The Appendages, Anatomy, and Relationships
of Trilobites
BY
PERCY E. RAYMOND, Ph.D.
ASSOCIATE PROFESSOR OF PALAEONTOLOGY, AND CURATOR OF INVERTEBRATE
PALAEONTOLOGY IN THE MUSEUM OF COMPARATIVE ZOOLOGY,
HARVARD UNIVERSITY
NEW HAVEN, CONNECTICUT
PUBLISHED BY THE
CONNECTICUT ACADEMY OF ARTS AND SCIENCES
AND TO BE OBTAINED ALSO FROM THE
YALE UNIVERSITY PRESS
THE TUTTLE, MOREHOUSE & TAYLOR COMPANY
TO THE MEMORY OF
CHARLES EMERSON BEECHER
SKILLFUL WITH HAND, BRAIN, AND PEN; REVEALER OF THE MYSTERIES OF TRILOBITES;
THIS MEMOIR IS DEDICATED
FOREWORD.
By CHARLES SCHUCHERT.
Trilobites are among the most interesting of invertebrate fossils and have long attracted the attention of amateur collectors and men of science. These "three-lobed minerals" have been mentioned or described in books at least since 1698 and now several thousand species are known to palæontologists. To this group of students they are the most characteristic animals of the seas of Palæozoic time, and even though they are usually preserved as dismembered parts, thousands upon thousands of "whole ones" are stored in the museums of the world. By "whole ones" perfect individuals are not meant, for before they became fossils the wear and tear of their time and the process of decomposition had taken away all the softer parts and even most of the harder exterior covering. What is usually preserved and revealed to us when the trilobites weather out of the embrace of their entombing rocks is the test, the hard shell of the upper or dorsal side. From time to time fragments of the under or limb-bearing side had been discovered, first by Elkanah Billings, but before 1876 there was no known place to which one could go to dig out of the ground trilobites retaining the parts of the ventral side.
Students of trilobites have always wanted specimens to be delivered to them weathered out of the rock by nature and revealing the ventral anatomy without further work than the collecting, but the wish has never been fulfilled. In the Utica black shales, near Rome, New York, there was finally discovered in 1892 a layer less than ten millimeters thick, bearing hundreds of Triarthrus becki with most of the ventral anatomy intact. The collector's first inkling that such were present in the Utica formation came to him in a chance find in 1884, and for eight years he sought off and on for the stratum whence this specimen came. His long search was finally rewarded by the discovery of the bed, and lo! here were to be had, in golden color, prostrate specimens with the breathing and crawling legs and the long and beautifully curved feeling organs all replaced by iron pyrites. Fool's gold in this case helped to make a palæontologic paradise. The bed contained not only such specimens of Triarthrus becki, but also, though more rarely, of Cryptolithus tessellatus and exceptionally of Acidaspis trentonensis. This important discovery, which has figured so largely in unraveling the evolution of the Crustacea and even has a bearing on that of most of the Arthropoda, was made by Mr. W. S. Valiant, then curator of the Museum of Rutgers College.
There were, however, great material difficulties to overcome before the specimens revealed themselves with all of their information exposed for study. No surgeon was needed, but a worker knowing the great scientific value of what was hidden, and with endless patience and marked skill in preparation of fossils. Much could be revealed with the hammer, because specimens were fairly abundant. A chance fracture at times showed considerable portions, often both antennæ entire, and more rarely the limbs protruding beyond the test, but the entire detail of any one limb or the variation between the limbs of the head, thorax, and tail was the problem to be solved. No man ever loved a knotty problem more than Charles E. Beecher. Any new puzzle tempted him, and this one of Triarthrus becki interested him most of all and kept him busy for years. From the summer of 1893. when he quarried out two tons of the pay stratum at Rome, until his death in 1904, his time was devoted in the main to its solution by preparing these trilobites and learning their anatomical significance.
