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Investigations on the Echinoids of the Coasts of Provence

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Investigations on the Echinoids of the Coasts of Provence René Koehler INVESTIGATIONS ON THE ECHINOIDS OF THE COASTS OF PROVENCE Translated by John M. Lawrence Herizos Press Herizos Press, Tampa This is a translation of Recherches sur les Échinides des côtes de Provence by Réne Kœhler. 1883. Annales du Musée d’Histoire Naturelle de Marseille. – Zoologie. Mèmoire No. 3. Translation copyright © 2008 by John M. Lawrence 4 5 Translator’s note Jean Baptiste François René Kœhler was born 7 March 1860 in Saint-Dié and died 19 April 1931 in Lyon. The change in the spelling of his name must have been prior to 1891 because in that year Lucien Cuénot, in his memoir (Études morphologiques sur les Échinodermes), wrote both Kœhler and Koehler. Koehler defended theses in both natural sciences and in medicine at Nancy in 1883. He entered the faculty of sciences at Nancy as an assistant in zoology and then at Lyon in 1889. He was named to the chair of zoology at Lyon in 1894. Koehler was a specialist in echinoderms but also worked with isopods. He was president of the Zoological Society of France in 1911. His extensive collection of echinoderms and his library are in the Collections de Zoologie, part of the Collections Patrimoniales Universitaires of the Université Claude Bernard (Lyon). Much of the work for his thesis in natural sciences was done at the Station Endoume in Marseille. The station was still new, having been founded in 1869. Antoine-Fortuné Marion, director of the station, founded the Annales du Musée d’Histoires Naturelles de Marseille in 1883. Koehler’s memoir appeared in the series that founding year. The memoir is in three parts. The first concerns the anatomy of echinoids and considers all the major systems. The second part concerns the echinoid species found on the coast of Provence, with a description of their anatomy, habitat and distribution. The third part is a short addendum with limited observations on early development of hybrids. I have been consistent with following Koehler’s memoir, including whether or not a species name is italicized or diacritical marks are given or misspelled. I have retained the species names and authors used by Koehler. Because some of the names have changed, the present names of all species are given here. The pagination in the memoir is given in bold, e.g. 12. I thank David Pawson for providing the photograph of Koehler from the Zoological Archives, Université Claude Bernard, Lyon. 6 Species’ names and authors used by Species’ names at present1 Koehler Dorocidaris papillata, A. Ag. Cidaris cidaris, Linnaeus Arbacia pustulosa, Leske Arbacia lixula, Linnaeus Centrostephanus longispinus, Peters Centrostephanus longispinus, Philippi Echinus melo, Lam. Echinus melo, Lamarck Echinus acutus, Lam. Echinus acutus, Lamarck Psammechinus microtuberculatus,Ag. Psammechinus microtuberculatus, Blainville Strongylocentrotus lividus, Brandt Paracentrotus lividus, Lamarck Sphærechinus granularis, A. Ag. Sphaerechinus granularis, Lamarck Echinocyamus pusillus, Gray Echinocyamus pusillus, O.F. Müller Spatangus purpureus, Muller Spatangus purpureus, O.F. Müller Echinocardium flavescens Echinocardium flavescens, O.F. Müller Echinocardiium cordatum, Gray Echinocardium cordatum, Pennant Echinocardium mediterraneum, Gray Echinocardium mediterraneum, Forbes Schizaster canaliferus, L. Ag. and Desor Schizaster canliferus, Lamarck Brissopsis lyrifera, Ag. and Desor Brissus lyrifera, Forbes Brissus unicolor, Klein Brissus unicolor, Leske 1 I thank Michel Jangoux for providing the species’ names at present. 7 8 9 INVESTIGATIONS ON THE ECHINOIDS OF THE COASTS OF PROVENCE ------------------------------ INTRODUCTION AND HISTORY --------------- The goal that I proposed to myself in beginning this work was to make as complete a study as possible of the different systems that constitute the internal anatomy of the irregular urchins living on the coast of Provence. The works published on this subject are, in fact, very few and the two most recent studies by Hoffmann and Teuscher address, moreover, only the single genus Spatangus. They also are contradictory regarding a large number of points, particularly on all that was reported about the circulatory system. For that reason it appeared necessary to return to this study anew The internal anatomy of the regular urchins, which had been the object of more extensive and much more numerous works, seemed to me more advanced and known in a satisfactory fashion, especially after the research of Perrier on the circulatory system. Until then it was described in a very incomplete and vague fashion and often the subject of debate. After having finished the study of the spatangoids, I was led to occupy myself with the anatomy of regular urchins by the difficulty that I encountered in comparing the circulatory system of spatangoids with that of urchins. Moreover, the research of Teuscher published in 1876 did not agree in certain points with those of 6 Perrier. The histology of certain organs, the digestive tube and the tube