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PSYCHE

A Journal of Entomology

Volume 65 1958

Editorial Board

Frank M. Carpenter, Editor P, J. Darlington, Jr. W. L. Brown, Jr. H. Levi

E. 0. Wilson

Published Quarterly by the Cambridge Entomological Club Editorial Office; Biological Laboratories Harvard University Cambridge, Mass., U. S. A.

The numbers of Psyche issued during the past year were mailed on the following dates :

Vol. 54, no. 4, Dec., 1957 : January 26, 1959

Vol. 65, no. 1, March, 1958 : May 22, 1959

Vol. 65, nos. 2-3, June-Sept., 1958: Decembers, 1959

S^n ^ i

PSYCHE

INDEX TO VOL. 65, 1958

INDEX TO AUTHORS

Adams, P. A. The Relationship of the Protoperlaria and the Endoptery- gota. 115

Blake, Doris H. Some New West Indian Eumolpid Beetles. 91

Brown, W. L., Jr. A Formica Slave-maker Raiding the Nest of a Myrmicine Ant. 39

The Indo-Australian Species of the Ant Genus Stramigenys Fr. Smith: Group of S. godeffroyi in Borneo. 81

Carpenter, F. M. Mexican Snake-Flies (Neuroptera: Raphidiodea). 52

Christiarisen, Iv. The Colletmbola of Lebanon and Western Syria. Part III. Family Isotomidae. 59

Deichmann, Elisabeth. Elizabeth Bangs Bryant. 1

Parsons, Margaret C. Cephalic Glands in Gelastocoris (Hemiptera- Heteroptera) . 99

Wilson, E. O. A Chemical Releaser of Alarm and Digging Behavior in the Ant Fogonomyrmex badius (Latreille). 41

Patchy Distributions of Ant Species in New Guinea Rain Forests. 26

Wilson, E. O., N. I. Durlach and L. M. Roth. Chemical Releasers of Necrophoric Behavior in Ants. 108

Woodland, J. T. Oogenesis and Fertilization in Thermobia do?nestica (Packard). 11

131

INDEX TO SUBJECTS

All new genera, new species and new names are printed in Capital type.

Ants, 26, 39, 41, 81, 108 Anurophorus asfouri, 60 Anurophorus coifjaiti, 59 Aphaenogaster rudis, 39 Ballistura levantina, 70 Ballistura schotti, 70 Cephalic Glands in Gelastocoris (Hemiptera-Heteroptera) , 99 Chalcosicya alayoi, 91 Chemical Releaser of Alarm and Digging Behavior in the Ant Pogonomyrmex badius (Latreille), 41

Chemical Releasers of Necrophoric Behavior in Ants, 108 Collembola of Lebanon and Western Syria. Part III. Family Isoto- midae, 59

Distribution of Ant Species in New Guinea Rain Forests, 26 Elizabeth Bangs Bryant, 1 Endoptervgota, 115 Eumolpidae, 91

Fertilization in Thermobia domes- tica, 16

Folsomia, 63, 69 Folsomia Candida, 68 Folsomia cavicola, 69 Folsomia penicula, 65 Folsomia quadrioculata, 63 Folsomides americanus, 63 Formica Slave-maker Raiding the Nest of a Mvrmicine Ant, 39 Formica subintegra, 39 Indo-Australian Species of the Ant Strumigenys Fr. Smith; Group of S. godeffroyi in Borneo, 81 Inocellia pilicornis, 56 Inocelliidae, 56 Isotoma notabilis, 78 Isotoma viridis, 78

Isotomidae, 59 Isotomina bitub erculala, 71 Isotomina pontica, 71 Isotomina salaymehi, 72 Isotomuris palustris, 79 Lepismatids, 20 Metachroma bredeni, 94 Metachroma cornutum, 93 Metachroma grande, 98 Metachroma mulipunctatum, 96 Metachroma nanum, 96 Metachroma zayasi, 92 Mexican Snake-Flies (Neuroptera ; Raphidiodea) , 52

Oogenesis and Fertilization in Thermobia domestica (Packard),

11

Patchy Distributions of Ant Species in New Guinea Rain Forests, 26 Pogonomyrmex badius, 41, 108 Proisotoma minima, 70 Proistoma minuta, 69 Protoperlaria, 115 Pseudisotoma anamola, 76 Rain Forests, 26 Raphidia Americana, 54 Raphidiidae, 52

