I J 5P30. 573 ZOOLOGICA SCIENTIFIC CONTRIBUTIONS of the NEW YORK ZOOLOGICAL SOCIETY VOLUME 35 1950 Numbers 1-20 Published by the Society The Zoological Park, New York CONTENTS Part 1. April 17, 1950. PAGE 1. Proceedings of the Second Conference on the Biology of Normal and Atypical Pigment Cell Growth. Abstracts of 23 Papers 1 2. The Tipulidae (Order Diptera) of Rancho Grande, North-central Venezuela. By Charles P. Alexander. Text-figures 1-39 33 3. Migration of Danaidae, Ithomiidae, Acraeidae and Heliconidae (Butterflies) at Rancho Grande, North-central Venezuela. By William Beebe. Text-figure 1 57 4. Home Life of the Bat Falcon, Falco albigularis albigularis Daudin. By William Beebe 69 5. The Vampryomorpha (Cephalopoda) of the Bermuda Oceanographic Expedi- tions. By Grace E. Pickford. Text-figures 1-8 87 Part 2. August 1, 1950. 6. Some Variations in Grouping and Dominance Patterns Among Birds and Mam- mals. By N. E. Collias. Text-figure 1 97 7. Fish that Live as Inquilines (Lodgers) in Sponges. By E. W. Gudger. Text- figures 1&2 121 8. Five New Races of Birds from Southeastern Asia. By H. G. Deignan 127 9. A Survey of Pasteurella tularensis Infection in the Animals of the Jackson Hole Area, by Mitsuru Nakamura 129 10. Neotropical Chilopods and Diplopods in the Collections of the Department of Tropical Research, New York Zoological Society. By Ralph V. Chamberlin. Text-figures 1-23 133 11. The Relative Position of the Cetacea Among the Orders of Mammalia as Indi- cated by Precipitin Tests. By Alan Boyden & Douglas Gemeroy. Text-figure 1 145 Part 3. November 27, 1950 12. Factors Influencing the Establishment of Residence in Shells by Tropical Shore Fishes. By C. M. Breder, Jr. Plates I & II; Text-figures 1-5 153 13. Notes on the Behavior and Morphology of Some West Indian Plectognath Fishes. By Eugenie Clark. Plates I & II ; Text-figures 1-7 159 14. The Effect of Alloxan on the Pancreas, Liver and Kidney of the Teleost, Lebistes reticulatus, with Notes on the Normal Pancreas. By Leonard L. Grosso. Plates I-IV '. 169 15. An Ecological Study of Helminths of Some Wyoming Voles ( Microtus spp.) with a Description of a New Species of Nematospiroides (Heligmosomidae: Nematoda). By Merle L. Kuns & Robert Rausch. Text-figui’es 1-7 181 16. Migration of Pieridae (Butterflies) Through Portachuelo Pass, Rancho Grande, North-central Venezuela. By William Beebe. Plate 1 189 17. Moths New to Trinidad, B.W.I. By Norman Lamont & E. McC. Callan. Edited by William Beebe 197 18. The Euchromiidae (Moths) of Kartabo, British Guiana, and Caripito, Venezuela. By Henry Fleming 209 Part 4. December 30, 1950. 19. Eastern Pacific Expeditions of the New York Zoological Society. XLII. Mollusks from the West Coast of Mexico and Central America. Part IX. By Leo George Hertlein & A. M. Strong. Plates I & II 217 20. Comparative Biology of Salticid Spiders at Rancho Grande, Venezuela. Part V. Postembryological Development of Color and Pattern. By Jocelyn Crane. . . 253 Index to Volume 35 263 ZOOLOGICA SCIENTIFIC CONTRIBUTIONS of the NEW YORK ZOOLOGICAL SOCIETY VOLUME 35 Part 1 Numbers 1-5 Published by the Society The Zoological Park, New York April 17, 1950 CONTENTS Page 1. Proceedings of the Second Conference on the Biology of Normal and Atypical Pigment Cell Growth. Abstracts of 23 Papers 1 2. The Tipulidae (Order Diptera) of Rancho Grande, North-central Venezuela. By Charles P. Alexander. Text-figures 1-39......... 33 3. Migration of Danaidae, Ithomiidae, Acraeidae and Heliconidae (Butterflies) at Rancho Grande, North-central Venezuela. By William Beebe. Text-figure 1 57 4. Home Life of the Bat Falcon, Falco albigularis albigularis Daudin. By William Beebe 69 5. The Vampyromorpha (Cephalopoda) of the Bermuda Oceano- graphic Expeditions. By Grace E. Pickford. Text-figures 1-8. 87 Abstracts : Pigment Cell Conference 1 1. Proceedings of the Second Conference on the Biology of Normal and Atypical Pigment Cell Growth. Foreword. The second conference on the Biology of Normal and Atypical Pigment Cell Growth was called in direct response to the requests of a group of biologists, biochemists, bio- physicists and medical research workers who wanted another opportunity for an exchange of ideas between specialists in different fields. They wanted to learn firsthand the results of recent experiments and observa- tions. The second conference was held at the New York Zoological Park on November 18- 19, 1949. The publication of “The Biology of Mela- nomas” by the New York Academy of Sci- ences, which sponsored the first conference in 1946, established Pigment Cell Growth as a united topic and brought its subject matter into focus and prominence. The many new contributors to the second conference testify to the growth of interest in the nature and behavior of the pigment cells. Among these contributors are some whose researches in the field go back a quarter of a century. The pool of information has been greatly enriched and the present publica- tion will make it accessible to an ever- widening group of investigators. The abstracts published here differ some- what from those originally submitted for distribution prior to the conference. They have been expanded and contain references to the most recent papers dealing with pig- ment cell growth and related topics. This report also includes discussions and the in- troductory remarks made by the chairmen of the four sessions of the conference. The next conference will be organized in response to the future needs of investigators in the broad field of pigment cell growth. Suggestions and inquiries will be welcomed at all times by the organizing chairman. Acknowledgments. The conference is indebted to Dr. George M. Smith of the Yale University School of Medicine for his continuing interest in these meetings. As Medical Advisor to the Anna Fuller Fund, he recommended that a grant be made to the New York Zoological Society to defray part of the costs of arranging and conducting the conference. In this connec- tion, the organizing chairman wishes to ac- knowledge the many valuable suggestions made by Dr. Smith that were put in action before and during the meeting. The thanks of the conference are also due to the New York Zoological Society for a grant in aid of publication of these abstracts. — Myron Gordon, Genetics Laboratory of the New York Zoological Society, the American Mu- seum of Natural History, New York 24, N . Y. &PR 2 4 I960 2 Zoologica: New York Zoological Society [35: 1 Abstracts. Table of Contents. Page First Session: Clinical and Pathological. Introduction. Harry S. N. Greene 3 Malignant Melanoma Associated with Pregnancy. Isabel M. Scharnagel & George T. Pack 4 Benign and Malignant Neoplasia of Mel- anoblasts Through the Eyes of the Der- matologist. S. William Becker 5 The Problem of Acanthosis Nigricans. Helen Ollendorff Curth 7 The Junction Nevus, Forerunner of Malig- nant Melanoma and Its Differential Diagnosis from the Standpoint of the Dermatologist. Eugene F. Traub 8 The Development of Epidermal Pigmenta- tion in the Negro Fetus. Arnold A. Zim- mermann 10 A Comparative Study of Malignant Mel- anoma Among Negro and White Pa- tients. Rudolph J. Muelling, Jr. & Wal- ter J. Burdette 12 Second Session: Genetical. Introduction. W. E. Heston 13 Significance of Quantitative Histological Studies of Pigment Found in the Coat Color Mutants of the Mouse to Ques- tions of Normal and Atypical Cell Growth. Elizabeth Shull Russell. ...... 13 Melanotic Tumors in Drosophila. Walter J. Burdette 15 Genetic Pigment Mosaics in the Pigeon. W. F. Hollander 16 Pigment Formation in the Eye of Ephestia and Its Genic Determination. Ernst Caspari 17 Problems of Origin and Migration of Pig- ment Cells in Fish. H. B. Goodrich. ... 17 The Origin of Modifying Genes that In- fluence the Normal and Atypical Page Growth of Pigment Cells in Fishes. Myron Gordon 19 Third Session: Physiological. Introduction. H. W. Chalkley 20 Pigment Spread in Guinea Pigs. P. B. Medawar 21 Atypical Pigment Cell Differentiation in Embryonic Teleostean Grafts and Iso- lates. Jane M. Oppenheimer 22 Regeneration of Neural Retina and Lens from Pigment Cells in the Eyes of Adult Salamanders. L. S. Stone 23 X-ray Effects on Mouse Pigmentation as Related to Melanoblast Distribution. Herman B. Chase & Virginia W. Smith. 24 A Comparative Colorimetric Study of Dopa-melanin Formation by Melanomas and Pigmented Skins. Ruth E. Shrader & Carroll A. Pfeiffer 25 Fourth Sessioji : Biochemical and Biophysical. Introduction. Jesse P. Greenstein 26 Mammalian Melanin Formation. I. Bio- chemical Studies. A. Bunsen Lerner & Thomas B. Fitzpatrick 27 Mammalian Melanin Formation. II. Histo- chemical Studies. Thomas B. Fitzpatrick & A. Bunsen Lerner 28 Observations on the Structure, Deriva- tion and Nature of Melanin. E. Meirow - sky, L. W. Freeman & R. B. Fischer. . . 29 Evidence for the Mitochondrial Nature and Function of Melanin Granules. Mark Woods, Herman DuBuy & Dean Burk 30 Oxidative Activities of Mouse Melanomas with Reference to Melanization. Marie L. Hesselbach, Mark Woods & Dean Burk 31 Chromatographic Separation of Melanin Granules. Vernon T. Riley, Mark Woods & Dean Burk 32 1950] Abstracts : Pigment Cell Conference 3 First Session: Clinical and Pathological. Introduction. Harry S. N. Greene. Yale University School of Medicine, New Haven, Conn. With increasing knowledge of the natural histories of tumors, it has become more and more apparent that there are many neo- plastic diseases rather than one. Conferences for the separate consideration of specific tumor types are now the fashion and in no instance is this more fully justified than in the case of the melanomas. The well marked peculiarities of such tumors, with their fail- ure to conform to the more or less conven- tional lines of behavior established by the more orthodox carcinomas and sarcomas, set them apart as black sheep of the neoplastic family and engender the necessity for in- dividual study and special examination. The source of the neoplastic cell, its de- velopment to biological autonomy and mode of dissemination, constitute problems differ- ing in their fundamental conditions from those associated with other tumors. For- tunately, the differing conditions tend to facilitate investigation and, by their anomo- lous nature, may suggest fertile approaches to the general cancer problem. The occur- rence of melanoblasts in heterotopic aggre- gates called nevi, presents an unparalleled situation in neoplastic disease, for here, assembled by some dysontogenetic process in convenient, lasting foci, are the cells in- trinsic to the disease. By virtue of location in readily accessible, non-vital regions, the conversion to neoplastic elements and the development of the properties of invasibility and metastasizability are open to morpho- logical and biological investigation as well as to clinical study. Further, the indiscrimi- nate incidence of nevi with occurence in dif- ferent locations, sexes and races, and their persistence throughout the different age periods and endocrine disturbances of an individual’s life, allow an assessment of the significance of constitutional, intrinsic fac- tors in the biological progress of the lesion. One of the more marked variations from the behavior pattern of other tumors con- cerns the dissemination of the melanomas. The mode of spread in general neoplastic disease is by lymphatic extension or metas- 1 tasis and it is customary to explain the occurrence of secondary foci of melanomas as a result of one of these processes. There is no question that fully developed, autono- mous melanomas follow such routes of spread, but there is both clinical and experi- mental evidence to suggest that secondary foci may occur before the primary tumor attains the ability to metastasize. For ex- ample, when a secondary focus is found as far distant from a primary tumor in the scalp as a popliteal node, it is generally con- sidered to be a blood-borne metastasis. Yet, if such a node is removed, the patient may survive for a number of years with no fur- ther evidence of tumor spread. Inasmuch as blood-borne metastasis entails the entrance of tumor cells into the blood stream with eventual lodgement and growth in multiple foci, it would appear either that the popliteal node represented the rare phenomenon of solitary metastasis or that in actuality it was not a metastasis at all. Evidence favor- ing this latter possibility is obtained from transplantation experiments. It appears to be a rule that all tumors capable of metas- tasis (and therefore all metastases) are hetero-transplantable ; yet, it has been a frequent experience that nodes of the type cited above fail to grow when transferred to guinea pigs. Successful transplantation portends death, the survival period after heterologous growth averaging six months. It must be concluded from this either that the rule of transplantability applicable to other tumors does not hold in the case of melanomas, or that a different mechanism of dissemination exists. The point should not be labored in the present context and is mentioned only to emphasize an unex- plored possibility of potential significance to an understanding of the tumors to be dis- cussed. The problems presented by the behavior of the melanotic tumors pertain to many different disciplines. The embryologist, the biochemist, the geneticist, the endocrinolo- gist, the clinician, and even the pathologist, recognize fields of interest, and a definite hazard exists that the fruits of such interest may be limited by scope or publication to the individual specialty. It would appear to be a primary purpose of this conference to counteract the biased inclination of special interests and to emphasize that while the melanoma provides an excellent tool for the biochemist to study biochemistry; the em- bryologist, embryology; the geneticist, ge- netics, etc., the central problem concerns the melanoma as a malignant tumor. 4 Zoologica: New York Zoological Society [35: 1 Malignant Melanoma Associated with Pregnancy. Isabel M. Scharnagel & George T. Pack. Memorial Hospital, New York. More than a thousand cases of malignant melanoma have been observed and treated in the Memorial Hospital and in other hospitals by the staff of the Pack Medical Group dur- ing the past thirty-two years (since 1917). The primary melanomas are located on all parts of the body integument, including the skin, palms of hands, soles of feet and nail beds. In addition, there ai’e malignant mel- anomas involving the paranasal sinuses, oral mucosa, anus and rectum, meninges, and choroid of the eye. We have been especially interested in studying the hormonal influences on pig- mented tumors. The development of pig- mented nevi at puberty is well known, and the peculiar benign nature of pre-pubertal or juvenile melanomas in young children, in spite of their malignant histological struc- ture, has been made a subject of special study. , Pregnancy is another period of altered hormone balance when many temporary and permanent pigmentations of the skin de- velop. There is often growth of a pre-existing pigmented nevus, fortunately not always to a malignant melanoma, as we have proved by excising them for histologic study. In 28 of our patients with malignant melanoma, how- ever, the melanoma has seemed to develop in a pre-existing nevus during pregnancy, or else a pregnancy has come in a patient with malignant melanoma and has hastened the growth. Eight of our 28 patients with malig- nant melanoma gave history of the growth of a pre-existing pigmented tumor during preg- nancy. Ten others presented themselves for treatment during the pregnancy with the malignant melanoma present and in some stage of development. We both remember very graphically a young 28-year-old woman who very early in the course of her disease presented a picture of a very advanced melanomatosis, showing multiple dark blue-black cutaneous nodules in the region of the right chest wall, shoulder, arm and back. This young woman was in the fifth month of her pregnancy when she no- ticed the small lesion on the back in the right scapular region, which became red and sore. She visited a local hospital about a month later, where it was removed by a small exci- sion, leaving a scar 3 cm. long. About four weeks later it grew back in the scar and she called attention to it when reporting for pre- natal care. It was again removed with a skin segment about twice as big as the first. When she reached the eighth month of gestation, the pregnancy was interrupted by bag induc- tion. A normal five-pound baby was delivered spontaneously. About a month later, a radical right axillary dissection was done and the specimen showed extensive metastases in the lymph nodes. Four months later, and eight months after the original excision, the mul- tiple skin nodules appeared and the skin of the face and chest had a bluish cast. There were numerous shiny blue-black nodules and marked melanuria. She did not yet have demonstrable visceral metastases, but a few months later the spleen was markedly en- larged and very hard. When she died, sixteen months after her admission to us and four- teen months after the delivery, postmortem examination showed the most extensive cu- taneous, subcutaneous and visceral involve- ment that one can imagine. Black nodules studded absolutely all the tissues. The end-result has been fatal in fourteen, or 50%, of these patients, in three years or less. Of the patients treated over five years ago, only three survived, and only two are known to live over a five-year period, as one was lost to observation after four years. Those two are living eight and thirteen years, respectively, and are considered cured. One patient has had two normal pregnancies and deliveries at a safe interval after the radical surgical removal of the malignant melanoma which had not metastasized to the regional lymph nodes when they were dis- sected out. All the patients treated within fairly re- cent years have had radical surgical treat- ment, including not only the wide excision of the site of the primary melanoma, but dissec- tion continuously to the regional lymph nodes and radical removal of the lymph-node-bear- ing area. A 35-year-old woman may be used to illustrate this method of treatment. She presented herself for treatment with a lesion on the back and giving the following history : For four years she noticed something on her back. Four months before, while she was pregnant, it began to ulcerate and have a dis- charge. She consulted her family physician, who removed it by electrodesiccation. It grew back within a month. The following month she delivered a normal child. When we saw her, she had a raised, firm, bluish to brown growth 10 by 15 mm. in the margin of a scar situated in the midline of the back between the scapulae. At that time she had two small lymph nodes palpable in one axilla and larger hard ones in the opposite one. The malignant melanoma on the back was excised with a skin segment 5 to 6 cm. wide above and below and extending continuously to the axilla on each side. The subcutaneous fat and fascia over the muscles were removed more widely across the back and to each axilla. The axil- lary veins were dissected free of all fat and lymphoid tissue up to a level behind the pec- toralis minor muscle. The axillary tissue on either side was removed in continuity with the tissue from the back. The microscopical examination confirmed the clinical diagnosis of metastatic melanoma in the lymph nodes from both axillae. In spite of this very radical surgery, a year later this patient had pal- pable hard nodules in both breasts. Both breasts were removed completely and there were two masses representing metastatic 1950] Abstracts: Pigment Cell Conference 5 melanoma in the right one and five in the left breast. Metastatic nodules next made their appearance in the neck and then in the liver, and in less than two years after her delivery and treatment, she died of malig- nant disease. We have eleven patients in this series liv- ing and well at the present time, but most of them have been operated upon within the past two years. These original observations are clinical, but may be summarized as follows : (11 The benign or premelanotic nevus is definitely stimulated by hormonal changes of puberty and pregnancy. (2) Pre-pubertal melanomas possessing all the histologic features of malignant mel- anoma except blood and lymph vessel inva- sion do not metastasize until puberty (with rare exception). (3) Certain benign nevi are prone to change to malignant melanoma during preg- nancy. (4) Malignant melanomas grow with great rapidity, metastasize early and widely, are more malignant and have a lower rate of curability during pregnancy. (5) Dormant, silent, I'esidual melanoma may flare into active recurrence during preg- nancy, subsequent to operative removal. (6) Prophylaxis is removal of all suspi- cious nevi in women before or at least early during pregnancy. Bibliography. Scharnagel, Isabel M. Treatment of Malig- nant Melanoma of the Skin and Vulva at Radiumhemmet, Stockholm. Acta Radiol., 14:473, 1933. Melanoma of the Skin. J.A.M.W.A., June, 1946. Scharnagel, Isabel M., G. T. Pack and Mason Morfit. Principle of Excision and Dis- section in Continuity for Primary and Metastatic Melanoma of the Skin. Surgery, 17, 6:849-866, June, 1945. Pack, George T. Subungual Melanoma. Bull. Memorial Hospital, 2:24-28, Feb., 1930. A Clinical Study of Pigmented Nevi and Melanomas. In: The Biology of Mel- anomas, New York Academy of Sciences, IV, pp. 52-62, Jan., 1948. Questions of Herman Charache, Brooklyn Cancer Institute, New York. 1. In your experience with over one thou- sand cases of malignant melanoma, did you find whether melanuria is of any prognostic value? Answer : We believe that melanuria is very frequently present in cases of malignant mel- anoma. There are other conditions also in which this reaction may be demonstrated. We do not believe that it has any important prognostic value. 2. Is the color of pigmented nevi, such as black, brown or blue, of any prognostic sig- nificance? Answer : It has long been our practice to excise all blue-black moles, no matter how benign they appear. Of course many of these moles are Jadassohn neuronevi. These nevi are considered not to undergo malignant de- generation very often, but we have cases in which it has occurred. Otherwise, there is no great significance in the color of the moles, as a certain percentage of malignant melano- mas contain no pigmentation at all. These non-pigmented melanomas may, however, produce pigmented metastases in lymph nodes. The color, therefore, has no great prognostic significance. 3. Would you recommend sterilization of young women with malignant melanoma, or interrupt early pregnancy in women with malignant melanomas? Answer: The sterilization of young women with malignant melanoma seems to us a rather drastic procedure, as we believe that with radical surgery in early stages, we are able to cure these patients. I have cited one case where a woman is living and well for thirteen years and has had two normal preg- nancies with deliveries since her malignant melanoma was treated, although it developed during the course of her first pregnancy. This woman had a spontaneous interruption of the first pregnancy, and I personally be- lieve that if the malignant melanoma is diag- nosed early enough in pregnancy so that therapeutic abortion can be carried out safely, it should be done. Benign and Malignant Neoplasia of Melanoblasts Through the Eyes of the Dermatologist. S. William Becker. University of Chicago School of Medicine. Benign neoplasia of melanoblasts (1) is seen in prenatal pigmented nevi, which are either present at birth or appear during the first few years of life. Human beings aver- age 18 to 20 such lesions (Pack). They vary from small macules to large, elevated, non- hairy or hairy plaques. Most nevi are pig- mented; a few small elevated lesions are non-pigmented, especially on the face and scalp. Acquired nevi appear in late childhood or early adult life in the form of flesh- 6 Zoological New York Zoological Society [35: 1 colored papules which later become brown (Ebert). Clinical diagnosis of pigmented nevus is made by dermatologists in 80 to 90% of cases. Of 710 lesions diagnosed as nevi (2), microscopic diagnosis was: nevus in 80%, seborrheic keratosis 5%, papillo-epithelioma 3%, carcinoma 2%, verruca vulgaris 1%, and other diagnoses 9%. Reversing the anal- ysis, of 649 lesions diagnosed microscopi- cally as nevi, nevus had been diagnosed clinically in 87%, seborrheic keratosis 2%, carcinoma 2%, fibroma molle, verruca vul- garis, lentigo maligna, papillo-epithelioma, adenoma sebaceum and cyst, 1% each, and other diagnoses 3%. Microscopically, nevi are divided into type A (nevus cells in masses associated with the epidermis) ; B (nevus cells in groups or cords in the dermis) ; and C (elongated nerve type cells in the deeper dermis) . Vari- ous combinations of the different types occur. In childhood, types A and AB nevi are most common, in adult life types B and BC predominate. Of 741 specimens of nevi, mostly removed from adults, the classifica- tion was as follows (2): Type of Percentage Number Nevus of Cases of Cases A 2.1 16 A and B 8.7 65 B 16.4 122 B and C 64.3 477 A, B and C 6.6 49 C 1.6 12 Total 99.7 741 Acquired nevi are of the A and AB types. Because of their activity, they may be con- fused with lentigo maligna, but may be dis- tinguished from the latter by the fact that the nevus cells only penetrate into the der- mis and do not work their way to the surface in the epidermis. Treatment of nevi depends on the variety, the location and the state of growth. Any nevus which is actually growing or is irri- tated or inflamed, should be excised surgi- cally. All specimens should be biopsied. If melanoma has supervened, treatment should be carried out for melanoma. Quiescent nevi in sites subject to constant friction should be removed prophylactically. Other nevi are treated for cosmetic reasons. Treatment is carried out by figuration, electric cautery or solid carbon dioxide. A portion of an elevated nevus should be removed for biopsy, after which treatment is directed toward obtaining a smooth scar. In this way unsus- pected degeneration of nevi into melanoma can be kept at an irreducible minimum. Large nevi are excised surgically and re- placed by grafted skin. Malignant neoplasia of melanoblasts (1) usually occurs in form of lentigo maligna, a premalignant or, better, early malignant lesion, starting as an asymptomatic brown macule which enlarges irregularly and be- comes slightly elevated. After a few months or years a tumor appears, usually in the center of the plaque, often erythematous (non-pigmented melanoma) or pigmented (pigmented melanoma). In the author’s series, melanoma arose as lentigo maligna in 77% of cases, and in true nevi containing nevus cells in only 23%. This contrasts with the statement in most reports that melanoma arises in nevi in more than 50% of instances. Diagnosis of melanoma is more difficult than that of nevus. Of 76 specimens removed from lesions diagnosed clinically as mela- noma, microscopic examination revealed melanoma in only 45%, pigmented nevus and blue nevus each 11%, lentigo 8% and seborrheic keratosis and melanotic basal- celled carcinoma, each 5%. Reversing the analysis — of 59 lesions diagnosed melanoma microscopically, only 51% had been correctly diagnosed clinically. Nevus pigmentosus had been diagnosed in 25%, seborrheic keratosis in 9%, carcinoma in 7% and other diagnoses in 9%. Diagnosis of non-pigmented melano- ma is made only by biopsy. Microscopic diagnosis of lentigo maligna (3, 4, 5) is made by the presence of three features: first, neoplastic proliferation of melanoblasts at the epidermo-dermal junc- tion ; second, round cell infiltrate, often pro- nounced, in the superficial dermis; and, third, sometimes absent, casting off of small masses of neoplastic cells through the epi- dermis. The cells are ordinarily round or oval, although fusiform melanoma cells are sometimes seen. Microscopic diagnosis of frank melanoma is made by the anaplastic nature of the cells forming the tumor, which may be either round or oval or fusiform. Melanoma originating in a nevus shows malignant neoplasia of melanoblasts at the epidermo-dermal junction. Non-pigmented melanoma is diagnosed by the proliferation of cells at the epidermo-dermal junction with the usual lack of cohesion between the cells, as contrasted to epidermal carcinoma. Treatment of early lentigo maligna con- sists of local surgical excision. For later lesions, regional lymph node removal should be strongly considered, and, if a frank mela- nomatous tumor is present, regional lymph node dissection should be done unless con- tra-indicated by the presence of extensive metastases. Prognosis of early melanoma is excellent, but failure to biopsy early lesions and carry out adequate treatment has doomed many persons. References. 1. Becker, S. W. and Obeumayee, M. E. Mod- ern Dermatology and Syphilology. 2nd ed., Philadelphia, J. B. Lippincott Company, 1947. 2. Becker, S. W. Diagnosis and Treatment of Pigmented Nevi. Arch. Dermal. & Syph., 60:44 (July), 1949. 3. Becker, S. W. Cutaneous Melanoma. Arch. Dermat. & Syph., 21:818 (May), 1930- 1950] Abstracts : Pigment Cell Confereyice 7 4. Becker, S. W. Melanotic Neoplasms of the Skin. Amer. J. Cancer, 22:17 (Sept.), 1934. 5. Becker, S. W. Dermatological Investigations of Melanin Pigmentation. The Biology of Melanomas, New York Academy of Sciences, 1948. Discussion by H. Z. Lund, Western Reserve University, Cleveland, O. The contrast of appearance of a pigmented nevus of a child and that of an adult is generally known. We have recently made a systematic study of the appearance of be- nign pigmented nevi at all ages and find that there are, in most instances, progres- sive changes which roughly correlate with the age of the patient. The most important variables are junctional proliferation which is present to a moderate to marked degree in about 90 per cent, of children under 10 years of age. The subsequent decades show a marked diminution in this percentage but there is no age at which it abruptly ceases. Slight degrees can be found in many nevi in adult life. Mitotic figures are found rarely but likewise show a higher incidence in the first decade and then a steady drop. Mitotic figures are uncommon in adult life. In later life the nevus cells are more fusiform and associated with fibrils. Fibrils progressively increase with age, as do the structures re- sembling tactile corpuscles. Thus the nevus cells which early in life resemble epithelium gradually differentiate to tissue resembling neuroid structures. There are exceptions to the above trend which cannot be discussed here. Reference. Lund, H. Z. and Stobbe, G.D. Am. J. Path., 25:1117, 1949. Discussion by Eugene F. Traub, New York Medical College. I am in accord with Dr. Becker that an accurate diagnosis of pigmented nevi is pos- sible only after microscopic examination. Not only are a number of other lesions fre- quently mistaken for pigmented nevi but the differentiation of one pigmented nevus from the other is often impossible without micro- scopic examination. Dr. Becker has used a classification for the study of nevi based upon that of Miescher and von Albertini. It is not entirely clear to me just exactly what all may be included in each of his three types. Furthermore, to remember them as types “A,” “B” and “C,” is rather meaningless unless fully explained by the accompanying text or a specially appended name. But it would seem that each of his types may be a hodge-podge of several varieties of nevi, which is never good. For example, type “A” is exemplified by lesions of an intra-epithel- ial growth while type “B” includes the “common mole” or intra-dermal nevus of my classification as well as the blue nevus and one would have to specify which of these, or any mixed types, was meant. Dr. Becker admits that depending on where and how the sections are cut, one might find the lesion fitting into type “A,” “B” or even “C.” This would not occur in a classification like mine, except in the case of a mixed nevus as, in most instances, there would be no difficulty in recognizing at once if its exact type is intra-epidermal, intra-dermal, junction or blue nevus. I do not agree with some of the methods of the treatment suggested by Dr. Becker. He suggests that large macular pigmented nevi could be destroyed with solid carbon dioxide, electrocoagulation or by figura- tion. I believe some of these procedures are satisfactory for some of the rather small lesions but where larger pigmented nevi are involved, I believe the hair line scar of a surgical excision gives a better cosmetic re- sult than can be accomplished by making a scar of the entire area. This is particularly true with those nevi complicated with hair that requires extensive electrolysis after the pigment is first destroyed with either solid carbon dioxide, electrocoagulation or fulgu ration. In these cases, it is much simpler to excise the lesion at one sitting, unless it is so large that closure is utterly impossible and even then sometimes skin grafting is preferable to many repeated treatements by electrocoagulation, electrolysis, etc. Further- more, as Dr. Becker pointed out, diagnostic errors can only be prevented or corrected on study of the tissues and this can best be done only by excising and examining the entire lesion. The Problem of Acanthosis Nigricans. Helen Ollendorff Curth. College of Physicians and Surgeons, Columbia University, New York. Acanthosis nigricans is a benign derma- tosis showing hyperpigmentation, mostly in the basal cell layer, as well as other charac- teristic epidermal changes. Warts and pig- mented spots or moles may accompany these lesions. About 450 cases of the disease have been reported. The dermatosis is regionally and symmetrically distributed, the axillas being the most commonly involved area. Melanomas do not develop from the lesions. Acanthosis nigricans occurs in man and dog and is found in both sexes. Its significance lies in its association with internal cancer. This combination is seen in 50% of cases of acanthosis nigricans. The type of acan- thosis nigricans associated with internal 8 Zoologica: New York Zoological Society [35: 1 cancer is falsely called malignant. The type of the disease not so associated is called benign. The two types look alike but can, I believe, be differentiated by careful anal- ysis. The cancers associated with acanthosis nigricans are highly malignant, non-pig- mented adenocarcinomas originating in the stomach or other abdominal organs in 92%, and in the breast, lung, etc., in 8%. Not a single patient has been cured by treatment of the cancer. The cancers occur mostly in the middle-aged but also in the very young and very old. Acanthosis nigricans may pre- cede the cancer by many years, may start simultaneously with, or it may follow can- cer. It appears, therefore, that neither acan- thosis nigricans nor cancer causes the other. An activating influence of cancer on acan- thosis nigricans, however, cannot be denied. Acanthosis nigricans not associated with cancer may assume the form of a unilateral pigmented verrucous nevus. It starts at birth, in childhood or at puberty. Sex hor- mones seem to have a stimulating influence on this benign type of the disease. As in the case of melanoma or certain physiologi- cal pigmentations, these patients are sub- jected at puberty to a physiologic barrage of sex hormones which result in hyper- pigmentation and other epithelial changes. Studies of the 17 keto-steroids in young and middle-aged men and women suffering from acanthosis nigricans gave normal val- ues. The parallel action of the cancer agent and the sex hormones on the two types of the dermatosis seems noteworthy. If an individual develops acanthosis nigri- cans after puberty he will sooner or later show an internal cancer and the cutaneous disease will become more intense with the manifestation of cancer. Young people, how- ever, may have “malignant” acanthosis ni- gricans before puberty. Treatment of the malignant type consists