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О Филогении Ранних Cardioceratidae (Ammonoidea) И Среднерусских Представителях Cadoceratinae На Рубеже Бата И Келловея © 2016 Г

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О Филогении Ранних Cardioceratidae (Ammonoidea) И Среднерусских Представителях Cadoceratinae На Рубеже Бата И Келловея © 2016 Г ПАЛЕОНТОЛОГИЧЕСКИЙ ЖУРНАЛ, 2016, № 4, с. 42–51 УДК 56(116.2):564.53(470.1/.6) О ФИЛОГЕНИИ РАННИХ CARDIOCERATIDAE (AMMONOIDEA) И СРЕДНЕРУССКИХ ПРЕДСТАВИТЕЛЯХ CADOCERATINAE НА РУБЕЖЕ БАТА И КЕЛЛОВЕЯ © 2016 г. В. В. Митта Палеонтологический институт им. А.А. Борисяка РАН e%mail: [email protected] Поступила в редакцию 26.05.2015 г. Принята к печати 08.07.2015 г. На основе изучения морфогенеза раковины и скульптуры и анализа стратиграфического распро странения реконструирован филогенез среднеюрских Cardioceratidae. К подсемейству Arctocephal itinae (верхний байос – самые низы келловея) отнесена филолиния Cranocephalites → Arctocepha lites → Arcticoceras → Paracadoceras. Подсемейство Cadoceratinae (верхний байос – келловей), также ведущее начало от Cranocephalites, включает филолинию Greencephalites → Cadoceras → Longae viceras, и боковые ветви Chamoussetia и Platychamoussetia. Происхождение раннекелловейских Eck hardites, также отнесенных к Cardioceratidae, не установлено. Родовые названия Rondiceras и Cado chamoussetia предлагается считать младшими синонимами Cadoceras и Chamoussetia, соответствен но. Приведено описание раннекелловейского Cadoceras bellabimba sp. nov. DOI: 10.7868/S0031031X16040097 ВВЕДЕНИЕ ся, главным образом, укрупнением раковины в Древнейшие среднеюрские представители се ходе филогенеза. Отметим, что для последних мейства Cardioceratidae Siemiradzki обычно под (средний бат) представителей Arcticoceras харак разделяются (Callomon, 1985) на подсемейства терно уменьшение размеров раковины, заметное Arctocephalitinae Meledina (верхи байоса – низы укорочение жилой камеры и наличие выражен келловея) и Cadoceratinae Hyatt (верхний байос – ных ребер на жилой камере. Параллельно этой келловей), являясь важнейшей группой аммоно филолинии развивался род Greencephalites Repin, идей для биостратиграфии этого интервала геоло возникший в позднем байосе бореальных райо гического времени, особенно для бореальных и нов одновременно с Arctocephalites, также в ре суббореальных районов. В то же время, несмотря зультате дивергенции рода Cranocephalites Spath. на наличие большого числа опубликованных ра Для Greencephalites характерны кадиконические бот, система ранних кардиоцератид не устоялась, и раковины с низким сечением оборотов и сравни принадлежность отдельных родов к тому или ино тельно более широким пупком; рельефная и даже му подсемейству до настоящего времени не уста грубая скульптура, в том числе на длинной жилой новлена. В статье на основе анализа морфогенеза камере. Последние представители гринцефали раковины и скульптуры, а также стратиграфиче тов известны также из среднего бата (Митта, Аль ской последовательности таксонов реконструиро сен, 2013). ван филогенез ранних кардиоцератид, кратко оха Практически одновременно с исчезновением рактеризованный ранее (Митта, 2015а). Детально арктоцефалитов и гринцефалитов на рубеже рассмотрены систематический состав и филогене среднего и позднего бата в Арктике (Callomon, тические взаимоотношения среднерусских пред 1993) появляются роды Cadoceras Fischer и Para ставителей подсемейства Cadoceratinae. cadoceras Crickmay, в последующем расширившие свои ареалы до суббореальных и даже перитети ческих районов. Первые представители рода Ca ФИЛОГЕНИЯ РАННИХ CARDIOCERATIDAE doceras (C. calyx Spath, C. apertum Callomon et Bir Преимущественно бореальное подсемейство kelund) имеют кадиконовую раковину с широким Arctocephalitinae представлено родами Arctocepha низким сечением и рельефную на фрагмоконе lites Spath и Arcticoceras Spath, связанными отноше скульптуру, сглаживающуюся на длинной жилой ниями “предок–потомок”, соответственно. Оба камере. Род Cadoceras является типовым для под рода характеризуются узкопупковыми раковинами семейства Cadoceratinae. К этому подсемейству субовального сечения, со сходной скульптурой, относится обычно и род Paracadoceras, характери сглаживающейся на жилой камере, и различают зующийся относительно меньшими размерами 42 О ФИЛОГЕНИИ РАННИХ CARDIOCERATIDAE (AMMONOIDEA) 43 раковины и жилой камеры, более высоким сече представлена последовательными видами R. sokolovi нием оборотов, обычно менее грубой скульпту (Kiselev) → R. geerzense (Behrendsen) → R. tcheff рой, часто выраженной и на относительно корот kini (d’Orbigny) → R. milaschevici (Nikitin) → кой жилой камере (P. efimovi Mitta, P. nageli Mitta, → R. stenolobum (von Keyserling emend. Nikitin) P. keuppi Mitta). Перечисленные представители (Митта, 2000). Дериватом последнего вида, со Cadoceras и Paracadoceras известны из верхнего гласно Д.