Relationship of the Protoperlaria and the Endopterygota, 115 Snake-Flies, 52

Some New West Indian Eumolpid Beetles, 91

Strumigenys dyak, 81 Strumigenys forficata, 86 Strumigenys indag atrix , 86 Strumigenys mjoebergi, 83 Strumigenys sublaminata, 84 Tetracanthella pilosa, 63 Thermobia domestica, 11 Vertagopus aborea, 74 Vertagopus ciliatus, 75

133

PSYCHE

A JOURNAL OF ENTOMOLOGY

Established in 1874

Yol. 65 March, 1958 No. 1

Elizabeth Bangs Bryant. Elisabeth Deichmann ....

Oogenesis and Fertilization in Thermohia domestica (Packard).

J. T. Woodland 11

Patchy Distributions of Ant Species in New Guinea Rain Forests.

E. 0. Wilson ... 26

A Formica Slave-maker Raiding the Nest of a Myrmicine Ant.

W. L. Brown, Jr. . ......... 39

CAMBRIDGE ENTOMOLOGICAL CLUB

Officers for 1957-58

President	•	R.
V ice-President	•	S.
Secretary	•	M.
Treasurer	•	F. ' N.
Executive Committee <	H.

B. Willey, Harvard University Duncan, Boston University C. Parsons, Radcliffe College M. Carpenter, Harvard University Bailey, Bradford Junior College Levi, Museum of Comparative Zoology

at Harvard College

EDITORIAL BOARD OF PSYCHE

F. M. Carpenter (Editor), Professor of Entomology , Harvard University

P. J. Darlington, Jr., Head Curator of Recent Insects , Museum of Comparative Zoology

W. L. Brown, Jr., Associate Curator of Insects , Museum of Comparative Zoology

E. O. Wilson, Associate Professor of Zoology, Harvard University

H. Levi, Associate Curator of Arachnology , Museum of Compara- tive Zoology

PSYCHE is published quarterly, the issues appearing in March, June, September, and December. Subscription price, per year, payable in advance: $3.00 to subscribers in the United States and Canada: other subscriptions $3.35. Single copies, 85 cents.

Cheques and remittances should be addressed to Treasurer, Cambridge Entomological Club, Biological Laboratories, Harvard University. Cambridge, Mass.

Orders for back volumes, missing numbers, notices of change of address, etc., should be sent to the Editorial Office of Psyche, Biological Laboratories, Harvard University, Cambridge, Mass.

IMPORTANT NOTICE TO CONTRIBUTORS

Manuscripts intended for publication, books intended for review, and other editorial matter, should be addressed to Professor F. M. Carpenter, Biological Laboratories, Harvard University, Cambridge, Mass.

Authors contributing articles over 8 printed pages in length will be required to bear a part of the extra expense, for additional pages. This expense will be that

of typesetting onty. which is about S3.U0 per page. The actual cost of preparing cuts for all illustrations must be borne by contributors; the expense for full page plates from line drawings is approximately $8.00 each, and for full page half-tones, $10.00 each; smaller sizes in proportion.

AUTHOR’S SEPARATES

Reprints of articles may be secured by authors, if they are ordered before, or at the time proofs are received for corrections. The cost of these will be furnished by the Editor on application.

The December, 1957, Psyche (Vol. 64, No. 4) was mailed January 26, 1959

EATON PRESS INC., WATERTOWN, MASS.