in early recognition and removal of the tumor. In the benign type the lesions some- times regress spontaneously after puberty. When it was found that there were no recorded familial cases of the malignant type of acanthosis nigricans, such as had been found in the benign type, an investiga- tion of the genetics of the disease under the auspices of the American Cancer Society was undertaken. Furthermore, in this study an effort is being made to determine whether adenocarcinoma occurs in a higher per- centage in the families of patients with acanthosis nigricans than in the ordinary population. Particular difficulties were en- countered. In addition to all the known han- dicaps of genetic studies of cancer in man, one cannot be certain whether cancer victims in the older generations had also shown the cutaneous changes, possibly to a mild degree. If the individual has not yet reached puberty or cancer age, acanthosis nigricans is not likely to be noted. Are genetic carriers of the dermatosis those with a great number of pigmented moles, and what is the “normal” amount of moles? Genetic carriers of can- cer may not have been recognized because family members were not examined for any special metabolic feature on which cancer associated with acanthosis nigricans might be based. Those who transmit the disease might have exhibited such a disturbance to a lesser degree than patients with overt manifestations. If we should succeed in establishing malig- nant acanthosis nigricans as a separate disease, a dermatosis might emerge which would, not in 50% but in 100% of cases, be an indicator of internal cancer and might point the way to the biology of certain malig- nant tumors. Bibliography. Curth, Helen Ollendorff. Benign Type of Acanthosis Nigricans: Etiology. Arch. Dermat. & Syph., 34:353-366 (Sept.), 1936. Cancer Associated with Acanthosis Nigricans. Review of literature and report of a case of acanthosis nigricans with cancer of the breast. Arch, of Surgery, 47 : 517-552 (Dec.), 1943. Acanthosis Nigricans and its Asso- ciation with Cancer. Arch. Dermat. & Syph., 57:158-170 (Feb.), 1948. Acanthosis Nigricans und Krebs. Z. f. Krebsforschung, 56:307-328, 1949. Curth, Helen Ollendorff and Slanetz, C. A. Acanthosis Nigricans and Cancer of the Liver in a Dog. Am. J. Cancer, 37 :216-223, 1939. The Junction Nevus, Forerunner of the Malignant Melanoma and Its Differential Diagnosis from the Standpoint of the Dermatologist. Eugene F. Traub. New York Medical College. The pigmented hairy and warty nevi have always been difficult to classify and no logi- cal clinical classification has ever appeared feasible. Therefore, a simple anatomic basis on which to classify them has given us the best chance to simply describe these lesions and to gain definite knowledge as to which of them are potentially dangerous and which generally remain benign throughout their 1950] Abstracts : Pigment Cell Conference 9 entire course. We have therefore classified the pigmented hairy and warty nevi as 1. intra-epidermal Nevi 2. Junction Nevi 3. Intradermal Nevi 4. Blue Nevi 5. Mixed types The name “junction” was derived from the fact that the cells constituting this nevus arise at the epidermal-dermal junction. Of the group this is the only one which may give rise to a malignant melanoma or nevo- carcinoma. Clinically, however, any of the above types may resemble the junction nevus so closely in clinical appearance that only on microscopic examination can one be certain of the diagnosis. The mixed lesions are particularly confus- ing because they comprise combinations of any of the above types and generally this can only be recognized on microscopic ex- amination. Classically the junction nevus is usually small in size, not over 2-3 cm. as a rule, slightly elevated and smooth surfaced. Occa- sionally the surface may be soft, warty, or uneven, but it never is a hard warty lesion. The color may vary from light brown to brownish-black, or it may even be blue-black. Hairs are not present except in exceptional cases. Junction nevi may appear at any time in life or may be present at birth. Clinically and microscopically they are indistinguish- able regardless of the time of their appear- ance, but their behavior and prognosis is different. Those present from earliest in- fancy as a rule grow slowly or not at all, and if they become nevocarcinomas they metastasize more slowly and prognosis fol- lowing radical treatment is successful in a relatively high percentage of cases. Those appearing in adult life, after puberty, grow more rapidly, metastasize more promptly and the outlook in these cases is always ex- tremely grave. The percentage of cures is low in this group. Since junction nevi, like all other nevi, are not more prone to cancer development than normal skin, most of these lesions can and should be left alone. Only when they are located at sites of extreme trauma, hands and feet, etc., is it safer to remove them pro- phylactically. Such removal may be accom- plished with the sacrifice of a minimum of surrounding normal skin. This is ample in most cases but if early changes to nevo- carcinoma are found, a wider excision may then be deemed advisable. It should be emphasized that junction nevi are benign and that adequate removal is a perfectly safe procedure without danger of aggravating the local lesion or causing me- tastasis. It is because of believing that a lesion is benign, when actually it is already malignant, that trouble results from a re- moval predicated on this erroneous basis. Even then the error can be corrected follow- ing microscopic examination of the tissue. The danger of malignant melanoma or nevocarcinoma can be greatly minimized by the early removal of questionable or badly located junction nevi. To advise removal of all of them is ridiculous, but it is equally bad and far more dangerous to suggest leaving them all alone. We should continually stress that the highest percentage of cures results from prophylactic removal of the earliest lesions. There are two types of pigmented nevi most likely to be confused with the junction nevus. The first is the benign pigmented intra- epidermal nevus and the second the blue nevus. Less frequently, even the intra-dermal nevus may be confused with the junction nevus. The reason the pigmented intra- epidermal nevus is so frequently confused is because it is a smooth pigmented area devoid of hairs but often with deep black pigment and when located on the hands and feet, where thickness or elevation are difficult to make out, it is almost impossible to be certain whether the lesion is an intra-epidermal or junction nevus. The junction nevus usually shows some thickening but in certain areas this is not easily determined. This point was brought home to me recently by a patient who had about 60 small pigmented nevi on his trunk and extremities. One on the foot was removed and proved to be an intra-epidermal nevus while a similar mark on the back proved to be a junction nevus. Therefore, probably all of the others, not removed, and scattered over his body, belonged in one of these two categories and one would have had to remove each one individually to be certain of its exact type. Would it be advisable to treat indiscriminately all of these nevi with solid carbon dioxide, electrocoagulation, des- iccation, etc? It is only fair to state that our rare instances of the development of malig- nant melanoma or nevocarcinoma following inadequate destruction of junction nevi have resulted from this practice. The typical blue nevus, because of its blue- black color, can usually be differentiated clin- ically from the junction type but if the blue nevus is more brownish or brownish-black in color and rather superficially located in the cutis, confusion does frequently result. The benign pigmented intra-epidermal nevus does not terminate as nevocarcinoma or ma- lignant melanoma but in rare instances it has terminated as a carcinoma. Bibliography. Traub, Eugene F. The Pigmented, Hairy and Warty Nevi and Their Relationship to Malignancy. Southern Med. Jour., 40 (12) : 1000-1005, Dec., 1947. Early Recognition of Possibly Danger- ous Nevi (Moles) and the Best Procedure to Avoid Development of Malignant Mela- nomas (Nevocarcinoma). N. Y. State Jour . Med., 49 (14) : 1661-1664, July 15, 1949. The “Common Mole.” Its Clinico- pathologic Relations and the Question of Malignant Degeneration. Arch. Dermat. & Syph., 41: 214-249, Feb., 1940. 10 [35: 1 Zoologica: New York Zoological Society Congenital Anomalies (Nevi) and Their Relationship to Cancer and Melano- ma. Penn. Med. Jour., pp. 1-8, June, 1941. Discussion by H. Z. Lund, Western Reserve University, Cleveland, O. From the standpoint of the surgical path- ologist, recognition of early melanoblastoma and distinction from benign nevus are ex- tremely important. The first feature which is looked for is junctional proliferation be- cause most, if not all, melanoblastomas arise in the epidermis or at the dermo-epidermal junction. Traub and Keil have stressed this and awakened many to awareness of it. However, the emphasis has been insuffi- ciently critical and there has been an ac- ceptance by some dermatologists and path- ologists that the presence of junctional proliferation at any age and of any degree is evidence of a potentially malignant lesion. Such a conclusion is drawn from the article by Traub and Keil. Actually the feature of junctional proliferation becomes significant if it is unusual in degree for a given age, as, for example, the finding of a nevus of a middle-aged person Avhich has the histo- logical characteristics of a nevus of child- hood; i.e., the cells are largely junctional in distribution, resemble epithelium and there is little evidence of differentiation to ordinary nevus cells and cells associated with fibrils. Such an appearance leads to the suspicion that the lesion is incipient and growing. (The clinical evidence obtained in our own analysis of the duration of growth of such lesions was variable and conflicting, permitting no final conclusion. Additional study is needed.) The patholo- gist is to be guarded in excluding mel- anoblastoma in these cases, but until more conclusive evidence is obtained, a positive diagnosis must rest on this finding plus ad- ditional changes, including (a) excessively large, irregularly scattered or otherwise bizarre masses or nests of cells at the dermo- epidermal junction, (b) deep penetration of large cells without differentiation to small nevus cells and fibrillar forms, (c) more mitotic figures than are usual, (d) atypical and pleomorphic cells, (e) invasion, (f) trophic changes, and (g) inflammation, other than folliculitis, which is not accounted for by trauma. References. 1. Traub, E. F. and Keil, H. The “common mole,” its clinicopathologic relations and the question of malignant degeneration. Arch. Dcrmat. & Syph., 41, 214-252, 1940. 2. Lund, LI. Z. and Stobbe, G. D. The natural history of the pigmented nevus; factors of age and anatomic location. Am. J. Path., 25:1117, 1949. Discussion by S. WILLIAM BECKER, University of Chicago School of Medicine. The term “junction nevus” has been criti- cized because such designation includes both “A” and “AB” nevi, for which it could be legitimately used, and, in addition, the oblig- ative premalignant (or actually malignant, depending on the criteria used) disorder called “lentigo maligna” by Dubreuilh. My preference is to designate as pigmented nevi only those lesions composed of benign mel- anoblastic cells (nevus cells). Dr. Traub has again emphasized the neces- sity for microscopic examination of nevi and tumors with which they may be confused. He has shown with excellent colored photo- graphs, a “junction nevus” and an “intraepi- dermal nevus” which cannot be distinguished clinically. On the basis of his excellent col- ored photomicrographic slides of the two dis- orders, it would seem that the first satisfies fully all the requirements of “lentigo ma- ligna” of Dubreuilh. The second is composed entirely of epithelial cells and contains no. neoplastic melanoblasts, and would seem to belong in the group of benign epidermal neoplasms (Becker, S. W. : Benign Epider- mal Neoplasms, Arch. Dermat. and Syph., 26:838 (Nov.) 1932) and could be called “be- nign epithelioma.” At any rate, it seems more scientific to classify pathologically rather than to perpet- uate the older designation “nevus” in the sense of a “mark.” The Development of Epidermal Pigmenta- tion in the Negro Fetus. Arnold A. Zimmermann. Department of Anatomy, University of Illinois, College of Medicine, Chicago, III. Fetal Negro skin constitutes excellent ma- terial for the study of melanogenesis. Sur- prisingly, no systematic survey, throughout the fetal stages, had previously been made. The main advantages reside primarily in the high potentiality for melanin production of specialized cells which reveal their activity early in fetal life. They are the_ dendritic melanoblasts which remain distinct, both morphologically and functionally, from ordi- nary epithelial cells. They are the sole pro- ducers of melanin both in the epidermis and in the papillae of hair follicles. The distribu- tion and signficance of dendritic cells in the adult human skin, both white and pigmented, recently was studied by Billingham. The derivation of mammalian melano- blasts from the neural crest has been demon- strated by Rawles (1947) . In the fetal Negro 1950] Abstracts : Pigment Cell Conference 11 skin, however, the search for migratory mel- anoblasts in the dermis, previous to their identification in the epidermis, encounters difficulties. If potential melanoblasts are present in the embryonic or early fetal dermis they cannot be demonstrated either by a positive dopa-reaction or by the usual silver techniques. This might be due to in- sufficient amounts of a melanogenous sub- strate (tyrosine) or to incomplete, as yet inactivated oxidizing enzyme-systems in the precursors of epidermal melanoblasts. The latter assumption appears more probable. Experimental evidence in Amphibia has es- tablished the fact that the ectoderm plays an important role in melanin formation (Du- Shane, 1943). My observations in the fetal Negro skin, so far, appear to confirm the hy- pothesis that substances necessary for the synthesis of melanin may be supplied by epi- dermal cells. An appositional relationship of human melanoblasts to the epidermis seems to be essential for their full functional ac- tivity. The material for this histological study of fetal Negro skin consisted of skin- specimens from 75 abortions. The main re- sults may be summarized as follows: 1. The basal cells of the human epidermis do not produce melanin. 2. The true source of melanin l'esides solely in the pigmentary dendritic cells or melano- blasts which constitute an independent cell- type within the epidermis. 3. Dendritic melanoblasts first appear in the fetal Negi’O skin early in the third month. 4. There are no cytological or functional transitions between epidermal melanoblasts and ordinary epithelial cells. 5. The dendritic melanoblasts consistently show a positive dopa-reaction throughout fetal life. Basal and other epidermal cells remain dopa-negative. 6. Dendritic melanoblasts of the third fetal month contain melanin-precursors (premel- anin) which can be demonstrated by reduced silver methods. From the fourth fetal month to term they contain both premelanin and melanin granules. 7. The earliest melanoblasts in the epider- mis have ovoid, fusiform or stellate cell-bodies with few and short dendritic processes at their poles. In the fourth fetal month the total length of dendritic melanoblasts may reach 100 microns. Their primary dendritic processes extend through the epidermal in- tercellular spaces and begin to form an in- tricate syncytium. 8. During the fifth fetal month numerous secondary dendritic processes arise from the long primary processes of the melanoblasts. These secondary tufts approach and progres- sively surround the ordinary epithelial cells mainly on their distal pole. 9. In unstained sections, brown melanin granules are first identified within the mel- anoblasts in the late fourth and early fifth fetal months. 10. The transfer of melanin granules elab- orated in the dendritic melanoblasts to neigh- boring epithelial cells begins late in the fifth fetal month. This process is accentuated throughout the remainder of the fetal period. The transfer occurs mainly from the sec- ondary dendritic processes and leads to the accumulation of melanin in ordinary epi- dermal cells as “supra-nuclear caps.” The active melanoblasts gradually become con- cealed among the pigment-carrying epithe- lial cells. 11. Melanization of hair shafts and of the matrix of the papillae in all essentials is sim- ilar to that of the epidermis. Dendritic melanoblasts accumulate in the papillae and extend their processes directly into the grow- ing base of hair-shafts, thus melanizing them independently of a pigmented epithelial ma- trix. The latter receives melanin granules quite secondarily after melanization of the hair shaft is well under way. References. 1. Zimmermann, Arnold A. The development of epidermal pigmentation in the Negro fetus. Anat. Rec., 100, p. 96, April, 1948. 2. Zimmermann, Arnold and Th. Cornbleet. The development of epidermal pigmentation in the Negro fetus. Journ. Investig. Derma- tol., 11, pp. 383-392, Nov., 1948. 3. Rawles, Mary E. Origin of pigment cells from the neural crest in the mouse embryo. Physiol. Zool., 20, 248-266, 1947. 4. Origin of melanophores and their role in development of color patterns in Vertebrates. Physiol. Rev., 28, 383-408, 1948. 5. DuShane, Graham P. The embryology of Vertebrate pigment cells. Part I. Amphibia. Quart. Rev. Biol., 18, 109-127, 1943. 6. Billingham, R. E. Dendritic cells. Journ. Anat., 82, 93-109, April, 1948. 7. Dendritic cells in pigmented hu- man skin. Journ. Anat., 83, 109-115, 1949. Discussion by Hermann Pinkus, Wayne Univer- sity Medical College, Detroit, Mich. Dr. Zimmermann in his splendid investi- gation was unable to find evidence of multi- plication of fetal melanoblasts. Masson (The Biology of Melanomas, 1948, p. 31 and plate 15, fig. 1) has stated his belief that epidermal melanoblasts usually multiply by amitosis, but says that he saw one unquestionable mi- tosis. I would like to contribute the purely accidental observation of mitotic division of normal dendritic pigment cells in the basal layer of adult human skin. (Pinkus, H.: Mi- totic Division of Human Dendritic Melano- blasts, J. Investig. Dermat., 13: 309-311, Dec., 1949). The specimen was a common wart (verruca vulgaris) taken from the finger of a dark colored woman. Two unques- tionable and two suggestive instances of mi- tosis were seen in twenty-four routine histo- logic sections. Mitosis of Malpighian cells was common in this specimen. Without doing an actual count, it was my impression that the ratio of dividing and resting nuclei was similar for dendritic and Malpighian cells. 12 Zoologic a: New York Zoological Society [35: 1 This experience proves that human melano- blasts can multiply by mitosis in non-malig- nant conditions. I have also recently observed occasional instances of mitosis of benign nevus cells in ordinary papillomatous nevi. A Comparative Study of Malignant Mela- noma Among Negro and White Patients. Rudolph J. Muelling, Jr. & Walter J. Burdette. Departments of Pathology and Surgery, Louisi- ana State University School of Medicine, New Orleans, La. The admissions to Charity Hospital of Louisiana at New Orleans are nearly equal for white and Negro males and females. Dur- ing the period from 1937 to date, there have been 101 cases of malignant melanoma among 26,800 malignant tumors in 686,293 admis- sions. That is, approximately 4 cases of malignant melanoma occurred per 1,000 cases of malignancy. Approximately one-third (32%) of these melanomas occurred in Negro and two-thirds in white patients. The sex incidence in the Negro group was equal, whereas the inci- dence in white females was slightly higher than in white males. The majority of cases (88%) occurred between the ages of 30 and 80 years, those in Negroes having highest incidence some ten years later than those in white persons. The foot was the most common site for the primary lesion in the Negro (50%) while in the white race the trunk was most common (33%). A pre-existing mole was described in one-third of the cases, mostly white. A history of trauma was elicited in one-half of the Negroes and one-tenth of the white patients. It is not possible to tell how much of the trauma was coincidental and falls in the post hoc, ergo propter hoc class. Clinical evidence of regional lymph node metastasis was found in about one-half of the cases. Better results of therapy were obtained when it was administered in the hospital rather than as hospital outpatients or in a physician’s office and when the metas- tases as well as the primary lesions were included in the plan of treatment. Inade- quate excision with recurrence of the pri- mary lesion was recorded in 17 cases. Two cases of melanoma appearing during preg- nancy were found to be of anaplastic his- tological appearance and were fatally termi- nated in a short time. The mean duration of life was 4 years and 5 months for the entire group. Eighteen out of 42 patients had a family history of at least one other indi- vidual with cancer. None of these had mela- noma, however. The sex incidence of malignant melanomas found is in agreement with that reported by Pack and co-workers (8) in an analysis of 862 cases. Dawson also found a history of the presence of a mole in one-third of the cases. However Pack et al (8) and Broders and MacCarthy (4) record this finding in one-half of their cases. Perhaps this dis- crepancy may be explained by the low inci- dence of antecedent moles among Negroes. The incidence of concomitant trauma is similar to that given by Pack et al (8) but much lower than that given by Horwitz (5), 57%. The tumor may arise from any portion of the Negro’s skin or eye as Sutton and Mallia (10) have reported. However, when the lesion occurs, it is more apt to arise in the less pigmented areas of the body, such as the foot (7). The individuals in which they appeared were not all lightly pigmented but in many instances were exceedingly dark. The rapidity of the process in two Negro males followed to termination demon- strates that de Lignis’ (6) conclusion that malignant melanoma in the native of North- ern Transvaal is a less intense process does not always hold true for the Negro in America. The evidence given here would in- dicate that the American Negro is not as peculiarly immune to malignant melanoma as one might assume from the literature. (1, 2,3). Summary. 1. Malignant melanoma was approximately one-half as frequent in Negroes as in white persons in a population of patients equally divided as to race and sex. This is a higher incidence than that usually quoted for Ne- groes. 2. Malignant melanomas arose more fre- quently from a mole in white than in Negro patients. 3. Metastases to regional lymph nodes oc- curred frequently. 4. Early therapy is better performed by those capable of treating not only the pri- mary but also secondary lesion. Bibliography. 1. Bauer, J. T. Malignant melanotic tumors in the Negro. Bull. Ayer Clin. Lab. & Pa. Hosp., 10:5-11, 1925. 2. Becker, W. Cutaneous melanoma. Arch. Dermat. & Syph., 21:818-835, 1930. 3. Bishop, E. L. Melanoma in the Negro. Am. Jour. Cancer, 16:522-539, 1932. 4. Broders, A. C. and MacCarthy, W. C. Melano-epithelioma. Surg., Gyn., & Obstet., 23:28-32,1916. 1950] Abstracts : Pigment Cell Conference 13 5. Horwitz, A. Melanotic tumors. Ann. Surg., 87:917-933, 1928. 6. de Lignis, M. J. H. Tumors in North Trans- vaal. S. African Med. Jour., 1:102-107, 1927. 7. Muelling, R. J. Malignant melanoma. The Military Surgeon, 103:359-364, 1948. 8. Pack, G. T., Perzik, S. L. and Scharnagel, I. M. The treatment of malignant mela- noma, report of 862 cases. Cal. Med., 66: 283-287, 1947. 9. Spencer, N. A. Melanoma. Brit. Med. Jour., 2:907-913, 1923. 10. Sutton, L. A. and Mallia, W. M. Malig- nant melanoma in the Negro. Arch. Dermat. & Syph., 8:325-343, 1923. Second Session: Genetical. Introduction. W. E. Heston. National Cancer Institute, Bethesda, Md. Since the birth of his science at the turn of the century, the geneticist has had a pro- found interest in pigment. Color varia- tions readily attracted his attention. It was through the study of coat color characters in mice and other rodents, plumage color in the fowl and eye color in Drosophila that much of the basic information on gene segre- gation, linkage, sex-linked characters, etc., was derived. In his book on the Genetics of the Mouse, Griineberg lists coat color genes located at 10 different loci, and two series have 5 alleles each. To this list could be added several more recently discovered coat color mutants as well as three genes controlling distribu- tion of pigmented areas. Analyses of these characters by Cuenot, Little, Castle, Mor- gan, Dunn and others constituted a major portion of the mammalian genetics of the first quarter of the century. Beyond the study directed at the gene itself, the geneticist is next interested in linking the gene to the biological character He wants to analyze the physiological and biochemical steps between the primary gene action and the end result. Here again the study of pigment has offered especial ad- vantages as emphasized by the work of those reportirg at this conference and of others, particularly Beadle and Ephrusi, and Wright and his students. As compared to other gene action paths those concerned with color appear to be relatively simple. Further- more the processes occur in cells readily accessible, and the history of these processes may be accurately recorded in such struc- tures as the hair or feathers. It is in studies such as these that the geneticist meets face to face with the bio- chemist, for final analysis of the gene action paths is on the biochemistry level. On the other hand the geneticist can be of great aid to the biochemist for already the geneti- cist has identified many of the genes related to the enzymes involved. The pigmented tumors afford a special field of physiological genetics. Those in the fruit fly and the platyfish hybrids have been of particular value because of the adapta- bility of both organisms to genetic and cyto- logic analysis and also because of the vast knowledge of the genetics of both organisms but particularly of the fruit fly. Along with the physiology of pigment formation is that of the malignant transformation, thus pre- senting a picture in which there is some danger in confusing the two sets of proc- esses. Are the genes which are responsible for the presence of the pigment cell, also in part at least responsible for the abnormal growth of the cell, or is the inheritance of the pigment cell more in line with that of eye and hair color, and the inheritance of the control of the growth of the cell in line with the genetics of neoplasms derived from other types of cells? In either case, what are the processes involved? As will be indicated in the reports of this section, much progress has been made in answering the basic ques- tions in these fields of physiological genetics. Reference. Heston, W. E. Genetics of Cancer. Advances in Genetics. 2:99-125, 1948. Significance of Quantitative Histological Studies of Pigment Found in the Coat Color Mutants of the Mouse to Questions of Nor- mal and Atypical Cell Growth. Elizabeth Shull Russell. Roscoe B. Jackson Memorial Laboratory, Bar Harbor, Me. Differences in visual effect of pigments in mouse hair are determined by variations in seven attributes of the melanin granules deposited there. Quantitative histological studies have shown four key pigmentation characteristics, relatively independent of 14 Zoologica: New York Zoological Society [35: 1 each other. From tip to base each hair is the final record of a regulated series of events occurring at each moult in the pig- ment-forming cells of the hair follicle. Vari- ations must be due to differences in number structure, or behavior of melanoblasts or possibly, in some cases, to differences in the hair matrix cells receiving the pigment. The stage of hair growth causes non-genetic variations in the complex, degree of pigmen- tation, involving changes in both number and size of granules, especially apparent in types with moderate or small total amounts of pigment deposition. In these there is a considerable lag in the commencement of pigmentation. Analysis of changes in gran- ule attributes with this pigmentation lag have been very valuable in determining the changes produced by simple quantitative alterations. The major genes affecting coat- color have been shown to affect one or at most two of the key pigmentation characters. The agouti series controls a trigger mech- anism, reversibly shifting the nature of pigment from eumelanin to xanthin. Since eumelanotic and xanthic pigments are chem- ically different; since eumelanotic granules are extremely variable in size and shape, while xanthic granules are always round; and since under most conditions many more considerably larger granules are deposited in eumelanotic than in corresponding xan- thic regions, the differences between the two systems of pigmentation must be quite extensive. Nevertheless, the rapid shift from one type to the other in agoutis suggests, though it does not prove, that the same melanoblasts produce both pigments, the difference being due to an altered condition within the melanoblast or hair matrix cell. Since non-agouti and agouti hairs start pig- ment formation identically, the agouti series genes definitely do not act by altering the time of arrival or beginning of functioning of melanoblasts. The action of the albino series is a quanti- tative change in the degree of pigmentation, affecting all types of pigment. For each qualitatively different type of pigment, its effects closely parallel the effect of pigment lag related to stage of hair growth. The gene action here could be change either in number or in activity of melanoblasts. Black-brown substitution alters the na- ture of eumelanotic pigments, but usually not their number. It seems that the Bb pair of genes must alter the function of some type of cell, rather than changing number. Pink-eye substitution alters eumelanotic granule size (aapp has tiny irregular shreds) and also degree of pigmentation. The action could be due to a qualitative defect in the sites of pigment deposition or to a great deficiency of some substance essential to eumelanin formation but not to xanthin formation. The dilute gene causes strong granular clumping which is almost certainly an in- tercellular affair resulting from altered be- havior of melanoblasts. There is no real reduction in amount of pigment, only an extremely irregular arrangement in which % to % of all pigment is concentrated into large clumps which absorb little light. The W-genes appear to alter the top limit of gi'anule number, their effect differing from that of the albino series in that they reduce granule number much more than size. Since the effects of the W-genes on pigment are exactly paralleled by their ef- fects on erythrocyte number and size (a severe macrocytic anemia is produced in the black-eyed white genotypes), and the effects on gonad development are also very similar, it seems probable that this series of genes is able to affect several types of cells. This is in strong contrast to the strict local and follicle-limited action of the other genic series. Throughout it appears that a given genic substitution alters only one aspect of the pigmentation process. The logical assump- tion of the physiological geneticist is that the dominant allele usually actively affects a process which is deficient in the presence of the double recessive. If this assumption holds for pigmentation, six different proc- esses have been shown, all of them active in producing the pigment of wild-type mouse hair. A thought I would like to leave with a group studying pigmentation intensively is that since there are at least six indepen- dent processes involved in pigment deposi- tion, it is very possible that some of them may affect behavior of melanoblasts, some number of melanoblasts, some the structure and function of cells forming the hair shaft and still others could have to do with a sub- stance diffusing out from the hair papilla. Bibliography. Russell, E. S. A quantitative histological study of the pigment found in the coat color mu- tants of the house mouse. 1946. I. Variable attributes of the pigment granules. Gene- tics, 31 :327-346. 1948. II. Estimates of the total volume of pigment. Genetics, 33:228- 236. 1949a. III. Interdependence among the variable granule attributes. Genetics, 34:133-145. 1949b. IV. The nature of the effects of genic substitution in five major allelic series. Genetics, 34:146-166. 1949c. Analysis of pleiotropism at the W-locus in the mouse: Relationship be- tween the effects of W and Wv substitution on hair pigmentation and on erythrocytes. Genetics, 34:708-723. 1950] Abstracts : Pigment Cell Conference 15 Melanotic Tumors in Drosophila .* Walter J. Burdette. Louisiana State University School of Medicine, New Orleans, La. At least 24 different strains of Drosophila which develop tumors have been described (1). A few, such as me, probably represent the end result of a degenerative rather than a neoplastic change (4). All except one of the remainder have tumors which are mela- notic. They are usually present in only a portion of the animals, although those with- out tumors transmit the susceptibility to their progeny. In the 8 stocks we have studied the incidence ranges from less than 1% to 50%. The tumors appear early in larval life and become pigmented. With met- amorphosis the cellular elements regress, leaving a residue of pigment by which they are identified grossly. The tumors which we have observed are benign, their charac- teristic size and location varies from one strain to another, and multiple tumors are not uncommon. They also survive trans- plantation. Sections reveal that the tumors are com- posed of polyhedral cells in clusters and fusiform cells may also be present. The tissue of origin has been inferred from the location and superficial resemblance to ad- jacent structures, but such conclusions are open to question (6). In the adult, amor- phous pigment is all that remains. These stocks of Drosophila have been in- bred and isogenic strains isolated. It is pos- sible not only to find the chromosomes responsible for tumor susceptibility in Dro- sophila but also to determine the locations of the genes involved on a specific part of each chromosome in certain instances. The number, location, and action of genes varies among the strains. Also certain tumor genes from one strain may affect tumor suscepti- bility when introduced into another tumor stock. Genes affecting tumor susceptibility have been found on the X chromosome and the three autosomes. Some modifying genes may enhance susceptibility while others sup- press it. A single gene near the left end of the X chromosome is responsible for the lethal 7 tumor (2). On the other hand. Stark reports that there are genes on four chromo- somes affecting tumor incidence in be-3 (5). The second chromosome is largely re- sponsible for the presence of tumors in tu° and tu^K The incidence of these melanotic tumors is affected by the environment in which the flies are raised. There are usually fewer turners when cultures are crowded, and nu- trition, temperature, and irradiation also influence the number of tumors which ap- pear. Therefore it is very important to maintain uniform culture conditions when studying the tumors. * Aided by a grant from the National United States Public Health Service. Cancer Institute, We have also used Drosophila to investi- gate chemical mutagens and carcinogens. Reports of the parallelism between carcino- genic and mutagenic properties of certain chemicals has lent support to the hypothesis that mutations play an important role in the etiology of tumors (3). In our laboratory results to date indicate that carcinogens are not always mutagens although they may be in some instances. Methyl-6fs (betachloro- ethyl) amine hydrochloride and 20-methyl- cholanthrene have been administered to flies and tests made for lethal mutations on the X chromosome. Using nitrogen mustard, 149 mutations were found in 17,052 chromo- somes tested. Eight parents were respon- sible for 67 of these lethals which were probably carried over from a previous gener- ation since the material was administered serially. Methylcholanthrene was adminis- tered as a vaginal douche and also as an aerosol. There were 3 mutations in 4.660 chromosomes tested in the former and 10 in 10,108 in the latter. Untreated flies showed 2 mutations among 2.822 chromosomes tested. In contrast to the data for nitrogen mustard, these results do not support the idea that methylcholanthrene is a mutagen, although the low mutation rate might be due to strain or species differences in re- sponse or to insufficient dosage for the mode of administration. The use of Drosophila as a test animal in the study of atypical growth thus presents certain advantages. This is particularly true for the study of hereditary factors which are active in many and diverse types of neoplastic disease. Both hereditary and en- vironmental factors are relatively easily con- trolled, the chromosome number is small, the life cycle is short, and the mutation rate may be determined in an objective manner. Experiments with tumors in Drosophila mav have added significance because of certain similarities to mammalian atypical growth. In this animal we already have concise evi- dence of gene action in tumor formation, and further study may be equally informa- tive. References. 