The specimens of Triarthrus becki from Rome are pseudomorphs composed of iron pyrites, as has been said, and are buried in a gray-black carbonaceous shale. A little rubbing of the specimens soon makes of them bronze images of the former trilobite and while under preparation they are therefore easily seen. However, as the average individual is under an inch in length and as all the limbs other than the antennæ are double or biramous, one lying over the other, and the outer one fringed with a filamentous beard, the parts to be revealed by the preparator are so small and delicate that the final touch often obliterates them. These inherent difficulties in the material were finally overcome by endless trials on several thousand specimens, each one of which revealed something of the ventral anatomy. Finally some 500 specimens worthy of detailed preparation were left, and on about 50 of these Beecher's descriptions of Triarthrus and Cryptolithus were based.
The black shale in which the specimens are buried is softer than the pseudomorphous trilobites, a condition that is of the greatest value in preparation. With chisel and mallet the trilobites are sought in the slabs of shale and then with sharp chisels of the dental type they are revealed in the rough. At first Beecher sought to clean them further by chemical methods, and together with his friends, the chemist Horace L. Wells, and the petrologist Louis V. Pirsson, several solutions were tried, but in all cases the fossils were so much decomposed as to make them useless in study. Therefore Beecher had to depend wholly oh abrasives applied to the specimens with pieces of rubber. Much of this delicate work was done on a dental lathe, but in the final cleaning most of it was done with patient work by hand. Rubber has the great advantage of being tough and yet much softer than either specimen or shale. As the shale is softer than the iron pyrites, the abrasives (carborundum, emery, or pumice) took away the matrix more quickly than the trilobite itself. When a part was fully developed, the rubbers were cut to smaller and smaller dimensions and the abrading reduced to minute areas. So the work went on and on, helped along from time to time by the dental chisels. Finally Beecher became so expert with these fossils that after one side was developed he would embed the specimen in Canada balsam and fix it on a glass slide, thus enabling him to cut down from the opposite side. This was done especially with Cryptolithus because of the great scarcity of material preserving the limbs, and two of these revealed both sides of the individuals, though they were then hardly thicker than writing paper.
Then came illustrations, which at first were camera-lucida drawings in pencil smoothed out with pen and ink. "In some quarters," however, it has been said, "his
When the writer, in the autumn of 1904, succeeded Professor Beecher in the chair of Palæontology at Yale, he expected to find considerable manuscript relating to the ventral anatomy of the trilobites, but there was only one page. It was Beecher's method first to prepare and thoroughly study the material in hand, then to make the necessary illustrations, and between times to read what others had written. There was no written output until everything had been investigated and read, certain passages being marked for later reference. Then when all was assimilated, he would write the headings of topics as they came to him, later cutting them apart and arranging them in a logical sequence. When the writer visited him in his home in January 1904, he was primed for his final trilobite memoir, but the writing of it had not been begun.
The writer has never made the trilobites his special subjects for study as he has the brachiopods, and therefore felt that he should not try to bring to light merely the material things that Beecher had so well wrought out. It seemed at first an impossible task to find the specialist and friend to do Beecher justice, but as the years have passed, one of Beecher's students, always especially interested in trilobites, has grown into a full appreciation of their structures and significance, and to him has fallen the continuation of his master's work. If in the following pages he departs here and there from the accepted interpretation and the results of others, it is because his scientific training, in desiring to see with his own eyes the structures as they are, has led him to accept only those interpretations that are based on tangible evidence as he understands such. Furthermore, in seeking the relationship of the trilobites to the rest of the Arthropoda, his wide study of material and literature, checked up by the ontogeny of fossil and recent forms, has led him in places from the beaten path of supposedly ascertained phylogenies. His results, however, have been won through a detailed study of the interrelations of the Arthropoda, starting from the fact that the Trilobita are chronogenetically the oldest and most primitive. The trilobites are held by him to be the most simple, generalized, ancient Crustacea known, and the progenitors, directly and indirectly, of all Arthropoda.