Psyche, 1958

Vol. 65, Plate 1

Elizabeth Bangs Bryant

April 7, 1875 — January 6, 1953

PSYCHE

Vol. 65 March, 1958 No. 1

ELIZABETH BANGS BRYANT By Elisabeth Deichmann

Museum of Comparative Zoology

It was some years after I had come to Cambridge that I first met Miss Bryant. At that time (1926) the entomolo- gists had their own entrance, and simply going around and making calls was not encouraged. But after Thomas Barbour became director (1928) and after the Biology Department had moved across the street, she and I became neighbors on the fourth floor. She discovered that I made tea for my lunch and suggested that I come in and drink tea with her, and for more than 15 years I usually had lunch with her about three times a week. She must have been about sixty when we first met and she seemed to change very little with the years. She was of medium height, fairly stout, with regal carriage. She had un- wrinkled skin, clear blue eyes and white hair. In some quarters of the museum she was referred to, not unkindly, as “Queen Victoria” and although much taller and with a decidedly retrousse nose, her black dress, snow-white hair and somewhat pendulous cheeks gave her a certain re- semblance to that lady. She is the only person I have known who used the word “twaddle”, and that with as much emphasis as I imagine Queen Victoria did. Behind her sedate exterior she kept a rather youthful spirit. She was extremely well read with a rich vocabulary of old Yankee expressions and after she had used one of these she would suddenly pretend that she was embarassed : “Oh, Miss Deichmann, I really should not teach you such language !”

It is perhaps quite characteristic that I never had the slightest idea of what her father had been, while I was

3

SMITHSONIAN m.y * g «H|

UNSTlTUTiON

4

Psyche

March

extremely well acquainted with the famous cat of her child- hood, so dignified that she and her brother always referred to it as Mister Verdant Green. She belonged to a good substantial Boston family and seemed to be related to a large number of prominent persons, which did not prevent her from expressing her, not always high, opinion of these relatives. She had evidently had an extremely happy child- hood and a youth filled with trips to Europe and the typical Bostonian’s cultural interests. The family had even been so progressive that they had allowed her to go to Radcliffe, which definitely was the great adventure of her life, as for all women of her generation who suddenly were allowed to get the same education as their brothers. She belonged to the class of 1897 but she did not graduate. There may have been illness in her family, or it may have been just at that time that her family, as well as many other sub- stantial Bostonians, were rather hard hit financially. Her interests were with natural history and she became early acquainted with the men in the Boston Natural History Museum and those connected with the Agassiz Museum, and in the latter institute she was soon given the little division of spiders to take care of. In her younger days she was an eager field worker: there was a widespread interest in nature study in Boston and she took part in many botanical and zoological excursions and built up quite a collection of New England spiders. Her father died rather early and after that time she took care of her mother for many years, while she regularly appeared in the Agassiz Museum three times a week, but her outdoor activities became quite naturally restricted and her circle of acquaint- ances limited to men much older than herself, Mr. Samuel Henshaw, Mr. Faxon and especially Mr. J. H. Emerton, who was delighted to see a younger person take over in the spider field.

In the museum she had been fortunate to be allotted a small division. With the myriapods and mites in two neighboring rooms, she was able to have all the spiders around her. She learned to type and besides she wrote hundreds of cards and labels in her precise, clear hand. Her reprint collection was well cared for, with binders which

1958

Deichmann — Elizabeth Bangs Bryant

5

she bought herself. Her first paper came in 1908; it was merely a list of local species, but it had probably taken some persuasion from her good friend, Mr. Emerton, to let her to allow her name to appear in print. It was 15 years later that her Barbados-Antigua report was published, and then, around 1930, she really began to be a regular contributor. Very likely it was the stimulus she received when Dr. Barbour saw to it that this volunteer worker of almost 30 years standing at last received a small salary. She stead- fastly refused to be listed in American Men of Science, feeling that that was to intrude into the ranks of profes- sionally trained, and no argument could change her position on that point. It was a great joy to her when other arach- nologists visited her and she was always helpful to beginners who came to seek her advice. When somebody brought in a spider she would tell all about its habits and occurrence, and she never crushed the collector with a remark that it was one of the most common forms. “It is such a long time since I have had occasion to see a live specimen of this spider,” — so the person went off feeling that his efforts had been fully appreciated.

Her main scientific work was not started until she was 55 years old, and continued to her death. Several large papers on West Indian spiders were illustrated by her clean pen drawings. She was handicapped in her work by an inadequate microscope and light (a modified automobile spotlight). She worked independently of all but a few col- leagues in arachnology.

For several years her mother’s health was failing and she looked after her with unswerving fidelity. After her mother’s death she moved from the house in Allston to a pleasant top floor apartment in Brookline. It looked as if she was going to have more freedom and our long planned trip to St. Lucia in the West Indies, the type locality for so many of the West Indian spiders, seemed near to becom- ing reality, when the old housekeeper’s health gave out and Miss Bryant undertook to care for her, most scornful of the idea that the housekeeper should be put in a Nursing Home.