1. Burdette, W. J. Tumors and mutations in Drosophila. Texas Reports on Biol. & Med., 1950. In press. 2. Bridges, C. B. Non-disjunction as proof of the chromosome theory of heredity. Genetics, 1:107-163, 1916. 3. Demerec, M. Induction of mutations in Drosophila by dibenzanthracene. Genetics, 33:337-348, 1948. 4. Gowen, J. W. The inheritance of focal melanosis in Drosophila. Arch. Path., 17 :638- 647, 1934. 5. Stark, M. B. and Bridges, C. B. The linkage relations of a benign tumor in Drosophila. Genetics, 11:249-266, 1926. 6. Russell, E. S. A comparison of benign and “malignant” tumors in Drosophila melano- gaster. J. Exper. Zool., 84:363-385, 1940. 16 Zoological New York Zoological Society [35: 1 Genetic Pigment Mosaics in the Pigeon. W. F. Hollander. Department of Anatomy, Yale University School of Medicine, New Haven, Conn. Plumage pigment of wild-type Columba livia is black, but domestic varieties exhibit also such hereditary conditions as “brown” or “chocolate” (recessive, sex-linked), “ash- red” (dominant, sex-linked, allele of brown), “almond” (dominant, sex-linked, closely linked with “brown”), “faded” (dominant, sex-linked, allele of “almond”), “dilute” (re- cessive, sex-linked), “recessive red” (not sex-linked), “grizzle” (dominant, not sex- linked), and others. Occasionally striking patchwork or mosaic combinations (“chimeras”) of the above color types are found. These afford some opportunity for analysis of pigment physi- ology. Successive molts produce little altera- tion in such specimens, and adjacent feathers growing at the same time may differ com- pletely, so that horomonal control does not seem involved. On the other hand, feathers at the edge of such patches may change color at molting, indicating that melano- blasts rather than tissue cells govern the pigmentation and are capable of some mi- gration. There is a perfect analogy here with grafted (experimental) color mosaics. Gene- tic evidence, from pedigrees and progeny tests, indicates that a number of such speci- mens may actually be a sort of natural graft, as a result of autonomous tissue for- mation by supernumerary sperms in em- bryogenesis. Other cases may have more conventional origins, such as chromosome aberrations, or somatic gene mutation. Another type of mosaic effect, more com- monly observable, is referred to as “flecking” or variegation. This occurs usually in pig- eons of the “ash-red,” “almond,” “faded,” and “grizzle” color types. For example, flecks in an ash-red male are black if he is hetero- zygous for black, but brown if he is hetero- zygous for brown; flecks in a male almond heterozygous for brown are black, brown, and faded (side by side). Genetic control is very evident. The flecks vary in size from almost microscopic to entire feathers or even small patches of feathers. When large, they usually show a lengthwise orientation in the feather. Flecks appear to be local somatic “muta- tions” of the dominant gene involved, so that a recessive allele appears. The effects are apparently much the same as in the “posi- tion effect mottling,” extensively studied in Drosophila. Apparently the “mutations” may occur at any time in the life of a pigment cell, and in the case of extremely small flecks may occur after the final mitosis. Since there is sharp distinction in pigmenta- tion of the flecks, it seems probable that the melanoblast is diploid; mutation in a polyploid cell would probably result in graded changes. Both chimeric and variegated mosaic ef- fects are known among other domestic birds, and other animals. Specimens were displayed at the meetings. References. Hollander, W. F. Mosaic effects in domestic birds. Quart. Review Biol., 19 :285-307, 1944. Bipaternity in pigeons. Jour, of Heredity, 40:27i-277, 1949. Discussion by J. P. TRINKAUS, Osborn Zoological Laboratory, Yale University, New Haven, Conn. Role of Epidermis and Hormones in the Differentiation of Melanoblasts. The melanin pigments of feathers are deposited in the epidermal cells of the dif- ferentiating feather by branched pigment cells called melanophores. In the undifferen- tiated condition, prior to the synthesis of pigment granules, they are referred to as melanoblasts. In the Brown Leghorn fowl, grafting experiments have demonstrated that the melanoblasts are embryonic-type cells, in feather germs of the adult as well as of the embryo, with a dual potency; they may form either black or red melano- phores [Trinkaus, J. Exp. Zool., 109:135- 170 (1948)]. The differentiation of melano- blasts, within the limits set by their genic constitution, is markedly influenced by the tract-specific epidermis of each feather germ, and by estrogenic and thyroid hormone [Trinkaus, J. Exp. Zool., 113, in press (1950)]. The nature of the resulting pig- ment pattern in each feather is, therefore, a consequence of the complex interaction of a number of tissue and humoral factors [cf. review by Rawles, Physiol. Rev., 28 :383- 408 (1948), for studies on pigment patterns in other fowl, and in amphibians and mam- mals]. This analysis suggests, of course, that more attention might be directed to the pos- sible role of tissue and humoral factors, in other studies on the differentiation of nor- mal and atypical pigment cells. It would be of interest, for example, to determine whether the pigment mosaics in pigeon feathers, described by Dr. Hollander, are due to modifications of the melanoblasts, or of the epidermis (or of both). By grafting melanoblasts from the pigmented area of a regenerating mosaic feather to the wing bud of an embryo, the melanoblasts would dif- ferentiate in the presence of different epi- dermis. The nature of the resulting pigment pattern in the host wing feathers should give a critical answer to the question. 1950] Abstracts : Pigment Cell Conference 17 Pigment Formation in the Eye of Ephestia and Its Genic Determination. Ernst Caspari. Wesleyan University, Middletown, Conn. In the study of the eye pigments of the meal moth Ephestia, the integration of em- bryological, biochemical and genetic analysis has proceeded so far as to give a consistent picture of the interaction of these factors. Pigments occur in three types of cells, pri- mary pigment cells, accessory pigment cells, and retinula cells. The pigment in the latter two types of cells is deposited in small brown granules insoluble in water, while a yellow, water-soluble pigment is found in the pri- mary pigment cells. The chemical behavior of these pigments is known from the investigations of Becker (1942). The brown pigment is a substance with well-defined chemical characteristics different from melanin, which has been given the name “skotommin.” Skotommin is derived from tryptophane according to the scheme: Tryptophane -* kynurenin -» oxykynurenin -» pigment. Developmental^, pigment appears first in the retinula cells. Pigment formation starts in these cells in the prepupa, and spreads gradually over the eye in the first 9 days of pupal life. Pigment appears in the accessory pigment cells about 10 days after pupation, in the primary pigment cells a few days later. Pigment formation in the accessory pigment cells has been more thoroughly de- scribed by Hanser (1948). The first thing to appear are very small precursor granules which enlarge under deposition of pigment on their surface. These precursor granules persist as the core of the final pigment granule. By enzymatic digestion experiments it could be demonstrated that the precursor granules contain ribonucleoproteins (Cas- pari and Richards, 1948). Two genes affecting the pigmentation of the eye have been studied, a and wa. a inter- feres with the transformation of trypto- phane to kynurenin, and in this way reduces the amount of skotommin pigment by 80- 90%. The size of pigment granules is de- creased. wa inhibits the formation of pig- ment completely, but does not interfere with the production of kynurenin. Hanser found that in wa wa animals the precursor gran- ules are not formed. aa animals can form large amounts of skotommin if supplied with kynurenin either by transnlantation of wild type organs (Cas- pari, 1933) or by injection. In this way it can be shown that the different types of cells of the eye have different “sensitive periods’' in which they are competent to react on supnly of kynurenin with pigment formation (Hanser, 1948). These sensitive periods start somewhat earlier than the periods in which pigment formation can be observed, and continue through the time of visible pigment formation. In accessory pig- ment cells, the sensitive period seems to start at about the time when the precursor granules become visible. From these experiments the following pic- ture of pigment formation arises. For the formation of pigment granules, at least two conditions are necessary: the presence of kynurenin, the chemical precursor of the pigment, and the ribonucleoprotein contain- ing granules. While the latter are not neces- sary for the formation of kynurenin, they probably have some function in transform- ing it into the pigment. Kynurenin is formed throughout the larval and pupal stages in all organs investigated (Caspari, 1949). Different cells acquire at different stages of their development the ability to react on kynurenin supply with formation of pig- ment. This stage of competence seems to be correlated with the appearance of the pre- cursor granules, and forms the develop- mental basis of the sensitive period. Genes may interact with pigment formation by affecting either kynurenin formation (a) or the formation of precursor granules (wa) . Bibliography. Becker, E. Uber Eigensehaften, Verbreitung und die genetisch-entwicklungsphysiolo- gisehe Bedeutung der Pigmente der Om- matin-und Ommingruppe (Ommochrome) bei den Arthropoden. Z. ind. Abst.-Vererb., 80:157-204, 1942. Caspari, E. Uber die Wii’kung eines pleiotropen Gens bei der Mehlmotte Ephestia kiihniella Z. Arch. Entwmech. Org., 130:253-281, 1933. Physiological action of eye color mu- tants in the moths Ephestia kiihniella and Ptychopoda seriata. Quart. Rev. Biol., 24:185-199, 1949. and J. Richards. Genic action. Yb. Carnegie Inst. Wash., 47:183-189, 1948. Hanser, G. Uber die Histogenese der Augen- pigmentgranula bei verschiedenen Rassen von Ephestia kiihniella Z. und Ptychopoda seriata Schrk. Z. ind. Abst.-Vererb., 82:74- 97, 1948. Problems of Origin and Migration of Pigment Cells in Fish. H. B. Goodrich. Wesleyan University, Middletown, Conn. In two papers published in 1915 Charles Stockard observed the origin of yolk sac chromatophores in Fundulus from chromato- blasts which migrated from the region of the closure of the blastopore and from the 18 Zoologica : Ne w York Zoological Society [35: 1 side of the embryo. In the light of the more recent “fate” maps of teleosts, these cells migrating from the blastopore would prob- ably be considered to have come from the presumptive mesodermal region of the germ ring. There does not at present appear to be available critical evidence to demonstrate the origin of fish chromatophores from the neural crest as in amphibia, birds and mam- mals. Transplants by Oppenheimer (1938) from the germ ring 180° removed from the embryonic shield gave rise to chromato- phores. Hybridization experiments (by Good- rich, unpublished) between Fundulus and Oryzias yielded some embryos without neural plate but with abundant chromato- phores on the yolk sac. Lopashov’s (1944) experiments on later stages may indicate an origin of chromatophores from the neural region. It seems not unlikely, however, that yolk sac chromatophores in the teleost may arise from remnants of the germ ring in the region of the closing blastopore. Other problems of pigmentation involve the determination of various cell types such as the melanophores, xanthophores and iri- docytes. Also sharply differentiated types of melanophores have been described as existing in the same fish (Platypoecilus, Oryzias). Evidence seems to indicate that all such cell types are determined prior to arrival in their final location. Patterns may for convenience be classified under micro-patterns and macro-patterns. The micro-patterns have to do with the spacing and arrangements of cells observ- able only under the microscope. In many cases there is a uniform distribution of cells over large areas. There is evidence, especi- ally from the goldfish, Carassius auratus (Goodrich and Anderson, 1939), of the ex- istence in the dermis of a reservoir of chro- matoblasts. Unpublished observations by Goodrich on Xiphophorus hellerii have traced the history of individual cells over a period of nine months. As cells disintegrate others appear in their places. In the variety of goldfish known as the Shubunkin, clusters of new melanophores erupt from time to time irregularly over the surface of the body. There may, however, occur a period of general lysis of melanophores over large areas. After this no further cells arise in the affected areas. Apparently the reservoir of chromatoblasts has also been destroyed. Formation of such clusters may be stimu- lated by radiation with ultra-violet light (Goodrich and Trinkaus, 1939), but this can not be accomplished after such a period of lysis. Nevertheless, such areas are not after such a period of destruction unfavor- able for the existence of melanophores as they may be transplanted into these regions (Goodrich and Nichols, I, 1933). Cells in micro-patterns are often clustered. It is not known whether this is a phenomenon of aggregation or of common origin by cell division as in the Shubunkin. The macro-patterns are those seen with- out magnification. Sometimes patterns arise as in the goldfish by destruction of some of the cells in a region of previous uniform distribution. During this past summer a study was made at the Bermuda Biological Station on the slippery-dick, Halichoeres bivittatus. This fish has longitudinal dark and light stripes. Tissue exchanges were made between these stripes. When made from light to dark the melanophores in- vaded the transplanted tissue. In the reverse transplants, the melanophores degenerated. In scales regenerating in the dark stripe, cells first appeared at the anterior and pos- terior edges and later invaded central re- gions. Experiments were tried under various light conditions, such as in dark aquaria, in diffuse light, bright sunlight with light background, and with dark background. The chromatophores invaded the white trans- planted tissue least rapidly in the dark and more rapidly in the order named above, with the fastest rate in fish kept in the sunlight on a dark background. In transplants from dark to light the cells usually disintegrated at the same rate under all conditions. There appeared to be a reservoir of melanoblasts in the dark stripe which is absent in the white stripe. Bibliography. Goodrich, H. B. and Anderson, Priscilla L. Variations of color pattern in hybrids of the goldfish, Carassius auratus. Biol. Bull., 77(2) :184-191, 1939. Goodrich, H. B. and Nichols, Rowena. Scale transplantation in the goldfish, Carassius auratus. I. Effects on chromatophores. Biol. Bull., 65(2) :253-259, 1933. Goodrich, H. B. and Trinkhaus, J. P. The differential effect of radiations on mende- lian phenotypes of the goldfish, Carassius auratus. Biol. Bull., 77(2) : 192-199, 1939. Lopashov, G. V. C. R. (Doklady) Acad. Sci., U. S. S. R., 44:169, 1944. Oppenheimer, Jane M. Potencies for differen- tiation in the teleostean germring. Jour. Exp. Zool., 79(2) :185-212, 1938. Stockard, Charles R. The origin of blood and vascular endothelium in embryos without a circulation of the blood and in the normal embryo. Amer. Jour. Anat., 18(2):227- 327, i915. A study of wandering mesenchymal cells on the living yolk-sac and their devel- opmental products: chromatophores, vascu- lar endothelium and blood cells. Amer. Jour. Anat., 18(3) :525-594, 1915. Discussion by PRISCILLA RASQUIN, American Mu- seum of Natural History, New York. It would be interesting to know whether the transplanted white scales would acquire melanophores if the host fish were kept in total darkness. It has been shown (1) that certain blind cave fishes ( Anoptichthys hubbsi) acquire melanophores after some months of living in the light, not only in the dermis and in the meninges but also in the peritoneum. No such peritoneal chromat- ophores have been noted in this species, 1950] Abstracts : Pigment Cell Conference 19 either when they are kept in darkness in the laboratory or when examined in the condition in which they are found in the total darkness of their natural habitat. It is difficult to see how these cells could have appeared in the peritoneum by a migration process, and they certainly have not ap- peared by an infective process such as Dr. Medawar has described, for they are com- pletely isolated cells. Dr. Goodrich’s idea of a “reservoir” of chromatoblasts in the fish dermis may also apply to the perineural and coelomic pigmentation. It is also interesting to note that the appearance of melanophores in these blind fish kept in the light is not mediated optically in any way. The fish lack a lens and retina, and many specimens also lack an optic nerve. Reference. 1. Rasquin, P. Progressive pigmentary re- gression in fishes associated with cave en- vironments. Zoologica, 32(4):35-42, 1947. The Origin of Modifying Genes that Influ- ence the Normal and Atypical Growth of Pigment Cells in Fishes.* Myron Gordon. New York Aquarium, New York Zoological Society. In natural populations of platyfishes and swordtails no melanotic tumors were dis- covered in more than 10,000 specimens. Under laboratory conditions, when a spotted member of one natural population is mated to that of another, tumors develop in their spotted hybrids. When a platyfish, Platypoecilus macula- tus, with the spotted (macromelanophore) pattern is mated with the related genus, the swordtail, Xiphophorus hellerii (with which it is sympatric in southern Mexican rivers), the spotted intergeneric hybrids develop melanomas. When the spotted Platypoecilus maculatus is mated with another platyfish species (whose natural habitat is a thousand miles to the north), Platypoecilus couchia- nus, from the Rio Grande, the spotted inter- specific hybrids also develop melanomas. When a spotted Platypoecilus maculatus is mated to a member of a nearby species, Platypoecilus variatus, from the Rio Pa- * Aided bv a grant from the National Cancer Institute of the United States Public Health Service. nuco, the degree of atypical pigment cell growth in the spotted hybrids is less severe. When a spotted Platypoecilus maculatus from the Rio Coatzacoalcos is mated to a member of the same species, Platypoecilus maculatus, from the Rio Jamapa, the intra- specific spotted hybrids show a definite mela- nosis very much like those between P. macu- latus and P. variatus. The significant point is that the parents in each type of mating (intergeneric, interspecific, intraspecific) represent different genetic populations. Me- lanomas or melanoses develop in hybrids in response to a genic imbalance between the genes for macromelanophores, five of which have been identified, and their growth regulators, specific macromelanophore modi- fying genes, two of which are known. These results may be explained by the same genetic principles that apply in ex- plaining the origin of organic diversity and the mechanisms of the origin of species. When a population of individuals belonging to a single species is separated into two aggregations, as by a geological accident or by other forces, and that separation is maintained rigidly so that interbreeding be- tween them is prevented, the two popula- tions will, in time, become genetically distinct. The isolated populations become different owing to the ever-occurring, muta- tional changes. The genic changes that ap- pear in members of the first population are not likely to be the same as the mutations that occur in the second. In time the random mutations accumulate in each of the two populations and eventually they make the two groups recognizably different. The ra- pidity of the genic changes depends on the size of the breeding population, the environ- ment, as well as upon time. As genetically interpreted, the piling up of these small changes, each meeting the challenge of na- tural selection successfully, constitutes the very beginning of the speciation process. There are six known natural populations of the platyfish, Platypoecilus maculatus, in the Atlantic coast rivers of Mexico, Gua- temala and British Honduras. Each of these six populations differs genetically in the frequencies of five macromelanophore pat- tern genes. Although these populations have been isolated geographically for more than 300,000 years, their morphological differ- ences are insufficient to indicate an evolu- tionary change at the subspecific level. They are, however, genetically distinct and these genetic differences can account for the genetic imbalance in the hybrid offspring of members representing different river popu- lations. Far back in geological time all platyfishes were probably like most fishes of their kind that are uniformly colored, unmarked by macromelanophores. In time the first “spot- ted” mutant appeared in a platyfish popu- lation, and transformed some previously ex- isting cells into macromelanophores. The nature of the mutation process is not yet 20 Zoologica: New York Zoological Society [35: 1 clear, but it is believed that mutations act like enzymes and affect the body chemically. Like most mutations, the macromelanophore mutation was probably harmful and most likely lethal, because most mutations upset an organism’s established genetic balance and this, in turn, upsets the biochemical processes in the developing organism. Prior to the spotted mutation’s successful establishment in the platyfish population, mutations which were to serve, in part, as modifiers and controllers of the activity of macromelanophores must have accumulated first, and then neutralized the lethal effects of the spotted mutants when they reoccurred. The platyfish had to be made ready geneti- cally for the coming of the spotted mutations by first accumulating macromelanophore controlling genes. When ready, the platy- fish “accepted” macromelanophores without danger to themselves. However, when a genetically balanced spotted platyfish from the Rio Coatzacoalcos is mated with another normal member from the Rio Jamapa, the new recombination of genetic modifiers of macromelanophore growth is not in balance in the spotted hybrids, and these large pig- ment cells grow atypically. The hereditary effects of macromelanophores can be demon- strated genetically, as well as the effects of gene modifiers which influence macromelano- phore growth patterns. The macromelano- phores are genetically labile cells subject to normal or typical growth by a series of modifying genes. References. Dobzhansky, Th. Genetics and origin of spe- cies. Columbia Univ. Press, 1941. Gordon, Myron. Biology of Melanomas. N. Y. Acad. Sci., Spec. Pub., 4:216-268, 1948. Animal Kingdom, 52:118, 1949. Endeavour, 9:26-34, 1950. Discussion by W. E. HESTON, National Cancer Institute, Bethesda, Md. The observations of Dr. Gordon are of especial interest to one studying the inheri- tance of cancer in mice because of the paral- lels to his observations that have occurred with mice. In general one can expect a greater variety of tumors to occur in hybrids between two inbred strains of mice than occur in either inbred strain. Little reported that in the F] hybrids resulting from cross- ing Mus bactrianus with strain C57 black (Mus musculus ) the incidence of non-epi- thelial tumors was 39.7%, whereas that for the C57 black mice was 13.2% and no such tumors were recorded for the Mus bactri- anus stock. In the history our experimental colony, 10 mice with tumors of the Har- derian gland have occurred, and all of these have been hybrids from a specific cross (strain CSH X strain C57 black). While this is a rare tumor, its occurrence in these spe- cific hybrids and not in either parent strain or any of the other strains of the colony is remarkable, The variation in degree of atypical pig- ment cell growth from melanosis to true melanomas effected by the genotypes of the different types of hybrid platyfish might also be compared with the variation in de- gree of expression of different genotypes affecting tumor formation in the mouse. This is particularly well illustrated with in- duced lung tumors. One observes from few to many nodules per animal, depending upon its genotype. Dr. Gordon’s observation of melanosis in certain types of hybrids compared with true melanomas in other types suggests that with the carcinogen 5, 9, 10 — trimethyl — 1, 2- — benzanthracene that produces pigmented foci in the skin of mice of certain strains, one might be able to pro- duce true melanomas in other strains or hybrid types with a more suitable genotype. Third Session: Physiological. Introduction. H. W. Chalkley. National Cancer Institute, Bethesda, Md. The papers in this group hold a place of basic interest. We have had the abnormal pigment cell and its destructive effect on the organism discussed from the pragmatic em- pirical medical approach. We have had the genetic control of the cell within the organ- ism discussed. Now we turn to the normal cell and its biologic behavior, considered as acting as an individual entity functioning cooperatively with other cells within the organism, its growth differentiation and regeneration, and the part it plays as a nor- mal component of the organism. It is this behavior that in the end will serve as a criterion for all theories of its structure or of the physical and chemical mechanisms posited as inherent in the cell. This is how it behaves as a biological entity and it is this behavior that we are seeking to understand by the biochemical or bio- physical approach. Its abnormal behavior, as for instance in cancer, will eventually also be comprehended as a part of this story. 1950] Abstracts : Pigment Cell Conference 21 Pigment Spread in Guinea Pigs. P. B. Medawak. University of Birmingham, England. It has been known for more than fifty years that the pigmentation of the super- ficial epidermis creeps from the colored into the white areas of recessively spotted guinea pigs, whether from grafts or, during normal life, across natural color boundaries. Analy- sis of the phenomenon has led to the sugges- tion that “pigment spread” is due to a permanent and serially propagable trans- formation of the dopa-negative, non-pig- mentary dendritic cells of white skin areas into the pigmentary dendritic cells (epi- dermal melanophores or melanoblasts) char- acteristic of colored areas. According to this hypothesis, pigment spread is made possible, anatomically, by the fact that dendritic cells form a partially syncytial web throughout the basal layer of the epidermis, whatever its color pattern, and whether it is pig- mented or not; and physiologically, by the fact that the epidermal dendritic cell is “cytocrine” in activity (Masson’s term) and can presumably “infect” neighboring den- dritic cells through a cytoplasmic anastomo- sis much as (in a trivial sense) it “infects” the Malpighian cells of the epidermis with melanin granules. It is a shortcoming of this hypothesis that the non-pigmentary dendritic cells (leuco- phores or “white melanophores”) of the colorless areas of spotted guinea pigs, on whose existence the truth of the hypothesis clearly depends, have not yet been seen in living skin. They are revealed by impregna- tion with acid gold chloride. This shortcom- ing is to some extent remedied by the fact that black pigmentation will encroach upon the red areas of spotted red-black or tricolor guinea pigs, though much more slowly than upon white. Red melanophores are visible in the living epidermis and are present in the same number and distribution as are black melanophores in black skin. Red skin trans- formed by pigment spread into black con- tains black melanophores only: either the red melanophores have been converted into black melanophores, or they have been by some unknown mechanism individually re- placed and destroyed by them. Epidermal epithelia lacking dendritic cells are non-infectable. Pigmentation may be propagated from a black skin area of one guinea pig to a white skin area of another guinea pig if a minute quantity of a suspension in Ringer’s solution of the basal layer cells of the black epidermis is “grafted” in such a way as to give the donor melanophores direct access to the “white” melanophores of the recipient. So initiated, foreign pigmentation has been ob- served to spread over an area as great as 500 mm2, and in certain proportion of cases appears to survive indefinitely. But foreign pigmentation of any area or time of stand- ing may be bleached out at will by trans- planting skin from the donor animal whose cells started the foreign pigmentation to the recipient in which it is spreading. This response is thought to be the consequence of an active immunization, and it is believed that foreign pigmentation, though certainly initiated by living foreign cells, is main- tained and propagated by the multiplication within host cells of some antigenically foreign cytoplasmic ingredient. “Infective” behavior by constituents of the cytoplasm cannot but be a phenomenon of the greatest rarity : if the authors’ inter- pretation is correct, it is made possible in the present instance by the cytocrine activity of dendritic cells, their semi-syncytial lay- out, and the co-existence of dendritic cells of different pigmentary activity in a single spotted guinea pig. References. Billingham, R. E. and Medawar, P. B. Pig- ment spread and cell heredity in guinea pigs’ skin. Heredity, 2:29-47, 1948. “Infective” transformations of cells. Brit. J. Cancer, 2:126-131, 1948. Pigment spread in mammalian skin: Serial propagation and immunity reactions. Heredity (in the press), 1950. Discussion by Morris Foster, Columbia University, New York If the semi-syncytial network provides the path by means of which infectively trans- forming cytoplasmic particles pass from black-pigment-producing dendritic cells into “white” or into red-pigment-producing cells, and if the same type of transmission mech- anism is responsible for “inoculating” Mal- pighian cells with melanin granules, then some infectively transforming particles should pass into the Malpighian cells along with the melanin granules, thereby infective- ly transforming the Malpighian cells as well. Since, however, Malpighian cells are not in- duced to form melanin, it is necessary to make subsidiary hypotheses. For example, one could assume that the cytoplasm in the Malpighian cell contains substances hostile to the action of the infectively transforming particles passing into it from the dendritic cell. If this be the case, however, then the cytoplasmic flow must always be in one direc- tion only, into the Malpighian cell, since a reverse flow would inhibit pigment produc- tion in the dendritic cell. Discussion by Ernst Caspari, Wesleyan University, Middletown, Conn. I was most interested to learn that the spreading phenomenon also occurs at the border of black and red areas, since this sit- utation might well provide critical evidence for the theory of transmission of pigment precursors. Under this theory it would be expected that cells containing both red and black granules would exist in the border re- gion. I wonder particularly whether it would be possible to observe this phenomenon di- 22 Zoologica: New York Zoological Society [35: 1 rectly in tissue cultures of black and red mel- anophores. Reply by Medawar. The question of whether “infected” red melanophores contain both red and black pig- mentary systems, or black alone, is certainly of first-rate importance. Although we are not very optimistic about the use of tissue cul- ture, we hope that really detailed histological analysis will help to solve the problem. Atypical Pigment Cell Differentiation in Embryonic Teleostean Grafts and Isolates. Jane M. Oppenheimer. Department of Biology, Bryn Mawr College, and Osborn Zoological Laboratory, Yale University, New Haven, Conn. The occasional development of red blood corpuscles and of chromatophores is a con- dition which sometimes obtains in otherwise nondifferentiating isolates and grafts from young teleostean embryos (Oppenheimer, 1949). Examples of such atypical differentia- tion have occurred in three separate series of experiments (Oppenheimer, 1936a, 1938) to be discussed below. Fundulus germ-ring grafts. In this series of experiments, portions of the germ-ring located 90° or 180° from the midline of the embryonic shield of Fundulus heteroclitus gastrulae (cf. fig. IB, Oppenheimer, 1938) were grafted to the embryonic shield or to the extra-embryonic membrane of gastrulae of the same species. In 37 cases either the whole grafts or portions of them continued development but failed to undergo typical histogenesis and differentiated no axial structures. In 16 of these, however, red blood corpuscles differentiated, and in 10 of these 16 grafts melanophores differentiated. In some cases, only a single melanophore was differentiated within the grafts, in others a larger number was present. Only those grafts are included among the 10 positive cases which contained melanophores located more or less centrally in the mass of cells ; grafts to which melanophores were applied only at the surface are excluded from consideration in the possibility that in these cases host melanophores might have migrated to cover the grafts. Only heteroplastic transplantation or the grafting of materials from vitally stained embryos can furnish direct proof that the melanophores are foi'med by graft rather than host cells in such experiments as these. Fortunately, however, supplementary data are available from other types of experiment which demonstrate that chromatophores are also similarly differentiated in some cases by isolates rather than transplants from young teleostean embryos. Epiplatys germ-ring isolates. The embryos of Epiplatys fasciolatus can be divided at late gastrula stages in such a way (cf. fig. 1A, Oppenheimer, 1938) that the portion of the germ-ring most remote from the embryonic shield can be isolated with part of the yolk from the portion of the egg containing the shield. Such isolates develop for several days in Ringer’s solution. Though in some cases they exhibit certain phenomena of growth which simulate the form of the tail, they undergo no histogenesis of axial structures. Of 12 such aggregates studied, one developed red blood corpuscles, one formed a single mel- anophore and a third differentiated a number of xanthophores. One explant of prospective tailbud region, isolated immediately after the closure of the blastopore, formed both melanophores and xanthophores although otherwise undifferentiated. Fundulus isolated blastoderms. Blasto- derms of Fundulus heteroclitus separated from the yolk during cleavage stages and cultivated in double-strength Holtfreter’s solution under some conditions form hyper- blastulae, masses of nondifferentiated cells provided with a large vesicle at one pole (Oppenheimer, 1936a). Of approximately 75 hyperblastulae studied, only one developed a single melanophore of typical size and con- figuration. Discussion and conclusions. The produc- tion of chromatophores by the grafts and iso- lates described above suggests that in the teleosts under certain experimental condi- tions pigment cells can be differentiated by cells which normally do not contribute to the teleostean counterpart of the neural crest. The 90° germ-ring of Fundulus normally con- tributes primarily mesoderm to the embryo (Oppenheimer, 1936b) ; and the 180° germ- ring is unrelated to nervous system forma- tion except insofar as it contributes to the tailbud blastema from which neural deriv- atives may later arise. While it is theoreti- cally possible, it is highly improbable that the chromatophores in these cases have been dif- ferentiated only by the particular cells des- tined to form them later after their passage through the tailbud blastema. The differen- tiation of pigment cells under these condi- tions is therefore presumably the result of what the experimental embryologists call bedeutungsfremde Selbstdiff erenzierung . This interpretation, however, is not in- compatible with the possibility that in tele- osts the chromatophores normally arise from cells corresponding to those of the neural crest of other vertebrates, as suggested by the results of the transplantation experi- ments of Lopashov (1944) on three species of teleosts, and in line with the neural crest 1950] Abstracts : Pigment Cell Coyiference 23 origin of pigment cells in amphibians (Du- Shane, 1935), birds (Dorris, 1938) and mammals (Rawles, 1947). It merely signifies that under certain abnormal conditions, othi:r than the usual cells can take over the function of pigment cell formation. Bibliography. Dorris, F. The production of pigment in vitro by chick neural crest. Arch. Entw.