feet, for example, required new investigations. The coasts of Provence offered me in addition very rich material for study that I was fully able to take advantage of thanks to the generous hospitality I received at the Laboratory of High Studies of the Faculty of Sciences of Marseille. The regular urchins are, in fact, represented in the Gulf of Marseille by seven species, of which four are always found in very great abundance (Echinus acutus, Strongylocentrotus lividus, Sphærechinus granularis and Psammechinus microtuberculatus). The three other species (Echinus melo, Dorocidaris papillata and Arbacia pustulosa) are much rarer. The spatangoids are represented there by seven species belonging to the five genera Spatangus, Echinocardium, Schizaster, Brissus and Brissopsis. Spatangus and Ech. flavescens are very common; the other species are in contrast very rare. But, thanks to those specimens preserved in alcohol placed at my disposition by M. Marion, I have been able to study the anatomy of these latter three genera. As to the class of clypeasteroids, they are represented in the Gulf only by Echinocyamus pusillus. The very small size of 10 this species does not lend itself to anatomical investigations and I have completely set aside its study. The most important knowledge we have on the anatomy of echinoids, especially on spatangoids, is not very old. Until 1870 the history of this group was still little known although we had the old works of Monro, Tiedemann, Delle Chiaje, and the more recent works of Milne Edwards, Krohn, Valentin and J. Muller. The most important works on echinoids were particularly those of zoology and paleontological zoology. The paleontologists are presented, in fact, with fossil echinoids that abound in the earth and whose remains are in a quite remarkable state of preservation. The result of these investigations, accompanied most frequently with analogous observations on living animals, has been to make the external morphology of these beings known in a very detailed manner. One has a very complete knowledge of the studied types about the test, but in reality the internal anatomy of various echinoids still remains very obscure. Nevertheless these investigations of paleontological zoology have given very important information on the path that this type has followed in its differentiation. They show us the ancient echinoids of the primitive earth (Palæchini) that by their obscure characters that approach the cystoids, acquire little by little the characters of echinoid types that begin with cidaroid forms. They differentiate then in a manner to offer a multiplicity of types unknown in the present. Several are very interesting because 7 they fill the gaps that seem to exist if we examine only living forms. It is thus that we know all the intermediary types between the regulars and irregulars that are so different in our time. We can follow step by step the irregularity of the echinoid organism that began by this curious migration of the anus and which, in fossil genera, is shown in all its phases. It is possible to suppose that, according to the measure of irregularity attained, the various anatomical systems whose equilibrium was strongly disturbed have shown considerable modifications made necessary by the great changes related to the arrangement of the plates of the test. We can recognize traces of these in living spatangoids. A comparative and very detailed study on the plates of the tests of living and fossil echinoids allowed Lovén2 (1872-1874) to establish their morphology in a definitive fashion, to find numerous points of relation of the various plates between regulars and irregulars, and to show, in living urchins, evident traces, although transitory, of the relationship of echinoids with crinoids. Important discoveries since Lovén’s work resulted from dredges in the deep sea. They have allowed the study of certain interesting types, sometimes with a strange and supposedly extinct form. They were able to be used by Lovén, and thus complete the beautiful series of studies of the Swedish scholar. We can thus say present knowledge of the test of echinoids studied in living animals and fossils is as complete as possible. This study, because of its scientific importance, is not the least interesting to us in echinoderms. It is a question that I must however lay aside because it is completely outside my subject. When I shall study the test in echinoids, I shall occupy myself exclusively only with the external appendages. Their 2 LOVEN. Etudes sur les Echinoïdes. Kong. Svenska Vetenkaps-Akademiens Handilingar. Bd 11, no 7. Stockholm. 11 study has some interest from the anatomical and zoological points of view. I shall also lay aside all morphological discussion that, only addressing facts already known, will not find its place here. I shall content myself with very rapidly reviewing the principal works published on the anatomy of echinoids. These works will be analyzed in the different chapters of this work. After that I shall occupy myself with such questions. This will allow me to consider them later rather than summarize them at this time.
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