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Gasteromycetes of the Genus Scleroderma (Sclerodermataceae) in Russia

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Gasteromycetes of the Genus Scleroderma (Sclerodermataceae) in Russia МИКОЛОГИЯ И ФИТОПАТОЛОГИЯ Том 44 2010 Вып.4 УДК 582.284.52(471) © Ю. А. Ребриев ГАСТЕРОМИЦЕТЫ РОДА SCLERODERMA (SCLERODERMATACEAE) В РОССИИ REBRIEV Yu.A. GASTEROMYCETES OF THE GENUS SCLERODERMA (SCLERODERMATACEAE) IN RUSSIA Благодаря достаточно четким морфологическим признакам род Scleroderma был выделен из рода Lycoperdon Микели (Micheli, 1729) под названием Lycoperdastrum. Позже Персон (Persoon, 1801) легализовал этот род под новым названием Scleroder- ma. В большинстве систем второй половины XX в. род был помещен в семейство Sclerodermataceae порядка Sclerodermatales. Однако полученные в последнее время данные секвенирования рибосомальной ДНК указывают на принадлежность семейст- ва к порядку Boletales (Hibbert et al., 1997; Moncalvo et al., 2003, и др.). Известно не ме- нее 27 видов этого космополитного рода в мире (Sims et al., 1995). Вариабельность наиболее таксономически важных признаков (размеры и орна- ментация оболочек спор, толщина, цвет и характер поверхности перидия) явилась причиной богатой синонимики, а также различного понимания объема видов разны- ми авторами. В монографии по этому роду Гусман (Guzmбn, 1970) говорит о 150 ви- дах и вариациях, но подробно рассматривает только 21 вид. Другим крайним вариан- том явилось объединение девяти известных на то время в СССР видов в качестве воз- растно-экологических форм или синонимов Scleroderma aurantium Pers. (Васильков, 1956). Причинами такого заключения кроме значительной вариабельности морфологи- ческих признаков явились значительные разногласия в разделении видов у различных авторов и недостаточно хорошо обработанный гербарный материал, использовав- шийся Б. П. Васильковым. В указанной работе были рассмотрены также отношения с близкородственным родом Pompholyx (единственный представитель — P. sapidum Corda). Почти одновременно с М. Я. Зеровой (1955) и независимо от нее был сделан вывод о несостоятельности рода. После детального изучения литературных описаний и доступных образцов вид P. sapidum (Scleroderma sapidum (Corda) Zerova) был пере- веден в синонимы S. citrinum как ранняя стадия развития. Нужно заметить, что в ино- странной литературе эта точка зрения была принята только Гусма�ном (Guzmбn, 1970). Сведение Б. П. Васильковым малоизученного Scleroderma fuscum в синонимы S. aurantium не соответствует современным данным.1 В работах Шебека (Љebek, 1953, 1958) не придается большого таксономического значения такому важному признаку, как орнаментация спор. В итоге вид с сетчатыми спорами S. bovista помещен в качестве подвида в S. verrucosum, и наоборот, S. cepa, характеризующийся несетчатыми спорами, понижен до ранга вариации S. citrinum. На каком основании одним видам придан статус подвида, а другим — вариации, оста- лось неясным. 1 В современном понимании S. fuscum принимается как синоним S. bovista (Pegler et al., 1995). 