-mech., 138: 323-334, 1938. DuShane, G. P. An experimental study of the origin of pigment cells in Amphibia. J. Exp. Zool., 72: 1-31, 1935. Lopasiiov, G. V. Origin of pigment cells and visceral cartilage in teleosts. C. R. ( Do - klady) Acad. Sci. U. S. S. R., 44: 169-172, 1944. Oppenheimer, J. M. The development of iso- lated blastoderms of Fundulus heteroclitus. J. Exp. Zool., 72: 247-269, 1936 (a). Processes of localization in developing Fundulus. J. Exp. Zool., 73: 405-444, 1936 (b). Potencies for differentiation in the teleostean germ-ring. J. Exp. Zool., 79: 185-212, 1938. Atypical pigment-cell differentiation in embryonic teleostean grafts and isolates. Proc. Nat. Acad. Sci., 35:709-712, 1949. Rawles, M. E. Origin of pigment cells from the neural crest in the mouse embryo. Physiol. Zool., 20: 248-266, 1947. Regeneration of Neural Retina and Lens from Pigment Cells in the Eyes of Adult Salamanders.* L. S. Stone. Anatomical Laboratory, Yale University School of Medicine, and the Osborn Zoological Labora- tory, Yale University, New Haven, Conn. The neural retina of adult salamander eyes degenerates when deprived of its blood sup- ply either by severing the vessels which vas- cularize it (Stone and Chace, 1941) or by transplanting the entire eye (Stone and Zaur, 1940; Stone and Cole, 1943; Stone and Ellison, 1945). The only retinal cells which survive are those of the pigmented epithelial layer from which a new neural retina re- generates (Stone, 1949). This has been studied in detail in many grafted adult uro- dele eyes. Degeneration spreads throughout the neural retina during the first three weeks. It often proceeds at different rates in various parts of the same eye, allowing early stages of regeneration to begin in some areas * Aided by grants from the James Hudson Brown Fund and the United States Public Health Service. while adjacent regions are still degenerating. The true origin of the new neural retina is therefore more easily followed in cases where degeneration is simultaneous in all parts or where the entire neural retina is detached from the underlying retinal pigment layer by a gentle stream of Ringer’s solution and then removed intact through a broad dorsal slit in the eye. Degeneration of the neural retina in graft- ed eyes usually spreads rapidly until the ret- inal pigment cells are finally denuded and the cellular debris is absorbed above them. The pigment cells then become dense black oval or flattened bodies resembling those found soon after surgical removal of the en- tire neural retina. Now they take on tem- porarily another function, entering the crit- ical stages which can be followed step by step as they determine the origin of the new neu- ral retina. These pigment cells increase markedly in size. Their nuclei become distinct as they un- dergo mitosis. One daughter cell migrates in- ward, loses its pigment and with similar ones forms a sharply defined layer, which by fur- ther cell division gives rise to a new neural retina. The other daughter cell retains its pigment and later takes on the status of a retinal pigment cell when the neural retina above it has fully regenerated. The func- tional capacity of this cell is finally expressed by the migration of its pigment granules in a light-adapted eye as soon as the rods and cones above it are differentiated. When a small portion of the neural retina is excised, or is detached as a permanent elevated fold, regeneration is also called forth from the underlying retinal pigment cells. On the other hand the initial reaction of the pigment cell to a small retinal injury is one of rapid mitosis and mass migration in- to the wound, somewhat similar to that found in the retinal wounds of other vertebrates. However, in the urodele retina these cells later lose their pigment and differentiate in- to neural retinal tissue. Retinal pigment epi- thelium transplanted into the eye chambers also gives rise to neural retina. Since the early pigment cell changes in- volve replacement of a lost tissue the term “dedifferentiation” might be considered as it applies to them. This term is often loosely and obscurely applied to cells which are as- sumed to be taking part in regeneration, al- though up to that moment they are recog- nized as highly differentiated elements with special morphological and functional char- acteristics. Under the proper stimulus they are supposed to lose their special features and take on a role of supplying new cells that will later develop into similar or different elements. If the term is defined in this sense it can be said that the retinal pigment cell in the salamander eye is capable of dedifferen- tiation. Another pigment cell in the eye of Triturus salamanders may be considered in the same category, for when the lens is removed the 24 Zoologica: New York Zoological Society [35: 1 heavily pigmented cells along the free pupil- lary margin of the dorsal iris increase in size, become depigmented, undergo mitosis and form a vesicle that develops into a lens (Wachs, 1914, and others). When the latter becomes detached the cells in the dorsal iris regain their normal appearance. If varying amounts of dorsal iris are replaced by non- lens regenerating ventral iris, two widely separated lenses develop from the remaining dorsal pupillary margins after removal of the original lens. Secondary pupils experi- mentally produced in various regions of the dorsal iris by the insertion of pieces of plio- film or cornea show that potentiality for lens formation is quite widely distributed and not confined to the free pupillary margin (Stone and Vultee, 1949). These changes in the pigment cells can be experimentally inhibited by the following: 1 ) the presence of a transplanted normal lens of the same or another species (Stone, 1945) ; 2) the presence of a thirty-day lens regen- erate (Stone, 1943) ; 3) injections of aqueous humor from eyes containing lenses (Stone and Vultee, 1949) ; 4) the presence of some carcinogens. Literature Cited. Stone, L. S. Factors controlling lens regenera- tion from the dorsal iris in adult Triturus viridescens eyes. Proc. Soc. Exp. Biol, and Med., v. 54, 102-103, 1943. Heteroplastic lens grafts related to factors inhibiting lens regeneration in Tri- turus. Proc. Soc. Exp. Biol, and Med., v. 60, 10, 1945. - Return of vision and functional polar- ization in the retinae of transplanted sala- mander eyes. Exp. Cell Research, Sup. 1, 582-584. {Proc. 6th Intern. Cong. Exp. Cytol., 191+7) , 1949. Stone, L. S. and Chace, R. R. Experimental studies on the regenerating lens and the eye in adult Triturus viridescens. Anat. Rec., v. 79, 333-348, 1941. Stone, L. S. and Cole, C. H. Grafted eyes of young and old adult salamanders (Ambly- stoma punctatum) showing return of vi- sion. Yale J. Biol, and Med., v. 15, 735-756, 1943. Stone, L. S. and Ellison, F. S. Return of vi- sion in eyes exchanged between adult sala- manders of different species. J. Exp. Zool., v. 100, 217-227, 1945. Stone, L. S. and Vultee, J. H. Inhibition and release of lens regeneration in the dorsal iris of Triturus v. viridescens. Anat. Rec., 103, 144-145, 1949. Stone, L. S. and Zaur, I. S. Reimplantation and transplantation of adult eyes in the sala- mander ( Triturus viridescens ) with return of vision. J. Exp. Zool., v. 85, 243-269, 1940. Wachs, H. Neue Versuche Zur Wolff’schen Linsenregeneration. Arch. f. Entwmech. d. Org., Bd. 39, 384-451, 1914. X-Ray Effects on Mouse Pigmentation as Related to Melanoblast Distribution. Herman B. Chase & Virginia W. Smith. Department of Biology, Brown University, Providence, R. 1. Several features of the x-ray-induced grey- ing response suggest certain properties of the pigmentation system. Follicles react largely as independent units, producing fully pigmented hairs (no apparent gross effect), white hairs, or infrequently mosaics. The ef- fect is permanent or toward more white hairs in successive hair generations of the partic- ular follicle (Chase, Quastler, and Skaggs, 1947; Chase, 1949). The percentage of fol- licles producing only white hairs after treat- ment increases with dose (200-1000 r) and with dose-rate. The percentage for a given dose is greater for follicles treated in catagen and telogen stages and for follicles of the smaller hair types (Chase, 1949; Chase and Rauch, in press) . In a mosaic some of the medullary and cortical cells lack pigment granules entirely, some have a full comple- ment, and some have a reduced number of granules. These various cells are arranged irregularly throughout the length of the hair. With increasing size of hairs (such as awls in the mouse, most of the hairs in rabbits and cats, all of the hairs in guinea pigs), the fol- licles are less “sensitive” to a given dose of x-radiation, produce a greater frequency of mosaics, and display less difference in re- sponse when treated in telogen or in anagen phases. It would seem that susceptible ele- ments must be few in number, very few in smaller follicles. At beginning of anagen there would be a moderate increase of these elements but little, if any, further increase during the anagen phase (2 to 17 days post- plucking) . After the initial supply for the original follicle invagination, there would be no new invasion for subsequent hair genera- tions. Following Masson’s (1948) definitions the dendritic cells which eventually produce pig- ment are termed melanoblasts whereas the recipient cells, if any, are termed melano- phores. Melanoblasts generally ai*e not ob- served because of their fragility with certain standard histological methods and because they are largely obscured after the matrix cells become pigmented. Phase microscopy with the Spencer B minus contrast low, oil immersion objective on unstained frozen sec- tions has proved most revealing. In young mice, melanoblasts of the basal layer of the skin epidermis can be seen to be incorporated in the original invaginations of newly- forming follicles. In early anagen stages of subsequent hair generations, melanoblasts are found in the permanent external sheath (continuous with basal layer of skin epi- dermis) or in the derivative basal layer of the bulb. They become melanogenic with fine dispersed granules and send long dendritic processes to the matrix cells. Later, begin- ning about 6 days post-plucking, the “inoc- 1950] Abstracts : Pigment Cell Conference 25 ulated” matrix cells (— melanophores) ex- hibit characteristic dense pigmentation. The keratinized hair shaft arises from these matrix cells. Pigmented matrix cells seldom divide, but new cells arise from the prolifer- ating lower bulb and receive pigment as they pass by the melanoblast processes of the up- per bulb. As few as 4 melanoblasts have been observed to supply the stream of matrix cells of a small follicle (zigzag) and the presence of only one or two results in a mosaic. From histological and x-radiation evidence it appears that the “reservoir” of melano- blasts is within the epidermal follicle (cf. Taylor, 1949). Dermal melanophores appar- ently do not contribute to pigmentation in the hair follicle. The epidermal melanoblasts are the original granule-forming cells, but in the recipient matrix cells, the characteristic shape and color of the phenotype are further imposed upon the granules. Whereas biotin deficiency, for instance, causes a failure of melanin-formation on these granule sites (Chase and Rauch, in press), x-radiation destroys the source of supply of granules, namely, the epidermal melanoblasts of the follicle. (Work supported by grant-in-aid from American Cancer Society, recommended by Committee on Growth of National Research Council). References. Chase, H. B. Greying of hair. I. Effects pro- duced by single doses of x-rays on mice. J. Morph., 84: 57-80, 1949. Chase, H. B., H. Quastler and L. S. Skaggs. Biological evaluation of 20 million volt roentgen rays. II. Decoloration of hair in mice. Am. J. Roent. and Rad. Ther., 57 : 359-361, 1947. Chase, H. B. and H. Rauch. Greying of hair. II. Response of individual hairs in mice to variations in x-radiation. In press. Masson, P. Pigment cells in man. Spec. Publ. N. Y. Acad. Sci., 4: 15-37, 1948. Taylor, A. C. Survival of rat skin and changes in hair pigmentation following freezing. /. Exp. Zool., 110: 77-111, 1949. A Comparative Colorimetric Study of Dopa-melanin Formation by Melanomas and Pigmented Skins. Ruth E. Shrader & Carroll A. Pfeiffer. Department of Anatomy, Yale University School of Medicine, New Haven, Conn. The costo-vertebral pigmented skin spot of the Syrian hamster, which contains both intradermal dentritic pigmented cells and exceptionally large pigmented hair roots1 has proved to be consistently dopa-negative in histological sections prepared according to several standard technical procedures2-3. A method utilizing photo-electric colorimetric evaluation of dopa-melanin has been evolved which permits comparison of this pigmented tissue with other tissues which can be shown by histological means to be dopa-positive. Skin specimens were surgically removed, freed of hair and unpigmented skin, chilled to 4° C., minced in a few drops of phosphate buffer, and rapidly weighed on small squares of clean coverslip using a microtorsion bal- ance. Increasing weights of minced tis- sues were placed in centrifuge tubes contain- ing 2 ml. of freshly prepared 1:1000 1-dopa solution, buffered to pH 7.4, and thoroughly mixed. After four hours’ incubation at 37° C. the tubes were centrifuged at 25,000 rpm. for 15 minutes. The supernatant fluid was de- canted, diluted to a standard volume, and kept in chilled colorimeter tubes until read on a Klett-Summerson photo-electric colorime- ter using a KS-42 filter. The auto-oxidative rate of dopa was determined for each experi- ment by means of a control tube containing only buffered dopa. The control tube was given a value of 100% and the values ob- tained for the different tissue samples were compared with it. Using this technique, experiments were done on tissues from the Cloudman mouse melanoma and from human melanomas. These tumor tissues gave almost straight line curves with the activity (color forma- tion) increasing directly with the weight of the tissue samples. The results agree favor- ably with those obtained from Warburg studies of the enzyme activity of such mela- noma tissues4. A number of types of skin were studied by this method. These included skin from the neck, flank and pigment spot of the hamster, pigmented nipples from guinea pigs, neck and flank skin from mice, adult and infant human skin, including depigmented and hyper-pigmented negro skin. In contrast to the straight line activity curve of the tumor tissue, these skins showed a biphasic activity curve with the final inclination in the down- ward direction. This characteristic of pig- mented skins has been interpreted to indicate the presence of inhibitory substances which interfere with the formation of dopa-mela- nin. This observation is in agreement with previous work showing such inhibitory sub- stances to be present in the skins of guinea pigs5, rabbits6-7 and man8-9. When the conditions of the experiment were changed so that the weight of the tissue was held constant and the concentration, but not the volume, of the dopa solution was in- creased, a curve was obtained which is sug- gestive of the classic “substrate competition” curve. The addition of a sulfhydril substance, glutathione10 increased the inhibition of color formation in dilute dopa solutions but did not significantly affect higher concentra- tions. 26 Zoologica: New York Zoological Society [35: 1 The effect of hormonal stimulation of pig- mentation upon the ability of the stimulated skin to form dopa-melanin was investigated, using both the pigmented nipple of the guinea pig and the hamster pigment spot. In both instances, tissues which had been caused to hypertrophy and blacken under hormonal influence gave increased color formation with dopa as contrasted with untreated control tis- sues, but the essential biphasic shape of the curves was unchanged. References. 1. Shrader, R. E. Development of the di- morphic pigment spot of the Syrian ham- ter. Anat. Rec., 105:561, 1949. 2. Laidlaw, G. F. The dopa reaction in normal histological technic. Anat. Rec., 53 :399, 1932. 3. Becker, S. W. Cutaneous melanoblasts as studied by the paraffin dopa technic. J. In- vest. Derm,., 5:463, 1942. 4. Greenstein, J. P. and Algire, G. H. Com- parative oxidase activity of melanotic -+• amelanotic melanomas. J. Nat. Can. Inst., 5:35, 1944. 5. Schaaf, F. Manometrische Vergleichsunter- suchungen mit Presssaften aus Weisser und Pigmentierter Meerschweinchenhaut. Arch, f. Dermatol., u. Syph., 176:646, 1938. 6. Pugh, C. E. M. Tyrosinase from the skin of certain black rabbits. Bioch. J., 27 :475, 1933. 7. Roller, P. On pigment formation in the D- black rabbit. J. of Genetics, 22:103, 1930. 8. Rothman, S., Krysa, H. F. and Smiljanic, A. M. Inhibitory action of human epidermis on melanin formation. Proc. Soc. Exp. Biol, and Med., 62:208, 1946. 9. Flesch, P. Inhibitory action of extracts of mammalian skin on pigment formation. Proc. Soc.. Exp. Biol, and Med., 70:136, 1949. Discussion by MORRIS FOSTER, Columbia University, New York. Two important sources of error are in- herent in the techniques used: (1) The col- orimetric method, whereby pigment produc- tion rate is equated with oxidase activity. Such reasoning could be erroneous. For example, a reducing agent reacting with a substance produced after the oxidation of dopa could inhibit pigment production with- out affecting dopa oxidase activity ; although, on the basis of colorimetric criteria, inhibi- tion or lack of enzyme activity would be inferred. (2) The use of dopa as a substrate, since dopa can be oxidized via both the phen- olase and cytochrome systems, whereas tyro- sine is not affected by the latter. Thus enzymatic oxidation of dopa could be wholly or partly attributed to the cytochrome sys- tem rather than to the phenolase system. (See paper by Hesselback et al.) A more direct measure of phenolase ac- tivity could be obtained by using tyrosine as the substrate and by measuring the rate of oxygen consumption in a Warburg respir- ometer. In regard to the hamster pigment spot, an explanation of increasingly inhibited oxida- tive activity with increasing weight of pig- ment spot sample is difficult to make, since larger samples should also give larger amounts of enzyme, resulting in increased rather than decreased activity. Reply by Shrader 8s Pfeiffer. We would like to strongly emphazise that we do not know the enzyme system or systems involved in the production of dopa-melanin in these experiments. Our experiments were not designed primarily to elucidate the mech- anisms of melanin formation but were in- tended to explain why the dopa reaction could not be obtained on sectioned hamster pigment spot. What we measured was the amount of dopa-melanin that was produced under experimental conditions as similar as possible to those of histological methods. The use of tyrosine as a substrate, or the War- burg apparatus as a method, would not have served to answer our original question. We do, however, feel that the results obtained justify the assumption that the colorimetric measurement of dopa-melanin is a valid cri- teria for an estimation of the pigment form- ing ability of the tissue. In regard to the increase in inhibition which occurs as hamster pigment spot tissue weight is increased, we can only say that this is a characteristic of all skins tested and that we have postulated that it represents the re- sults of some type of competitive action be- tween the enzyme and inhibitor systems for utilization of the substrate. This suggested explanation gains further support from the fact that such inhibition is more marked when the quantity of available substrate is low and can be overcome by the addition of more substrate. Fourth Session: Biochemical and Biophysical. Introduction. Jesse P. Greenstein. National Cancer Institute, Bethesda, Md. Melanin chemistry may be roughly divided into two general approaches: (a) the char- acter of the pigment noted in natural sources, i.e., the intracellular granules in skin, hair, plants, etc., and (b) the nature of the poly- meric pigments obtained by the in vitro oxi- dation of such amino acids as tyrosine and dihydroxyphenylalanine, and the relation, if not the identity, of such pigments to those noted in natural sources. As yet, no clearcut 1950] Abstracts : Pigment Cell Conference 27 evidence has linked the synthetic pigments under (b) with the pigments derived from natural sources, although the presence of dopa oxidase in melanin-containing melano- blasts, and the absence of this enzyme from amelanotic tissues are more than suggestive of a connection between the two. The natural pigment is generally associ- ated with protein in the form of cytoplasmic granules. These granules themselves contain dopa oxidase activity, and an essential prob- lem which requires clarification is whether the pigment may be preformed from small molecular weight precursors followed by con- jugation with one or more proteins in the cytoplasmic complex or whether it is derived from certain potentially chromophoric groups (tyrosine residues) existing at the surface of the protein molecules of the com- plex. It is a matter of some difficulty to free the pigment completely of protein, and even when this is accomplished the vigorous meth- ods required raise some doubt as to whether the melanin has not been altered to some extent. Indeed, the characterization of the particulate, sub-unit components of any cell, and the separation of the various members of these particulate components, demand techniques which for the most part are not yet adequately developed. However, advances in this, as in other fields, may come from many able quarters. That such advances have been made in several approaches to the prob- lem of pigment chemistry are shown in the excellent papers which follow. Mammalian Melanin Formation: I. Biochemical Studies. A. Bunsen Lerner, University Hospital , University of Michigan, Ann Arbor, Mich. & Thomas B. Fitzpatrick, Fellow in Dermatology and Syphilology, Mayo Foundation, Rochester, Minn. Investigation of the biochemistry of mel- anin formation in plants, insects and marine animals had shown that the enzyme, tyro- sinase, catalyzes the oxidation of L-tyrosine to dihydroxyphenyl-L-alanine (dopa), and then the oxidation of dopa to melanin. Until recently, however, the presence of tyrosinase in mammalian tissue had not been demon- strated conclusively, and it was believed that melanin in mammalian tissue is formed by a mechanism different from that operating in other species. Histochemical evidence indi- cated that mammalian skin contains an en- zyme, “dopa oxidase,’’ which catalyses the oxidation of dopa, but not tyrosine, to mel- anin. Largely as a result of these beliefs, two separate hypotheses of melanogenesis evolved: 1) melanin formation in insects and plants was associated with tyrosinase, while 2) melanin production in mammalian skin was associated with dopa oxidase. Re- cent evidence has shown that these separate concepts can now be merged into a single hy- pothesis to account for melanin formation in man, lower animals, insects and plants (1). It is now known that the original distinction between tyrosinase and dopa oxidase is no longer valid. Hence, it is suggested that the single term, tyrosinase, should be used in- stead of the separate terms, tyrosinase and dopa oxidase (2). While tyrosinase obtained from different species has some unique properties, depend- ing on the particular source, three character- istics are common to tyrosinase, under proper conditions, regardless of its origin: 1) all catalyze the oxidation of tyrosine to melanin, 2) the enzymatic reaction with the monohydroxyphenyl compound is catalyzed by some orthodihydroxyphenyl compound (dopa, catechol, etc.), and 3) copper is associated with the activity of the enzyme. When tyrosine and tyrosinase are allowed to react in the presence of oxygen, there is often a lag period before oxidation of tyro- sine begins. This lag interval is referred to as the “induction period.’’ Small amounts of dopa are very effective in shortening the in- duction period in the tyrosine-tyrosinase re- action. For mammalian tyrosinase there is a linear relationship between the negative logarithm of the dopa concentration and the induction period. Compounds related struc- turally to dopa, such as epinephrine, catechol and the like can shorten the induction period, but not nearly so effectively as dopa does. When dopa itself is used as a substitute for tyrosine, there is no induction period. Dopa participates in the tyrosine-tyrosin- ase reaction in at least three ways: 1) dopa is formed from tyrosine, 2) dopa catalyses the tyrosine-tyrosinase reaction, and 3) some amount of dopa is reformed during the conversion of dopa to melanin. Various substances inhibit melanin form- ation in vitro and in vivo. The mechanism of inhibition is dependent upon the particular step in the tyrosinase-catalyzed series of re- actions in which the conversion of tyrosine to dopa and eventually to melanin is blocked. References. 1. Lerner, A. B. and Fitzpatrick, T. B. Bio- chemistry of Melanin Formation, Physiol. Rev., 30:91-126, Jan., 1950. 2. Lerner, A. B., Fitzpatrick, T. B., Calkins, Evan and Summerson, W. H. Mammalian Tyrosinase: Preparation and Properties. J. Biol. Chem., 178:185-195 (Mar.), 1949. 28 Zoological New York Zoological Society [35: 1 Mammalian Melanin Formation: II. Histochemical Studies. Thomas B. Fitzpatrick, Fellow in Dermatology and Syphilology, Mayo Foundation, Rochester, Minn. & A. Bunsen Lerner, University Hospital, University of Michigan, Ann Arbor, Mich. As early as 1901 von Fiirth advanced the hypothesis that melanin formation is the re- sult of the action of an intracellular oxidase on aromatic or chromogen groups in certain protein molecules. Bloch, a Swiss derma- tologist, stimulated by this hypothesis, attempted to prove it by experimental methods. He selected the naturally occur- ring amino acid, dihydroxyphenyl-L-alanine (dopa), as the substrate for his histochemi- cal studies. Bloch immersed frozen-fixed sections of human skin in a 1 :1,000 solution of dopa buf- fered to pH 7.3, and noted that after twenty- four hours at room temperature black (mel- anin) granules were deposited in the cyto- plasm of cells located in the basal layer of the epidermis. These specialized cells, which he called “ melanoblasts,” are located at the epidermal-dermal junction, and were consi- dered by Bloch to be the site of melanin form- ation. Further evidence was provided by Bloch and his co-workers to support the hy- pothesis that the melanoblasts contained an enzyme, dopa oxidase, which catalyzed the oxidation of dopa to melanin within the cell. The dopa oxidase hypothesis of human mel- anin formation as advanced by Bloch never has been generally accepted as the explana- tion for the chemical mechanism underlying melanin formation. The main criticism has been that dopa never has been demonstrated in mammalian tissue. In the past eight years it has been firmly established that extracts from mouse, horse and human melanomas contain tyrosinase and dopa oxidase activities (1) .Furthermore, it has been shown that under certain condi- tions no true distinction can be made between tyrosinase and dopa oxidase activities in mammalian tissue. At the time Bloch carried out his important histochemical studies, little was known about the optimal conditions for the enzymatic oxidation of tyrosine. This may account for the fact that Bloch, working with mammalian tissue slices, obtained mel- anin formation from dopa but not from tyro- sine. In some recent histochemical experiments the authors, in collaboration with S. William Becker, Jr., demonstrated the formation of melanin from tyrosine in human white skin which had been irradiated with ultraviolet radiant energy for seven days before ex- cision. Tissue slices cut from the biopsy material were incubated in tyrosine solutions at pH 7.1 for twenty-four to forty-eight hours. In paraffin sections of this material, there were seen large dendritic melanoblasts containing melanin granules in their cyto- plasm, identical in their morphology to the “dopa-positive” cells obtained by Bloch. The catalytic effect of these cells on the oxidation of tyrosine to melanin is absent when the tis- sue slices are heated for ten minutes at 100°C. or when the tissue slices are incubated with sodium diethyldithiocarbamate, a cop- per-binding compound, for 6 hours prior to incubation in tyrosine. Since tyrosine, in con- trast to dopa, which readily auto-oxidizes, is a stable amino acid which does not oxidize spontaneously to melanin in vitro, it is likely that the melanoblasts of human skin contain an intracellular oxidase, tyrosinase, similar to the enzyme described previously. The en- zyme apparently exists in human skin in a partially inhibited state, and can be activated by ultraviolet radiant energy. The mech- anism of this activation is not fully under- stood, but the inactivation of epidermal sulf- hydryl by the ionizing radiation appears to play an important part. The data offer strong evidence that the pigment precursor in human melanin formation is tyrosine and not dopa. The elaboration of melanin pigment in the epidermal melanoblast ordinarily depends on the available concentration of three sub- stances : 1) The enzyme tyrosinase: a copper-pro- tein complex attached to particles in the cytoplasm of the melanoblast. 2) A suitable substrate : usually tyrosine or dopa. 3) Molecular oxygen. If any of these substances is absent, the for- mation of melanin is impaired. The reaction of the three basic substances, tyrosinase, substrate and molecular oxygen, is controlled by several physico-chemical fac- tors which determine the rate of melanin formation : 1 ) A catalytic substance, usually dopa, which can accelerate the tyrosine- tyrosinase reaction. 2) Chemical groups which normally in- hibit copper enzymes, for example, sulfhydryl groups, normally found in the epidermis. 3) Physical and chemical factors, such as temperature, hydrogen ion concen- tration and oxidation-reduction po- tentials. The quantity of melanin produced by the cell depends on the overall balance of these dif- ferent forces. Reference. 1. Lerner, A. B. and Fitzpatrick, T. B. Bio- chemistry of Melanin Formation. Physiol. Rev., 30:91-126, Jan., 1950. 1950] Abstracts : Pigment Cell Conference 29 Observations on the Structure, Derivation and Nature of Melanin. E. Meirowsky, L. W. Freeman & R. B. Fischer. Indiana University School of Medicine, Indianapolis, Ind. Developments in enzyme and cellular chemistry have clearly demonstrated that the nucleus is the center of metabolism of nu- cleoproteins. The most intense production of nucleoprotein appears to take place in the nucleolus, and the material diffuses to, and through, the nuclear membrane into the cyto- plasm. As shown by DuBuy and his co- workers, the melanin of the S91 melanoma contains approximately 20% nucleic acids. This would lead to the suspicion that melanin is closely allied to, or identical with, nucleo- proteins, especially when the morphological picture is taken into account. A series of color slides of pertinent cells will be presented, enlarged 540 times, which give a pictorial representation of the follow- ing pattern: Melanin first appears in the nucleolus and spreads to the linin framework of the nucleus. It appears in intranuclear vacuoles and finally involves the entire nu- cleus and the membrane. It leaves the nucleus by way of the chromidial derivatives. We are of the opinion that this demonstra- tion clearly indicates that the source of mel- anin formation is within the nuclear mate- rial. It appears that the life history of the melanin — as demonstrated — closely parallels that of chromatin and that nucleoprotein is replaced by melanoprotein. It can be noted from the pictures to be pre- sented that melanized nuclei show a consider- able variation in size, by measurement, from 7X7 micra up to 16 X 36 micra. This raises the question as to whether these larger nuclei result from further growth of the melanized chromatin, or from propagation of individual melanin granules. Their main features, so far observed, in- clude the characteristics of membranes, an inner body showing amitotic division; “X” bodies which could not be identified, which, however, are an apparatus of propagation, connected with the process of budding and of buckling of the membrane. This process leads to buds connected with the mother-body by fine strands or a solid stem. These buds also show the “X” structure and may undergo severance from the mother body ; free buds which no longer can be differentiated from melanin granules; and cloudy masses spread- ing from melanin granules in which new granules make their appearance. In general, the youngest buds are colorless and show all transitions to deep coloring, just as described by Smith in his cultures of the pigmented epithelium of the eye. We have, on occasion, succeeded in differentially stain- ing the body and the bud by neutral red as a basic dye and light green as an acid dye. The buds have been seen to stain acidophilic with light green. Supravital staining with borax- methylene blue applied to unfixed specimens by Scott’s method often shows the young buds to be colorless. These young buds ap- pear to be without a membrane and as they develop, they stain blue and a membrane makes its appearance. The most striking feature of all of these buds is the presence of the “X” structure. Entirely the same buds with “X” structures are also found, in very great numbers, sepa- rated from melanin granules freely in the specimen. While further work is warranted, in order to understand the foregoing observations in their entirety, we believe that the outstand- ing features of melanin granules in mel- anomas and mammalian choroids are com- patible with those of a living, dividing, and budding structure derived from living, divid- ing and budding nuclear material. References on the Main Investigations of the Relationship: Chromatin and Melanin. A. Nucleolus and Melanin. Roessle, R. Der Pigmentierungsvorgang im Melanosarcom. Ztschr. Krebsforsch., 2:291- 332, 1904. “The most remarkable observation is the ex- trusion of nucleolar material into the cyto- plasm and its transformation into mela- nin.” Meirowsky, E. Die Entstehung des Oberhaut- pigments aus der Substanz der Kernkoer- perchen. Monatshefte Prakt. Derm. 43:155- 163, 1906. Ueber den Ursprung des Mela- notischen Pigments der Haut und des Auges. Publ. W. Klinkhardt, Leipzig, 1908. “Formation of pigment is preceded by an augmentation of nucleolar material irre- spective of the material under observation (skin of man and rabbits after irradiation, embryonic skin, triton larves, pigment of the eye, nevi melanomas). The nucleolar material is transformed into melanin.” Goda, T. Cytoplasmic Inclusions of Amphibian Cells with Special Reference to Melanin Formation. J. Faculty of Science Imperial University, Tokyo. Section IV, Zoology, First Part, Vol. II, 51-122, 1928-1931. “The melanin granules arise from the nu- cleus by the following process: nucleolar bodies — nucleoliquid drops — melanogran- ules.” Schultz, O. T. The Formation of Pigment by the Dermal Chromatophores. J. Med. Res., 26 (new series 21) : 65-77, 1912. “. . . chromatin is thrown out of the nucleus into the cytoplasm, leading to the formation of a functional chromatin net. The chroma- tin present in the cytoplasm becomes changed into a material which has the staining reactions of nucleolar substance. Further change leads to the formation of this material into pigment.” “(Pigment) ... is the product of the spe- cialized physiological activity of the cell by which nuclear derivates are transformed directly into pigment.” 