ebook img

Paleoneurology 1804–1966: An Annotated Bibliography PDF

256 Pages·1975·20.541 MB·English
Save to my drive
Quick download
Download
Most books are stored in the elastic cloud where traffic is expensive. For this reason, we have a limit on daily download.

Preview Paleoneurology 1804–1966: An Annotated Bibliography

Advances in Anatomy, Embryology and Cell Biology Ergebnisse der Anatomie und Entwicklungsgeschichte Revues d'anatomie et de morphologie experimenLale 49/1-6 Editors A. Brodal, Oslo' W. Hild, Galveston' J. van LimlJoryh, Amsterdam R. Ortmann, Koln· T. H. Schiebler, Wurzbury·G. TOndury, Zurich·E. Wolff. Paris Tilly Edinger Paleoneurology 1804-1966 An Annotated Bibliography Springer-Verlag Berlin Heidelberg New York 1975 t Dr. Tilly Edinger Museum of Oomparative Zoology Harvard University OamlJridge/Mass. 02138 U.S.A. ISBN-13:978-3-540-07060-3 e-ISBN-13:978-3-642-66029-0 DOl: 10.1007/978-3-642-66029-0 Library of Congress Cataloging in Publication Data. Edinger, Tilly, 1897-1967. Paleoneurology 1804-1966. (Ad vances In anatomy, embryology, and cell biology; v. (9) Bibliography: p. Includes Index. 1. Vertebrates, Fossil-Bibliography. 2. BraIn-Evolutlon-Bibliography I. Title. II. Series. [DNLM: 1. Brain-Bibliography. 2. Evolution-Bibliography. 3. Paleontology-Bibliography. Wl AD433k v. 49/ZWL300 E23p 1804-1966] QL801.E67 vol. 49 [Z6033.P2] [QE841] 574.4'08s [016.566] 75-12819 This work Is subject to copyright. All rights are reserved, whether the whole or part of the material Is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under § 54 of the German Copyright Law where copies are made for other than private use, a fee Is payable to the publisher, the amount of the fee to be determined by agreement with the publisher. © by Springer-Verlag Berlin' Heidelberg 1975 The use of registered names, trademarks ete. in this publication does not Imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use, Contents Foreword (by Bryan Patterson). 7 General Catalogue . . . . . . 12 Appendix: Hominidae Catalogue 183 Systematic Index (by Joan Echols) 235 Foreword Dr. Tilly Edinger's first published paper dealt with a brain cast-in more exact terms an endocast of the cranial cavity-of Noth08auru8, a Triassic relative of the plesiosaurs. With this she embarked on a working lifetime of devotion to paleoneurology, a field of study that she was to transform. A daughter of the famous neurologist Ludwig Edinger, it was appropriate as well as fortunate that her early interest in fossil vertebrates should have become focused upon the recovery of such information concerning the history of the central nervous system as could be obtained from fossil material. Her father evidently had no direct influence upon her choice of· this then obscure and difficult subject, although within the family circle she presumably absorbed from him some appreciation of neoneurology. Indirectly, however, through his accumulation in Frankfurt of an outstanding collection of recent brains, he provided the comparative material essential to her studies during the years she spent there. Early in her career she published Die FOBsilen Gehirne (1929). Here was gathered together for the first time nearly all the widely scattered information on the topic. It had an immediate effect. As one author justly remarked, this "invaluable review ... serves not only as a basis for continuing and systematizing research on brain casts but also as an indication of the more serious gaps in present knowledge" (Simpson, 1933). The bibliography appended to it listed 250 titles. A bibliography she published in 1937 included 160 additional titles. Some of these were older papers that she had not previously encountered but 115 of them-a number equal to nearly half of the total listed in Die FOBsilen Gehirne had appeared in less than a decade after pUblication of that work. It may fairly be claimed that many, perhaps a majority, of these had been inspired in one way or another by it. The groundwork for a science of paleoneurology had been laid. During the decade that followed, the political situation in Germany made life increasingly difficult for her, although she continued to work and publish. The most interesting of her papers during this period dealt with sirenian brain casts (1933). She was able to show that the brain in this order, while increasing some what in size, had changed very little in form from the late Eocene to the present. This study, the first demonstration of an evolutionary series, however incomplete, over a very long stretch of time, might have served as a basis for erroneous con clusions as to the general course of brain evolution in the Mammalia had it con tinued to stand alone. It was to be superseded by a truly revolutionary study, one that was to bring paleoneurology into a new dimension. Fortunately for herself and for science Dr. Edinger was able to leave Germany in 1939 and to come to the Museum of Comparative Zoology in 1940. There she began work on a project that she had long believed should be carried out, a study