It is now twenty-six years since Professor Beecher began his publications on the class Trilobita, and in commemoration of him and his work, Professor Percy E. Raymond of Harvard University presents this memoir, to bring to fruition the studies and teachings of his honored guide. It has been with Professor Raymond a labor of love, and it is for the writer of this foreword a long-desired memorial to the man to whose position in the Museum and University he had the privilege of succeeding.
Yale University, New Haven, Connecticut.
PREFACE.
The primary object of this memoir is, as has been stated by Professor Schuchert, to. rescue from oblivion the results of the last few years of Professor Beecher's investigations on the ventral anatomy of trilobites. Since he left his data in the form of drawings and photographs, without even rough notes, it became necessary, in order to write a text to accompany the plates, to restudy the entire subject. Under these circumstances, it seemed best to include all that is known about the appendages of trilobites, thus bringing together a summary of present information on the subject.
The growth of the memoir to its present size has been a gradual one. As first completed in 1917, it contained an account of the appendages only. Thoughts upon the probable use of the appendages led to the discussion of possible habits, and that in turn to a consideration of all that is known or could be inferred of the structure and anatomy of the trilobite. Then followed an inquiry into the relationships to other Arthropoda, which ultimately upset firmly established preconceptions of the isolated position of the group, and led to a modification of Bernard's view of its ancestry.
During the progress of the work, I have had the opportunity of examining most of the known specimens retaining appendages. From the Marsh collection in the Yale University Museum were selected the forty-six specimens showing best the appendages of Triarthrus, Cryptolithus, and Acidaspis. Dr. Charles D. Walcott very kindly returned to the Museum of Comparative Zoology the slices of Ceraurus, Calymene, and Isotelus which were the basis of his paper of 1881, and which had been loaned him for further study. He loaned also eight of the more important specimens of Neolenus serratus, and two of Triarthrus becki. At the United States National Museum I saw the specimens of Isotelus described by Mickleborough and the isolated limbs of Calymene from near Cincinnati. The Isotelus at Ottawa I had already studied with some care while an officer of the Geological Survey of Canada.
This memoir consists, as shown in the table of contents, of four parts. The appendages of Neolenus, Isotelus, Ptychoparia, Kootenia, Ceraurus, Calymene, and Acidaspis are discussed, as fully as circumstances warrant, in the first part, and new restorations of the ventral surfaces of Neolenus, Isotelus, Triarthrus, Ceraurus and Cryptolithus are included It is not supposed that these restorations will be of permanent value in all of their detail, but they are put forward as the best approximations to the real structure that the writer is able to present from the materials so far discovered. I am greatly indebted to Doctor Elvira Wood for the care and skill with which she has worked up these restorations from my rather sketchy suggestions. She has put into them not only a great amount of patient work, but also the results of considerable study of the specimens.
Part II is a discussion of the internal anatomy of the trilobite and a brief statement of some of the possible habits and methods of life of these animals. Part III, which begins with a survey of the relationships of the trilobites to other Arthropoda, is largely taken up with an attempt to demonstrate the primitive characteristics of the former, and their probable ancestral position. The form of the ancestor of the trilobite is deduced from a study of the morphology, ontogeny, and phylogeny of the group, and evidence adduced to indicate that it was a depressed, flattened,
In Part IV are included somewhat detailed descriptions of a few of the best specimens of Triarthrus and Cryptolithus. Professor Beecher, while an observer of the minutest details, believed in publishing only the broader, more general results of his investigations. This method made his papers brief, readable, and striking, but it also resulted in leaving in some minds a certain amount of doubt about the correctness of the observations. In a matter so important as this, it has seemed that palæontologists are entitled to the fullest possible knowledge of the specimens on which the conclusions are based. The last part is, therefore, a record of the data for the restorations of Triarthrus and Cryptolithus.
The illustrations in the plates were nearly all made by or under the supervision of Professor Beecher, as were also text figures 45 and 46.