30 Zoologica: New York Zoological Society [35: 1 B. Melanization of Nuclei, Chromidia and Mitoses. VON Szily, A. Ueber die Entstehung des Melano- tischen Pigmentes im Auge der Wirbeltier- embryonen und im Choroidealsarkomen. Arch. Mikr. Anat., 77: 1-70, 1911. “The colorless stromata of the melanin of metazoa originate in all cases examined ex- clusively in the nucleus. Their direct origin from the chromatin of the nucleus and their transformation into the cytoplasm can ex- actly be followed up. These ‘chromidia’ change over into melanin under the influ- ence of ferments.” Jeliaskowa Paspalewa, A. Cytologische Un- tersuchungen ueber die Entstehung des Melanotischen Pigments. Ztschr. Wissen- schaftl. Zoologie, 137: 365-402, 1930. “The observation of a total melanization of chromosomes makes it impossible to con- test the origin of melanin from chromatin.” Ludford, R. J. Nuclear Activity during Melano- sis with Special Reference to Melanin For- mation in a Melanotic Sarcoma. J. Royal Micr. Soc., 13-28, 1924. The General and Experimental Cytol- ogy of Cancer. J. Micr. Soc., 249-292, Sept. 1925. “Melanosis occurring during mitosis and following chromatin extrusion, nuclear fragmentation, nuclear budding, karrhyor- rhexis, and pycnosis.” Ludford believes that the greatest part of the melanin is formed in the cytoplasm under the influ- ence of the Golgi apparatus which is con- cerned with enzyme formation. C. Melanization of Intranuclear Vacuoles. Meirowsky, E. Die Entstehung des Oberhaut- pigments aus der Substanz der Kernkoer- perchen. Montsh. Prakt. Derm., 43: 155- 163, 1906. “A cavity is formed in the nucleus with melanin granules at its wall. Through the ruptured wall pigment is extruded into the protoplasm.” Ludford, R. J. Nuclear Activity During Melano- sis with Special Reference to Melanin For- mation in a Melanotic Sarcoma. J. Roy. Micr. Soc., 13-28, 1924. Summary 8. “Melanin is often formed in- side the nucleus, generally in intranuclear vacuoles.” Apitz, K. Ueber die Pigmentbildung in den Zellkernen Melanotischer Geschwueslte. Virchow Arch., 300: 89-112, 1937. “The nucleus forms the mother substance of melanin.” “The origin of the nuclear melanin takes place within vacuoles owing to the retention of physiological secretion product of the nucleus.” D. Biochemistry of Nucleoproteins. DeRobertis, E. D. P., Nowinski, W. M. and Saez, F. A. General Cytology. Publ. W. B. Saunders, Philadelphia and London, 1938. E. Budding Processes in Individual Melanin Granules. As far as known, no references are available. Evidence for the Mitochrondrial Nature and Function of Melanin Granules. Mark Woods, Herman DuBuy & Dean Burk. National Institutes of Health, Bethesda, Md. Melanin occurs in the cytoplasmic granules of the melanoblasts of the Harding-Passey and Cloudman S91 mouse melanomas, and also, of course, in the phagocytes of these tumors. In both tumors, cells occur in which all, or nearly all, of the visible cytoplasmic granules are melanized. Colorless granules of similar size occur in amelanotic or par- tially amelanotic cells of the S91 tumor and also in the derived Algire partially amelan- otic S91A melanoma. Some colorless granules usually occur in the perikaryon of the Hard- ing-Passey melanoblast. The S91A amelan- otic melanoma sometimes contains cells in which the cytoplasmic granules are very slightly melanized. The cytoplasmic granules (melanized and non-melanized) are the only structures which stain with Janus Green B in the absence of nuclear staining by this dye. The granule staining is reversibly dependent upon oxygen tension. No other structures resembling mitochondria are visible in the cells. On alka- line hydrolysis, both melanized and non- melanized granules yield solutions with strong ultraviolet absorption at 2580A, and contain organic phosphorus and pentose. Centrifugally isolated cytoplasmic granules of amelanotic melanoma cells possess en- zymic activities characteristic of mitochon- dria of other origin (e.g., from liver, kidney, heart). Aerobically, these activities include the cytochrome oxidase and succinic oxidase systems. Anaerobically, the cytoplasmic granules possess glycolytic activities also comparable to those of typical mitochondria. Centrifugally isolated melanized granules from both Harding-Passey and S91 mela- nomas possess not only all of the enzymic ac- tivities found in the amelanotic granules, but also dopa oxidase activity. On the basis of the foregoing morphologic, chemical and en- zymic data it is concluded that the melanized and non-melanized granules of these mouse melanomas are mitochondria. Mitochondria are fundamental structures in both animal and plant cells, and in plants it is now well established that they derive from pre-existing mitochondria, possess a complex hereditary system (chondriogenes) , and are capable of mutation. In mutant states plant mitochondria display abnormal en- zymic activities. Self-duplication of these ab- normal mitochondria results in development of neoplasia. Plant mitochondria may be spe- cifically modified by certain viruses that re- sult in neoplasias very similar to those caused by mutant chondriogenes. The behavior of normal, mutant or virus-modified mitochon- dria may also be affected by specific nuclear genes. 1950] Abstracts: Pigment Cell Conference 31 Bibliography. 1. Woods, M. W., DuBuy, H. G., Burk, Dean and Hesselbach, M. L. Cytological studies on the nature of the cytoplasmic particulates in the Cloudman S91 mouse melanoma, J. N. C. Inst., 9: 311-323, 1949. 2. DuBuy, H. G., Woods, M. W., Burk, Dean and Lackey, Mary D. Enzymatic activities of isolated amelanotic and melanotic gran- ules of mouse melanomas and a suggested relationship to mitochondria. Ibid., 9: 325- 336, 1949. 3. DuBuy, H. G. and Woods, M. W. A possible common mitochondrial origin of the varie- gational and virus diseases in plants and cancers in animals. In Am. Assoc. Advance- ment Sc. Res. Conf. on Cancer, pp. 162-169. Lancaster press (1945). 4. Woods, M. W. and DuBuy, H. G. Cytoplas- mic diseases and cancer. Science, 102: 591- 593, 1945. Oxidative Activities of Mouse Melanomas with Reference to Melanization. Marie L. Hesselbach, Mark Woods & Dean Burk. National Institutes of Health, Bethesda, Md. Dopa Oxidase. In the absence of added cytochrome C, Harding-Passey mouse mela- noma homogenates oxidized tyrosine, dopa and catechol, with consumption of oxygen gas and formation of dark brown or black col- oration. Cloudman S91 melanoma homogen- ates acted likewise on dopa and catechol. Tyrosine was oxidized either not at all or fol- lowing a variable lag period. Similarly pre- pared homogenates of the derived Algire partially amelanotic melanoma S91A did not measurably oxidize any of the phenols or pro- duce browning, nor did they retard the oxi- dation of dopa by S91 homogenates to which they were added. Normal liver homogenates behaved like the S91A extracts. Cytochrome Oxidase. In the presence of added cytochrome C, oxidation of dopa was markedly increased in the S91 melanoma homogenates, and also now occurred exten- sively in the S91A amelanotic melanoma and liver homogenates. This second type of dopa oxidation took place via cytochrome oxidase, not only because cytochrome C was required but because the action could be eliminated en- tirely by pretreatment of the homogenates with 70% ethyl alcohol, a treatment that did not decrease true dopa oxidase activity, in fact, often enhanced it, suggestive of elimi- nation of a dopa oxidase inhibition. In no in- stance was tyrosine oxidized by the cyto- chrome system. A third type of melanization was ob- served in S91AB derivatives of S91A tumors that were obtained by prolonged transfer of the latter in brown dba mice instead of white C mice. Such tumors eventually became high- ly pigmented, but their homogenates at this stage oxidized dopa only in the presence of added cytochrome C. However, homogenates of later transfers gradually came to show, without added cytochrome C, endogenous dopa oxidation enhanced by the alcohol treat- ment, indicating eventual development of true dopa oxidase activity in addition to oxidation via cytochrome oxidase. None of the amelanotic tumors, or their secondarily melanized derivatives, oxidized tyrosine ex- cept the S91AB tumor that had undergone prolonged passage (35 generations) in dba mice. In this instance oxidation was not ma- nometrically detectable within the first 4 hours although the extracts to which tyrosine had been added blackened after ca. 24 hours. Even though the S91AB tumor, main- tained by prolonged transplantation in dba mice, assumed the enzymic pattern of the original metastatic S91 tumor, it remained biologically distinct. Thus like the original S91A amelanotic tumor, from which it was derived, it gave no evidence of producing metastases. Bibliography. 1. Woods, M. W., DuBuy, H. G., Burk, Dean and Hesselbach, Marie. Cytological studies on the nature of the cytoplasmic particulates in the Cloudman S91 mouse melanoma. J. N. C. Inst., 9: 311-323, 1949. 2. DuBuy, H. G., Woods, M. W., Burk, Dean and Lackey, Mary D. Enzymatic activities of isolated amelanotic and melanotic gran- ules of mouse melanomas and a suggested relationship to mitochondria. Ibid., 9: 325- 336, 1949. 3. Burk, Dean, Algire, G. H., Hesselbach, Marie L., Fischer, Clara E. and Legallais, Frances Y. Tissue metabolism of trans- planted mouse melanomas, with special ref- erence to characterization by paraphenylene- diamine. Special Publications of the New York Academy of Sciences, IV, 437-446, 1948. 4. Hesselbach, Marie L. Cytochemical studies of certain melanotic and amelanotic mouse tumors. Master’s thesis. Dept, of Chem- istry, Georgetown University, Washington, D. C. May, 1949. 32 Zoologica : New York Zoological Society [35: 1: 1950] Chromatographic Separation of Melanin Granules. Vernon T. Riley, Mark Woods & Dean Burk. National Institutes of Health, Bethesda, Md. Separation of cytoplasmic particulate con- stituents from cells has heretofore been ac- complished principally by differential cen- trifugation which exploits the size and spe- cific gravity of the particles. A chromato- graphic system has recently been obtained which permits the separation of some of the cytoplasmic constituents from mouse mela- nomas by exploiting still other properties of the particulates, presumably their surface characteristics as expressed by their differ- ent adsorption affinities. By utilizing the ad- sorption and elution reversibility of some of the particulates, melanin granules have been separated from the Cloudman S91 and the Harding-Passey mouse melanomas in a high- ly active state with respect to succinoxidase, cytochrome oxidase and dopa oxidase enzyme activities. Relatively mild adsorption sys- tems, varying between physiological saline a1'^ distilled water as the solvent extremes, and involving diatomaceous silica (Celite) columns, have been employed in order to avoid enzymic inactivation. No gross mor- phologic changes have been observed in the granules whether separated chromatograph- ically or centrifugally, A systematic study of the effects of vari- ous ions at graded concentrations in adsorb- ing and eluting melanin granules is under investigation, especially with reference to the possibility of obtaining granules of vary- ing composition, either pre-existing in the tumor or induced by column passage. The adaptation of chromatography to sub- c llular particulates provides a new approach to the problem of separating from the cell morphologically similar units possessing dif- ferent physical or chemical surfaces. Bibliography. 1. Riley, Vernon T. Application of chroma- tography to segregation studies of the agent of chicken tumor I (Rous sarcoma virus). Science, 107: 573-575, 1948. 2. Riley, Vernon T., Hesselbach, Marie L., Fiala, Silvio, Woods, M. W. and Burk, Dean. Application of chromatography to the separation of subcellular, enzymatically ac- tive granules. Science, 109: 361-364, 1949. 3. Riley, Vernon and Woods, M. W. A short column procedure for separating cytoplasmic components from normal and tumor tissue. Proc. Soc. Exp. Biol. & Med., 73:1, 92-95, 1950. Alexander : Tipulidae of Rancho Grande, Venezuela 33 2. The Tipulidae (Order Diptera) of Rancho Grande, North-central Venezuela.1 Charles P. Alexander. Professor of Entomology, University of Massachusetts. (Text-figures 1-39). [This is one of a series of papers resulting from the 46th, 46th and 47th Expeditions of the Department of Tropical Research of the New York Zoological Society, made during 1945, 1946 and 1948 under the direction of Dr, William Eeebe, with headquarters at Rancho Grande in the National Park of Aragua, Venezuela. The expeditions were made possible through the gen- erous cooperation of the National Government of Venezuela and of the Creole Petroleum Cor- poration. [The characteristics of the research area are in brief as follows: Rancho Grande is located in north-central Venezuela (10° 21' N. Lat., 67° 41' W. Long.), 80 kilometers west of Caracas, at an elevation of 1,100 meters in the undisturbed montane rain forest which covers this part of the Caribbean range of the Andes. Adjacent ecological zones include seasonal forest, savan- na, thorn woodland, cactus scrub, the fresh- water lake of Valencia and various marine littoral zones. The Rancho Grande area is gen- erally subtropical, being uniformly cool and damp throughout the year because of the preva- lence of the mountain cloud cap. The dry season extends from January into April. The average humidity during the expeditions, including parts of both wet and dry seasons, was 92.4% ; the average temperature during the same period was 18° C; the average annual rainfall over a five-year period was 174 cm. The flora is marked by an abundance of mosses, ferns and epiphytes of many kinds, as well as a few gigantic trees. For further details, see Beebe and Crane, Zoologica, Vol. 32, No. 5, 1947. Unless otherwise stated, the specimens discussed in the present paper were taken in the montane cloud forest zone, within a radius of one kilometer of Rancho Grande.] I am very greatly indebted to Dr. William Beebe and Mr. Henry Fleming, of the De- partment of Tropical Research of the New York Zoological Society, for the opportunity of studying a large and important series of crane-flies taken in 1945, 1946 and 1948 at and near Rancho Grande. A most interesting account of the region is provided by Dr. Beebe’s recently published book, High Jun- gle.2 The physiography and ecology of the 1 Contribution No. 870, Department of Tropical Research, New York Zoological Society. 2 Beebe, William. High Jungle, 379 pages (Duell, Sloan & Pearce) . area have been detailed in two important papers by Beebe and Crane.3 The Tipulidae of Venezuela are still very insufficiently known despite somewhat in- tensive collecting and study of the group during the past several years. The writer has published a total of nine reports that sum- marize what is known of this fauna to the year 1948. These papers bear the general title of “New or little-known Tipulidae from Venezuela (Diptera),” and were published in the Boletin de Entomologia Venezolana, Caracas, between 1943 and 1947, as follows: Part I— Vol. 2: 17-26; 1943. Part II— Vol. 2: 125-144, 5 figs.; 1943. Part III— Vol. 3: 35-50, 8 figs.; 1944. Part IV— Vol. 3: 143- 160. 5 figs.; 1944. Part V— Vol. 3; 171-192, 5 figs. ; 1944. Part VI— Vol. 4 : 59-80, 6 figs. ; 1945. Part VII— Vol. 6: 37-54, 5 figs.; 1947. Part VIII— Vol. 6 : 55-74, 14 figs. ; 1947. Part IX— Vol. 6: 74-106, 13 figs.; 1947. The above-cited papers record a total of 180 species of Tipulidae from Venezuela, in- cluding several from Rancho Grande that have been incorporated in the present report to assure completeness. Various additional species, not included in the paper, are from other stations in Aragua, specifically Cho- roni. It may be expected that several of these latter species will be found at Rancho Grande and that many further novelties may be dis- covered as a result of continued collecting. Despite the earlier work done at Rancho Grande, it is of interest to note that no fewer than 24 species are characterized as new at this time out of a total of 76 reported from Rancho Grande and vicinity. I wish to express my sincere thanks to Dr. Beebe, Mr. Fleming, and others who have co- operated actively in securing the Tipulidae herein considered. I am particularly indebted to Dr. Beebe for permitting me to retain the specimens upon which this report is based, including the types of the novelties. 3 Beebe. William. & Jocelyn Crane. 1947. Ecology of Rancho Grande, a subtropical cloud forest in northern Venezuela. Zoologica , 32: 43-60, 6 pis., 10 text- figures ; bibliography. 1948. Ecologia de Rancho Grande, una selva nublada subtropical en el Norte de Venezuela. Bol. Soc. Venez. Cien. Nat., 11 : 217-258, 5 pis., 10 text-figures. 34 Zoologica: New York Zoological Society [35: 2 TlPULINI. Brachypremna Osten Sacken. 1. Brachypremna cfispeffens (Walker, 1860). Tipula dispellens Walker; Trans. Ent. Soc. London, (n.s.) 5: 834; 1860. Rancho Grande, August 27, 1944, collected by Lichy. A species having an unusually wide distribution in the New World, ranging from the United States (New Jersey, In- diana and Illinois), through Mexico and Central America to Brazil. It is known from Trinidad but is lacking elsewhere in the West Indian islands, in the Greater Antilles being replaced by Brachypremna unicolor Osten Sacken. 2. Brachypremna similis Williston, 1900. Brachypremna similis Williston; Biol. Centr.-Americana, Diptera, I, Supple- ment: 229; 1900. Rancho Grande, May 20, 1946. Elsewhere in Venezuela from Cano del Tigre, Merida, September, 1943 (Anduze). Type from Teapa, Tabasco, Mexico, collected by H. H. Smith. Known from Mexico, Nicaragua, Costa Rica, Panama and Venezuela. 3. Brachypremna triangularis Alexander, 1945. Brachypremna arcuaria triangularis Alexander; Bol. Ent. Venezolana, 4: 60-61; 1945. Rancho Grande, May 11-July 20, 1946; one pair taken in coitu while flying. Mt. Limon, 1500 meters. May 23, 1948. Known only from this vicinity, where the type was taken at Rancho Grande on August 27, 1944, by Lichy. Most nearly allied to Brachy- premna arcuaria Alexander, of Amazonian Ecuador. Tanypremna Osten Sacken. 4. Tanypremna ITanypremna I kadeni Alex- ander, 1941. Tanypremna ( Tanypremna ) kadeni Al- exander; Ann. Ent. Soc. America, 34: 232-233; 1941. Rancho Grande, June 24, 1945. The type was from Venezuela, without more exact geographical data, collected in August, 1857, by Carl Gotthelf Kaden. Also known from Rio Chacaito, Miranda, July 16, 1939 (Vivas- Berthier). Still known only from Venezuela. Holorusla Loew. 5. Holorusia (Holorusial plagifera Alexander, 1943. Holorusia {Holorusia) plagifera Alex- ander; Bol. Ent. Venezolana, 2: 125- 127; 1943. Rancho Grande, March 27, 1946, 29; June 22, 1946, S ; July 18, 1946, $ ; May 3, 1948, 8. The type was taken here on June 24, 1944, by Lichy. Still known only from the type locality. Nephrotoma Meigen. 6. Nephrotoma medioligula Alexander, 1945. N ephrotoma medioligula Alexander; Bol. Ent. Venezolana, 4: 69, fig. 2; 1945. Rancho Grande, May 17, 1948, 9; May 24, 1948, 8 ; July 17, 1948, 2. The type was taken here on August 27, 1944, by Lichy. Still known only from the type locality. Tipula Linnaeus. 7. Tipula IBellardinal theobromina Edwards, 1920. Tipula theobromina Edwards ; Mem. Mus. Paris, Arc de Meridien Equatorial, Dipteres Nematoceres, 10 (2) : 159- 160, fig. 21 (ovipositor) ; 1920. Rancho Grande, June 18, 1946, 8. Now known from Venezuela, Ecuador, Peru and Bolivia. The type, from Ecuador, had no dis- tinct pale ring on the femora but regional material shows such an annulus that varies in width and distinctness in different speci- mens. Because of the rather constant struc- ture of the male hypopygium, I am referring all such specimens to the present species. The identity of Tipula obliquefasciata Macquart, 1846, still remains in question and may well be found to pertain to the present fly. 8. Tipula IMierotipulal liehyana Alexander, 1945. Tipula ( Microtipula ) liehyana Alexan- der; Bol. Ent. Venezolana, 4: 71, figs. 3, 4; 1945. Rancho Grande, May 23-June 14, 1948, mi- grant. The type was from here, taken May- August, 1944, by Lichy, for whom the species was named. Still known only from the type locality. 9. Tipula IMierotipulal regressa, sp. n. Mesonotum obscure brownish-yellow, vaguely patterned with pale brown; anten- nae with flagellum black, the extreme tips of the more proximal segments restrictedly pale ; wings with a weak brownish tinge, cells C and Sc, with the stigma, darker brown; male hypopygium with caudal margin of ninth tergite unequally trilobed, the lateral lobes slender, the median lobe low and broad ; eighth sternite yellow, sheathing, narrowed to the broadly obtuse tip. Male : Length about 11 mm. ; wing 12 mm. ; antenna about 5 mm. Frontal prolongation of head yellow ; nasus short and stout, distinct; palpi yellowish testaceous. Antennae (male) relatively long, as shown by the measurements ; scape, pedi- cel and base of first flagellar segment yellow; flagellum black, the extreme tips of the more proximal segments restrictedly pale; flagel- lar segments very weakly incised, much longer than the verticils. Head with the front whitened, the posterior portions brownish- yellow, the center of the posterior vertex still darker brown ; sides of vertex with numerous 1950] Alexander: Tipulidae of Rancho Grande , Venezuela 35 black setae; vertical tubercle lacking or bare- ly indicated. Pronotum light brown, paling to yellow on sides. Mesonotal praescutum obscure brownish-yellow, vaguely patterned with pale brown, the markings irregular and scarcely apparent; scutum brownish-yellow; posterior sclerites of notum somewhat clearer yellow. Pleura and pleurotergite clear light yellow. Halteres with stem yellow, knob infuscated. Legs with the coxae and trochanters pale yellow; femora brownish-yellow to pale brown, the bases clear yellow; tibiae and tarsi slightly darker brown; claws (male) toothed. Wines with a weak brownish tinge, cells C and Sc, with the stigma, darker brown; prearcular field relatively pale; re- stricted pale areas before stigma and across cell 1st Mn‘, veins brown. No macrotrichia on squama or in wing cel’s. Venation : Rs gently arcuated, about one-half longer than Rw, vein R^+2 entire; petiole of cell M i less than two times m; m-cu at near three-fourths the length of ATm; ceil 2nd A of moderate width. Abdom°n with the basal segments ob- scure yellow, the third and succeeding seg- ments chiefly infuscated, the caudal margins restrictedly pale ; subterminal segments, in- cluding six, seven and base of eighth sternite dark brown to form a ring; hynopygium yel- low. Male hypopygium (Text-fig. 1) with the ninth tergite, 9f, subequal in length and width; caudal margin unequally trilobed, in- cluding slender lateral lobes that are tipped with about four black spinous setae, and a low broad median lobe with more numerous setae ; in the slide mount, these lobes are bent backward or cephalad, possibly not a natural condition; proctiger a slender pale lobe, curved and slightly enlarged at tip. Both dis- tistyles blackened and sclerotized, shaped about as in figure d; beak of inner style slender, lower beak acute. Eighth sternite, 8s, yellow, sheathing, narrowed to the broad- ly obtuse tip, the outer part with pale incon- spicuous setae. Appendage of ninth sternite, 9s, appearing as two pale elongate lobes or blades, at anex more expanded, the surface, and especially the margins, with numerous fimbriations. Holotype, A Rancho Grande, April 23, 1946 (Beebe-Fleming) . The most similar described species are Tipula ( Microtipula ) inaequilobata Alexan- der and T. ( M .) prolixisterna Alexander, which differ particularly in the structure of the male hypopygia. 10. Tipula IMicrotipulal paralenta, sp. n. General coloration of thorax pale brown, the praescutum with four scarcely differenti- ated more brownish-yellow stripes; antennae (male) elongate, flagellar segments beyond the first black; male hypopygium with the caudal border of tergite produced into a de- pressed-flattened median lobe, on either side of the apex of which with about a dozen blackened spinous setae ; appendage of ninth sternite appearing as two flattened divergent lobes, densely fimbriate. Male: Length about 13.5-14 mm.; wing 13 mm. ; antenna about 9.5 mm. Frontal prolongation of head obscure yel- low; nasus long and conspicuous; palpi elongate, yellow, the first segment and apex of the last a trifle more darkened. Antennae (male) elongate, as shown by the measure- ments; basal three segments yellow, succeed- ing segments black; flagellar segments elon- gate, very gently incised ; longest verticils a little more than one-third the segments. Head brown; anterior vertex broad, about four times the diameter of the scape; orbital setae numerous, short and black, directed late rad. Pronotum yellowish-brown. Mesonotal praescutum pale brown, with four scarcely differentiated more brownish-yellow stripes; posterior sclerites of notum more uniformly brown. Pleura and pleurotergite pale yellow, unpatterned. Halteres elongate, stem brown, knob a trifle darker. Legs with the coxae and trochanters pale yellow; femora brownish- yellow, tibiae and tarsi darker, brownish- black; claws (male) with a very small, acute tooth at near midlength. Wings with a weak brownish tinge; stigma oval, dark brown; cell Sc brownish-yellow; a scarcely indicated dark cloud over the anterior cord ; veins dark brown. Venation: Sco ending about opposite midlength of Rs, the latter subequal in length to m-cu; R 1+2 preserved; petiole of cell Mi a little longer than m ; basal section of A/4 relatively long, perpendicular; cell 2nd A moderately wide. Abdominal tergites brown, the subtermin- al ones darker to form a more or less distinct ring; remaining sternites and the hypopy- gium chiefly yellow. Male hypopygium with the tergite subequal in length and breadth, the caudal margin gently concave on the cen- tral portion, very unequally trilobed ; lateral lobes very slender, each tipped with one or two strong spinous setae ; median lobe broad, the apex slightly produced, on either side with about a dozen similar blackened spinous setae, the actual apex glabrous. Proctiger without blackened points. Basistyle with a blackened lobe on mesal face near cephalic end. Outer dististyle a flattened lobe, nar- rowed at base, the tip obtuse, the surface with strong scattered setae. Inner dististyle with the beak flattened, its tip obtuse; lower beak extended into an acute point. Eighth sternite relatively short, not sheathing. Ap- pendage of ninth sternite appearing as two separate short flattened lobes, expanded out- wardly, the apex of each subtruncate; outer surface and margin with very abundant elongate fimbriations. Holotype, $ , La Suiza, Turrialba, Costa Rica, May, 1919 (Pablo Schild) ; Alexander Collection through A. L. Melander. Paratype, $, Mount Limon, Aragua, Venezuela, 1500 meters, May 23, 1948; (Beebe-Fleming). The most similar of the described species 36 Zoologica: New York Zoological Society [35 : 2 is Tipula ( Microtipula ) neolenta Alexander, 1945 ( lenta Alexander, preoccupied), which has the antennae much shorter and with all details of the male hypopygium quite dis- tinct. In the general features of the ninth tergite, the fly also suggests species such as T. ( M . ) heterodactyla Alexander and T. ( M .) inaequilobata Alexander, but all other de- tails of structure of the hypopygium are dis- tinct. 11. Tipula I Eumicrotipula I aedon Alexander, 1947. Tipula ( Eumicrotipula ) aedon Alex- ander; Bol. Ent. Venezolana, 6:80-82, fig. 3; 1947. Rancho Grande, August 7, 1946. Type like- wise from here, taken June 10, 1945, by Lichy. Still known only from the type local- ity. 12. Tipula I Eumicrotipula I andromache, sp. n. Size small (wing, male, 9 mm.) ; antennae one-third the length of wing, scape and pedi- cel yellow, flagellum uniformly black; meso- notum yellowish-gray, the praescutum with a broad brown median stripe; lateral stripes paler, their anterior ends curved laterad into Text-figs. 1-8. (Symbols: a, aedeagus; b, basistyle; g, gonapophysis; t, tergite; yd, ventral dististyle). 1 — Tipula ( Microtipula ) regressa, sp. n.; male hypopygium, details. 2 — Tipula ( Eumicrotijmla ) araguensis, sp. n. ; male hypopygium, details. 3 — Limonia ( Limonia ) thamyris, sp. n.; venation. 4 — Limonia ( Rhipidia ) brevipetalia, sp. n.; vena- tion. 5 — Limonia (Dicranomyia) serratiloba, sp. n. ; venation. 6 — Limonia ( Limonia ) thamyris, sp. n.; male hypopygium. 7 — Limonia ( Rhipidia ) brevipetalia, sp. n. ; male hypopygium. 8 — Limonia ( Dicranomyia ) serratiloba, sp. n. ; male hypopygium. 1950] Alexander: Tipulidae of Rancho Grande. Venezuela 37 the margin ; legs blackened, the femora with a narrow yellow subterminal ring; claws (male) small and simple; wings cream-yel- low, marbled with pale brown, cell Sc with four darkened areas that are more extensive than the interspaces; distal ends of outer radial cells uniformly darkened ; outer cells of wing with sparse macrotrichia; male hy- popygium with the tergite transverse, the caudal margin truncate, on either side of midline with a small triangular point. Male : Length about 8 mm. ; wing 9 mm. ; antenna about 3 mm. Frontal prolongation of head above light yellow, infuscated on sides and beneath; nasus distinct ; palpi with basal segment dark brown, segments two and three weakly dark- ened basally, paler at tips, terminal segment whitened. Antennae with scape and pedicel pale yellow, the flagellum abruptly and uni- formly blackened ; antennae of moderate length, as shown by the measurements; fla- gellar segments subcylindrical, much longer than their verticils. Head above pale yellow- ish brown, clearer yellow on the front and an- terior vertex, the latter broad, its tubercle very low and inconspicuous. Pronotum obscure yellow medially, weak- ly infuscated on sides. Mesonotal praescutum with the ground yellowish-gray, with a broad brown median stripe that is more expanded in front; lateral stripes somewhat paler but extensive, at anterior ends curved laterad to the margin ; posterior sclerites of notum in- fuscated and pruinose, the scutellum more testaceous. Pleura chiefly gray, the pro- pleura and pteropleurite more yellowed. Halteres with stem yellow, knob infuscated. Legs with the coxae grayish, the fore pair pale yellow; trochanters yellow; femora yel- lowed basally, passing into brown, the tips black, preceded by a yellow ring of about one- half their width; tibiae and tarsi brownish black; claws (male) small and simple. Wings cream-yellow, marbled with pale brown ; stigma and subcostal interspaces darker brown; outer portion of cell C infuscated, paler nearer h; disk of wing chiefly pale, the darkened clouds being chiefly marginal in distribution, involving the distal one-third to one-half of the outer radial cells, this part of cell Rs uniformly dark; further pale brown areas crossing the disk at near midlength and again just before outer end of cells R and M; a darkened cloud over fork of M 1+2; cell 2nd A uniformly pale brown; veins brown, yellow in the subcostal interspaces which are smaller than the four darkened areas. Sparse macrotrichia in outer ends of cells R?, to Mi, inclusive, very restricted in cells M3 and Mi. Venation: Compared with immerens, R2+3 shorter and more oblique; cell 1st M2 smaller; m-cu on Mi some distance beyond origin of latter; cell 2nd A narrower. Abdomen with the basal tergite pale, the succeeding segments brown, the subterminal segments even darker; hypopygium weakly darkened. Male hypopygium with the tergite transverse, the caudal margin truncate, on either side of midline with a small triangular point, the small area between broadly U- shaped. Outer dististyle small, with long setae. Inner dististyle with the beak elongate, pale, the shorter lower beak blackened, nar- row; dorsal crest scarcely evident, weakly elevated, with pale setae and longitudinal furi'ows or impressed lines. Long slender dusky bladelike rods from the genital cham- ber appear to represent gonapophyses. Holotype, $, Rancho Grande, April 24, 1948, resting on tree trunk (Beebe-Flem- ing) ; No. 4873. Allied to Tipula ( Eumicrotipula ) immer- ens Alexander, likewise from Venezuela, dif- fering in the small size, uniformly darkened antennal flagellum, and in the pattern and degree of trichiation of the wings. 