of the history of the brain in the horse family from the then time of earliest appearance in the fossil record to the present. Published in 1948, the findings were spectacular: the brain of the early Eocene eohippus proved to be approximately on a level with that of Didelphis, one of the most primitive of living mammals, advances in brain structure, like those in other organs, had not proceeded at a constant rate within the family but had varied in tempo, the olfactory sense in 7 horses had not diminished in the course of time, and so on. The specializations achieved between eohippus and Equus had taken place after the family had be come distinct from others in the order Perissodactyla, and it followed from this, since most of the major groups of mammals were in existance by Eocene time, that a very great deal of parallel and convergent evolution in brain structure has gone on, particularly as regards the neocortical and neocerebellar areas. At a stroke, in fact, it had become obvious that the scala naturae approach to brain evolution could not be expected to reveal the history of events. In Die Fossilen Gehirne (p. 81), in agreement with an almost forgotten suggestion by Turner (J. Anat. Physiol., 25, p. 153, 1890), she had remarked "die Gyrifizierung also augen scheinlich in jeder Ordnung unabhangig entwickelt wird." Here was 'proof; she could now (1949, p. 21) state with confidence that "the material of paleoneurology demonstrates that the material of comparative anatomy, the series of extant brains, is not capable of revealing the actual courses of evolution which have resulted in these existing types." Evolution of the horse brain provided a standard against which much could be judged!. Looking back in time from the cerebra of eohippus and of the con temporary condylarth Phenacodus it became possible for her to interpret small protuberances on the otherwise smooth cerebra of Paleocene ungulates as being neopallial in nature (1956b, p. 19). The few known Mesozoic mammalian brain casts show nothing comparable to these little hillocks nor any suggestion of a rhinal fissure, from which (1964 b, p. 14) "one must conclude that non-olfactory cortex can hardly have been developed beyond the inherited 'primordium neo palii'." The neopallium, that hallmark of living Mammalia, did not make its appearance until long after the origin of the class, and its major elaborations were Cenozoic events. As regards neopallial specialization mammalian groups have varied widely. Some, including highly successful surviving ones, have under gone little. The horses are a group in which a good deal of advance went on throughout Tertiary time, others, such as camels, are more or less comparable to them in this respect, while yet others, cetaceans, elephants, hominids, have gone far beyond them. Early specialization was no guarantee of further progress. The early Eocene anaptomorphid prosimian Tetonius had the most advanced brain so far known for its time; its living relative Tarsius has one no different in essentials. Neocortical specialization went on at different rates even among closely related families. Palaeotheres and titanotheres, both of which became extinct in the Oligocene, were more advanced by that time than the contemporary M esohippus. Dr. Edinger took careful note of all this, and of more besides, and drew a bead (especially 1962a) on the hypotheses of O. C. Marsh, who in the 1870's and 80's had set forth various propositions under what he termed the "general law of brain growth." His views were based on quite insufficient evidence and were only partially correct, but they had a very pervasive influence and some of his more spectacular errors have been repeated ad nauseam. Their influence, she 1 Since this was written it has been reported (Soc. Vert. Pal. News Bull., no. 100, 1974, p. to) that L. Radinsky has "prepared an endocranial cast of Hyrawtherium that is signifi cantly more advanced than the specimen figured by Edinger in her monograph on the evolution of the horse brain; this suggests that Edinger's specimen was not from a perissodactyl." Dr. Edinger did consider this possibility, but decided against it on the evidence available at the time. Her conclusions as to the very rapid early evolution of the equid brain may thus require modification, but the rest of her views on the subject should not be affected. 