In conclusion, I wish to express my thanks to Mrs. Charles E. Beecher for the use of drawings which were the personal property of Professor Beecher; to Doctor Charles D. Walcott for photographs of the limbs of Calymene, and for his kindness in sending me the slices of trilobites from Trenton Falls and specimens of Neolenus and Triarthrus; to Doctor R. V. Chamberlin for suggestions and criticisms in regard to the relationship of trilobites to Insecta, Arachnida, Chilopoda, and Diplopoda; to Mr. Samuel Henshaw, Director of the Museum of Comparative Zoology, for permission to use the time which has been devoted to this work; and to Miss Clara M. Le Vene, for assistance in the preparation of the manuscript. My greatest debt is to Professor Charles Schuchert, to whom the work owed its inception, who has assisted in many ways during its prosecution, and who read the manuscript, and arranged for its publication. To him I can only express my warmest thanks for the favors which I have received and for the efforts which he has put forth to make this a worthy memorial to our friend and my teacher, Professor Charles Emerson Beecher.
Harvard University, Cambridge, Mass.
November, 1919.
TABLE OF CONTENTS.
Historical review 17 Part I. The appendages of trilobites 20 Terminology 20 The appendages of Neolenus 21 Historical 21 Neolenus serratus (Rominger) 21 Cephalon 21 Thorax 22 Pygidium 23 Epipodites and exites 23 Description of individual specimens 23 Restoration of Neolenus 30 Nathorstia transitans Walcott 31 The appendages of Isotelus 32 Historical 32 Isotelus latus Raymond 34 Isotelus maximus Locke 35 Restoration of Isotelus 37 Isotelus gigas Dekay 37 Isotelus arenicola Raymond 39 The appendages of Triarthrus (see alsoLIST OF ILLUSTRATIONS.
1 Triarthrus becki Green. Diagram of limb to show nomenclature employed 20 2 Neolenus serratus (Rominger). Two thoracic appendages 24 3 The same. An exopodite 26 4 The same. A so-called "epipodite" 26 5 The same. The so-called "exites" 29 6 The same. A cephalic limb 29 7 The same. Restoration of a transverse section 30 8 The same. Restoration of the ventral surface 31 9 Isotelus. Restoration of the ventral surface 38 10 Triarthrus becki Green. Restoration of the ventral surface 41 11 The same. Median appendage 44 12 Ceraurus pleurexanthemus Green. Slice showing an exopodite 49 13 Calymene senaria Conrad. Slice showing cephalic coxopodites 53 14 The same. Another similar slice 53 15 The same. Slice showing method of articulation of the appendages 53 16 The same. Restoration of the ventral surface 55 17 Ceraurus pleurexanthemus Green. Slice showing the method of articulation of the appendages 58 18 The same. Slice showing an exopodite above an endopodite 58 19 The same. Restoration of a transverse section 60 20 Cryptolithus tessellatus Green. Restoration of the ventral surface 63 21 Ceraurus pleurexanthemus Green. Slice showing the abdominal sheath 79 22 The same. Slice showing the large alimentary canal 79 23 Calymene senaria Green. Slice showing the large alimentary canal 79[15] 24 Ceraurus pleurexanthemus Green. Restoration of a longitudinal section 81 25 Cryptolithus tessellatus Green. Cheek showing the genal cæca 84 26 Illænus. Volborth's figure of the heart 85Frontispiece. Charles Emerson Beecher, 1896.
Plates 1-5. Photographs of Triarthrus becki, made by C. E. Beecher.
Plate 6. Photographs of Triarthrus becki (figs. 1-3), Acidaspis trentonensis (fig. 6), and Cryptolithus tessellatus (fig. 7), made by C. E. Beecher. Photographs of the endopodites of a probable species of Calymene (figs. 4, 5)
Plates 7-8. Photographs of Cryptolithus tessellatus, made by C. E. Beecher.
Plate 9. Drawings of Cryptolithus tessellatus, made by C. E. Beecher or under his direction.
Plate 10. Photographs of Isotelus latus and I. maximus, made by C. E. Beecher.
Plate 11. Drawing of a restoration of Ceraurus pleurexanthemus, made by Elvira Wood.