13. Tipula l Eumicrotipula I araguensis, sp. n. Belongs to the monilifera group ; mesono- tum almost uniformly light cinnamon brown, the normal stripes not or poorly indicated ; antennae (male) a little less than one-half the length of wing, the flagellar segments weakly bicolored; wings marbled; abdomen of male obscure orange or buffy, the basal segments very slightly patterned with darker, the subterminal segments deepening in color to form a darkened ring; male hypo- pygium with the tergal lobes produced into slender blades at their mesal angle; inner dististyle with the setae of the dorsal crest relatively weak, scattered, all straight; eighth sternite with the median lobe very small, low hemispherical in outline. Male: Length about 13-14 mm.; wing 15- 16.5 mm.; antenna about 7-8 mm. Female: Length about 21-22 mm.; wing 16-17 mm. Frontal prolongation of head light to me- dium brown, more pruinose at base; nasus distinct; palpi dark brown. Antennae (male) relatively long, as shown by the measure- ments ; scape and pedicel light yellow, flagel- lum weakly bicolored, the stems light brown, the abrupt basal swellings brownish-black. Head light gray, with a conspicuous capil- lary brown vitta on vertex, the sides of the posterior vertex more extensively paler brown; vertical tubercle scarcely developed. Mesonotum almost uniformly light cinna- mon brown, more or less gray pruinose, praescutal stripes not or scarcely indicated ; posterior sclerites of notum and the pleura more yellowed. Halteres with stem pale, knob infuscated. Legs brownish-yellow, the femor- al tips more infuscated; remainder of legs passing through brown to dark brown. Wing of type whitish-subhyaline, with an extensive but weakly contrasted brown pattern that is arranged about as in other members of the group ; the other specimens of the type series have the dark pattern much heavier and con- trasted. Venation: Rs a little more than twice m-cu; tip of vein R1+2 pale but entire; 38 Zoologica : New York Zoological Society [35: 2 cell 1st M2 relatively small, irregularly pen- tagonal; m-cu before fork of M3+4; petiole of cell Mi shorter than m. Basal abdominal segments of male chiefly obscure orange to light brownish yellow, not or scarcely patterned, the sixth and succeed- ing segments deepening in color to form a dark brown ring; outer portion of hypopyg- ium somewhat paler, brownish-yellow. In the female, abdomen more elongated; ter- gites with a conspicuous sublateral dark brown stripe on either side, these more widened behind. Male hypopygium (Text- fig. 2) with the tergal lobes, 9f, at mesal angle produced into a slender flattened blade ; dorsal surface of tergite furrowed. Dorsal dististyle with outer half dilated. Inner dis- tistyle, id, relatively long and narrow; setae of the scarcely developed dorsal crest rela- tively weak, scattered, all straight, not an- gularly bent as in allied species. Gonapo- physes, g, with apical lobes unequal, the lower one a slender decurved spinous point. Appendage of ninth sternite an oval lobe, the longest setae exceeding the diameter of the lobe. Eighth sternite, 8s, transverse, the caudal margin gently and evenly convex, the median lobe very small, low-hemispherical in shape, the setae much longer than the lobe itself. Holotype, $, Rancho Grande, July 18, 1946 (Beebe-Fleming). Allotopotype, 9, May 3, 1948. Paratopotypes, 1 $, in copula and pin- ned with allotype; other $Q, May 9-16, 1948; paratype, of S, Mount Limon, 1500 meters, May 23, 1948 (Beebe-Fleming). One pair, taken in copula, were resting on the exposed root of a tree growing on a steep bank. There are several similar regional mem- bers of the monilifera group, including Tipula ( Eumicrotipula ) cliacopata Alex- ander, T. ( E .) inaequidens Alexander, T. ( E .) infinita Alexander, and T. ( E .) palenca Alexander, all of which differ from the pres- ent species and among themselves in the structure of the male hypopygium, particu- larly the tergite, inner dististyle, gonapo- physis and lobe of the eighth sternite. 14. Tipula I Eumicrotipula) cristata Alexander, 1945. Tipula ( Eumicrotipula ) cristata Alex- ander; Bol. Ent. V enezolana, 4: 72-74, fig. 5; 1945. Rancho Grande, September 13, 1944, col- lected by Lichy; types. Still known only from the type locality. 15. Tipula l Eumicrotipula) infinita Alexander, 1945. Tipula ( Eumicrotipula ) infinita Alex- ander; Bol. Ent. V enezolana, 4: 76-78, fig. 6; 1945. Rancho Grande, April, 1944, collected by Lichy ; type. Still known only from the type locality. 16. Tipula l Eumicrotipula! tovarensis Alex- ander, 1947. Tipula ( Eumicrotipula ) tovarensis Alexander; Bol. Ent. V enezolana, 6: 40-42, fig. 1; 1947. Rancho Grande, August 24, 1944, collected by Lichy. Still known only from the type lo- cality. Limoniini. Limonia Meigen. 17. Limonia I Limonia) alfaroi (Alexander, 1922). Dicranomyia alfaroi Alexander; Proc. U. S. Nat. Mus., 60, Art. 25 : 2-3 ; 1922. Rancho Grande, Aug. 8, 1946. The type was from San Jose, Costa Rica, taken October 7, 1920, by Alfaro. The typical form of the spe- cies ranges from southern Mexico to Vene- zuela. 18. Limonia I Limonia) fumosa (Alexander, 1912). ‘ IFurcomyia fumosa Alexander; Can. Ent., 44: 364, fig.; 1912. Rancho Grande, June 9, 1937, collected by Vivas-Berthier. The type was taken at Amatuk, British Guiana, July 14, 1911, by F. E. Lutz. Still known only from Venezuela and British Guiana. 19. Limonia I Limonia) onoma, sp. n. Thoracic pleura yellow, with a broad brown longitudinal stripe; front of head sil- very, the posterior part dark brownish-gray; legs dark brown; wings subhyaline, conspic- uously patterned with brown, including a quadrate area at origin of Rs, stigma and broad seams over cord and outer end of cell 1st M 2; no darkening at arculus but with one at near one-fourth the length of cell R; Sci ending about opposite midlength of Rs; basal section of vein Ri+s angulated; vein 2nd A bent strongly into the margin. Female: Length about 7 mm.; wing 8.3 mm.; antenna about 1.5 mm. Rostrum black; palpi brownish-black. An- tennae black throughout; basal flagellar seg- ments short-subcylindrical, the outer ones passing through oval to elongate; terminal segment nearly twice as long as the penulti- mate; longest verticils unilaterally arranged, exceeding the segments, excepting the ter- minal one. Head dark brownish-gray; front and anterior vertex silvery, the latter re- duced to a linear strip. Pronotum brownish-yellow. Mesonotum light brown, the lateral portions of prae- scutum restrictedly paler. Pleura yellow, with a broad brown longitudinal stripe extending from the cervical region to the base of abdo- men, passing beneath the root of halteres. Halteres brown, base of stem restrictedly brightened, knob brownish-black. Legs with all coxae and trochanters pale yellow; re- mainder of legs dark brown, the femoral bases not or scarcely brightened. Wings with the ground subhyaline, with a conspicuous brown pattern that includes a quadrate area at origin of Rs, stigma, and broad seams 1950] Alexander: Tipulidae of Rancho Grande, Venezuela 39 over cord and outer end of cell 1st M i; other darkenings include a circular area over fork of Sc, not touching the one at origin of Rs; spots at ends of both Anal veins, that at 2nd A larger; a cloud at about one-fourth the length of cell R, in transverse alignment with the end of vein 2nd A; no darkening at arcu- lus; small marginal clouds on veins Rs, Ri+5, M3, Mi and Cu, scarcely evident on the veins near wing-tip; in outer radial field with a narrow subapical cloud in cells R 2 and R3', veins brown, Sc brighter. Venation: Sc long, Sci ending beyond midlength of Rs, Sci near its tip; Rs long, square at origin; inner end of cell Ri lying basad of that of 1st Mi, the basal section of Ri*s being angulated; cell 1st Mi elongate, subequal to vein M 1+2 be- yond it; m-cu at fork of M, longer than the distal section of Cui ; vein 2nd A bent strong- ly into the wing margin. Abdomen dark brown, the sternites a trifle paler. Ovipositor with cerci slender, up- curved; hypovalvae longer and straight. Holotype, 9, Rancho Grande, May 12, 1946 (Beebe-Fleming). The most similar regional species are Li- monia ( Limonia ) eiseni (Alexander) and L. ( L .) macintyrei Alexander, both of which are quite distinct in the wing pattern and details of venation. 20. Limonia ILimonial pampoecila (Alexan- der, 1922), var. Dicranomyia pampoecila Alexander; Proc. U. S. Nat. Mus., 60, Art. 25: 1- 2; 1922. Rancho Grande, June 28, 1946 ; a frag- mentary female. Close to the typical form but with the darkened femoral ring terminal in position whereas in typical pampoecila this is subterminal. The material is too poor for more exact determination. The type of pampoecila was from Tiribi, Costa Rica, col- lected October 9, 1920, by Alfaro. What ap- pears to represent a single species ranges from Costa Rica to southeastern Brazil but this complex requires further study. 21. Limonia (Limonial thamyri s, sp. n. General coloration gray, patterned with brown, the markings on the praescutum ir- regular in distribution; femora black, yellow basally, with a very narrow and indistinct yellow subterminal ring; wings whitish-sub- hyaline, with a heavy reticulated brown pat- tern ; cell Mi open by the atrophy of m; male hypopygium with the ventral dististyle large and fleshy, its rostrum with two long spines, the more proximal one from a small basal tubercle. Male: Length about 5.5 mm.; wing 6 mm. Rostrum brownish-black, sparsely prui- nose; palpi black; rostrum in direct align- ment with the front. Antennae with the short scape black, pedicel light yellow; flagellum black, the first segment paler; flagellar seg- ments oval, slightly longer than the verticils. Head buffy brown, more pruinose medially behind, the center of the posterior vertex with a blackened area. Pronotum brownish-yellow, pruinose. Mes- onotal praescutum with the ground grayish pruinose, patterned irregularly with dark brown, the broader median stripe entire on the posterior third, in front branching into four narrow lines; lateral stripes similarly narrow; scutal lobes dark brown, the median area narrowly more silvery, the line con- tinued caudad onto the scutellum; remainder of the latter brown, margined with darker brown; central portion of mediotergite dark brown, the lateral margins yellow. Pleura and pleurotergite striped longitudinally with brownish-yellow and dark brown, the sur- face pruinose to produce a variegated effect. Halteres with stem yellow, knob dark brown. Legs with the coxae obscure yellow, the fore pair darkened basally; trochanters obscure yellow, the fore pair darkened apically; fe- mora obscure yellow on about the proximal third or fourth, the remainder brownish- black, enclosing a very narrow and indistinct obscure yellow ring some distance back from the blackened tip; tibiae dark brown; tarsi more yellowish-brown, the outer segments blackened, the posterior tarsi more uniform- ly so; claws (male) with a strong, nearly ba- sal spine, with a smaller acute point still more basad. Wings (Text-fig. 3) with the re- stricted ground whitish-subhyaline, with a heavy reticulated brown pattern, paler brown in the apical and posterior cells of wing; cell Sc and the prearcular field more yellowed; a series of about a dozen brown dashes in cell C, narrower than the inter- spaces, cell Sc clearer yellow; three major darker brown areas in cell R, the first post- arcular, the third at origin of Rs; a similar slightly paler brown cloud in cell M just be- fore the level of origin of Rs; further major areas at stigma and anterior cord and near the outer end of vein R3 ; reticulated areas in apical and posterior cells paler and few in number but distinct; veins brown, Sc and R more yellowed. Venation: Sc moderately long, Sci ending just before midlength of Rs, Sc2 at its tip ; Rs angulated and short-spurred at origin ; vein R3 slightly arcuated on basal half; cell M2 open by atrophy of m; m-cu shortly before fork of M, long and very oblique, about one-fourth longer than the distal section of vein Cui. Abdomen brown , the hypopygium yel- lowed. Male hypopygium (Text-fig. 6) with the tergite, 9f, transverse, the caudal mar- gin very gently emarginate, the low lateral lobes with long pale setae. Basistyle, b, with the ventromesal lobe large, obtuse at apex. Dorsal dististyle a strong rod, the long apical spine nearly straight. Ventral disti- style, vd, large and fleshy, its area about one- third greater than the total of the basistyle; rostral prolongation stout, the two spines relatively long, subequal in length, placed one behind the other, the more basal one from a slightly larger tubercle. Gonapophy- sis, g, with mesal-apical lobe elongate, pale, terminating in a small darkened knob that is directed laterad. 40 Zoologica: New York Zoological Society [35: 2 Holotype, $ , Rancho Grande, June 27, 1946 (Beebe-Fleming) . Generally similar to Livionia ( Limonia ) pampoecila (Alexander) and certain allied species, differing in the pattern of the legs and wings, the venation, especially the open cell M2, and in the details of structure of the male hypopygium. 22. Limonia I Neolimnobia) diva (Schiner, 1868). Limnobia diva Schiner; Novara Reise, Diptera, p. 46; 1868. Rancho Grande, May 7-11, 1946, August 1-7, 1946; also September 5, 1942, collected by Lichy. Specimens taken between July 10 and 18, 1946, have the pattern of the femora much obscured, as noted below. The species diva, as now restricted, has a wide range in tropical America, including the Greater Antilles (Cuba, Puerto Rico), Mexico, Venezuela and Brazil. Further races or closely allied species are found in Ecuador and Peru. The variability in the distinctness of the leg pattern in what seems to represent a single species has been discussed by the writer in another paper (Notes on the Trop- ical American species of Tipulidae (Dip- tera). VI. Rev. de Entomologia — in press). As there indicated, some individuals have two or, in cases, three dark rings on the fe- mora while other specimens that are refer- red to diva without question have the femoral pattern much obscured, the darkened annuli being broader and more diffuse, in the ex- treme cases being so vague and extensive as to eliminate the yellow interspaces. The ex- treme specimens from Rancho Grande above mentioned do not quite represent the condi- tion described but individuals from elsewhere in Venezuela have this leg pattern. 23. Limonia (lihipidial brevipetalia, sp. n. Praescutum reddish-brown, with indica- tions of darker lines and dashes; pleura and pleurotergite more gray pruinose, narrow- ly lined with darker; antennae dark, the two subterminal segments white; flagellar seg- ments short-bipectinate; femora darkened subterminally, the actual tip narrowly yel- low; wings pale yellow, with a very abund- ant pale brown dotted pattern, the marks chiefly confluent; male hypopygium with the caudal margin of the tergite very gently emarginate ; rostral prolongation of the ven- tral dististyle unusually long and slender, the two spines subequal in length and size. Male: Length about 7 mm.; wing 8 mm. Rostrum brownish-black, shiny, the mouthparts conspicuous; palpi black. An- tennae black, the long pedicels of the flag- ellar segments pale yellow ; subterminal two segments whitened; flagellar segments one to three strongly produced but simple; seg- ments four and five each with two conspicu- ous branches that slightly exceed in length the remainder of organ (the succeeding seg- ments broken, the color pattern having been noted before the breakage occurred) . Head dark gray; anterior vertex reduced to a lin- ear strip that does not exceed in width two rows of ommatidia. Pronotum dark brown, the scutellum and pretergites more reddish-brown. Mesonotal praescutum chiefly reddish-brown, with indi- cations of dark lines and dashes, as well as the posterior end of the usual median stripe; scutal lobes dark; scutellum yellowish-gray with a black central line; mediotergite gray pruinose, the central part more heavily so, the lateral borders paling to yellow. Pleura and pleurotergite gray pruinose, narrowly lined longitudinally with dark brown, the dorsal stripe most conspicuous, the ventral line a short dash on the sternopleurite; the dorsal stripe begins at the propleura, nar- rowed behind at the root of the halteres. Hal- teres with stem dirty white, more yellowed at base, knob infuscated. Legs with the coxae brown, pale at tips; trochanters yel- low; femora light brown, paler basally, deepening to a somewhat more intense sub- terminal ring, the actual tip narrowly yellow; tibiae and tarsi obscure yellow, the terminal two segments black; claws (male) with a strong subbasal spine, with a smaller one a little more basad. Wings (Text-fig. 4) with the restricted ground pale yellow, with a very abundant pale brown dotted pattern, the areas being so abundant as to be exten- sively confluent; very small darker dots be- yond midlength of vein R, origin of Rs, and fork of Sc; cord and outer end of cell 1st M2 narrowly seamed with darker; a more or less evident series of two or three brown spots along vein Cu in cell M; veins yellow, varie- gated with darker in the more heavily pat- terned areas. Venation: Sci ending opposite or just beyond midlength of Rs, Sc2 at its tip; m-cu more than one-third its length before the fork of M. Abdominal tergites reddish-brown, the lateral borders narrowly blackened; stern- ites a trifle more yellowed; hypopygium and segment eight yellow. Male hypopygium (Text-fig. 7) with the tergite, 9 1, transverse, the caudal margin very gently emarginate, the surface of the low lobes with abundant setae. Basistyle, b, subequal in area to the ventral dististyle, the ventromesal lobe large, with a small lateral lobule. Dorsal dististyle a gently curved blackened rod, the tip acute. Ventral dististyle, vd, with the rostrum un- usually long and slender, the two spines slightly separated, the outermost at near mid- length of the prolongation; spines straight, subequal in size. Gonapophysis, g, with the mesal-apical lobe stout, gently curved, heav- ily blackened, the tip acute or subacute. Aedeagus, a, stout. Holotype, $, Rancho Grande, June 7, 1946 (Beebe-Fleming) . In its abundantly dotted wings, the present fly most resembles species such as Livionia ( Rhipidia ) monoxantha Alexander, which differs conspicuously in the coloration and in the wing pattern. The male sex of mono- xantha is still unknown. 1950] Alexander: Tipulidae of Rancho Grande, Venezuela 41 24. Limonia IRhipidial domestica (Osten Sacken, 1859). Rliipidia domestica Osten Sacken ; Proc. Acad. Nat. Sci. Philadelphia, 1859: 208, pi. 3, figs. 8-9; 1859. Rancho Grande, June 26-28, July 10, Au- gust 3, 1946. Described from the eastern United States; very widely distributed, ranging from Con- necticut, New York and New Jersey, south- ward throughout the southern states; An- tilles (Cuba, Jamaica, Puerto Pico, Lesser Antilles) ; Venezuela; Brazil. 25. Limonia IRhipidial fiabelliformis Alexan- der, 1934, var. Limonia ( Rhipidia ) fiabelliformis Alex- ander; Ann. Ent. Soc. America, 27: 59-60; 1934. Rancho Grande, May 29, 1946. The type was from Chiriqui, Panama, altitude 5400 feet, taken September 13, 1932, by Lawlor. The typical form ranges from Panama to Peru. 26. Limonia IBicranomyial brevicubitalis Al- exander, 1947. Limonia ( Dicranomyia ) brevicubitalis Alexander; Bol. Ent. Venezolana, 6: 87-90, figs. 7, 8; 1947. Rancho Grande, May 5, June 16-28, July 23, 1946; May 13, 1948. Type from the Rio Chacaito, Miranda, Venezuela, 980 meters, September 18, 1938, collected by Vivas-Ber- thier. Other specimens from Los Venados, D. F., 1520 meters, February 26, 1939, Vivas- Berthier. Still known only from Venezuela. 27. Limonia l Dicranomyia I brevivena capra Alexander, 1947. Limonia ( Dicranomyia ) capra Alex- ander; Bol. Ent. Venezolana, 6 : 57-59, figs. 1, 5; 1947. Rancho Grande, April 21, 1946. Type from here, collected by Lichy on August 27, 1944. Elsewhere in Venezuela known from the Rio Chacaito, Miranda, 980 meters, September 18, 1938, Vivas-Berthier ; Los Canales, Nai- guata, D. F., 720 meters, September 24, 1938, Vivas-Berthier. I am considering this as representing a race of the more northern and unusually widespread brevivena (Osten Sacken, 1869). 28. Limonia IBicranomyial serratiioba, sp. n. Allied to mutata; general coloration gray, the praescutum and scutal lobes extensively more infuscated; legs chiefly black, the fe- moral bases obscure yellow; wings with a strong brownish tinge, the very small sub- circular stigma darker brown; male hypopy- gium with the tergite narrowly transverse, the caudal margin nearly truncate; rostral prolongation of the ventral dististyle stout, the two spines relatively short, straight, from small basal tubercles; mesal-apical lobe of the gonapophysis long and slender, the margin with conspicuous serrulations. Male : Length about 5.5 mm. ; wing 6 mm. Rostrum and palpi brownish-black. Anten- nae black throughout; flagellar segments oval. Head gray; anterior vertex very nar- row, reduced to a linear strip. Pronotum brownish-gray. Mesonotum chiefly dark brownish-gray, the praescutum and scutal lobes extensively more infuscated. Pleura blackened, heavily pruinose. Halteres relatively long, stem dirty white, knob in- fuscated. Legs with the coxae dark brown, sparsely pruinose; trochanters obscure yel- low; femora obscure yellow basally, passing into black; remainder of legs black; claws (male) with at least two basal teeth, the out- ermost strongest. Wings (Text-fig. 5) with a strong brownish tinge, the very small sub- circular stigma darker brown; exceedingly vague and narrow seams over the cord; pre- arcular field a little more whitened; veins brown. Venation: Sci ending opposite origin of Rs, Sco near its tip ; Rs slightly more than twice as long as the more arcuated basal sec- tion of R4+5; cell 1st M 2 a trifle longer than vein Mi\ m-cu a short distance before the fork of M. Abdomen, including hypopygium, dark brown. Male hypopygium (Text-fig. 8) with the tergite, 9 1, narrowly transverse, the cau- dal margin nearly truncate; scattered pale setae along the margin, with an additional discal pair. Basistyle, b, with the ventrome- sal lobe large, obtuse, provided with long yellow setae. Dorsal dististyle a curved blackened rod, narrowed very gradually in- to a long acute spine. Ventral dististyle, vd, fleshy, its total area somewhat greater than that of the basistyle; rostral prolongation stout, its tip obtuse; rostral spines two, aris- ing from equal small tubercles, the spines straight and relatively short. Gonapophysis, g, with the blade pale, the mesal-apical lobe long and slender, gently curved, the margin with several conspicuous serrulations. Holotype, 8, Rancho Grande, June 16, 1946 (Beebe-Fleming) . The present fly is generally similar to spe- cies such as Limonia ( Dicranomyia ) mutata Alexander, differing in the coloration of the body and wings and in the structure of the male hypopygium, particularly the tergite, ventral dististyle and gonapophyses. 29. Limonia I Peripheropferal angustifasciata (Alexander, 1922). Peripheroptera angustif asciata Alex- ander ; Trans. Ent. Soc. London, 1922 : 35-36; 1922. Rancho Grande, April 17-27, June 25, July 15, 1946; July 10, 1945; March 26, 1948. Two further specimens of the migrant series, May 23, 1948, 9 (No. 48569) and July 21, 1948, 8 (No. 481275). The type was from “Venezuela,” with no further data, contained in the collection of the Vienna Museum and to this time no fur- ther specimens had been taken. The wings in 42 Zoologica: New York Zoological Society the present series agree well with those of the type female except in the sexual charac- ters, as found in most species in the subgenus Peripheroptera Schiner. I regard the iden- tification as correct. 30. Limonia IGeranomyial callinota Alexan- der, 1941. Limonia ( Geranomyia ) callinota Alex- ander; Ann. Mag. Nat. Hist., (11) 8: 318-320; 1941. Rancho Grande, June 28, 1946; 1 2. The type was from Carpapata, Junin, Peru, 2600 meters, May 15, 1940, collected by Woytkow- ski. Known only from these two stations. I see no reason to question the present deter- mination even though it is based on the fe- male sex. 31. Limonia IGeranomyial furor Alexander, 1944. Limonia ( Geranomyia ) furor Alexan- der; Bol. Ent. V enezolana, 3: 183-185, fig. 5; 1944. Rancho Grande, June 26, 1946. Type from Rio Chacaito, Miranda, Venezuela, 980 meters, September 18, 1938, Vivas-Berthier. Still known only from Venezuela. 32. Limonia IGeranomyial opinator, sp. n. Size small (wing, male, 5.5 mm.) ; rostrum relatively short; praescutum light gray, with three brown stripes additional to the dark- ened lateral borders; femora with a narrow brown subterminal ring; wings grayish-sub- hyaline, with a very restricted darker pat- tern, including a common cloud over the or- igin of Rs and fork of Sc; male hypopygium with the ventral dististyle large and fleshy, the prolongation slender, with two spines that arise from slightly unequal tubercles; gonapophysis with mesal-apical lobe a slend- er pale horn. Male: Length, excluding rostrum, about 5 mm. ; wing 5.5 mm. ; rostrum about 2.5 mm. Rostrum of moderate length, approximate- ly oneffialf the remainder of body, brownish- black throughout. Antennae black, relatively short; flagellar segments oval, with short verticils. Head gray, the posterior vertex with a dark brown longitudinal stripe on either side of the subequal median ground line. Pronotum buffy, weakly infuscated medi- ally and on the sides. Mesonotal praescutum light gray, with three dark brown stripes on the disk, the intermediate pair convergent behind and becoming confluent at the suture, the shorter median stripe becoming obsolete some distance before the suture ; lateral prae- scutal borders less heavily darkened, humer- al region yellowed; scutum chiefly brownish- gray, including the lobes ; posterior sclerites of notum dark brown, pruinose. Pleura with a dark brown dorsal stripe, the ventral pleurites yellow. Halteres with stem obscure yellow, its base brighter, knob dark brown. Legs with the coxae yellow, the fore pair with the basal half darkened; trochanters yellow; femora obscure yellow, passing into light [35: 2 brown outwardly, with a narrow darker brown subterminal ring, subequal in extent to the yellow tip; tibiae and tarsi pale, the outer tarsal segments dark brown; claws slender, with a single strong basal spine. Wings (Text-fig. 9) grayish-subhyaline, the prearcular and narrow costal region more yellowed ; a very restricted dark pattern, in- cluding the stigma; a small common cloud over the origin of Rs and fork of Sc; and vague narrow seams over the cord and outer end of cell 1st M,; veins brown, pale in the yellowed areas. Venation: Sc short, Sci end- ing immediately beyond the origin of Rs, Sc2 opposite this origin ; cell 1st M? long, nearly equal to the distal section of vein A/1+2; m-cu shortly beyond the fork of M. Abdomen, including hypopygium, dark brown. Male hypopygium (Text-fig. 16) with the tergite, 9t, transverse, the caudal border broadly emarginate, the relatively low lateral lobes with numerous long coarse setae. Basi- style, b, small, its total area a little less than one-third that of the ventral dististyle; ven- tromesal lobe simple. Dorsal dististyle a slender curved rod, the slightly upcurved tip acute. Ventral dististyle, vd, large and fleshy ; rostral prolongation slender, the two spines straight, slightly unequal in length from tubercles of unequal size, the longer spine lying more basad on the longer tubercle. Gonapophysis, g, with the mesal-apical lobe a slender nearly straight pale horn. Aedea- gus, a, with apical lobes obtuse. Holotype, S, Rancho Grande, June 26, 1946 (Beebe-Fleming) . While generally similar to species such as Limonia ( Geranomyia ) recisa (Alexander), the present fly differs evidently in the struc- ture of the male hypopygium. 33. Limonia IGeranomyial stenophallus Alex- ander, 1944. Limonia ( Geranomyia ) stenophallus Alexander; Ann. Ent. Soc. America, 37: 310-311; 1944. Rancho Grande, June 26, 1946. The type was from Abitagua, Ecuador, 1100 meters, March 21, 1940, collected by Macintyre. The species ranges from Venezuela to Ecuador and Peru. 34. Limonia IGeranomyial subvirescens Alex- ander, 1930. Limonia ( Geranomyia ) subvirescens Alexander; Journ. N. Y. Ent. Soc., 38: 112; 1930. Rancho Grande, July 23, 1946. The type was from the Trinidad Mountains, Cuba, 1000 feet, taken March 25, 1925, by J. G. Myers. Ranges from the Greater Antilles (Cuba) to Venezuela. 35. Limonia IGeranomyial tibialis (Loew, 1851). Aporosa tibialis Loew; Linnea Ento- mologica, 5: 397; 1851. Rancho Grande, July 5-31, 1946. The type was from Brazil, without further data, col- lected by Sellow. This is one of the most 1950] Alexander: Tipulidae of Rancho Grande, Venezuela 43 widely distributed of all tropical American Tipulidae, ranging from the Greater Antilles (Jamaica, Puerto Rico) ; Lesser Antilles (Dominica, Grenada) ; Central America; British Guiana, into Paraguay. It may be noted that this is the only species of crane- fly so far taken on the Galapagos Islands. 36. Limonia IGeranomyial timens, sp. n. Size small (wing under 6 mm.) ; rostrum short, only a little more than one-third the length of wing; praescutum gray with three narrow brown stripes, additional to the paler sublateral darkening; femora with a very poorly indicated pale brown subterminal ring; wings weakly tinged with brown, with a very restricted pale brown pattern, includ- ing a common area over the origin of Rs and fork of Sc; male hypopygium with the ven- tral dististyle large and fleshy, the prolonga- tion relatively short but slender, more or less constricted basad of the spines, the latter arising from strong tubercles. Male: Length, excluding rostrum, about 5 mm.; wing 5.8 mm.; rostrum about 2.1 mm. Rostrum short, only a little more than one- Text-fig. 9-18. (Symbols: a, aedeagus; b, basistyle; d, dististyles; g, gonapophysis; od, outer dististyle; t, tergite; vd, ventral dististyle). 9 — Limonia (Geranomyia) opinator, sp. n.; venation. 10 — Limonia ( Geranomyia ) timens, sp. n.; venation. 11 — Orimarga ( Orimarga ) dichroptera, sp. n.; venation. 12 — Helius ( Helius ) bitergatus, sp. n.; vena- tion. 13 — Protohelius venezolanus, sp. n.; venation. 14 — Paradelphomyia ( Oxyrhiza ) venezolana, sp. n.; venation. 15 — Helius ( Helius ) bitergatus, sp. n.; male hypopygium. 16— Limonia ( Geranomyia ) opinator, sp. n. ; male hypopygium. 17 — Limonia (Gerano- myia) timens, sp. n.; male hypopygium. 1&— Paradelphomyia ( Oxyrhiza ) venezolana, sp. m; male hypopygium 44 Zoologica: New York Zoological Society [35 : 2 third the length of the wing, black through- out. Antennae black; flagellar segments sub- cylindrical, with inconspicuous verticils. Head above gray, with a blackish line on either side of the median ground line of the posterior vertex. Pronotum buffy gray, dark brown medi- ally. Mesonotal praescutum gray, with three narrow but conspicuous intermediate dark brown stripes that are subequal in width and nearly twice as broad as the interspaces; sub- lateral stripes broader but less intense than the three central ones ; posterior sclerites of notum gray, each scutal lobe with two dark- ened areas. Pleura chiefly pale reddish- brown, the surface vaguely pruinose, the more ventral portions paler. Halteres with stem pale, knob dark brown. Legs with the coxae and trochanters pale yellow; femora yellow, with a narrow and very poorly in- dicated pale brown subterminal ring, the yel- low apex subequal in extent; remainder of legs brownish-yellow, the tarsi not or scarce- ly darkened. Wings (Text-fig. 10) with a weak brownish tinge, the prearcular field and relatively broad costal border more yellowed, the latter pattern continued almost to the wing tip; a very restricted pale brown pat- tern, including the stigma, a smaller com- mon area over the origin of Rs and fork of Sc, and a still smaller cloud at the super- numerary crossvein in cell Sc; veins brown- ish-yellow, slightly darker in the patterned areas. Venation: Sc relatively short, Sci ending about opposite one-third the length of Rs, Sc 2 a short distance back from tip; r-m reduced in length by the approximation of veins #4+5 and Mi+2; cell 1st M2 slightly long- er than vein M3; m-cu at fork of M. Abdomen, including hypopygium, pale yellowish-brown. Male hypopygium (Text- fig. 17) with the tergite, 9 1, transverse, the caudal border with a median notch to form two low rounded lobes that are provided with abundant long setae. Basistyle, b, relatively small, its total area about one-third that of the ventral dististyle; ventromesal lobe sim- ple. Dorsal dististyle a gently curved rod, the tip gradually narrowed into a slender spine. Ventral dististyle, vd, large and fleshy, the rostral prolongation relatively short but slender, more or less constricted basad of the spines; spines two, from strong tubercles, slightly unequal in length, the shortest sub- equal in length to the prolongation. Gona- pophysis, g, with the mesal-apical lobe pale, appearing as a gently curved flattened horn, the tip acute. Holotype, $ , Rancho Grande, June 26, 1946 (Beebe-Fleming) . Related to species such as Limonia ( Gera - nomyia) recisa Alexander, L. ( G .) scolopax (Alexander) and L. (G.) opinator, sp. n., differing from all chiefly in the structure of the male hypopygium. 37. Limonia IGeranomyiai vindicta Alexan- der, 1943. Limonia ( Geranomyia ) vindicta Alex- ander; Bol. Ent. Venezolana, 2: 21- 22; 1943. Rancho Grande, May 10, 1946. Type from Rio Chacaito, Miranda, Venezuela, 980 meters, September 18, 1938, collected by Vivas-Berthier. The typical form is still known only from Venezuela; the race, dilu- cida Alexander, from Ecuador. 38. Limonia IGeranomyiai v/alkeri Alexander, 1930. Limonia ( Geranomyia ) ivalkeri Alex- ander; Ann. Ent. Sos. America, 23: 730-732; 1930. Rancho Grande, September 5, 1937, Vivas- Berthier. The type was from Vista Nieve, Mount Santa Marta, Colombia, 5000 feet, August 8, 1926, collected by Fred W. Walker. Other typical material from Jaragua, Santa Catharina, Brazil, August 29, 1929. The spe- cies occurs from Colombia and Venezuela to southeastern Brazil. Orlmarga Osten Sacken. 39. O rimarga lOrimargal dichroptera, sp. n. Allied to niveitarsis ; general coloration black, the sides of the thorax with two nar- row silvery longitudinal stripes; legs black, the femoral bases of the middle and hind legs yellowed; tarsi extensively white; wings strongly bicolored, the cephalic half brown- ish-black, the posterior half and the pre- arcular field whitened; vein Sc short, Sci ending some distance before origin of Rs; cell M3 approximately twice its petiole; vein 2nd A short; abdomen elongate. Female: Length about 9.5-10 mm.; wing 5.5 mm.; abdomen alone 7.5-8 mm. Rostrum and palpi black. Antennae black throughout, short; flagellar segments oval. Head above black, sparsely dusted with gray. Thoracic dorsum black, the praescutum with a narrow silvery gray longitudinal stripe. Pleura black, with a slightly broader silvery gray longitudinal stripe, extending from the cephalic portion of the sternopleu- rite to the abdomen. Halteres black. Legs with the fore coxae and trochanters black, the remaining coxae black basally, their apices broadly yellow; trochanters yellow; fore legs black, middle and posterior legs black, the femoral bases extensively yellow; outer two-fifths of basitarsi and remainder of tarsi excepting the last segment snowy white. Wings (Text-fig. 11) strongly bicol- ored, the cephalic half brownish-black, the posterior part more whitened; darkened areas include the anterior third of wing, most of cells R and M, bases of Cu and 1st A, and all of cell 2nd A; prearcular field chiefly white; veins brownish-black. Venation: Sc short, Sci ending a distance before origin of Rs greater than the length of m-cu, the latter opposite the base of Rs; E2+3 and R2 subequal, about one-half R1+2] inner end of cell Rr, a little more distad than those of cells R3 and 1950] Alexander: Tipulidae of Rancho Grande, Venezuela 45 Mz‘, cell M 3 about one and one-half to two times its petiole; vein 2nd A short. Abdomen elongate, black; valves of ovi- positor horn-yellow. Holotype, 9, Rancho Grande, July 9, 1946 (Beebe-Fleming) . Paratopotype, 1 9. Readily distinguished from Orimarga ( Orimarga ) niveitarsis Alexander, and its larger race majuscula Alexander, by the strongly bicolored wings, with the venational details distinct. 40. Orimarga (Orimarga) excessive Alexan- der, 1926. Orimarga ( Orimarga ) excessiva Alex- ander; Ann. Ent. Soc. America, 19: 380-381; 1926. Rancho Grande, May 10, June 26-28, 1946. Type from Tachira, Venezuela, April 4, 1920, Williamsons & Ditzler. Still known only from Venezuela. Hellas St. Fargeau. 