THE UNIVERSITY OF ILLINOIS LIBRARY NATURAL HISTORY SURVEY 5705 ILL v.3cop.4 ILLINOIS BIOLOGICAL MONOGRAPHS Vol. IX January, 1924 No. 1 THE CALCIFEROUS GLANDS OF LUMBRICIDAE AND DIPLOCARDIA WITH TWELVE PLATES BY FRANK SMITH Price $1.25 Published by the University op Illinois under the auspices of the graduate school Urbana, Illinois f«T"u Siii-VfrV UNIVERSITY OF ILLINOIS STUDIES PUBLISHED ILLINOIS BIOLOGICAL MONOGRAPHS The Illinois Biological Monographs is a serial published quarterly by the Univer- sity of niinois. It is the purpose of the publication to present in monographic form research contributions of especial merit and of somewhat extended character. Each volume will include about 500 pages and 20 or more full page plates. The series may be obtained under subscription price of three dollars yearly. Individual numbers will be sold separately at prices determined by the size and amount of illustration contained in the particular number. 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ILLINOIS BIOLOGICAL MONOGRAPHS PUBLISHED QUARTERLY UNDER THE AUSPICES OF THE GR.'\DUATE SCHOOL BY THE UNIVERSITY OF ILLINOIS VOLUME IX Urbana, Illinois 1924 Editorial Committee Stephen Alfred Forbes William Trelease Henry Baldwin Ward i>2 1 ^o5 P. ^ TABLE OF CONTENTS VOLUME DC NUMBERS PAGES 1. The calciferous glands of Lumbricidae and Diplocardia. By Frank Smith. With 12 plates 1-76 2-3. A biologic and taxonomic study of the Microsporidia. By Roksabro Kudo. With 27 plates 77-344 4. Animal ecology of an Illinois elm-maple forest. By A. O. Weese. With 7 plates 345-438 73217^ NatVftl History Surv«f TJbriict-v ILLINOIS BIOLOGICAL MONOGRAPHS Vol. IX January, 1924 No. i Editorial Committee Stephen Alfred Forbes William Trelease Henry Baldwin Ward Published under the Auspices of the Graduate School by the University of Illinois Copyright 1923 by the University of Illinois Distributed November 20, 1924 THE CALCIFEROUS GLANDS OF LUMBRICIDAE AND DIPLOCARDIA WITH TWELVE PLATES BY FRANK SMITH Contributions from the Zoological Laboratory of the University of Illinois No. 233 TABLE OF CONTENTS Introduction J Historical f 8 Leo, J. 1820 Morren, C. F. A. 1826 W Henle, F. G. J. 1835 W Lankester, E. R. 1864 H 11 D'Udekem, J. 1865 Claparede, E. 1869 12 Perrier, E. 1874 13 Horst, R. 1876 13 Danvin, C. 1881 14 Vejdovskv, F. 1884 If Marshall and Hurst. 1887 1* Kulagin, N. 1888 15 Rosa, D. 1887-1895 15 Beddard, F. E. 1895 15 Moore, H. F. 1895 16 Harrington, N. 1899 JJ Ribaucourt, E. 1901 17 Combault, A. 1909 18 20 Stephenson, J. and Prashad, B. 1919 Zoological text-books 20 21 Resume ; Types of calciferous glands in Lumbricidae 24 Esophageal pouches wanting 25 Hclodrilus oculalus Hoffmeister 25 Hdodrilus vnidus var. hortensis (Michaelsen) 26 Helodrilus venclus (Rosa) 27 Hdodrilus foelidus (Savigny) 27 Helodrilus lonnbcrgi (Michaelsen) 28 Esophageal pouches present; no other lateral enlargements 29 Helodrilus paluslris (Moore) 29 Hdodrilus gicsdcri (Ude) 30 Helodrilus brddardi (Michaelsen) 3t Hdodrilus parvus (Eisen) 31 Helodrilus roscus (Savigny) 32 Hdodrilus letrocdrus (Savigny) 32 Helodrilus chlorolicus (Savigny) 33 Hdodrilus ailigiiiosus i. Irapczoides (Duges) 34 Octolasium laeteum (Orley) 34 Esophageal pouches and other lateral enlargements present 36 LumbricHs Icrrcslris Linnaeus 36 Lumhricus rubdlus Hoffmeister 36 Lumbrii us lasloncus (Savigny) 36 Helodrilus octaedrus (Savigny) 36 Calciferous glands in abnormal earthworms 3/ Mode of origin of calciferous glands 39 Descriptions of new species of Diplocardia 41 Diplocardia mississippicnsis n. sp 41 Diplocardia jloridana n. sp 45 Summary 50 51 Literature cited .• Explanation of plates 54 GLANDS OF LUMBRICIDAE AND DIPLOCARDIA—SMITH INTRODUCTION Certain organs related to the posterior part of the esophagus in earth- worms belonging to the family Lumbricidae, and also in those of certain other S3'stematic groups, have been studied and described at various times since the early part of the nineteenth century. These organs have received various names, of which calciferous glands, esophageal glands, and glands of Morren are among those more commonly used in the recent literature on the subject. The purpose in the preparation of this paper has been to bring together the more important results of the work of previous investigators who have studied the anatomy of such organs in the Lumbricidae; to add some facts gained from my own study of worms of this group; and to describe similar organs found in two new species of the genus Diplocardia which belongs to another family of earthworms. This involves the description of these two new species, but it is hoped that a better basis for an outline of the probable history of the development of the complicated structures in the Lumbricidae will result from the dis- cussion of the interesting series of such organs found in the different species of Diplocardia. Among these species we find organs of a degree of complexity similar to that in Lumbricidae; others of a very simple type in which we have merely a few longitudinal folds instead of a complicated set of longitudinal tunnellike chambers; and several intermediate stages between these two extremes. Preliminary to a discussion of the literature on the subject and to an attempt to trace the principal steps in the attainment of the present state of our knowledge of the anatomy of the calciferous glands, it is necessary to briefly summarize the principal facts of their structure, and explain the terminology used. In the Lumbricidae more commonly studied, the part of the esophagus which is included in somites 10-14, and is just anterior to the crop, is characterized by an extensive glandular development of the wall. This includes a pair of evaginated pouches in somite 10 which communicate through large apertures with the lumen of the esophagus. In the wall of the somites next following is a series of longitudinal tunnel- like chambers which have their cavities continued into the posterior part of 10 where they are in communication with the large cavities of the paired pouches in that somite. Figure 17 illustrates the appearance of a trans- verse section, and figure 50 of a sagittal section through the wall of one side in this region. Longitudinal blood vessels extend along the margins of the lamellae which separate the longitudinal chambers, both along the inner 8 ILLINOIS BIOLOGICAL MONOGRAPHS [8 margins in close relation to the lining epithelium of the esophagus and also along the outer margins in close relation to the muscle layers of the esopha- gus. These vessels pro\acle an extensive blood supply and account for frequent references to the reddish color. The chambers are much reduced in size in somite 14 and the diameter of the esophagus is correspondingly reduced. In a very few species including the much described Lumbricus terreslris, the lateral walls of the esophagus are considerably expanded in each of the somites 11 and 12 and this has led to the common practice of referring to three pairs of glands in somites 10, 11 and 12, when in reality we are dealing with what would more reasonably be considered as a single organ.