Н. Киселеву (2005), является Longae бата Русской платформы (Mitta, 2005). viceras alpha Kiselev из верхов среднего келловея, раковины которого показывают отчетливую тен Эволюция рода Cadoceras продолжается и в денцию к повышению высоты оборотов и суже фазу elatmae нижнего келловея – C. frearsi (d’Or нию пупка. Видимо, от “L.” alpha и происходит bigny) → C. falsum Voronetz → C. elatmae (Nikitin) → следующая (позднекелловейская) итерация кадо → C. tschernyschevi Sokolov. Все эти виды имеют цератин, завершающаяся узкопупковыми оксико крупную раковину с более или менее низким се новыми раковинами: “Cadoceras” allae Kiselev → чением оборотов и умеренно широким пупком, → “C.” patruum (Eichwald) → “Chamoussetia” длинную (до одного оборота) жилую камеру, хо funifera (Phillips). Последний вид обозначен как рошо выраженную скульптуру. Но уже у вида тип рода Funiferites Kiselev et al., 2003, в свою оче C. tschernyschevi на жилой камере ребра редуци редь, являющегося, повидимому, младшим си руются до изогнутых умбиликальных бугорков. 1 Следующее звено этой филолинии – C. stu нонимом рода Platychamoussetia Repin, 2002 . pachenkoi Mitta – характеризуется, в дополнение Остальные позднекелловейские кардиоцера к дальнейшей редукции вентральных ребер на тиды (Quenstedtoceras Hyatt и близкородственные жилой камере, увеличением высоты сечения и за роды), вероятно, лучше относить к подсемейству метным сужением пупка на жилой камере так, что Cardioceratinae Siemiradzki или использовать для она полностью перекрывает предшествующий них название Quenstedtoceratinae Meledina. При оборот фрагмокона. Это начало нового тренда, конвенционном использовании последнего вари выражающегося в дальнейшем сужении пупка, анта собственно кардиоцератины появляются с постепенном превращении кадиконовой ракови начала оксфорда, с рода Vertumniceras Buckman. ны в пахиконовую и субоксиконовую (род Cado chamoussetia Mitta: C. surensis (Nikitin) → C. subpatr Стратиграфическая последовательность пере uus (Nikitin)), и далее в оксиконовую, с редуциро численных выше таксонов родового ранга ранних ванной скульптурой, сохраняющейся в виде кардиоцератид в обобщенном виде показана на вентральных “зубчиков” [род Chamoussetia R. Dou рис. 1. Рассмотрим представленную там же схему villé: C. stuckenbergi (Lahusen), C. buckmani Cal филогенетических взаимоотношений. lomon et Wright]. Постепенно сокращается и жи Стратиграфическая и филогенетическая пре лая камера. На последнем виде, имеющем жилую емственность Cranocephalites → Arctocephalites → камеру не более 0.6 оборота, эта ветвь заканчива → Arcticoceras после работ Дж. Калломона (Cal ется (Mitta, 1999). lomon, 1985, 1993) не вызывает сомнений. По Аналогичная короткая ветвь кардиоцератид с следние представители Arcticoceras – A. crassipli субоксиконовой раковиной, с жилой камерой, catum Callomon (in litt.) и A. cranocephaloide Cal занимающей до 0.6 оборота, и скульптурой, на lomon et Birkelund демонстрируют отчетливую последних оборотах представленной лишь в вен тенденцию к уменьшению размеров раковины, тролатеральной части [род Eckhardites Mitta: расширению пупка и сохранению скульптуры на E. menzeli (Mönnig) → E. pavlowi (Smorodina) → жилой камере – т.е., являются переходными к ро → E. dietli Mitta], появляется и раньше, в низах ду Paracadoceras. Сравнение микроконхов аркти нижнего келловея (Митта, 2009). Некоторые ис коцерасов (см. Rawson, 1982) и паракадоцерасов следователи относят виды этого рода (в ранге (рис. 2), имеющих крупные для микроконхов ра подрода) к Macrocephalites (Гуляев, 2015а). ковины с толстыми оборотами и относительно редкими и грубыми ребрами, также показывает В верхах нижнего и в среднем келловее субборе принадлежность этих двух родов к одной филоли альных и бореальных районов широко распростра нии. Соответственно, род Paracadoceras следует нены кадоцератины, относимые отечественными рассматривать в составе подсемейства Arctoceph исследователями к роду Rondiceras Troizkaya. Рако alitinae. Более детально эта филолиния, с привле вины этого рода представлены крупными кадико чением печорского и гренландского материала, нами и сфероконами с длинной жилой камерой. будет рассмотрена в отдельной статье. Видовые различия проявляются преимуществен но в вариациях сечения оборотов и пупковой во С обособлением рода Greencephalites (с типо ронки, степени рельефности ребер на внутренних вым видом Cadoceras freboldi Spath) (Репин и др., оборотах. Основным трендом развития этой груп 1 Этот род как предположительно новый был указан в: Мит пы видов является постепенное сглаживание ре та, 2000, с. 59, в объеме видов “Ammonites” patruus Eichwald бер, начиная с молодых оборотов. Эта филолиния и “Ammonites” funiferus Phillips. ПАЛЕОНТОЛОГИЧЕСКИЙ ЖУРНАЛ № 4 2016 44 МИТТА ia et ss ou Cardioceratidae m Ярус a ch ty Подъярус a Pl Qu en ste dto ce ras Верхний Platychamoussetia Longaeviceras tia Cardioceratinae se Cadoceras s ou m ha C es Средний t Келловей di ar kh ? Ec Cadoceras Chamoussetia Нижний Eckhardites Бат Cadoceratinae Arctocephalitinae Greencephalites Нижний Средний Верхний
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