There was one unusual activity in which Miss Bryant

6

Psyche

March

indulged, and that was taking care of her investments. After her father’s death, her mother, mindful of the debacle of some years before, made arrangements, so that she got a fixed income and divided the family fortune between the two children. That was in the happy days before the income tax had been invented. The usual course would have been that Miss Bryant’s share be nut in a trust fund, but here her old friend, Mr. Henshaw, intervened. She took, as always, his advice and developed into an extremely shrewd and careful investor. Through her interest in her invest- ments, by reading newspapers and magazines, she acquired an unusual understanding of what was going on in the United States and in the world as a whole, and her down to earth realism and a total lack of sentimentality made conversations with her extremely interesting.

In addition to her indisputable business ability, which would have made her a gift to a brokerage firm, she pos- sessed also the virtue, thrift. She saw to it that nothing was wasted in her house, got the utmost wear out of her few garments and she kept all unnecessary expenditures down to zero. She subscribed to a few magazines which she knew she could manage to read, and the back numbers were quickly passed on to some other person for whom she was happy to save the cost of a subscription. While despis- ing people who made themselves miserable by being “penur- ious” she enjoyed her own little pet economies. For years she would happily trot down a few blocks so that she could get home on the 5 cent fare, and until her last illness it was our monthly joke when she handed me her check for the Faculty Club and asked me to take it over to Harvard Square when I was paying my own bill, adding with mock seriousness: “I just can’t hear spending 3 cents on such a short distance, and the bus no longer sets me off in Harvard Square.”

She felt her responsibility toward the needy, particularly children and old people, and she insisted on knowing where the money went. She saw to it that nothing was lost between her hand and that of the recipient. She lived for years in a rather poor district where as she once expressed it: “They are counting on my contribution” — and they got it. At

1958

Deichmann — Elizabeth Bangs Bryant

7

one time an over eager Community Chest agent tried to persuade her to give her large contribution through the Museum instead of just the token which she usually gave. But she was adamant, her contribution was going to where it had been going for years, long before the Community Chest had been established. The argument became rather heated: “It was her duty toward the Museum.” Then Miss Bryant got up: “My resignation shall be on the Director’s desk this afternoon!” The subject was quickly dropped.

When she had been fifty years out of college she received an honorary Phi Beta Kappa membership from Radcliffe as the one of her class who had made the most out of her education, and this was a great pleasure to her. But she appreciated it even more when, at about the same time, she was taken into the Radcliffe Sigma Xi, for this enabled her to get into contact with young people, a pleasure which her duties toward her mother had prevented for years. Contrary to her custom of never going out in the evening, she would attend all their evening meetings.

She continued working after her retirement in 1950, it seemed almost with even greater vigor than before. Several years before that time the difficulties the museum had, and still would have for years, regarding decent pay to the curators were clearly seen by us all; she had also seen how the Boston Natural History Museum had packed several collections aside and had given others away to where they could be used, and she realized that a similar fate might well befall the collections which she was in charge of. Sud- denly she realized that she was able to act so that this should not happen. It was with deep emotion that she one day came to me and told me that she had radically altered her will so that the work could go on after her death and she was a changed person from that moment. Her plans for her work became bolder and she decided to get a new and better microscope even at this late time of her life.

It was when I came home from a trip in the fall of 1952 that she informed me that she was not feeling well and that an operation was necessary because of stomach cancer. After the operation I saw her two or three times a week at her home and our conversations went on almost as if she

8

Psyche

March

were still in her office in the Museum. She admitted she was feeling oddly “lighthearted” and there were a few moments when she almost believed that she might get well enough to come back for a short time and do a little work. But when her new microscope at last arrived she sent it straight back to the Museum. At first she was well enough to get up and come out and wave goodby to me on the stair case, later she asked to be excused. The end came on January 6, 1953.