8 believed, was due to their fitting in with the thinking of many neoneurologists in whose eyes the brain, and especially the neopallium, would naturally be the all important organ of the body. By her forceful presentation of the facts revealed by paleoneurology and by the very uneven levels of neopallial development exhibited by living mammals she placed the brain in proper perspective by implicitly stressing that it is simply one organ among the many on which evolution can act in various ways, and that specialization of it was not by any means the only road to evolutionary success. Unfortunately, having done this she went further and came in the end to question whether a specialized brain was of any advantage at all. In one of her last papers (1965) she-made the following remarkable statement: "The series [of camelid braincasts] also testifies against the concept that a brain thus advanced is an advantage in natural selection. Extinct in North America where they flourished for some 40 million years, wild Camelids now exist only in small areas of two other continents." As if advances in brain structure had not been selectively advantageous to camels during their 40 million year occupancy of the North American continent and as if a specialized brain, any more than specialized teeth or feet, should if really adaptive ensure a lineage possessing it against decline or extinction. In fact the statement is not one that a person familiar with current evolutionary thinking would be at all likely to make. And for this there is, I think, an explanation. During her lehrjahre there was no generally accepted body of evolutinary theory. Biologists agreed on the fact of evolution but not on much more; represen tatives of the disciplines involved were at swords' points and confusion reigned. Under these circumstances most people simply went on with their work while waiting for the dust to settle. As she passed into her meisterjahre she came to suffer increasingly from a form of deafness for which a hearing aid did not ade quately compensate. This she kept turned off much of the time, switching it on for purposes of conversation. And a conversation with her could be as taxing as it was fascinating. She would start to pursue a main topic, branch off to deal with a secondary one, perhaps branch off again from that to take up a tertiary one, exhaust this and return to the secondary, exhaust that and at last get back to the main track, often repeating the process once or twice more before reaching the end. Her listeners while following these discussions within discussions had also to keep the branching points firmly in mind if they were not to become lost. Once this key to her discourse was grasped her expositions could be followed but-and this is the point-it was largely one way communication. To venture intruding upon and breaking the thread of these conversational arabesques seemed almost improper; comments and queries that came to a listener's mind as she talked remained all too often unspoken. She was thus progressively cut off from the give and take that is so pleasant and useful a part of scientific life and which helps so much in the formulation and testing of ideas. Her affliction was already far advanced by the time that a synthesis was being hammered out in evolutionary theory. Absorbed as she was in her own investigations, much of this work may simply have passed by her unnoticed. She did not, at any rate, look at brain evolution in terms of the theory; had she done so I believe that aspects of it she found puzzling might have been clearer to her. Paleoneurology, as Dr. Edinger repeatedly stressed is just getting into its stride and an enormous amount of work lies ahead. There are whole orders, even of mammals, for which not one braincast is known. ("May this bibliography make 9 my colleagues aware of the ... gaps to be filled merely by having endocasts made of the crania in their collections.") Even for the Equidae, the best evolutionary sequence yet available, only a good outline is so far in hand and a great deal remains to be learned. Other lineages demand investigation. It would be inter esting for example to know when the cerebrum began to enlarge markedly in the Cetacea and the Proboscidea, and if the increases were gradual or initially rapid. A fascinating and wide open area of investigation lies in searching for clues to the times of origin of aspects of behavior, particularly behavior of what may be called the bread and butter kind-the ways in which animals cope with and make a living in the environments to which they are adapted-rather than of the more spectacular sorts-courtship, agonistic and the like-that' engage so much of the attention, of behaviorists. Behavior of this basic sort is intimately correlated with structure and function and may be reflected in the gross morphol ogy and surface details of the br8.in.. A case in point-an obvious one-is provided by the pterosaurs. From braincast evidence alone (Edinger, 1927, 1941) it should have been quite clear that these reptiles were fully capable of sustained flight, the long-tailed rhamphorhynchoids of stable, the short-tailed pterodactyloids of the more demanding unstable flight (cf. J. M. Smith, 1952). The-braincast of Archaeo pteryx, especially as reinterpreted by Jerison (Nature, 219, p. 1381, 1968), is consistent with the adaptations to stable flight shown by the skeleton and feathers. Continued progreBB in electrophysiological mapping should enable much progreBS to be made in tracing back somatic sensory specializations in various lineages. The problems and the possibilities flow on from the thresold to which Dr. Edinger has conducted us. Dr. Edinger had at one time hoped to round out her life's