41. Helius IHelius) biter gatus, sp. n. Praescutum and scutal lobes brown; pos- terior sclerites of notum and the pleura light- er brown; legs dark brown, the outer tarsal segments paling to yellowish-brown; wings with a strong brownish tinge, stigma darker brown; male hypopygium with the tergite produced into two conspicuous earlike lobes that are directed beneath into the straight tergal spines; basistyle without lobes; dis- tistyle terminal, the outer spine of outer style reduced ; aedeagus beyond the enlarged base more or less bifid. Male: Length about 6.5 mm.; wing 6.8 mm. Rostrum and palpi black, the former sub- equal in length to the remainder of head. An- tennae black throughout; flagellar segments oval, with long conspicuous verticils. Head black; eyes very large, reducing the anterior vertex to a linear strip. Pronotum brown, the pretergites more testaceous yellow. Mesonotal praescutum and scutal lobes brown, the humeral region more yellowed; median region of scutum and the scutellum brownish-yellow; postnotum light brown, the central area of mediotergite dark- er. Pleura and pleurotergite more reddish- brown. Halteres with stem obscure yellow, knob infuscated. Legs with coxae brownish- yellow; remainder of legs dark brown, the outer tarsal segments paling to light brown or yellowish-brown, not whitened as in most allied species. Wings (Text-fig. 12) with a strong brownish tinge, the prearcular field more whitened; stigma elongate-oval, darker brown; veins dark brown. Venation: Branches of Rs gradually divergent, cell R2 at margin only a trifle more extensive than cell R3; r-m short but present; m-cu at near two-thirds the length of vein M 3+4 and op- posite r-m. Abdomen, including hypopygium, brown- ish black. Male hypopygium (Text-fig. 15) with the tergite, 9 1, produced into two con- spicuous triangular earlike lobes that are further armed beneath with the usual lateral tergal arms, the spine of the latter straight to gently sinuous, its tip acute; entire pos- terior border of tergite, including the lobes, glabrous. Basistyle, h, unarmed with lobes but with a small concentration of setae on mesal face near proximal end. Dististyles, d, terminal, the outer style small, glabrous, its outer spine reduced to a triangular tubercle, the axial spine strong, decurved ; inner dis- tistyle longer, slightly dilated on basal half and here provided with scattered tubercles, each bearing a small seta. Aedeagus, a, be- yond the enlarged base more or less bifid, curved into a circle, the tip pale. Holotype, $ , Rancho Grande, July 31, 1946 (Beebe-Fleming). The yellowish-brown tarsi are quite dif- ferent from those of the various species that center around Helius ( Helius ) albitarsis (Osten Sacken), which have the tarsi snowy white. The structure of the male hypopygium of the present fly is distinctive. 42. Helius (Helius) rec tispina Alexander, 1947. Helius ( Helius ) rectispina Alexander; Bol. Ent. Venezolana, 6: 43-44, fig. 2; 1947. Rancho Grande, August 27, 1944, collected by Lichy; type. Still known only from the type locality. Protohellus Alexander. 43. Protohelius venezolanus, sp. n. General coloration of body dark brown to black; antennae unusually long, nearly one- third the length of wing; legs brownish- black, the outer tarsal segments paling to brown ; wings with a strong blackish tinge, the prearcular and costal fields a trifle darker; Sc long, Sci ending about opposite the fork of the long straight Rs; m-cu just beyond the fork of M. Female : Length about 7 mm. ; wing 8 mm. ; antenna about 2.5 mm. Rostrum short and inconspicuous; palpi comparatively long, black. Antennae unusu- ally long for the female sex, as shown by the measurements, black, the pedicel a trifle brightened; flagellar segments cylindrical, becoming shorter and more slender out- wardly; first segment long, nearly equal to segments two and three combined; verticils short and inconspicuous. Head black, sparse- ly pruinose ; eyes large, the anterior vertex reduced to a narrow strip. Pronotum reduced, hidden from above by the forward projecting praescutum; pre- tergites testaceous yellow. Mesonotum uni- formly dark brown; setae long and con- spicuous, especially on scutellum. Pleura dark brown. Halteres dark brown, the extreme base of stem brightened. Legs brownish- black, the trochanters more testaceous ; outer tarsal segments paling to brown; claws (fe- male) long and slender, simple. Wings (Text- fig. 13) with a strong blackish tinge, the pre- 46 Zoologica: New York Zoological Society [35: 2 arcular and costal fields a trifle darker; veins brownish-black. All veins beyond cord with abundant macrotrichia. Venation: Sci end- ing about opposite fork of the long straight Rs, Sc2 near its tip ; R2 and R1+2 subequal in length, the former pale but strong; Ri+n about four-fifths as long as Rs; inner end of cell 1st M2 arcuated; m-cu just beyond the fork of M. Abdominal tergites brown, the borders narrowly more blackened, sternites more uni- formly paler brown. Ovipositor with valves very long, the cerci slender, gently upcurved. Holotype, 9, Rancho Grande, July 27, 1946 (Beebe-Fleming) . The only other neotropical species of the genus is Protohelms cisatlanticus Alexander, of Ecuador. This differs in the much paler color of the body, legs and wings and in the details of venation. From the length of the antennae in the female sex of the present fly it is suspected that this organ in the male may be considerably longer than in cisat- lanticus. Each of the two species mentioned is known from a single specimen, cisatlan- ticus a male, venezolanus a female, and in these the antennae are virtually equal in both cases. Almost invariably in the Tipulidae, species having elongate antennae in the male sex have these much shorter in the corre- sponding females. Hexatomini. Paradelphomyla Alexander. 44. Paradelphomyia I Oxyrhiia I venezolana, sp. n. General coloration of thorax almost uni- formly medium brown; wings with a weak brownish tinge, stigma oval, pale brown; macrotrichia of wing cells very sparse, in the extreme outer ends of cells R3 to Mi inclu- sive; wings (male) widest nearly opposite the termination of vein 2nd A; male hypo- pygium with the arms of the ventral fork slender, their tips very feebly dilated, sub- acute ; gonapophysis with the blade triangu- lar, the mesal angle unusually slender and pointed. Male: Length about 4 mm. ; wing 4.4 mm. Rostrum black; palpi dark brown Anten- nae dark brown throughout; basal flagellar segments oval, the outer ones passing into cylindrical; verticils elongate. Head above dark. Thorax almost uniformly medium brown, the posterior sclerites somewhat more testa- ceous; praescutal setae very sparse but long. Halteres pale. Legs with the coxae brownish- yellow; trochanters light yellow; remainder of legs brown, the tarsal segments paling to yellowish brown ; tibial spurs present. Wings (Text-fig. 14) with a weak brownish tinge, the prearcular and costal fields a trifle more yellowed ; stigma oval, pale brown ; scarcely apparent dark seams over cord and outer end of cell 1st M2, best evidenced by a slight deepening in color of the veins, the remain- ing veins pale brown, yellowed in the bright- ened fields. Wings (male) conspicuously widened nearly opposite the termination of vein 2nd A. Macrotrichia of cells very sparse and restricted, in the extreme outer ends of cells R3 to Mi, inclusive, most numerous in cell Ri where they total 12 or 13. Venation: Sci ending about opposite fork of Rs, Scz nearly opposite two-thirds the length of this vein; veins Rz and Ri nearly parallel to one another, not as conspicuously divergent as in costaricensis, cell R3 at margin narrower; m-cu only a little more than its own length beyond the fork of M. Abdomen pale brown, the hypopygium and eighth segment brownish black. Male hypo- pygium (Text-fig. 18) with the outer dis- tistyle terminating in two slightly unequal spines, the additional ventral spine unusual- ly long and slender. Gonapophysis, g, with the apical blade triangular, the mesal angle unusually slender and pointed. Each arm of the ventral fork slender, the tip very feebly to scarcely dilated, the apex subacute. Holotype, 8, Rancho Grande, July 25, 1948 (Beebe-Fleming) ; migrant No. 481374. The most similar regional species is Para- delphomyia ( Oxyrhiza ) costaricensis (Alex- ander), which differs in the details of vena- tion, as described. Unfortunately, the male sex of costaricensis is still unknown and the important hypopygial features cannot be compared at this time. The tropical American species of the genus have been considered in a paper by the writer (Rev. de Entomol- ogia, 19: 151-153, fig. 2; 1948). Austrolimnophila Alexander. 45. Austrolimnophila l Austrolimnophila) viva s- berthieri Alexander, 1938. Austrolimnophila ( Austrolimnophila ) vivas-berthieri Alexander; Rev. de Entomologia, 9: 436-437; 1938. Rancho Grande, September 5, 1937, Vivas- Berthier; type. The species is still known only from the unique type. The tropical American species of Austro- limnophila are discussed by the writer else- where (Rev. de Entomologia, 19: 153-168, figs. 3, 4, 7-18; 1948). Eplphragma Osten Sacken. 46. Epiphragma lEpiphragma) enixa Alexan- der, 1939. Epiphragma ( Epiphragma ) enixa Alex- ander; Ann. Mag. Nat Hist., (11) 3: 190-192; 1939. Rancho Grande, March 15, May 8, 1946 (Beebe-Fleming). The types were from Abitagua, Ecuador, 1200 meters, March 29, 1937, collected by Clark-Macintyre. The spe- cies is still known only from Ecuador and Venezuela. 47. Epiphragma lEpiphragma) persancta Al- exander, 1938. Epiphragma ( Epiphragma ) persancta Alexander; Rev. de Entomologia, 9: 248-249; 1938. Rancho Grande, March 28, April 18, May 1950] Alexander: Tipulidae of Rancho Grande, Venezuela 47 8-23, 1946; April 10, May 8, 1945; April 25- May 3, 1948. Part of the type material was taken here on September 5, 1947, by Vivas- Berthier. The species occurs from Venezuela to southeastern Brazil. 48. Epiphragma lEpiphragmal so latrix (Osten Sacken, 1859). Limnophila ( Epiphragma ) solatrix Osten Sacken; Proc. Acad. Nat. Sci. Philadelphia, for 1859: 238; 1859. Rancho Grande, July 18-23, August 5, 1946. Also taken here August 29, 1937, by Vivas-Berthier. The species was originally described from the eastern United States. It ranges from southern New York through Mexico and Central America to northern Argentina. The neotropical species of Epiphragma have been discussed by the writer {Rev. de Entomologia, 19: 168-175, fig. 5; 1948). Polymera Wiedemann. 49. Polymera I Polymerodesl conjunctoides Al- exander, 1920. Polymera {Polymer odes') conjunctoides Alexander; Ent. News, 31: 74-75; 1920. Rancho Grande, June 28, 1946; 2 $$. The type was from Itacoatiara, Amazonian Bra- zil, October 16, 1919, collected by Parish. New to Venezuela; formerly fi'om Amazon- ian Brazil, Ecuador and Peru. The neotropical species of Polymera are considered in a paper by the writer {Rev. de Entomologia, 19: 182-190, figs 27-33; 1948). Limnophila Macquart. 50. Limnophila guttulatissima Alexander, 1913. Limnophila guttulatissima Alexander; Proc. U. S. Nat. Mus., 44: 546-547, fig. 38; 1913. Rancho Grande, July 17, 1946. Type from Totonicapan, Guatemala, collected by Eisen. Known hitherto from Guatemala and Costa Rica. The neotropical species of Limnophila are discussed in a paper by the writer {Rev. de Entomologia, 19: 513-518, figs. 6, 7; 1948). Shannonomyla Alexander. 51. Shctnnonomyia araguae Alexander, 1947. Shannonomyla araguae Alexander; Bol. Ent. Venezolana, 6: 48-50, figs. 3-4; 1947. Rancho Grande, July 25, 1948 (Beebe- Fleming) ; migrant No. 481374. The type was taken at this station on August 24, 1944, by Lichy. Still known only from the type locality. 52. Shannonomyla lathraea (Alexander 1926). Pilaria lathraea Alexander; Ann. Ent. Soc. America, 19: 386-387; 1926. Rancho Grande, May 9, 1948; a broken $, taken over a pool of water by a steep mossy rock cliff. Type from San Lorenzo Mt., Co- lombia, taken December 13, 1922. Known hitherto only from Colombia. 53. Shaimonomyia providers, sp. n. Mesonotal praescutum light brown with three darker stripes, the median one more evident; antennae short, dark brown; fe- mora and tibia obscure yellow, the tips weak- ly infuscated; wings with a weak brown tinge, restrictedly patterned with darker; vein R‘2 a short distance befoi'e fork of R 3+4; male hypopygium with the tergal lobes truncate, separated by a broad U-shaped notch; outer dististyle and gonapophysis bi- dentate. Male: Length about 4.5-5 mm.; wings 4.8- 5.5 mm.; antenna about 0.9-1 mm. Female: Length about 5.5 mm.; wing 6.2 mm. Rostrum dark brown, gray pruinose; palpi black. Antennae short, dark brown ; flagellar segments oval, shorter than the verticils. Head gray, somewhat clearer gray on front; a narrow blackish central line on posterior vertex; anterior vertex about two and one- half times the diameter of scape. Pronotum gray. Mesonotal praescutum light brown, the surface pruinose ; three fair- ly evident brown stripes, the lateral pair less distinct, crossing the suture onto the scutal lobes; posterior sclerites of notum brownish- gray. Pleurotergite and dorsal part of pleura brownish-gray, the ventral pleurites and sternum light yellow. Halteres dusky, the large knobs still more darkened. Legs with the coxae and trochanters pale yellow, the fore coxae a trifle more darkened; femora and tibiae obscure yellow, the tips weakly in- fuscated ; tarsi brownish-yellow, the termin- al segments darker. Wings (Text-fig. 19) with a weak brownish tinge, the prearcular and costal fields more yellowed ; a restricted darker brown pattern, including the small oval stigma and vague seams over the cord, outer end of cell 1st M2 and origin of Rs, best indicated by a slight deepening in color of the veins. Venation: Sc moderately long, Sci ending about opposite three-fourths to four- fifths Rs, Sc2 a short distance from its tip; Rs relatively long, about one-third longer than i?2+3+4j vein R2 a short distance be- fore fork of 72.3+4; vein Ri long, approxi- mately three times R3; cell 1st M2 a little longer than vein M4; m-cu shortly beyond the fork of M. Abdominal tergites brown, with a darker brown subterminal ring involving segments six and seven, segments eight and nine more yellowed, the styli dark; basal sternites yel- low. Male hypopygium (Text-fig. 25) with the tergal lobes, 9t, truncated, separated by a broad U-shaped notch. Outer dististyle, d, unequally bidentate at apex, the tip black- ened ; outer surface of style with long setae. Inner dististyle shorter, the outer surface with abundant erect setae. Gonapophysis, g, bidentate, the outer spine longest. Holotype, $, Rancho Grande, June 17, 1946 48 Zoologica: New York Zoological Society [35: 2 (Beebe-Fleming). Allotopotype, 9, July 22, 1948; migrant No. 481365, in copula with a paratype. Paratopotypes, 1 broken $, with allotype; 1 $, July 17, 1948. The present fly is most similar in its gen- eral appearance to species such as Shannon- omyia adumbrata Alexander and S. protuber- ans Alexander, differing in the coloration of the body and wings and in the structure of the antennae and the male hypopygium. The tropical American species of Shan- nonomyia have been recorded in a paper by the writer (Rev. de Entomologia, 19: 518- 523, figs. 8, 9, 11-13; 1948). Text-figs. 19-27. (Symbols: a, aedeagus; b, basistyle; d, dististyles; g, gonapophysis; s, sternite; t, tergite). 19 — Shannonomyia providens, sp. n.; venation. 20— Atarba ( Atarba ) quasimodo, sp. n. ; venation. 21 — Atarba ( Ischnothrix ) brevisector Alexander; venation. 22 — Atarba (Ischnothrix) spinituber, sp. n.; venation. 23 — Hexatoma (Erio- cera) beebeana, sp. n. ; venation. 24 — Atarba (Atarba) quasimodo, sp. n.; male hypopyg- ium. 25 — Shannonomyia providens, sp. n. ; male hypopygium. 26 — Atarba (Ischnothrix) spinituber, sp. n. ; male hypopygium. 27 — Atarba (Atarba) cincticomis thamia, subsp. n. ; male hypopygium. 1950] Alexander : Tipulidae of Rancho Grande, Venezuela 49 Hexatoma Latreille. 54. Hexatoma lErioceral beebeana, sp. n. Size relatively small (wing, male, 9 mm.) ; general coloration black, the praescutum and scutal lobes with metallic blue reflections; fore and middle femora with about the basal third yellow, the remainder blackened, en- closing a narrow obscure yellow subterminal ring; posterior femora clear yellow basally, the tip conspicuously blackened ; wings with the restricted ground whitened, much re- stricted by broad brown seams to the veins; cell R. i unusually short, vein Rz before the cell subequal in length to vein #3+4; a super- numerary crossvein at near midlength of cell R5 ; m-cu close to midlength of cell 1st M2. Male: Length about 10 mm.; wing 9 mm.; antenna about 2.5 mm. Rostrum brownish-black, yellow pollinose; palpi black, the first segment paler. Antennae (male) short, 7-segmented; scape and pedi- cel dark brown, flagellum black; first flagel- lar segment exceeding the combined second and third segments and a little shorter than the fourth; fourth segment nearly three times as long as the fifth. Head dark brown, sparsely pollinose on genae; vertical tubercle orange, deeply split medially. Pronotal scutum and pleura heavily light gray pruinose, the scutellum brownish-black. Mesonotal praescutum and the scutal lobes black with distinct metallic blue reflections; remainder of notum intensely black, very sparsely pollinose, the pleurotergite more heavily so; praescutal vestiture very short and sparse. Pleura black, sparsely gray pruinose, more heavily so on the pteropleur- ite; dorsopleural membrane brownish-black. Halteres black. Legs with the coxae black, heavily gray pruinose; trochanters black; fore and middle femora with about the basal third yellow, the remainder blackened, en- closing a narrow and inconspicuous obscure yellow subterminal ring; tibiae brown, the tip and the tarsi more blackened; posterior femora clear yellow basally, the tip conspicu- ously blackened, with vague indications of a broad brownish-yellow subterminal ring, the narrow area before the apex clearer yel- low; tibiae and tarsi black. Wings (Text-fig. 23) with the ground whitened, much re- stricted by broad brown seams to the veins ; costal border and bases of cells R and M, to- gether with the apex, more extensively and uninterruptedly brown ; both Anal cells uni- formly paler brown; veins brown, arculus conspicuously light yellow. Outer radial veins with abundant macrotrichia, these greatly reduced in number to vii’tually lacking in the medial field. Venation: Sci ending about opposite the fork of Rs; vein R3+4 subequal to vein R2, both a little shorter than R 1+2, cell Rn thus unusually short; a supernumer- ary crossvein at near midlength of cell .Rs; m-cu close to midlength of cell 1st M2. Abdominal tergites black, basal sternites medium brown, the outer ones intensely blackened; hypopygium small, obscure brownish-yellow. Holotype, 8, Rancho Grande, March 26, 1945 (Beebe-Fleming) . This striking crane-fly is respectfully ded- icated to Dr. William Beebe. There are no very close allies so far made known and the fly will be readily recognized. The presence of supernumerary crossveins in the wings is very uncommon in species of Eriocera, in the neotropical fauna including only Hexatoma ( Eriocera ) acunai Alexander, of Cuba, among the described species. This latter fly has the cross-vein in cell R4 instead of in cell R5, as in the present species. 55. Hexatoma I Eriocera) bifurcafa Alexander, 1947. Hexatoma ( Eriocera ) bifurcata Alex- ander; Bol. Ent. Venezolana, 6 : 97-99 ; 1947. Rancho Grande, July 10, 1945, 9; May 6, 1946, 9; July 9, 1946, 8. The type was from Choroni, Km. 25, taken September 12, 1945, by Lichy. Still known only from the state of Aragua, Venezuela. 56. Hexatoma lErioceral candidipes (Alexan- der, 1923). Penthoptera candidipes Alexander ; Ent. News, 34 : 20 ; 1923. Rancho Grande, July 18, 1946. Type from Tachira, Venezuela, taken April 9, 1920, by the Williamsons and Ditzler. Still known only from Venezuela. The tropical American species of Hexa- toma ( Eriocera ) have been discussed by the writer in another paper (Rev. de Entomo- logia, 19: 528-535, figs. 16-20; 1948. Atarba Osten Sacken. 57. Atarba llschnofhrix I spinifuber, sp. n. Color variable, the mesonotum ranging from reddish-brown to dark brown ; antennae ( male) elongate, exceeding the wing in length, the segments with unusually long erect setae scattered over their whole length ; wings with a weak brownish tinge; Sc short; Rs short, weakly angulated; vein Rs short, arcuated, less than one-third the petiole of cell R 3 ; male hypopygium with the appendage of the eighth sternite an elongate lobe, of the ninth sternite a conspicuous lyriform structure; basistyle on mesal face near proximal end with a cylindrical lobe bearing spinous setae ; aedeagus sinuous to the acute apex. Male: Length about 5.5-5. 7 mm.; wing 6- 6.2 mm.; antenna about 6-7 mm. Rostrum yellow pollinose; palpi brownish- yellow. Antennae (male) elongate, exceeding the wing in length, pale brown ; flagellar seg- ments very elongate-cylindrical, with dense erect pale setae, these unusually long and conspicuous, the longest approximately two- thirds the length of the segment; in addition to the longest vestiture there is an even more abundant shorter type of seta that is about one-fourth the segment. Head dark brown- ish gray. Thorax of type dark reddish-brown to brown, of the paratype much darker brown 50 Zoologica: New York Zoological Society [35: 2 to brownish-black. Pleura of type light yel- low, of the paratype more brownish-yellow, sparsely pruinose. Halteres infuscated, the base of stem narrowly pale yellow. Legs with the coxae and trochanters yellow; remainder of legs yellowish-brown (type) to brownish- black (paratype) . Wings (Text-fig. 22) with a weak brownish tinge, the oval stigma darker brown, inconspicuous; veins brown, those near wing base a trifle brighter. Ven- ation: Sc short, Sc i ending just beyond ori- gin of Rs, Sc2 before this origin; Rs short, weakly angulated at origin; vein Rs short, arcuated, less than one-third the petiole of cell Rs ; cell Rs at margin about one-third to two-fifths the extent of cell Rs; m-cu at near one-third the length of cell 1st M s. Abdomen yellowish-brown to brown, the subterminal segments still darker to form a ring; hypopygium and sternites more yel- lowed. Male hypopygium (Text-fig. 26) with the appendage of the eighth sternite, 8s, a depressed-flattened elongate lobe, narrowed to obtuse tip, with setae over the whole sur- face. Appendage of ninth sternite, 9s, pro- foundly lyriform, the branches unusually long and slender, near apex with weak re- trorse points or roughenings. Basistyle, b, on mesal face of proximal half with a finger- like lobe that is tufted with strong black spinous setae, those at apex even stouter. Outer dististyle, d, a slender blackened gently curved rod, provided with scattered spines along the outer face, the terminal one sub- equal to the apex and slightly more slender. Inner dististyle subequal in length but ex- tending beyond the dorsal dististyle due to its place of insei'tion, gently arcuated. Aedeagus, a, sinuous to the acute apex. Holotype, $, Rancho Grande, May 5, 1945 (Beebe-Fleming) . Paratopotypes, S$, May 7, 1945, May 27, 1948. The closest relative of the present fly is Atarba ( Ischnothrix ) brevisector (Alex- ander) , described from the cloud forest zone of the Silla de Caracas, 1900 meters. This is a larger fly (wing, $, nearly 8 mm.) that is still known only from the female sex. Be- sides the large size, this species has the an- tennal flagellum black, with short vestiture, the latter undoubtedly a sexual character. The venational details (Text-fig. 21) are dis- tinct, particularly the outer radial field where vein Rs is normally oblique and relatively long, exceeding one-half the length of the petiole of cell Rs; cell Rs at margin scarcely one-third that of cell Rs. The tropical American species of Atarba have been considered by the writer in another paper {Rev. de Entomologia, 19; 536-552, figs. 21-35; 1948). 58. Atarba I Atarba I cincticornis thamia, subsp. n. Male : Length about 5 mm. ; wing 5.5 mm. ; antenna about 4.5 mm. Antennal flagellum almost uniformly dark- ened, only the incisures of the more prox- imal segments restrictedly yellow; on outer segments the amount of pale color is some- what increased ; basal flagellar segments long-cylindrical, with conspicuous outspread- ing verticils and erect setae; outer segments progressively much shorter, the outer four or five scarcely more than four times as long as the diameter across base. In typical cincti- cornis, the antennae are longer, the segments more evidently bicolored, their bases dis- tinctly thicker than their tips. Mesonotum almost uniformly dark brown, the postnotum and pleura more pruinose. Halteres infuscated. Legs yellow, the outer tarsal segments darker. Wings with a weak brownish tinge, the costal border more yel- lowed; stigma small; veins pale brown, Sc more yellowed. Venation : Sci ending virtual- ly opposite the origin of Rs, Sc2 a short dis- tance from its tip; Sci nearly as long as the short Rs, the latter subequal to the basal sec- tion of Rs; cell 1st Ms nearly rectangular in outline, the second section of Mi+ 2 approx- imately twice the basal section; m-cu about one-third its length beyond the fork of M. In typical cincticornis , Rs is longer; cell 1st M 2 very small and short, the second section of vein M1+2 being subequal to or actually shorter than the basal section; m-cu at or close to the fork of M. Abdomen brown, with a dark brown sub- terminal ring; hypopygium yellow. Male hypopygium (Text-fig. 27) generally as in the typical race, particularly as regards the outer dististyle and the aedeagus. Outer dis- tistyle, d, dilated on outer half, the outer margin with conspicuous black spines, those at near midlength of the group more crowd- ed and in double rows. Inner dististyle with conspicuous setae, especially along the outer : margin. Gonapophysis simple, appearing as an elongate blade, the top subacute. Aedeagus, a, very long and slender, longer than the com- 1 bined basistyle and dististyles, the apex not or scarcely expanded. In typical cincticornis, the outer dististyle has the central spines shorter and arranged in several rows. Inner dististyle shorter, without conspicuous setae. Gonapophysis even more slender and elongate. Holotype, $, Rancho Grande, June 26, 1946 (Beebe-Fleming) . The discovery of more material will prob- ably result in elevating the present fly to full specific rank. Typical cincticornis Alexander is known from British Guiana and Ecuador. 59. Atarba I Atarba l quasimodo, sp. n. Thorax almost uniformly brownish-yellow, unpatterned; antennae (male) about one- third the length of wing, flagellum black ; legs yellow; wings with a strong yellow tinge, stigma very pale brown, inconspicuous; Sci ending about opposite one-third the length of Rs; cell 1st Ms pentagonal in outline; male hypopygium with the outer lateral angles of the appendage of the ninth sternite I produced into very slender spines; outer dis- tistyle with about nine spines on outer mar- gin; gonapophyses appearing as bilobed smooth blades; aedeagus elongate, stout, the 1950] Alexander : Tipididae of Rancho Grande, Venezuela 51 apex a little dilated, on margin at near mid- length with a very conspicuous obtuse tu- bercle or hump. Male: Length about 5 mm.; wing 4. 5-4. 6 mm.; antenna about 1.4-1. 5 mm. Rostrum dark brown ; palpi more reddish- brown. Antennae (male) of moderate length, about one-third the length of the wing; fla- gellar segments long-subcylindrical, exceed- ing the verticils, the latter stout, about four per segment ; the entire segment with a dense erect pale pubescence, these approximately one-half the length of the verticils. Head dark gray; anterior vertex broad. Text-figs. 28-39. (Symbols: a, aedeagus; b, basistyle; bd, basal dististyle; d, dististyle; p, phallosome; t, tergite). 28 — Sigmatomera beebei, sp. n.; venation. 29 — Trentepohlia (Paramongoma) fuscolimbata, sp. n ; venation. 30 — Gonomyia ( Gonomyia ) spiculistyla, sp. n. ; venation. 31 — Erioptera (Empeda) abitaguae Alexander; venation. 32 — Erioptera (Erioptera) beebeana, sp. n.; venation. 33 — Molophilus ( Molophilus ) compactus, sp. n.; venation. 34 — Sigmatomera beebei, sp. n.; male hypopygium. 35 — Gonomyia ( Gonomyia ) spiculistyla, sp. n.; male hypopygium. 36 — Gonomyia ( Lipophleps ) vindex Alexander; male hypopygium 37 — Erioptera ( Erioptera ) beebeana, sp. n.; male hypopygium. 38 — Molophilus ( Molophilus ) compactus, sp. n. ; male hypopygium. 39 — Molophilus ( Molo- philus) flemingi, sp. n.; male hypopygium. 52 Zoologica : New York Zoological Society [35: 2 Thorax almost uniformly brownish-yel- low, unpatterned, the pleura more yellowed. Halteres with stem yellow, knobs weakly darkened. Legs with the coxae and tro- chanters reddish-yellow; remainder of legs yellow, the terminal tarsal segments more infuscated. Wings (Text-fig. 20) with a strong yellow tinge, the prearcular and costal fields clearer yellow; stigma oval, very pale brown, inconspicuous; veins yellow. Vena- tion: Sc moderately long, Sc\ ending about opposite one-third the length of Rs, Sc2 faint, a short distance before this origin ; branches of Rs very gently divergent; cell 1st M 2 pen- tagonal, less than one-half the basal section of M: s; m-cu variable in position, from a short distance before the fork of M, as illus- trated, to a little beyond this fork. Abdomen light brown, the bases of the seg- ments paler; subterminal segments dark brown to produce a conspicuous ring; hypo- pygium yellow. Male hypopygium (Text-fig. 24) with the appendage of the ninth sternite, 9s, short and broad, the caudal margin nearly truncate, very slightly emarginate at the midline; outer lateral angles produced lat- erad into unusually slender spines; surface of appendage with relatively few coarse setae. Outer dististyle, d, relatively slender, the outer margin with teeth over virtually the whole length, totalling nine or ten, the more basal ones smaller and appi'essed, the outer spines longer and more conspicuous; terminal spine subequal to the subterminal one and slightly more curved; on ventral margin before apex with three or four deli- cate spinules. Inner dististyle a little longer than the outer style, appearing as a darkened slender rod, gently curved, the obtuse tip a little enlarged. Gonapophysis, g, appearing as bilobed smooth blades. Aedeagus, a, elon- gate, stout, the apex a little dilated and flar- ing; on margin at near midlength with a very conspicuous obtuse dilation or tubercle. Holotype, 8, Rancho Grande, May 12, 1945 (Beebe-Fleming) . Paratopotype, 8, pinned with type. The present fly is quite distinct in the structure of the male hypopygium. In the prominent tubercle on the aedeagus, it sug- gests Atarba ( Atarba ) tuberculif era Alex- ander, but is an entirely different fly, being one of the smallest members of the genus whereas tuberculif era is one of the largest. Elephantomyia Osten Sacken. 60. Elephantomyia I Elephantomyia I setuiistyla Alexander, 1938. Elephantomyia ( Elephantomyia ) setu- listyla Alexander; Ann. Mag. Nat. Hist., (11) 1: 351-352; 1938. Rancho Grande, May 12-JuIy 8, 1946; July 17, 1948. The type was from Abitagua, Ecuador, 1200 meters, April 7, 1937, col- lected by Clarke-Macintyre. Known only from Ecuador and Venezuela. The tropical American species of Elephan- tomyia have been listed in a further paper by the writer (Rev. de Entomologia, 19 : 552- 556, figs. 27, 28, 36; 1948). Eriopterini. Sigmatomera Osten Sacken. 61. Sigmatomera heehei, sp. n. General coloration reddish-yellow, unpat- terned; halteres and legs yellow; wings uni- formly rich yellow, all the veins deep yellow; cell 1st M2 elongate, subequal in length to the distal section of vein M1+2; male hypo- pygium with the dorsal lobules of the ninth tergite very small, suboval in outline; inner dististyle stout, the lobe on its outer margin very low and obtuse; aedeagus without elon- gate spines at apex. Male: Length about 14 mm.; wing 14.5 mm. Head broken. Prothorax and mesothorax almost uniformly reddish-yellow, the prae- scutum with three such stripes on a slightly more yellowed ground. Halteres uniformly yellow. Legs yellow, only the terminal tarsal segment brown. Wings (Text-fig. 28) uni- formly rich yellow, the prearcular and costal borders more saturated yellow; all veins deep yellow. Venation: Sci ending about opposite one-third the length of the outer or angu- iated section of Rn 3+4, SC2 opposite this angu- lation; E2+3+4 angulated just before mid- length, with a long spur jutting basad into cell Rl at the angle; cell 1st M 2 elongate, widened outwardly, subequal in length to the distal section of vein M 1+2; basal section of vein Ms, with a weak spur jutting basad into cell 1st M 2; m-cu at or close to the fork of M. Abdomen brownish-yellow, hypopygium yellow. Male hypopygium (Text-fig. 34) with the dorsal lobules of the ninth tergite, 9t, very small, suboval in outline, separated by a space that is subequal to the diameter of either lobe. Inner dististyle stout, the lobe on outer margin very low and obtuse. Aedeagus, a, stout, terminating in a small central point, not produced into elongate spines, as com- mon in the genus; if present, these latter are microscopic. Holotype, 8, Rancho Grande, May 2, 1945 (Beebe-Fleming) . I am very pleased to name this species for Dr. William Beebe. The fly is most similar to other species with unpatterned wings, such as Sigmatomera flavipennis Osten Sacken (genotype) and S. occulta Alexander. It dif- fers from these and from all other known forms in the uniformly bright yellow veins and in the structure of the male hypopygium. In the latter respect, the species differs very markedly from flavipennis, which has the inner dististyle long and slender. The species that have this style most like that in the present fly include S. occulta and S. shan- noniana Alexander, which differ evidently in the pattern of the wings and in the details of the hypopygium, including the tergite, in- ner dististyle and aedeagus. The tropical American species of Sigma- tomera have been listed in another report by 1950] Alexander: Tipulidae of Rancho Grande , Venezuela 53 the writer {Rev. de Entomologia, 18: 68-69, figs. 3, 4; 1947). Trentepohlia Bigot. 62. Trentepohlia IParamongomal fuscolimbata, sp. n. Mesonotum and pleura almost uniformly reddish-brown; legs medium brown, the tip of the tibia and the tarsi paling to cream- yellow; wings grayish-subhyaline; stigma relatively large and conspicuous, subcircular to short-oval in outline ; veins of the proximal three-fourths of wing extensively seamed with paler brown; vein Rs oblique. Female: Length about 10 mm.; wing 8.2 mm. Rostrum yellow. Antennae with the scape and pedicel dark brown; flagellum broken. Head brown ; anterior vertex reduced to a linear strip that is subequal in width to a single row of ommatidia. Cervical region dark brown above, paler on sides. Pronotum dark brown above, paling to yellowish-brown laterally. Mesonotum and pleura almost uniformly reddish-brown, the scutellum and especially the postnotum sparsely gray pruinose. Halteres pale. Legs with the coxae reddish-brown; trochanters yellow; legs medium brown, the genua, espe- cially the tibial base, very narrowly and almost insensibly whitened; tip of tibia and the tarsi paling to cream-yellow. Wings (Text-fig. 29) grayish-subhyaline; stigma relatively large and conspicuous, subcircular to short-oval; veins before level of outer end of cell 1st M 2 extensively seamed with paler brown, such cloudings on Rs, Rs* 3+4, cord, outer end of cell 1st M2, outer end of M and M: 5+4, distal half of Cu, and at and near the bases of the Anal cells; wing tip even more vaguely suffused; veins brownish-yellow, somewhat clearer yellow in the outer costal field. Venation : Vein Rs more oblique than in suffuscipes; cell 1st M 2 broader; m-cu only a short distance before the fork of M. Abdominal tergites dark brown, sternites yellow; genital shield brownish-yellow. Cerci relatively long and very slender. Holotype, 9, Rancho Grande, May 11, 1945 (Beebe-Fleming) . The most similar regional species is Tren- tepohlia ( Paramongoma ) suffuscipes Alex- ander, which differs in the coloration of the body, legs and wings, as well as in slight details of the venation. The tropical American species of Trente- pohlia have been considered earlier by the writer {Rev. de Entomologia, 18: 69-72, figs. 5-7; 1947). Teucholabis Osten Sacken. 63. Teucholabis ITeucholabisi anfhracina Al- exander, 1921. Teucholabis anthracina Alexander ; Proc. Acad. Nat. Sci. Philadelphia, for 1921: 85-86; 1921. Rancho Grande, May 12, 1945; 1 9 only. The type was from the Napo River, Ama- zonian Peru, taken June 14, 1920, by Parish. The fly was known hitherto from Ecuador and Peru. Despite the lack of the male sex I can see no reason to question the determi- nation. The tropical American species of Teucho- labis were discussed by the writer in an earlier report (Rev. de Entomologia, 17: 375-400, figs. 1-14; 1946). Gnophomyia Osten Sacken. 