lawrence: Geol. Soc. America; Univ. Kansas Press, 2000b. P. 200–207.</mixed-citation></ref><ref id="B42"><label>42.</label><mixed-citation>Popov L.Ye., Williams A. Kutorginata // Treatise on Invertebrate Paleontology. Part H, Brachiopoda (Revised). Vol. 2: Linguliformea, Craniiformea, and Rhynchonelliformea (part) / Ed. Kaesler R.L. Boulder, Lawrence: Geol. Soc. America; Univ. Kansas Press, 2000. P. 208–215.</mixed-citation></ref><ref id="B43"><label>43.</label><mixed-citation>Reed C.G., Cloney R.A. Brachiopod tentacles: ultrastructure and functional significance of the connective tissue and myoepithelial cells in Terebratalia // Cell and Tissue Res. 1977. V. 185. P. 17–42.</mixed-citation></ref><ref id="B44"><label>44.</label><mixed-citation>Rowell A.J., Caruso N.E. The evolutionary significance of Nisusia sulcata, an early articulate brachiopod // J. Paleontol. 1985. V. 59. P. 1227–1242.</mixed-citation></ref><ref id="B45"><label>45.</label><mixed-citation>Rudwick M.J.S. Filter–feeding mechanisms in some brachiopods from New Zealand // Zool. J. Linn. Soc. 1962. V. 44. № 300. P. 592–615.</mixed-citation></ref><ref id="B46"><label>46.</label><mixed-citation>Rudwick M.J.S. Living and Fossil Brachiopods. L.: Hutchinson &amp; Co., 1970. 199 p.</mixed-citation></ref><ref id="B47"><label>47.</label><mixed-citation>Shi G.R., Shen S. Asian–western Pacific Permian Brachiopoda in space and time: biogeography and extinction patterns // Devel. in Palaeontol. and Stratigr. 2000. V. 18. P. 327–352.</mixed-citation></ref><ref id="B48"><label>48.</label><mixed-citation>Shu-Zhong S., Shi G.R. Paleobiogeographical extinction patterns of Permian brachiopods in the Asian–western Pacific region // Paleobiology. 2002. V. 28. № 4. P. 449–463.</mixed-citation></ref><ref id="B49"><label>49.</label><mixed-citation>Steele-Petrović H.M. The physiological differences between articulate brachiopods and filter-feeding bivalves as a factor in the evolution of marine level-bottom communities // Palaeontology. 1979. V. 22. Pt 1. P. 101–134.</mixed-citation></ref><ref id="B50"><label>50.</label><mixed-citation>Storch V., Welsch U. Elektronenmikroskopische und enzymhistochemische Untersuchungen über die Mitteldarmdrüse von Lingula unguis L. (Brachiopoda) // Zool. Jb., Abt. für Anat. und Ontogenie der Tiere. 1975. V. 94. S. 441–452.</mixed-citation></ref><ref id="B51"><label>51.</label><mixed-citation>Strathmann R. Function of lateral cilia in suspension feeding of lophophorates (Brachiopoda, Phoronida, Ectoprocta) // Mar. Biol. 1973. V. 23. № 2. P. 129–136.</mixed-citation></ref><ref id="B52"><label>52.</label><mixed-citation>Tappan H. Primary production, isotopes, extinctions and the atmosphere // Palaeogeogr., Palaeoclimatol., Palaeoecol. 1968. V. 4. № 3. P. 187–210.</mixed-citation></ref><ref id="B53"><label>53.</label><mixed-citation>Tappan H., Loeblich Jr A.R. Evolution of the oceanic plankton // Earth-Sci. Rev. 1973. V. 9. № 3. P. 207–240.</mixed-citation></ref><ref id="B54"><label>54.</label><mixed-citation>Temereva E.N., Kuzmina T.V. The first data on the innervation of the lophophore in the rhynchonelliform brachiopod Hemithiris psittacea: what is the ground pattern of the lophophore in lophophorates? // BMC Evol. Biol. 2017. V. 17. P. 1–19.