332 П. Е. Сосин (1952) описал новый вид S. poltaviense Sosin (как poltaviensis), вклю- ченный впоследствии в «Определитель гастеромицетов СССР» (Сосин, 1973). Приве- денные признаки четко подходят под описание вида S. cepa, на что обращал внимание и Б. П. Васильков (1956). Однако голотип данного вида, хранящийся в Национальном гербарии Украины (KW) при Институте ботаники им. Н. Г. Холодного НАН Украи- ны, соответствует S. bovista, что согласуется с мнением В. Демулена, изучившего го- лотип в 1975 г. Еще один описанный П. Е. Сосиным вид — S. sapidiforme Sosin (Со- син, Мелик-Хачатрян, 1959). Предположительно типовое описание S. sapidiforme сде- лано по очень молодым плодовым телам. В настоящей работе мы не рассматриваем этот вид. Для разрешения вопроса о его состоятельности необходимо изучение типо- вого образца, местонахождение которого нам неизвестно. Во время работы в Микологическом гербарии БИН РАН (LE) нами обнаружены образцы, отнесенные предыдущими исследователями к ранее не приводимым для России видам: Scleroderma michelii (Lйv.) De Toni, S. reae Guzmбn, S. nitidum Berk. Од- нако эти образцы после детального изучения отнесены нами к довольно обычным ви- дам — Calvatia lilacina (Berk. et Mont.) Henn., Scleroderma verrucosum и S. bovista со- ответственно. Таким образом, по обобщенным данным, в России произрастает 7 ви- дов, включая сомнительные и малоизученные. Ниже приводится описание рода, составленное на основании литературных дан- ных и собственных наблюдений, аннотированный список обнаруженных в России ви- дов рода Scleroderma и идентификационный ключ к этим видам. Материалом для ис- следований послужили образцы, собранные на территории Российской Федерации, Казахстана, Грузии, Украины, Великобритании, Швеции, США и других государств, в том числе 56 образцов из 11 серий эксикат, хранящиеся в Микологических гербари- ях БИН РАН (в конспекте указаны с акронимом LE), ВИЗР (LEP), а также в личной коллекции автора (акроним YuR). Всего изучено около 400 образцов. В обработке коллекций БИН и ВИЗР в разное время участвовали известные российские микологи А. С. Бондарцев, Б. П. Васильков, А. И. Лобик, П. Е. Сосин, В. А. Траншель, А. А. Ячев- ский, а также коллекторы, упомянутые в аннотированном списке. При обработке ма- териалов нами были использованы современные определители и статьи (Demoulin, 1966; Guzmбn, 1970; Pegler et al., 1995; Jeppson, 1998). Микрофотографирование проводилось на базе Южного научного центра РАН (ЮНЦ) с использованием сканирующего электронного микроскопа Carl Zeiss EVO-40 XVP. Распространение видов в пределах России приведено на основании изученного гербарного материала и литературных данных. При этом приняты следующие сокраще- ния географических названий: Адыг. — Республика Адыгея, Белгород. — Белгород- ская обл., Брян. — Брянская обл., Волг. — Волгоградская обл., Воронеж. — Воронеж- ская обл., Иван. — Ивановская обл., Краснодар. — Краснодарский край, Краснояр. — Красноярский край, КЧР — Карачаево-Черкесская Республика, Лен. — Ленинград- ская обл., Липецк. — Липецкая обл., Мар. — Республика Марий Эл, Моск. — Мос- ковская обл., Пенз. — Пензенская обл., Примор. — Приморский край, Ростов. — Рос- товская обл., Ряз. — Рязанская обл., Самар. — Самарская обл., Сарат. — Саратов- ская обл., Смол. — Смоленская обл., Среднерус. — Среднерусская возвышенность, Ставр. — Ставропольский край, Тамбов. — Тамбовская обл., Твер. — Тверская обл., Том. — Томская обл., Тул. — Тульская обл., Удм. — Удмуртская Республика. Род SCLERODERMA Pers., Syn. Meth. Fung.: 