Her will was a model of careful consideration of the needy in the community and her family. Two of the larger bequests, of equal amounts, went to Radcliffe College, which had given her her education, and to the Museum, where she had been able to make use of it. With her usual reticence she had in the latter case succeeded in keeping her name hidden. The bequest was simply named the “Emerton Fund” in honor of the old arachnologist who had taught her to draw and encouraged her in her work.

There were also two small bequests each of 500 dollars to the two Radcliffe Honor societies. In the case of Phi Beta Kappa the money helped to hasten the completion of the $10,000 Scholarship Fund which this old and fairly wealthy chapter had been working on, and her name was duly added to the Memorial Roll. In the case of the much younger, smaller, and anything but wealthy Chapter of Sigma Xi, the sum, with interest and smaller gifts added, was some years later voted to be used as the nucleus of a much needed loan for Radcliffe science students, and was named the “Elizabeth Bangs Bryant Loan Fund of Sigma Xi.”

The following is the known list of her publications:

Bibliography of Elizabeth Bangs Bryant

1908 List of Araneina in Fauna of New England, 9. Occ. Pap. Boston Soc. Nat. Hist., vol. 7, pp. 1-105.

1923 Report on the spiders collected by the Barbados- Antigua Expedition from the University of Iowa in 1918. Univ. Iowa Stud, in Nat. Hist., vol. 10, pp. 10-16.

1958] Deichmann — Elizabeth Bangs Bryant 9

1930 A revision of the American species of the genus Ozyptila. Psyche, vol. 37, pp. 375-391.

New species of the genus Xysticus, (Arachnida). Ibid., vol. 37, pp. 132-140.

1931 Notes on North American Anyphaeninae in the Museum of Comparative Zoology. Ibid., vol. 38, pp. 102-126.

1933 New and little known spiders from the United States. Bull. Mus. Comp. Zool., vol. 74, pp. 171-193.

Notes on types of Urquhart’s spiders. Rec. Canter- bury Mus., vol. 4, pp. 1-27.

1934 New Lycosidae from Florida. Psyche, vol. 41, pp. 38-41. '

1935 Notes on some of Urquhart’s species of spiders. Rec. Canterbury Mus., vol. 4, pp. 53-70.

Some new and little known species of New Zealand spiders. Ibid. vol. 4, pp. 71-94.

A few southern spiders, Psyche, vol. 42, pp. 73-83. A rare spider. Ibicl,. vol. 42, pp. 163-166.

1936 New species of southern spiders. Ibid., vol. 43, pp. 87-100.

Descriptions of some new species of Cuban spiders. Mem. Soc. Cubana Hist. Nat., vol. 10, pp. 325-332.

1938 Spiders in the Museum of Comparative Zoology, Harvard Alumni Bull., Feb. 25.

1940 Notes on Epeira pentagons Hentz. Psyche, vol. 47, pp. 60-65 (with A. F. Archer).

Cuban spiders in the Museum of Comparative Zoology. Bull. Mus. Comp. Zool., vol. 86, pp. 247-532.

1941 Notes on the spider fauna of New England. Psyche, vol. 48, pp. 129-146.

1942 Notes on the spiders of the Virgin Islands. Bull. Mus. Comp. Zool., vol. 89, pp. 317-363.

Additions to the spider fauna of Puerto Rico. Jour. Agr. Univ. Puerto Rico, vol. 26, pp. 1-19.

Descriptions of certain North American Phidippus (Araneae) . Amer. Midland Nat., vol. 28, pp. 693-707.

10 Psyche [March

1943 The salticid spiders of Hispaniola. Bull. Mus. Comp. Zool., vol. 92, pp. 443-522.

Notes on Dictyolathys maculata Banks (Araneae: Dictynidae) . Psyche, vol. 50, pp. 83-86.

1944 Three species of Coleosoma from Florida (Araneae: Theridiidae). Ibid., vol. 51, pp. 51-58.

1945 The Argiopidae of Hispaniola. Bull. Mus. Comp. Zool., vol 95, pp. 357-418.

Notes on some Florida spiders. Trans. Connecticut Acad. Sci., vol. 36, pp. 199-213.

Some new and little known southern spiders, Psyche, vol. 52, pp. 178-192.

1946 The genotype of Mimetus Hentz. Ibid., vol. 53, p. 48.

1947 A list of spiders from Mona Island with descriptions of new and little known species. Ibid., vol. 54, pp. 86-89.

Notes on spiders from Puerto Rico. Ibid., vol. 54, pp. 183-193.