</mixed-citation></ref><ref id="B55"><label>55.</label><mixed-citation>Thayer C.W. Are brachiopods better than bivalves? Mechanisms of turbidity tolerance and their interaction with feeding in articulates // Paleobiology. 1986. V. 12. № 2. P. 161–174.</mixed-citation></ref><ref id="B56"><label>56.</label><mixed-citation>Vargas C. de, Audic S., Henry N. et al. Ocean plankton. Eukaryotic plankton diversity in the sunlit ocean // Science. 2015. V. 348. № 6237. P. 1261605.</mixed-citation></ref><ref id="B57"><label>57.</label><mixed-citation>Westbroek P., Yanagida J., Isa Y. Functional morphology of brachiopod and coral skeletal structures supporting ciliated epithelia // Paleobiology. 1980. V. 6. № 3. P. 313–330.</mixed-citation></ref><ref id="B58"><label>58.</label><mixed-citation>Williams A., Carlson S.J. Affinities of brachiopods and trends in their evolution // Treatise on Invertebrate Paleontology. Part H, Brachiopoda (Revised). Vol. 6: Supplement. Lawrence: Univ. Kansas Press, 2007. P. 2822–2900.</mixed-citation></ref><ref id="B59"><label>59.</label><mixed-citation>Williams A. Carlson S.J., Brunton C.H.C. et al. A supra–ordinal classification of the Brachiopoda // Phil. Trans. Roy. Soc. London. Ser. B: Biol. Sci. 1996. V. 351. № 1344. P. 1171–1193.</mixed-citation></ref><ref id="B60"><label>60.</label><mixed-citation>Williams A., James M.A., Emig C.C. et al. Anatomy // Treatise on Invertebrate Paleontology. Part H, Brachiopoda (Revised). Vol. 1: Introduction. Lawrence: Univ. Kansas Press, 1997. P. 7–151.</mixed-citation></ref><ref id="B61"><label>61.</label><mixed-citation>Zezina O.N. Biogeography of the recent brachiopods // Paleontol. J. 2008. V. 42. № 8. P. 830–858.</mixed-citation></ref><ref id="B62"><label>62.</label><mixed-citation>Zhang Z., Han J., Zhang X. et al. Soft-tissue preservation in the Lower Cambrian linguloid brachiopod from South China // Acta Palaeontol. Pol. 2004. V. 49. № 2. P. 259–266.</mixed-citation></ref><ref id="B63"><label>63.</label><mixed-citation>Zhang Z., Han J., Zhang X. et al. Note on the gut preserved in the Lower Cambrian Lingulellotreta (Lingulata, Brachiopoda) from southern China // Acta Zool. 2007a. V. 88. № 1. P. 65–70.</mixed-citation></ref><ref id="B64"><label>64.</label><mixed-citation>Zhang Z., Holmer L.E., Skovsted C.B. et al. A sclerite-bearing stem group entoproct from the early Cambrian and its implications // Sci. Rep. 2013. V. 3. № 1. P. 1066.</mixed-citation></ref><ref id="B65"><label>65.</label><mixed-citation>Zhang Z.F., Li G.X., Holmer L.E. et al. An early Cambrian agglutinated tubular lophophorate with brachiopod characters // Sci. Rep. 2014. V. 4. № 1. P. 4682.</mixed-citation></ref><ref id="B66"><label>66.</label><mixed-citation>Zhang Z., Shu D., Han J. et al. A gregarious lingulid brachiopod Longtancunella chengjiangensis from the Lower Cambrian, South China // Lethaia. 2007b. V. 40. № 1. P. 11–18.</mixed-citation></ref><ref id="B67"><label>67.</label><mixed-citation>Zhang Z., Shu D., Emig C. et al. Rhynchonelliformean brachiopods with soft‐tissue preservation from the early Cambrian Chengjiang Lagerstätte of South China // Palaeontology. 2007c. V. 50. Pt 6. P. 1391–1402.</mixed-citation></ref></ref-list></back></article>
