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On the Morphology and Evolution of the Cucullograptinae (Monograptidae, Graptolithina)

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On the Morphology and Evolution of the Cucullograptinae (Monograptidae, Graptolithina) ACT A PAL A E 0 N '..TO . LOG I CAP 0 LON IC : A Vol. XI 1 9 6 6 ADAM URBANEK ON THE MORPHOLOGY AND EVOLUTION OF THE CUCULLOGRAPTINAE (MONOGRAPTIDAE, GRAPTOLITHINA) Contents Page Abstract 292 Introduction 292 Acknowledgements 294 G e n era 1 Par t 294 Origin of the material 294 Investigation methods 302 Terminology 304 Stratigraphic position of the fauna inv.estigated 306 Comparative anatomical analysis of the Cucullograptinae and the rules of their evolution 315 Evolution of siculae 315 Evolution of blastozooid thecae 326 Evolution 'of astogeny 343 Phylogenetic relationships and taxonomy of the Cucullograptinae 359 Phylogeny of the Cucullograptinae 359 Taxonomy of the Cucullograptinae . 374 Generic subdivision · of the Cucullograptinae 374 Origin of the Cucullograptinae 378 S y s t e mat i cPa r t 384 Order Graptoloidea Lapworth, 1875 384 Suborder Monograptina Lapworth, 1880 384 Family Monograptidae Lapworth, 1873 . 384 Subfamily Cucullograptinae Urbanek, 1958 384 Genus. Lobograptus Urbanek, 1958 384 Lobograptus progenitor n. sp. 384 Lobograptus simplex Urbanek, 1960 396 Lobograptus exspectatus exspectatus Urbanek, 1960 406 Lobograptus exspectatus bicomis n. subsp. 417 Lobograptus scanicus parascanicus (KUhne, 1955) 420 Lobograptus scanicus amphirostris n. subsp. 436 Lobograptus scanicus scanicus (Tullberg, 1883) sensu Bulman, 1953 444 Lobograptus imitator n. sp. 452 Lobograptus invertus n. sp, 458 LobograptuscirTifer n. sp. 468 Genus Cucullograptus Urbanek, 1954' . 480 Cucullograptus hemiaversus Urbanek, 1960 . 480 Cucullograptus aversus d . aversus (Eisenack, 1942) .492 Cucullograptus aversus rostratus Urbanek, 1960 495 Cucullograptus pazdroi Urbanek, 1954 519 292 ADAM URBANEK Abstract. - A description is given of the structure, astogenetic development and evolution of the Upper Silurian Cucullograptinae. It is based on the material contained in the core samples from the Mielnik on the Bug (Easter n Poland) deep-boring and from the Baltic drift. The graptolites were prepared by dissol­ ving the limestone and marly matrix in hydrochloric acid. For detailed study the specimens were bleached. The stratigraphic position and significance of the fauna investigated are discussed. The main trend in the evolution of sicula consists in its simplification and is interpreted as due to foetalization. In contrast with sicula, the evolution of thecae displays progressive changes which primarily consist in the development of apertural apparatus, connected, in most lines, with the elaboration of its secondary asymmetry. The biological significance of this pattern of evolution of thecal characters. is discussed. The Cucullograptinae sup­ ply convincing evidence for the distal introduction of phylogenetic novelties which makes up a prevailing mode of evolutionary changes of astogeny in this group of moriograptids. Problems, related to the mechanisms of evolution and orga­ nization of graptolite colonies are discussed. Using the data, now available, ten­ tative phylogenetic relationships within the group are established. An attempt is made to show some implications of these data in the problem of origin of the Cucullograptinae. Fourteen representatives (species and subspecies) of the Cu­ cullograptinae are described in the systematic part. Four species and two sub­ species are new. INTRODUCTION A detailed study of the structure and astogenetic development of the representatives of the subfamily Cucullograptinae Urbanek, 1958 (Mo­ nograptidae, Graptolithina), as well as an explanation of their evolution and stratigraphic importance constitute the aim of the present work. Despite the fact that the representatives of this subfamily have re­ cently been subject to frequent studies (Eisenack, 1942; Bulman, 1953; Walker, 1953; Urbanek, 1954, 1958, 1960; Kuhne, 1955) not all of the problems, related to this interesting group of the Upper Silurian mo­ nograptids have exhaustively been explained. In 1960, the accumulation of a new material,coming, in part, from the Baltic erratic boulders but chiefly that, prepared from the Mielnik on the Bug boring core which, to a considerable extent, supplemented the family tree of the Cucullograptinae and threw a new light on se­ veral related problems, has induced the present writer to investigate this group once more and in a yet more exhaustive manner. The idea of the present monograph may be summarized in the follo­ wing four items: 1) a detailed investigation of the structure of the the­ cae and the development of the colony, carried out on the basis of adequately bleached microscopic preparations with particular considera­ tion of the fusellar structure; 2) a comparison of the structure