work with a modern ized version of Die Fossilen Gehime. To this end she kept up to date her ever growing card file of the literature and began a catalogue of the known fossil brain casts. By one of life's ironies, however, she had wrought too well; the field she had revivified in 1929 had grown during the intervening years to an extent such that a work of comparable scope would have required many years for its com pletio~. Besides, she came to feel (MS, 1961), "there is now such an expansion of aspects and also of material, and of literature ... that the situation is just too fluid ... to write a book about it all." Accordingly, she decided to concentrate first on completing the annotations to the bibliography, publish that and then go on to the catalogue. Fate in the shape of a fatal accident intervened but fortu nately the bibliography could be brought into order for publication. It is here presented and is the last contribution we shall receive from her. She had planned an extensive Introduction for which four slim notebooks are among her effects. I have repeatedly but without SUCceBB gone through these in the hope of re constructing what she had intended to write. The entries in them consist of jottings, in part mnemonic, that would have served only their author as the skeleton for a narrative; they cannot do so for another. The bibliography records a nearly six-fold increase of literature on the subject over the combined lists of 1929 and 1937. This "is mainly the result of the recent increase in the field, but it is also due to the inclusion of older literature that I had not previously seen and of works of a general nature, such as textbooks, that contain original remarks on fossil brains and/or figures copied from other pUblications. All this should make the bibliography a historical record of paleoneur ogy, reflecting both its spread and the changing attitudes toward the value of 10 endocranial casts." The annotations refer strictly to paleoneurological content and therefore "[I must] warn that necessarily partial quotations may give a wrong impression. Picking [out] only data about and comments on paleoneurology is not an abstract of the entire publication, nor can brief quotes contain all of an author's ideas." Papers that she believed to contain errors of one sort or another tend to be disproportionately annotated. Somewhat paradoxically there fore" it is the best, the entirely reliable papers, which are listed with the briefest annotations-a mere pointing out that unobjectionable descriptions exist." Some contributions may have been missed: "much to my regret, papers published entirely in non-Latin alphabets, although I suspect they-contain paleoneuro logical data, have been passed over unless they contained figures of endocranial casts." In the course of her work she came across numerous references that might have been expected from the titles to discuss paleoneurology yet made no mention of it at all. In her notebooks she laments the time spent on these, but readers will nonetheless be grateful for the trouble taken over them. If an alluring title, such as "Phylogeny of the avian brain," does not occur in the bibliography it is a rather safe bet that, however interesting otherwise, the paper contains nothing paleoneurological. A large part of the literature on fossil hominid brain casts is, she felt, "utterly different from that on brains of extinct animals, which is anatomical"; much of it is "not really brain anatomy, except for [attempted] tracing of this or that sulcus-for which hominid endocasts, particularly those made with methods now obsolete, are most unsuitable objects." In one notebook she quotes with approval von Bonin's stricture (1963, p. 41) that "much that we have gathered from the literature is of little value, since it is based on mistaken premises and homologies which can hardly be expected to survive a more critical examination." In defer ence to her wishes nearly all the hominid literature is set apart and is not indexed. Here and there through the notebooks are scattered sorrowful entries com menting on discussions by some neoneurologists-even in the 1960's--of brain evolution based on series of living forms, and indignant ones castigating yet another repetition of one of Marsh's more sensational errors. For an end to such things we can look with confidence to the passage of time. Support for the preparation of the bibliography and of other work was pro vided her by the Milton Fund of Harvard University, the American Association of University Women, the American Philosophical Society and the National Science Foundation through Grants GB 1364 and G 23566. Aid was received from numerous individuals over the years; grateful acknowledgment is due to Dr. Joan Echols for the preparation of the Index to the General Catalogue, to her and to Dr. Clayton E. Ray for aid on the annotations, to Mrs. Eleanor Adam Gossen, Mrs. Donna Ray, Mr. Dean Smith and Mrs. B. Gail Browne for typing and checking, and to Mrs. Browne for final arranging and editing. It was Dr. Edinger's hope that this work would give renewed impetus to the subject she loved; this will surely prove to be the case. Museum of Comparative Zoology, Harvard University Bryan Patterson September, 1970 Papers cited in the foregoing by year only may be found in the bibliography. Quotations without indication of source have been taken from the notebooks. For obituary notices of Dr. Edinger see: H. Hofer (Morph. Jahrb., 113, pp. 303-313,1969), A. S. Romer (News Bull. Soc. Vert. Pal., no. 81, pp. 51-53, 1967) and H. Tobien (Palaont. Zeitsch., 42, pp. 1-2, 1968). II General Catalogue Abel, 0.: Grundziige der Palaeobiologie der Wirbeltiere. Stuttgart. 