1948 The spiders of Hispaniola. Bull. Mus. Comp. Zool., vol. 100, pp. 329-447.

Some spiders from Acapulco, Mexico. Psyche, vol. 55, pp. 55-77.

1949 The male of Prodidomus rufus Hentz. (Prodidomi- dae, Araneae) . Ibid., vol. 56, pp. 22-25.

A new genus and species of Theridiidae from eastern Texas (Araneae). Ibid., vol. 56, pp. 66-69.

The “Iris spider” of Kent Island. Bull. Ninth Annual Rep., Bowdoin Sci. Sta. no. 11, pp. 14-15 (mimeo.). Acanthepeira venusta (Banks) (Araneae). Psyche, vol. 56, pp. 175-179.

1950 The salticid spiders of Jamaica. Bull. Mus. Comp. Zool., vol. 103, pp. 161-209.

1951 Redescription of Cheiracanthium mildei L. Koch, a recent spider immigrant from Europe. Psyche, vol. 58, pp. 120-123.

OOGENESIS AND FERTILIZATION IN THERMOBIA DOMESTIC A (PACKARD)1

By John T. Woodland

State Teachers College, Salem, Massachusetts

The materials and methods used in this study were the same as those described in an earlier paper (Woodland, 1957).

Oogenesis

The primary oocytes in the vitellarium reach a length of about 420 microns just before yolk accumulation begins. They measure only about 45 microns wide. Each has a large central nucleus about 40 microns in diameter. It is a typical vesicular oocyte nucleus and usually has a visible nuclear membrane. A fine network is visible throughout the nucleus and there is a large, eccentric, irregular, granu- lar nucleolus which may be as long as 20 microns. Neither network nor nucleolus is Feulgen-positive. The finely- divided chromatin is so scattered throughout the large nucleus that it is barely visible. The cytoplasm appears very finely granular and contains a few inclusions. These are 10 to 15 microns in diameter and consist of a dozen or less clumped globules.

When each of the vitellaria contains four or five of these oocytes, yolk accumulation begins. It starts peripherally in all of the oocytes simultaneously and gradually proceeds toward the center. There appear scattered through the peripheral cytoplasm tiny globules which stain bright orange-red with Mallory’s triple stain. They greatly re- semble the globules of protein reserves scattered through the fat body surrounding the ovarioles, but are usually a little smaller. The negative images of small fat droplets also occur in the peripheral cytoplasm. The fat droplets and proteinaceous globules increase in size, forcing the diminishing cytoplasm into a network around them. By

!Tlie research was carried out at the Harvard Biological Laboratories, Cambridge, Massachusetts, in partial fulfillment of the degree of Doctor of Philosophy.

11

12

Psyche

March

the time that small fat droplets begin to appear in the center of the oocyte, the peripheral droplets and protein- aceous globules have greatly enlarged (3 to 7 microns in diameter) and are present in about equal numbers. The diminishing central layer of cytoplasm around the nucleus is connected by the fine network between the yolk globules to a thin peripheral cytoplasmic layer, the periplasm. By the time the perinuclear cytoplasm has become very thin, the fat droplets appear to have coalesced, for the visible proteinaceous yolk globules, some of which are now as large as 11 microns in diameter, stand out in fixed preparations against a clear background.

The oocyte has been growing only slightly meanwhile, and is still very narrow in proportion to its length. Before oviposition the maximum length of the egg increases by about two and one-half times, and its maximum width increases about eighteen times. Growth is accomplished chiefly by an increase in the proteinaceous yolk. Just before oviposition the egg contains relatively little fatty yolk.

At the start of the growth period the nucleus moves to the periphery. The nuclear network condenses at the peri- phery of the nucleus and the chromatin becomes distinctly visible with the Feulgen technique. Evidence of tetrads is seen. The interior of the large nucleus now appears homo- geneous and has little affinity for the stains used. Before oviposition there occur both a marked decrease in nuclear volume and further condensation of the chromatin into small compact chromosomes.