of dif­ ferent representatives with the application of the comparative-anatomi- MORPHOLOGY AND EVOLUTICN OF CUCULLCGRAPTINAE 293 cal methods; 3) a determination of the main evolutionary trends and of the ch aracter of morphological changes which took place in particu­ lar phyletic lineages; 4) an Investigation of stratigraphic ranges so as to be able to determine the significance of the Cucullograptinae as index fossils. The fact that the remains of the Cucullograptinae might be etched from the rock by chemical methods was an especially important factor in the studies on their evolution. Thus isolated specimens allowed one to study several anatomical characters, otherwise inaccessible to ob­ servation by standard methods of investigation usually applied to the graptolite remains preserved on the rock surface. To obtain transparent preparations, enabling one to examine the fusellar structure of thecae and, in particular, the fusellar structure of the apertural apparatus, the present author bleached the isolated specimens. A detailed analysis of the fusellar structure is one of the main methods, applied to the present work. In solving all these problems, the present writer aimed, in a consis­ tent way, at the integration of all data, based on the studies of the forms from the drift material, with those obtained from the boring at Miel­ nik on the Bug (Eastern Poland). The latter data constituted the main basis for concluding on the evolution and stratigraphic occurrence. The material, coming from the Baltic boulders served .m ostly for the in­ vestigation of the morphology and astogeny of particular species. The results of the studies on the following representatives of the Cucullograptinae are discussed in the present work: Genus Lobograptus Urbanek, 1958 Lobograptus progenitor n. sp. Lobograptus simplex Urbanek, 1960 Lobograptus exspectatus exspectatus Urbanek, 1960 Lobograptus exspectatus btcornis n. subsp, Lobograptus scanicus parascanicus (Kuhne, 1955) Lobograptus scanicus amphirostris n. subsp. Lobograptus scanicus scanicus (Tullberg, 1883) sensu Bulman, 1953 Lobograptus imitator n. Sp.1 Lobograptus invertus n. Sp.1 Lobograptus cirrifer n. Sp.1 Genus Cucullograptus Urbanek, 1954 Cucullograptus hemiaversus Urbanek, 1960, Cucullograptus aversus d . aversus (Eisenack, i942) Cucullograptus aversus rostratus Urbanek, 1960 Cucullograptus pazdroi Urbanek, 1954 I Thus marked forms are known on ly fr om ' the boring at Mielnik on the Bug. 294 ADAM URBANEK ACKNOWLEDGEMENTS In preparing the present monograph, I availed myself of much good advice, indications and stimulating suggestions, offered - during many discussions - by Professor Roman Kozlowski (Warszawa, Poland) to whom I would like to express my warmest gratitude. My thanks are due to Professor E. Ruhle, Director of the Geological Institute in Warsaw, for his consent to supply valuable materials from the Mielnik 1. G. 1 bore-hole for the scientific elaboration and to Dr. H. Tomczyk ,fl'Om this Institute, as well as to his wife, Dr. E. Tomczyk for a friendly cooperation and aid afforded during sampling and a sub­ sequent elaboration of the material. My gratitude is also extended to Professor Z. Kielan-Jaworowska, Director of the Palaeozoological Institute of the Polish Academy of Scien­ ces {Warsaw), for a friendly help and a kind assent to the technical assistance given to the present author by her scientific-technical staff. I feel particularly indebted to Mrs. K. Budzynska and to Mrs. E. Ga­ domska for a careful inking of my pencil drawings, to Mrs. J. Skarzynska and Mr. W. Skarzynski for a technical aid in the preparation of the ma­ terial, as well as to Miss M. Czarnocka for taking photographs. A warm gratitude is also due to all graptolitologists who, visiting Warsaw, looked through a part or a whole of the material prepared and, discussing it with the writer, offered their advice and suggestions. First of all, Professor O. M. B. Bulman (Cambridge, England), Dr. R. Thorsteinsson (Ottawa, Canada), Professor H. Whittington (Cambridge, U.S.A.), as well as Dr. S. Willefert (Rabat, Morocco), Dr. H. Jaeger (Berlin, the German Democratic Republic), Dr. B. Rickards (Cambridge, England) and Professor A. M. abut (Novosibirsk, U.S.S.R.) are r mong them. During his foreign journeys, involved in the preparation of this work, the present writer cooperated with and used a /friendly aid of Dr. F . Bro­ tzen (Stockholm, Sweden), Professor G. Regnell (Lund, Sweden), Dr. V. Zazvorka (Prague, Czechoslovakia), Dr. R. Horny (Prague, Czechoslovakia), Professor B. Boueek (Prague, Czechoslovakia) and Dr. J . Paskevieius (Vilnius, U.S.S.R.). The present author's gratitude is due to all of them. GENERAL PART ORIGIN OF THE MATERIAL The material of the Cucullograptinae, studied in the present mono­ graph, comes from: 1) the core of a deep boring at Mielnik on the Bug (Eastern Poland), drilled
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