150, 1801 Тип рода — Scleroderma verrucosum (Bull.) Pers. Плодовые тела наземные или полуподземные, редко — подземные, округлые, си- дячие или с ложной ножкой, часто с мощным пучком мицелиальных тяжей в основа- нии. Перидий жесткий, четко отделенный от глебы, желтый, желтовато-коричневый до темно-коричневого, чешуйчатый, бородавчатый или почти гладкий, неправильно или звездчато разламывающийся. Глеба мясистая, у очень молодых экземпляров бе- 333 лая, потом темноокрашенная со светлыми жилками трамы, зрелая — порошащая. Споры шаровидные, в незрелом состоянии коричневые, слабоорнаментированные, часто окружены покровом из бесцветных или светлоокрашенных питающих клеток (трофоцист). У многих видов из-за плотно прилегающих трофоцист образуется более или менее четкий сетчатый орнамент эписпория. Зрелые споры темно-коричневые, бородавчатые или игольчатые. Паракапиллиций (у Сосина, 1973 — зачаточный ка- пиллиций) хорошо развит или почти отсутствует, капиллиция нет.2 Представители рода образуют микоризу с широким спектром лиственных и хвойных пород, но спо- собны произрастать и как гумусовые сапротрофы, т. е. являются факультативными микоризообразователями (Шубин, 1988; Moreno, 2005). АННОТИРОВАННЫЙ СПИСОК ВИДОВ РОДА SCLERODERMA, ОТМЕЧЕННЫХ В РОССИИ 1. Scleroderma areolatum Ehrenb., Sylvae Mycol. Berol. 15: 27, 1818. — Scleroderma lycoperdoides Schwein., Schrift. Natьrf. Ges. Leipzig 1: 61, 1822. — Scleroderma verruco- sum Pers. subsp. typicum Љebek var. violascens Herink in Љebek, Sydowia 7: 176—177, 1953. — Scleroderma verrucosum Pers. var. violascens (Herink in Љebek) in Љebek, Flora ИSR. B-1: 570, 1958. — Склеродерма пятнистая. Распространение в России: Волг., И. В. Землянская (YuR 899, 917); Моск., Ю. А. Ребриев (YuR 1854); Пенз. (Иванов, 1993); Ростов., Ю. А. Ребриев (YuR 800, 838, 843, 867, 903-a, 914, 918, 924, LE 217348, 217349, 217350, 217351); Ряз., Л. Ф. Во- лоснова (YuR 1245); Краснодар., Б. П. Васильков (LE 226486); Примор. (Васильева, Сосин, 1959); Ставр., Ефимова (YuR 811), КЧР, Поспелова (LE 2608). Общее распространение: Африка, Европа, Северная Америка, Австралия (Sims et al., 1999). 2. Scleroderma bovista Fr., Syst. Mycol. 3: 48, 1829. — Scleroderma verrucosum (Va- ill.) Pers. subsp. bovista (Fr.) Љebek, Sydowia 7: 177—179, 1953. — Scleroderma verruco- sum Pers. var. bovista (Fr.) Љebek, Flora ИSR. B-1: 570—572, fig. 204: 5—6, 1958. — Phlyctospora fusca Corda in Sturm, Deutschl. Fl. 2: 51, tab. 16, fig. 1—5, 1841. — Sclero- derma fuscum (Corda) E. Fischer in Engler-Prantl, Die Natьrl. Pflanzenfam. 1/1: 336, fig. 175 D, 1900 et 1. c. 7a: 36, fig. 26 D, 1933. — Scleroderma texense Berkeley, J. Bot., 4: 308, 1845. — Scleroderma poltaviense Sosin [ut ‘poltaviensis’] in Not. syst. Sect. crypt. Inst. bot. Acad. Sci. U. S. S. R. 8: 126, 1952. — Склеродерма-порховка. Распространение в России:Белгород. (Беденко, 1984); Волг., И. В. Землян- ская (LE 217352); Лен., Н. Наумов (LEP), Ю. А. Ребриев (YuR 875); Пенз., Е. К. Шту- кенберг (LEP); Ростов., Е. Горохова (YuR 857), И. А. Матвеев (YuR 874), Ю. А. Реб- риев (YuR 799, 817, 819, 821, 822, 823, 834, 840, 858, 879, 888, 910, 922, LE 819, 217353, 217354, 217357, 217358, 217360), С. П. Чеботарева (YuR 884); Ряз., Л. Ф. Во- лоснова (YuR
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