The follicle cells start to secrete the endochorion at the time when yolk accumulation first begins. The process is best studied in preparations stained with phosphotungstic hematoxylin. Just before secretion of the endochorion starts, the brown-staining connective tissue sheath sur- rounding the ovariole becomes greatly thickened. It ap- pears that the follicle cells withdraw material from the sheath and secrete it to form the endochorion. Comparison with the accompanying process of yolk accumulation indi- cates that the sheath is reduced to its former size within a relatively short time. The secretion of the exochorion, which occurs much later, is mentioned below.

1958] Woodland — Thermobia domestica (Packard) 13

The external appearance of the newly laid egg of the firebrat has been described by Adams (1933), Sweetman (1938), and Remington (1948). The present study reveals that the endochorion (Fig. 5, EN) is about 2 microns thick. Its thickness does not vary appreciably over the whole egg. There adheres to its outer surface a single layer of spherical droplets, each about 11 microns in diameter immediately after oviposition. The droplets are rather irregularly placed, but tend to be in groups with large intervening spaces. In the spaces are droplets many times smaller than the large ones. Within a few hours a change occurs. The large droplets break up into rather evenly spaced ones about 3 microns in diameter (Fig. 5, CD). Usually a space of 2 microns now occurs between droplets, although pairs of contiguous droplets are sometimes seen. The very minute droplets are still scattered among the larger ones. The appearance of the droplets is essentially the same on shed chorions long after the nymphs have left them. Possibly the droplets are present as a thin continuous film before oviposition.

The droplets adhere firmly to the exochorion. A few individual droplets are separated from it by sectioning, but the other procedures to which the eggs have been subjected practically never dislodge them. In contrast, the exochorion is loosened from the endochorion when the egg contents shrink during fixation, owing to dissolution of the fatty elements. Moreover, the exochorion is normally loosened from the endochorion during development, as the volume of the egg contents is diminished slightly. The surface of the endochorion is then seen to be reticulated into hundreds of small polygonal areas. Most of these polygons are fairly regular hexagons with diagonals usually between 55 and 85 microns long. The wall separating adjacent areas is 5 microns thick.

Sweetman (1938) supposed this hexagonal reticulation to be produced by the cells of the embryonic tissues, but the present study showed it to be produced by the follicular cells that secreted the endochorion. The large size of the hexagons emphasizes the amount the egg has grown since the endochorion was secreted.

14

Psyche

March

The endochorion is hardly ever loosened at all from the yolk and no vitelline membrane has been found between them. Heymons (1897) reported a vitelline membrane in Lepisma saccharina, but Uzel (1898), who studied the eggs of the same lepismatid, did not mention it.

Between the exochorion and the endochorion at the an- terior end of the egg is the micropylar area, a circular thickening about 280 microns in diameter (Fig. 5). The thickening consists of as many as twenty or more concentric lamellae, each about as thick as the exochorion, with which the thickening is identical in staining reactions and to which it adheres if the latter becomes loosened from the endo- chorion. The thickening is thinner peripherally than cen- trally, since not all of the lamellae extend to the margin of the area.

In the micropylar area are found three small infundibular depressions which, if connected, would form the corners of an equilateral triangle. Two of the depressions are shown in figure 5 (DP). The depressions, which can rarely be demonstrated to penetrate the micropylar thickening com- pletely, are filled with folds of the exochorion and with the chorionic droplets described above. There sometimes ap-

Explanation of Plate 2

All figures represent portions of sections. Fixative: Maximow’s. Stain: Feulgen’s all figs, except 2 and 5; Mallory’s triple, figs. 2 and 5. Magnifi- cations approximate. Fig. 1. 20 minute old egg, showing sperm head ac- cumulating cytoplasm, 400 x. Fig. 2. 30 minute old egg, showing egg nucleus in anaphase of first maturation division, 500 x. Fig. 3. 2% hour old egg, showing egg nucleus in anaphase of second maturation division, 500 x. Fig. 4. 75 minute old egg, showing late telophase of first maturation divi- sion; secondary oocyte nucleus forming in upper left, first polar body in lower right; 500 x. Fig. 5. Longitudinal section through micropylar (an- terior) end of 2 hour old egg, 160 x; CD, chorionic droplet; DP, depres- sion; EN, endochorion; EX, exochorion; EXT, exochorionic thickening. Fig. 6. 90 minute old egg, showing late stage in the contraction of the sperm head, 1250 x. Fig. 7. Egg about 2 hours old, showing male pronuc- leus during its growth period; 1875 x. Figs. 8-11. 3 to 4 hour old eggs, 1250 x. Fig. 8. Early stage of union of male and female pronuclei. Fig. 9. Female pronucleus shown near top just enteriing the sperm plasm; male pronucleus (below). Fig. 10. Female pronucleus (below) and male pronucleus (above) about to unite. Fig. 11. Late stage of union of male and female pronuclei.