1912. [No brains men tioned.] In: "Special Adaptations to Life in Water", "The parietal organ" (pp. 463-465), enlarging upon Dollo 1905, finds that, as for example in Plioplatecarpus, the parietal organ "was best developed in those vertebrates whose eyes lie on the upper surface of the head" [which would exclude, e.g., therapsids!] "Likewise enigmatic is the physiological significance of the hypophysial foramen in ichthyosaurs (figs. 357, 358)" [from Broili 1909, with the respective opening labelled in the ventral view, "Austrittsstelle der Hypophyse"!] Abel, 0.: Die eocanen Sirenen der Mittelmeerregion. Erster Teil: Der Schadel von Eotherium aegyptiacum. Palaeontogr. 09, 289-360 (1963). pp. 350-351, "The Brain", uses three natural endocasts; pI. XXX, fs. 1, 5; pI. XXXII, f. 6; pI. XXXIII, fs. 3-5; detailed de scription of last specimen in legend Abel, 0.: Vorgeschichte der Cetaceen. Pp. 395-410, in Weber, 1928 a. P. 399, brain of Zeuglodon; notes retention of large olfactory nerve Abel, 0.: Embrithopoda. Pp. 440-445 in Weber, 1928b. P. 441, "The volume ofthe braincase is larger than in the amblypods." Fig. 273, A-C, Arsinoitherium brain, after Andrews 1906 Abel, 0.: Vorgeschichte der Sirenia. P.469-504 in Weber, 1928c. P. 497, Eotherium, known first only by an endocranial cast. Fig. 326, otic region and endocranial cast, lateral, after Abel 1912 Abel, 0.: Amblypoda. Pp. 690-695 in Weber, 1928d. P. 693, brain of Uintatherium extraordi narily small Abel, 0.: Saugetiere (Palaontologie). Handworterb. Naturwiss., 2nd ed., 8, 859-944, 1933. In general part lists, among" Some of the most important specializations in the course of phylogeny of the mammals": "brain capsule small, narrow~brain capsule broad, vaulted" (p. 862). In systematic part, introduction to Carnivora contains a paragraph entitled "The Specialization of the Brain of the Carnivores": "The oldest Carnivora (e.g., Arotooyon and Triisodon of the Paleocene, Didymictis and Sinopa of the Lower Eocene) had a small, little specialized brain whose evolutionary level is comparable to that of marsupials. In the course of Tertiary times there occurs, independently in the dif ferent branches, a conspicuous specialization and size increase of the brain" (p. 883). [Otherwise, this extensive memoir takes no notice of brain structure, nor post-Marshian literature; it merely re-states and generalizes Marsh's dicta in the following instances:] The family Elotheriidae is characterized as having a brain cavity "almost reptile-like small" (p. 920), the Condylarthra as having a very small brain and the cerebellum poste rior to the smooth cerebrum (p. 921). Brain in skull of Brontotherium is copied from Marsh "to show the extraordinary smallness of the brain in proportion to the skull" (fig. 112). "The brain of Goryphodon was extraordinarily small ... Also in the Dinoceratidae the brain cavity was extremely small" (p. 936) Agnew, J. D.: Cranio-osteological studies in Dicynodon grimbeeki with special reference to the sphenethmoid region and cranial kinesis. Palaeont. afro 6, 77-107 (1959). Two serially sectioned skulls establish that the preparietal forms the anterior border of the parietal foramen which is "well-developed in both specimens" (p. 85) [its length is almost half the breadth of the foramen magnum in fig. 34], and the presence of a distinct floccular fossa (pp. 102, 106, figs. 41-42) Ahl, E.: Xenopus stromeri Ahl n.sp. Pp. 141-142 in E. Kaiser, ed., Die Diamantenwiiste Siid westafrikas, 2, Berlin. 1926. All frontoparietals of this presumably Lower Miocene frog have a small, longitudinally oval foramen close to the anterior end (p. 142, fig. 22). [This canal, which encloses the connection of Stirnorgan and brain, has since been ascertained in extant species of Anura; see, e.g. Winterhalter 1931 in Acta zooi. 12, pp. 55-58, and Griffiths 1954 in Proc. Zooi. Soc. London 123, p. 788.] Aldinger, H.: tlber einen Eugnathiden aus der unteren Wolgastufe von Ostgronland. Meddei. om Gremland 86, no. 4, 1-52 (1932). "Brain Cavity and Brain" of Gaturus (pp.21-23) and its cranial "Vessels and Nerves" (pp.26-30) are described on the basis of sectioned crania (figs. 3-6) and an "endocranium" (pI. I, f.2) in which some of the natural endo- 12

See more

The list of books you might like

Most books are stored in the elastic cloud where traffic is expensive. For this reason, we have a limit on daily download.