Psyche, 1958

Vol. 65, Plate 2

Woodland

Thermobia domestica

16

Psyche

March

pears to be an opening through the exochorion where it dips into a depression (Fig. 5), but no opening through the endochorion was found. Other evidence suggests that there is none. Eggs completely immersed in tap water at 37.5° C. for a period of 3 or 4 days during incubation hatch into healthy nymphs in about the normal time. (Hatching itself occurs under water, but the nymphs cannot get their abdomens out of the shell. This is because they cannot get traction and also occurs in a small percentage of eggs hatching in an empty glass container.) If openings com- pletely penetrated the chorion, then water should be ex- pected to enter the egg by osmotic pressure and cause it to burst. If eggs are kept in tap water for as long as seven days, the incubation period is prolonged by several days and only about half of the eggs hatch at all. Those not hatching do not burst nor are they turgid. Presumably they die from oxygen lack, as the water was not mechanical- ly oxygenated.

The follicle cells appear to become a syncytium during the enlargement of the oocyte. This syncytium probably secretes the micropylar thickening and the thin structure- less exochorion just before the oocyte leaves the ovariole. The chorionic droplets are believed to be added as the egg passes through the lateral oviduct, for droplets of similar appearance have been seen in the cells of the oviducal epi- thelium of females with large oocytes in the ovarioles.

Fertilization

After the oocyte leaves the gonopore but before it pro- ceeds down the ovipositor, it is undoubtedly held at the base of the ovipositor for a few moments while sperms are dis- charged onto it by the spermatheca. The sperms presum- ably enter the probable openings in the exochorion over the depressions in the micropylar thickening. They can appar- ently penetrate the endochorion directly. They would not have to penetrate the endochorion directly under the micro- pylar thickening, but could pass between the lamellae of the thickening and enter nearer the equator of the egg. Whether the entire sperm penetrates the egg was not deter- mined. Within the freshly laid egg, however, may usually be identified from one to six of the very long sperm heads,

1958] Woodland — Thermobia domestica (Packard) 17

each of which is bent and coiled upon itself many times.

The contents of the living, freshly laid egg appear to consist of colorless yolk spheres in a colorless liquid of low viscosity. The spheres are about 10 to 15 microns in diameter. In sections of eggs fixed with Maximow’s fluid the negative images of the fat spheres measure 9 or 10 microns in diameter. Most of the more numerous, visible, proteinaceous yolk globules are 14 or 15 microns in diameter. Some of these globules appear structureless, while others appear finely granular. The globules appear only very slight- ly flattened at the poles with Maximow’s fixative. The other fixatives used always badly distorted the yolk of young eggs. The yolk spheres and liquid fill the entire egg. Cyto- plasm was not identified.

Perrot (1933) stated that the first maturation division occurs while the oocyte is still in the ovariole. He was not able to find any trace of the prophase of the division. He reported that the mitotic figure occupies a very small space at the periphery of the oocyte and figured the anaphase of the division. He stated further that after this first matura- tion division the nucleus enters a resting stage which he figured from a freshly laid egg. The present study did not confirm this part of Perrot’s work. The only mitotic figures ever found in sections through the vitellarium were those of the follicle cells. The anaphase of such follicular mitoses often resembles that shown in the first of Perrot’s figures just mentioned. Eggs fixed immediately after oviposition show the nucleus to be in the metaphase of the first matura- tion division. No nucleus resembling both in appearance and position that shown in the second of Perrot’s figures was found until the beginning of the formation of the primary epithelium.

The anastral type of meiosis occurs. The mitotic figure is located more or less equidistant from the two poles