64. Gnophomyia IGnophomyial digiliformis Alexander, 1941. Gnophomyia ( Gnophomyia ) digitifor- viis Alexander ; Journ. N . Y. Ent. Soc., 49: 144-145; 1941. Rancho Grande, March 27; April 8-24; June 18; July 1-18; August 16, 1946. Type fi”om San Esteban, Venezuela, taken Decem- ber 19, 1939, by Anduze. The species is still known only from Venezuela. The neotropical species of Gnophomyia have been discussed briefly in another paper by the writer (Rev. de Entomologia, 18: 72- 80, figs. 8-12; 1947). Neognophomyla Alexander. 65. Neognophomyia monophora Alexander, 1941. N eo gnophomyia monophora Alexander; Journ. N. Y. Ent. Soc., 49: 146-147; 1941. Rancho Grande, April 24, 1948 (No. 4872) ; July 25, 1948 (No. 481374), migrant series. The types were from San Esteban, Venezuela, December 28, 1939, Anduze, and Borburata, Venezuela, 500 meters, March 10-15, 1940, Anduze. Still known only from Venezuela. The tropical American species of Neogno- phomyia are listed by the writer (Rev. de Entomologia, 18: 81-82, fig. 16; 1947). Gonomyia Meigen. 66. Gonomyia (Gonomyial spieulistyla, sp. n. Mesonotum chiefly medium brown; scu- tellum obscure yellow; legs brownish-black; wings with a weak dusky tinge; Sci ending opposite origin of Rs, R2+3H subequal to Rs; male hypopygium with the outer disti- style a simple stout rod, the surface with abundant short spines or spicules, without setae. Male: Length about 4 mm.; wing 4.2 mm. Head broken. Pronotum brown, the preter- gites light yellow. Mesonotum chiefly medium brown, the praescutum darker medially ; scutal lobes dark brown; median region of scutum and the scutellum obscure yellow, the latter restrictedly darkened at base; postnotum brownish-gray, the lateral bor- ders of the mediotergite yellow; pleuro- tergite paler and pruinose. Pleura variegated dark reddish-brown and yellow, the latter color producing a vague stripe on the dorsal sternopleurite and pteropleurite. Halteres 64 Zoological New York Zoological Society [35: 2 dark brown. Legs with the coxae dark red- dish-brown; trochanters yellow; remainder of legs brownish-black. Wings (Text-fig. 30) with a weak dusky tinge, the stigmal area slightly darker, ill-delimited and vague; veins dark brown. Venation: Sci ending opposite origin of Rs, Scz at its tip; R2+3+4 subequal to or a trifle longer than Rs; vein R3 oblique; r-m long, subequal to m-cu, the latter at near one-third the length of cell 1st M2. Abdomen dark brown. Male hypopygium with the dististyles (Text-fig. 35, d ) dis- tinctive; outer style a simple stout rod, its surface densely covered with short spines or triangular points to produce a spiculose appearance, without setae. Inner dististyle appearing as a straight rod, before apex on outer margin with a strong arm that termi- nates in a flattened black plate, strongly recurved, its lower apical angle further pro- duced into a point; main axis of style bear- ing the usual two strong fasciculate setae, additional to a few other normal bristles. Phallosome broken and in part lost before mounting; a single strong slender spine per- sists in the slide mount; from a cursory pre- liminary examination made before the loss of this part, it is believed that there was a second spine and a further elongate pale rod that was expanded at its apex into a tri- angular blade, the apex shallowly emargi- nate. Holotype, 8, Rancho Grande, July 10, 1946 (Beebe-Fleming) . While similar in its general appearance to species such as Gonomyia ( Gonomyia ) juarezi Alexander, G. (G.) methodica Alex- ander, and others, the present fly is entirely distinct in the structure of the male hypopy- gium, including both the dististyles and the phallosome. 67. Gonomyia I Lipophleps I vindex Alexander, 1941. Gonomyia (Lipophleps) vindex Alex- ander; Journ. N. Y. Ent. Soc., 49: 142-143; 1941. Rancho Grande, May 12, 1945. The type was from Borburata, Venezuela, 500 meters, taken March 15, 1940, by Anduze. Still known only from Venezuela. In the present speci- men, the male hypopygium (Text-fig. 36), while being generally similar to that of the type, differs in slight details, especially the longest element of the phallosome, p, where the outer spine is longer and more conspicu- ous than in the type. 68. Gonomyia I Paralipophlepsi lemniscata Al- exander, 1931. Gonomyia (Lipophleps) lemniscata Alex- ander; Ann. Ent. Soc. America, 24: 634-635; 1931. Gonomyia (Paralipophleps) lemniscata Alexander; Rev. de Entomologia, 18: 97-98; 1947. Rancho Grande, July 10, 1946. The types were from Vista Nieve, Mount Santa Marta, Colombia, 5,000 feet, August 8, 1926, taken by Fred W. Walker, and from Jaragua, Santa Catharina, Brazil, taken September 20, 1929. The species is known from Colombia, Vene- zuela and southeastern Brazil. The tropical American species of Gon- omyia have been discussed in a paper by the writer (Rev. de Entomologia, 18: 83-99, figs. 17-29; 1947). Erloptera Meigen. 69. Erioptera I Erioptera I beebeana, sp. n. Allied to multiannulata ; general coloration of thorax and abdomen dark brown, the thor- acic pleura with a longitudinal silvery white stripe; femora with three dark brown and three china-white rings ; tibiae and basitarsi variegated with darkened annuli ; wings with a strong light brown tinge, unpatterned ; male hypopygium with both dististyles slender, the outer one terminating in several strong spines ; inner styles very slender ; phallosome with the apophyses bispinous. Male: Length about 2.5 mm.; wing 3 mm. Rostrum brown; palpi black. Antennae black throughout; flagellar segments oval, with very long verticils. Head above gray. Pronotum and mesonotum almost uni- formly dark brown, the surface with a very sparse pruinosity ; praescutum with two very faintly indicated darker brown stripes; an- terior pretergites obscure whitish, the region before the wing root pale. Pleura and pleuro- tergite dark brown, pruinose, with a narrow silvery white longitudinal stripe that is bor- dered both above and below by a slightly narrower more blackened line. Halteres clear light yellow. Legs with the fore coxae dark brown, the middle pair a little paler; poste- rior coxae and trochanters testaceous yellow; femora conspicuously ringed with dark brown and china white, there being three dark rings and three narrower white ones, the last terminal; tibiae yellow with a nar- row nearly basal dark ring and a more exten- sive subterminal one, the latter a little less than the pale tip ; tarsi yellow, the proximal end of the basitarsi narrowly darkened ; two outer tarsal segments infuscated; vestiture of legs including setae and normal linear scales. Wings (Text-fig. 32) with a strong light brown tinge, the prearcular and costal regions a trifle more brightened; veins brownish-yellow, the vestiture pale brown. Venation: Petiole of cell M3 from one-third to one-half longer than m-cu ; vein 2nd A gently sinuous on nearly the outer half. Abdomen, including hypopygium, dark brown, the ninth segment more yellowed. Male hypopygium (Text-fig. 37) with both dististyles, d, slender, terminal in position; outer style blackened, at apex with about three strong spines, with other smaller den- ticles back from tip, inner style subequal in length but even more slender, glabrous, the tip acute, near apex with two or three pale punctures. Phallosome, p, with powerful horn-colored spinous apophyses, one pair on either side, the strong inner spines nearly 1950] Alexander: Tipulidae of Rancho Grande, Venezuela 55 parallel to one another, their tips gently out- curved, acute; lateral arms approximately equal in length but a little more slender and sinuous. Holotype, $, Rancho Grande, July 25, 1948 (Beebe-Fleming) ; migrant series. This attractive crane-fly is named in honor of Dr. William Beebe. The most similar re- gional species include Erioptera ( Erioptera ) multiannulata Alexander, E. (E.) polydonta Alexander, and E. ( E .) polytriclia Alexander, all of which have the male hypopygium en- tirely distinct in structure. The neotropical species of Erioptera have been considered in an earlier report by the writer {Rev. de Entomologia, 18: 328-337, figs. 10-17; 1947). 70. Erioptera I Erioptera) celestis Alexander, 1940. Erioptera ( Erioptera ) celestis Alex- ander; Ann. Mag. Nat. Hist., (11) 5: 294-296; 1940. Rancho Grande, July 25, 1948 (Beebe- Fleming) ; migrant, No. 481374. The type was from Abitagua, Ecuador, 1200 meters, taken April 5, 1937, by Clarke-Macintyre. Known only from Ecuador and Venezuela. 71. Erioptera lEmpeda) abitagua! Alexander, 1941. Erioptera ( Empeda ) abitaguai Alex- ander; Ann. Mag. Nat. Hist., (11) 8: 131-132; 1941. Rancho Grande, May 12, June 11-28, July 8-10, 1946. The type was from Abitagua, Ecuador, 1100 meters, taken September 1, 1939, by Clarke-Macintyre. Known only from Ecuador and Venezuela. The wing venation is shown (Text-fig. 31). Molophilus Curtis. 72. Moiophilus I Molophilus) compactus, sp. n. Belongs to the plagiatus group; mesono- tum chiefly reddish-brown, unpatterned ; an- tennae (male) relatively long, nearly one- half the length of the body; wings with a strong brownish tinge; male hypopygium with the basal dististyle a slender nearly straight rod that narrows into a long spine, the lower face at near midlength with a com- pact group of erect setae, beyond this brush with a few strongly appressed spinous points. Male : Length about 3.2 mm. ; wing 4 mm. ; antenna about 1.5 mm. Rostrum and palpi black. Antennae rela- tively long, nearly one-half the body, black throughout; flagellar segments long-sub- cylindrical; longest verticils unilaterally dis- tributed, nearly twice as long as the seg- ments ; additional shorter erect pale verticils. Head gray. Pronotum brown, darker laterally ; preter- gites pale yellow. Mesonotum chiefly reddish- brown, the praescutum unpatterned. Pleura darker brown. Halteres yellow basally, darker beyond, with golden yellow vestiture. Legs with the coxae and trochanters testa- ceous yellow; remainder of legs medium brown, the outer tarsal segments more blackened. Wings (Text-fig. 33) with a strong brownish tinge, the prearcular and costal fields slightly more yellowed; veins and macrotrichia brown. Venation ; R„ lying slightly distad of the level of r-m; petiole of cell Ms about one-fourth longer than the slightly sinuous m-cu; vein 2nd A long, end- ing about opposite midlength of m-cu. Abdomen brown, the hypopygium brown- ish-black. Male hypopygium (Text-fig. 38) with the beak of the basistyle, b, slender. Outer dististyle with the two arms unequal, the more distal one slender. Basal dististyle, bd, a slender nearly straight rod that nar- rows into a long spine, on lower face at near midlength with a compact group of erect setae, beyond this brush with a few strongly appressed spinous points. Phallosomic plate unusually small, oval, the surface glabrous. Holotype, $ , Rancho Grande, June 28, 1946 (Beebe-Fleming). The most similar regional species include Molophilus {Molophilus) brownianus Alex- ander, M. (M.) capricornis Alexander, and M. (M.) paganus Alexander, all differing from the present fly and among themselves chiefly in the structure of the male hypo- pygium, particularly of the basal dististyle. 73. Molophilus I Molophilus) Renting i, sp. n. Belongs to the plagiatus group; thorax al- most uniformly black; antennae (male) relatively short; halteres yellow; legs black; wings grayish-subhyaline, vaguely cross- banded with pale brown, the bands lying at cord and beyond the arculus; male hypopyg- ium with the basal dististyle a long gently curved rod that terminates in a short spine; lower face of distal half of style with a series of strong appressed spines. Male : Length about 4 mm. ; wing 4.5 mm. ; antenna about 1 mm. Rostrum and palpi black. Antenna (male) relatively short, only about one-fourth the length of body, black throughout; flagellar segments subcylindrical or long-suboval, gradually decreasing in length and diameter outwardly ; verticils scattered, much exceed- ing the segments ; remaining vestiture short and inconspicuous. Head gray. Thorax almost uniformly black; preter- gites very narrow, testaceous yellow. Halteres yellow. Legs with the coxae infuscated bas- ally, paling to yellow outwardly; trochanters yellow; femora dark brown, yellow basally; remainder of legs black; fore tibia (male) with a linear glandular swelling. Wings gray- ish subhyaline, vaguely crossbanded with pale brown, including a band at cord and another across the postarcular region; pre- arcular field yellow; veins brown, yellow in the more brightened portions. Venation: Rz in approximate transverse alignment with r-m; petiole of cell Ms a little longer than m-cu; vein 2nd A relatively short, ending slightly beyond the posterior end of m-cu. 56 Zoologica: New York Zoological Society [35: 2: 1950] Abdomen, including hypopygium, brown- ish-black to black. Male hypopygium (Text- fig. 39) with the beak of the basistyle, b, slender. Outer dististyle elongate, the two arms unequal, the inner long and slender. Basal dististyle, bd, a long slender gently curved rod, the terminal spine short and stout; lower face of distal half of style with a series of strong appressed spines, on prox- imal half more scattered and finally becoming tuberculate ; outer face of style with a few small inconspicuous denticles back from apex. Holotype, $ , Rancho Grande, June 28, 1946 (Beebe-Fleming) . I take unusual pleasure in naming this distinct fly for Mr. Henry Fleming, Ento- mologist of the Department of Tropical Re- search of the New York Zoological Society, whose diligent collecting has added mate- rially to our knowledge of the crane-flies of Venezuela. The fly is quite distinct from the other members of the plagiatus group that have the basal dististyle of the male hypo- pygium of somewhat the same structure and appearance. These include Molophilus ( Molo - philus) brownianus Alexander, M. ( M .) cap- ricomis Alexander, M. ( M .) compactus, sp. n., M. ( M .) luxuriosus Alexander, M. (M.) paganus Alexander, and some others, all dif- fering among themselves in the pattern of the wings, length and structure of the an- tennae, and in the details of structure of the male hypopygium. The tropical American species of Molo- philus have been listed by the writer in an- other report (Rev. de Entomologia, 18: 338- 353, figs. 19-28; 1947). Styringomyia Loew. 74. Styringomyia americana Alexander, 1914. Styringomyia americana Alexander; Trans. Amer. Ent. Soc., 40 : 231 ; 1914. Rancho Grande, July 10, 1946. The type was taken at Mallali, British Guiana, on March 8, 1913, by Parish. This is the most widely distributed of the American species of the genus, occurring in British Honduras, Costa Rica, Colombia, Venezuela, British Guiana, Surinam and Ecuador. The tropical American species of Styrin- gomyia have been discussed in an earlier pa- per by the writer (Rev. de Entomologia, 18: 354-356, fig. 29; 1947). Toxorhlna Loew. 75. Toxorh ina IToxorhinal pergracilis Alexan- der, 1944. Toxorhina (Toxorhina) pergracilis Alexander; Ann. Mag. Nat. Hist., (11) 11: 181-182, fig. 13; 1944. Rancho Grande, June 28, July 10-27, Au- gust 1, 1946. The type was from Ayna, La Mar, Ayacucho, Peru, 2400 meters, collected April 27, 1941, by Woytkowski. Known only from Peru and Venezuela. 76. Toxorhina IToxorhinal stenophallus Alexan- der, 1937. Toxorhina ( Toxorhina ) stenophallus Alexander; Ann. Mag. Nat. Hist., (10) 20: 503-504; 1937. Rancho Grande, August 27, 1944, collected by Lichy. The type was from Nova Teutonia, southeastern Brazil, taken September 30, 1935, by Plaumann. Still known only from Venezuela and Brazil. The tropical American species of Tox- orhina have been considered by the writer in an earlier report (Rev. de Entomologia, 18: 356-360, figs. 30-32; 1947). Beebe: Migration of Butterflies at Rancho Grande, Venezuela 57 3. Migration of Danaidae, Ithomiidae, Acraeidae and Heliconidae ( Butterflies ) at Rancho Grande, North-central Venezuela.1 William Beebe. Director, Department of Tropical Research, New York Zoological Society. (Text-figure 1). [This is one of a series of papers resulting from the 45th, 46th and 47th Expeditions of the Department of Tropical Research of the New York Zoological Society, made during 1945, 1946 and 1948, under the direction of Dr. William Beebe, with headquarters at Rancho Grande in the National Park of Aragua, Venezuela. The expeditions were made possible through the generous cooperation of the National Govern- ment of Venezuela and of the Creole Petroleum Corporation. [The characteristics of the research area are in brief as follows: Rancho Grande is located in north-central Venezuela (10° 21' N. Lat., 67° 41' W. Long.), 80 kilometers west of Cara- cas, at an elevation of 1.100 meters in the un- disturbed montane rain forest which covers this part of the Caribbean range of the Andes. The migration flyway of Portachuelo Pass, which is also the water-shed between the Carib- bean and Lake Valencia, is 200 meters from Rancho Grande. Adjacent ecological zones in- clude seasonal forest, savanna, thorn woodland, cactus scrub, the fresh-water lake of Valencia and various marine littoral zones. The Rancho Grande area is generally subtropical, being uni- formly cool and damp throughout the year be- cause of the prevalence of the mountain cloud cap. The dry season extends from January into April. The average humidity during the expedi- tions, including parts of both wet and dry seasons, was 94.4%; the average temperature during the same period was 18° C.; the average annual rainfall over a five-year period was 174 cm. The flora is marked by an abundance of mosses, ferns and epiphytes of many kinds, as well as a few gigantic trees. For further de- tails see Beebe and Crane, Zoologica, Vol. 32, No. 5, 1947. Unless otherwise stated, the speci- mens discussed in the present paper were taken in the montane cloud forest zone, within a radius of one kilometer of Rancho Grande. [For an account of Portachuelo Pass, together with a general introduction to the groups of migrating insects and migrating factors, see “Insect Migration at Rancho Grande,” by William Beebe, Zoologica, 1949, Vol. 34, No. 12, pp. 107-110], Migration op Danaidae. Among the migrants through Portachuelo Pass are two members of the subfamily Danainae. The first is Danaus plexippus meg- 1 Contribution No. 871, Department of Tropical Research, New York Zoological Society. alippe, so close to our northern monarch, D. p. plexippus, that it is impossible to distin- guish them in flight. The second species is Danaus eresimus eresimus, corresponding to our northern queen, Danaus gilippus bere- nice. The monarch is a notable migrant in the United States, but its neotropical subspecies has been considered, as in the case of many tropical birds, to be a permanent resident. In the course of our Rancho Grande ob- servations throughout the years 1945, 1946 and 1948 we recorded 643 megalippe passing southward through the Pass. A much larger number of eresimus was recorded, 5,254 al- together, but these were concentrated within a comparatively brief space of time, more than 5,000 on July 21. Danaus plexippus megalippe (Hfibner). Field Name: Southern Monarch. Species Range : North and South America and the West Indies. Subspecies Range: Northern and western South America. Field Characters : Unmistakable to anyone who knows the northern monarch, D. p. plex- ippus. The chief subspecific distinction is the whiteness or the tawny appearance of the forewing spots. This is impossible to detect in flying insects. In every captured migrat- ing specimen and in those observed close at hand, at rest, the spots were megalippe- white. This also agrees with the geographical locale. The only possibility of confusion (and that hardly) is with the smaller and darker Danaus e. eresimus. Number: Total recorded, 643. Taken, 5. Sex: Both sexes seen and taken. Date : April 1 to September 1. Frequency : Mostly flying singly and fairly high, but a few closely associated groups, such as 28 on July 15, 55 with many eresimus on July 21, and a decided migration of 423 plus on September 1. Condition: Most of the butterflies seemed fresh and new, a few were worn. Non-migrant Record: Mr. Henry Fleming of the Department of Tropical Research staff reports megalippe as resident and breeding 58 Zoologica: New York Zoological Society [35: 3 in the low Valencia plain (circa 450 meters altitude) from Limon to Maracay, and to the north of the pass at sea-level around Ocu- mare. Record: 1945 — July 16 (16 through pass against high wind ; 12 low at km. 15) . 1946 — April 1(1); May 7 (1 male alighted) ; Sep- tember 1 (423 in fairly compact flock through pass) . 1948 — May 24 (5 seen) ; June 6 (male taken, 48737), 22 (17 seen twelve feet up), 30 (8 seen) ; July 5 (6 seen), 6 (15 seen), 8 (3 seen), 9 (10 very high), 10 (6 at pass; 1 taken km. 31), 15 (28 seen), 16 (12 sin- gly), 20 (21 singly), 21 (at least 55 with large numbers of eresimus; 2 females tak- en), 26 (female taken km. 16). Danaus eresimus eresimus (Cramer). Field Name : Southern Queen. Species Range : Central America to South Brazil. Subspecies Range : Venezuela, the Guianas and northern Brazil. Field Characters: Closely resembles Da- naus gilippus berenice, the queen of the United States. Smaller and darker but occa- sionally flying with D. plexippus megalippe. Number: Total recorded, 5,258. Taken, 16. Sex: Both sexes taken. Date: May 25 (1946); July 21 to 29 (1948). Frequency : Only one was taken in 1946, a season of infrequent observation. In 1948 a sudden heavy migration of at least 5,000 started on July 21. Several hundred passed in early morning, increasing to several thou- sand from 10 A.M. on. Many seemed tired and rested on foliage and in long grass. Dur- ing the succeeding eight days less numerous members of this movement were seen and taken. Condition : Although many were weary, no worn specimens were seen. Record: 1946— May 25 (2 taken, 46502), 28 (3 taken). 1948 — June 6 (1 taken) ; July 21 (5,000+ seen; 9 taken), 22 (210 seen), 23 (27 in grass), 26 (1 taken), 29 (4 seen). Migration of Lycorinae. A single member of this second subfamily of the Danaidae used Portachuelo Pass for purposes of north to south migration. In flight it was indistinguishable from sev- eral butterflies of other families, except on days of high winds, when flocks were forced down into the underbrush. The widely radi- ating scent hairs characterized it in the net or hand, rarely when resting on foliage. Ly corea ceres ceres (Cramer), form afergatis Doubleday and Hewitson. Field Name: Tuft-scented Ithomiid-like. Species Range: American neotropics and West Indies. Subspecies Range: Species range, except Cuba and Haiti. Field Characters: Considered the model for a number of mimics. Almost indistin- guishable in flight from two ithomiids ( Ti- Text-fig. 1. Map showing location of Rancho Grande, Portachuelo Pass and surrounding territory. thorea harmonia furia and Melinaea illis illis), a heliconid ( Heliconius numata) and a nymphalid ( Protogonius hippona trini- tata). The swift, bullet-like flight of the latter removes it from too close optical confusion. Once in the hand the almost invariable pro- trusion of three tufts of black scent hairs identifies Lycorea. Frequency : On June 6 a flurry of more than 90 large ithomiid-like butterflies drifted through the Pass, and three taken several minutes apart were all of this species. Four others showed their scent hairs while alighted on melanostomid blossoms, so it seems certain all were of the same species. Nine days later, on June 15, an identical occurrence on a smaller scale took place. A thirty-mile wind blew through the Pass and grounded a host of insects. Among these were 29 Lycoreas. On July 21 a dense flock of at least 32 passed through. Eight other flocks may well have been of this species. Eight were taken singly. Number: Total recorded, 185. Taken, 12. Date: April 30 to July 26. Condition: More than half of those taken or seen showed signs of wear and tear. Record: 1946 — May 5 (1 taken at Pass). The following were taken in 1946 at or near kilometer 21 : June 11 (female), 23 (female), 25 (male and female), 29 (female), 30 (fe- male). 1948 — April 30 (1 taken, 48458), 28 (1), 31 (1) ; June 6 (3 taken, 90 seen), 15 (2 taken, 27 seen), 26 (2) ; July 9 (2 taken, 10 seen), 10 (1 taken, 15 seen), 15 (1), 20 (1), 21 (1 taken, 32 in dense flock), 26 (1). 1950] Beebe: Migration of Butterflies at Rancho Grande, Venezuela 69 Migration of Ithomiidae. Thirty-one species and subspecies of Itho- miidae were recorded as migrants through Portachuelo Pass. Dr. Wm. T. Forbes lists forty -five itho- miids collected by Dr. P. J. Anduze in “Del Norte de Venezuela.” ( Boletin de las Socie- dad Venezolana de Ciencias Naturales, VI, 1949, pp. 308-317). In the relatively minute area represented by the narrow bottle-neck of the Pass we thus took seventy per cent, of the number of ithomiids so far recorded from northern Venezuela. In a report of the Ithomiinae of Rancho Grande by Dr. Richard M. Fox and Jean W. Fox ( Zoologica , 32, No. 20, 1947, pp. 173- 178) thirty species are listed. Of these ail but three have since been recorded by us as migrants, and as these three occur within two or three kilometers of the Pass they should doubtless take their place in the mi- grant list. I have, however, placed them at the end of the observed migrants. In addi- tion, four species occur as migrants which are not included in the paper by Dr. and Mrs. Fox. From the point of view of pattern and color the thirty-one species of migrant itho- miids are much more homogeneous than the preceding family of Papilionidae or the suc- ceeding family of Heliconidae. This makes for more difficult, or in a number of cases, impossible distinguishableness in flight. The species fall into two general types, those typically ithomiine in yellow, black and cinnamon-rufous patterns, and those which have more or less transparent wings, with black or rufous borders. The relative propor- tion of species is, opaque 14 and transparent 17. This superficial division is strongly re- flected and emphasized in habits. The opaque species fly in the open, at lower or higher altitudes, whereas the transparent forms are seldom found away from the shade of the jungle, usually threading their way through dense underbrush. This was especially ap- parent at the Pass, where a lateral shift of observation of only three or four meters, from open to heavy low growth, would mark the presence or absence of opaque-colored and of transparent patternless species. The reality of what I have called “typical- ly ithomiine” pattern and coloring is em- phasized when we consider such migrants as the danaid Lycorea ceres ceres, the nympha- lid Protogonius hippona trinitata, the helico- nid Heliconius numata numata, the pierid Charonias eurytale eurytale and even the arctiid moth Pericopis angulosa. Again and again these unrelated forms completely fooled us in the field, we mistaking one for the other, so remarkably similar are the pat- terns and colors of these models and mimics. As to relative comparison of species and individuals, of fifteen species, or about fifty per cent., one to ten individuals were re- corded. One hundred to foui'teen hundred specimens were observed in only five species, or twenty per cent Month after month there was noticed among passing migrants a large number of species comprising very few or solitary in- dividuals. Six examples of this type of oc- currence (so characteristic of many tropical organisms) are as follows, each record being made within a period of one to three hours ; and all six within forty days’ time. June 17, 12 specimens of 10 species July 8,12 it “ 12 “ July 15,35 it “ 13 “ July 16, 12 it “ 10 “ July 21, 50 a “ 14 “ July 26, 22 a “ 11 “ In 1945 and 1946 Mr. Fleming was interested in the composition and habits of dense flocks of ithomiids, in the vicinity of kilometers 20 and 21. The details of this in- teresting occurrence will soon be published. These flocks of butterflies appear earlier in the year than the first migrants and seem to be resident, but in a number of cases the comparative counts and times of occurrence link them so closely to contemporary passing migrants that I have recognized the ap- parent identity of origin by inclusion. Eutresis hyperla hyperia Doubleday and Hewitson. Field Name: Plain Hindwing. Species Range: Central America, Colom- bia, Peru, Ecuador and Venezuela. Subspecies Range: Venezuela, Colombia and northeast Peru. Field Characters: Large semi-clear fore- wing, plain, pale brown hindwing. These characters sometimes apparent in flight, easily identified in resting individuals. Number: Total recorded, 70. Taken, 24. Sex: Both sexes taken. Date: From April 24 to September 8. Frequency : Recorded on 16 days, usually singly. The largest number together 28, on September 28. Condition: Mostly fresh-appearing. Record: 1946— -April 8 (male, km. 21); July 1 (male, km. 21), 8 (male, km. 21; fe- male, Pass); August 13 (female, km. 21); September 8 (1, 461174 ; 28 seen driven down to foliage by high wind). 1948 — April 24 (1 taken, after being caught and dropped by blue-and-white swallow; 5 at Pass on fo- liage) ; May 1 (3) , 9 (2) ; June 6 (5) ,18 (1), 19 (1 taken, 3 seen) ; July 2 (1 seen), 3 (1, km. 27), 4 (1 taken, 481055; 2 seen), 10 (1 taken, 6 seen), 16 ( 1) , 21 (l);August2 (1). Tithorea h armonia furia Staud. Species Range: Trinidad, Guianas, Vene- zuela, Colombia, Ecuador, Bolivia and Ama- zons. Subspecies Range: Venezuela and Colom- bia. Field Characters : Large, typical ithomiid pattern, yellow and orange. Indistinguish- able on the wing from at least three others. Number: Total recorded, 9. Taken, 4. Sex : Both sexes taken. Date : From May 22 to June 16. 60 Zoologica : New York Zoological Society [35: 3 Frequency : A rare species. Only 9 record- ed in two years. Record: 1946 — May 22 (female taken). 1948 — May 24 (1 taken) ; June 15 (1 taken), 16 (1 caught from a group of 6, with one of which it was trying to mate. The group was all worn, alighted close together, so I am considering them of this species). O/yris crathis crathis Doubleday and Hewitson. Species Range : Central America and Ven- ezuela. Subspecies Range : Venezuela. Field Characters: Large, typical ithomiid pattern and color; hindwing plain red- brown with central, oblong, transparent window. Record: Only a single specimen taken. April 30, 1948. No. 48457. Taken on a sunny day in company of other species of ithomiids. Melinaea lilis tills (Doubleday and Hewitson). Species Range : Mexico to Colombia, Ven- ezula and Peru. Subspecies Range: Northern Venezuela and Colombia. Field Characters : Large, typical ithomiid pattern and color. Except for larger size (forewing length 43 mm. as compared with 38 mm.), indistinguishable in flight from Mechanitis doryssus veritabilis. Number: Total recorded, 55. Taken, 21. Sex : Both sexes taken. Date: April 13 to July 20. Frequency : Seen singly or in twos, ex- cept for two groups of 16 and 18. Usually high fliers, so we doubtless missed hundreds. Flight: Usually higher than most itho- miids. Condition: Mostly appeared freshly emerged. Record: 1946 — April 13 (female, km. 21) ; June 10 (female, km. 21) ; July 3 (female, 46725, km. 21), 8 (2 females, km. 21). 1948 —May 13 (1 taken, 48522), 15 (1), 20 (1 taken, 18 seen low and alighted), 23 (2), 26 (3) ; June 17 (1), 24 (1 taken with pierid and heliconid mimics) ; July 6 (3 taken, 16 seen close at hand in same group as those captured), 15 (1), 20 (1), 22 (1). Xanthocleis aedesia aedesia (Doubleday and Hewitson). Field Name: Black Hook Ithomiid. Species Range : Central America and northwestern South America. Subspecies Range: Northwestern South America. Field Characters: Large (forewing 38-40 mm.), with large black-bordered pale fore- wing spots, and black hook projecting down- ward into mid-hindwing orange. Hook easily identified in flight with naked eye or No. 7 binoculars. Number: Total recorded, 434. Taken, 104. Sex: Both sexes taken. Date: April 8 to August 4. Frequency: Decidedly gregarious, flocks recorded of 15, 19 and 41, up to 135. Record: 1946 — April 8 (male, km. 21) ; May 7 (male, km. 21), 21 (female), 25 (4 taken, 46502), 28 (9 taken) ; June 23 (3 fe- males, km. 21) ; July 3 (1), 15 (3 females) ; August 4 (male, km. 21). 1948 — April 30 (4 taken, 48455) ; May 4 (87 seen A.M., 48 P.M., slow, low flying), 6 (1), 9 (1), 21 (1), 23 (4), 24 (1), 29 (3) ; June 6 (15 taken), 7 (1), 10 (2 taken, 19 seen), 15 (2), 27 (2), 28 (6 taken, 68 seen) ; July 5 (1 taken, 12 seen) , 6 (3 taken, 32 seen) ,8 (1) , 9 (41 seen high), 10 (1), 13 (1 taken, 6 seen), 15 (2 taken), 19 (2 mating), 21 (8), 23 (1), 26 (1). Mechanitis doryssus veritabilis Butler. Species Range : Central America and northern South America. Subspecies Range: Venezuela and Trini- dad. Field Characters: Typical ithomiid pat- tern and color. Midway in size of forewing length (38 mm.) between the larger (44 mm.) Melinaea lilis lilis, and the smaller (28 mm.) Hypothyris euclea fenestella. Other- wise indistinguishable in flight. Number: Total recorded, 133. Taken, 98. Sex: Both sexes taken. Date: Api’il 29 to July 29. Frequency: Mostly singly, otherwise in small groups of 4, 6, 7 and 18. Condition: Very few worn specimens seen. Record: 1946 — May 28 (7 taken), 29 (5) ; June 23 (2 males, km. 21), 29 (male, km. 21), 30 (2 males) ; July 3 (1 at Pass, 46725; 3 females, km. 21), 7 (female, km. 21), 8 (female), 25 (female). 1948 — April 29 (1 taken); May 6 (1, 48497), 8 (3 taken, 3 seen), 15 (1 taken, 4 seen), 23 (4), 24 (7), 26 (16), 31 (5) ; June 6 (6), 7 (1), 8 (1), 15 (1), 17 (2), 18 (2), 19 (1), 22 (1), 23 (1),24 (4), 28 (4 taken from swarm of 18) ; July 3 (1), 4 (3), 5 (1 taken, 6 seen), 8(1), 9 (1) , 10 (1 at Pass, 1 taken, 4 seen, km. 31) , 13 (1), 15 (4), 17 (1), 21 (2), 23 (1), 24 (1 at Pass, 1 at km. 15), 26 (2), 29 (1). Hyalyris cana cana (Haensch). Field Name : Black-bordered Clearwing. Species Range: Venezuela and Colombia. Subspecies Range : Eastern Colombia and northern Venezuela. Field Characters: This rare ithomiid is very distinct and easy to distinguish both on the wing and alighted. The wide, white- dotted border surrounding all four wings, enclosing transparent central areas, is un- mistakable. Indistinguishable from the con- generic H. coeno. Number: Total recorded, 7. Taken, 7. Sex: Both sexes taken. Date: July 14 to 21. Frequency: Very remarkably, in 1948, seven specimens were taken, one a day, July 18 being the only exception to the regular sequence, no insect being captured on this date. They were low-flying, close to the top of the low growth like the other clearwings. 1950] Beebe: Migration of Butterflies at Rancho Grande, Venezuela 61 In addition to the 1948 migrants at the Pass, Mr. Fleming took 4 males and a female in the dry season, at kilometer 21. These seem- ed to be residents, judging by the dates of capture, and were associated with a large swarm of ithomiids of other species. They were taken on February 27 and March 4. Record: 1948 — July 14 (1 taken), 15 (1, 481174), 16 (1 taken), 17 (1), 19 (1), 20 (1 taken at ginger blossoms) , 21 (1). Hypothyris euclea fenestella (Hewitson). Field Name : Dwarf Typical Ithomiid. Species Range: Central America, Colom- bia, Venezuela, Ecuador and Peru. Subspecies Range: Venezuela and Trini- dad. Field Characters: Typical ithomiid yel- low, orange and black. Differs in its small size from other similarly colored species. From the equally numerous and equally sized Ithomia iphianassa it differs in lacking the transparent large spots and oblong windows. Number: Total recorded, 488. Taken, 123. Sex : Both sexes taken. Date: April 29 to August 2. Frequency : One of the most abundant spe- cies and decidedly gregarious, counts of 54 to 266 being made, and of course many thou- sands of butterflies missed. On a number of days this was the dominant species. Record: In 1946 Mr. Fleming collected 35 males and 56 females in the vicinity of kilo- meters 20 and 21. 1948 — April 29 (1,48449) ; May 21 (2), 23 (1), 25 (14 taken, 54 seen), 26 (2), 29 (1) , 31 (1) ; June 6 (14 taken), 15 (1), 16 (1 taken, 12 seen), 17 (1), 21 (3), 22 (1), 24 (2), 27 (2); July 3 (2), 4 (6 taken, 28 seen), 5 (5 at Pass, 3 taken km. 21), 8 (13 taken, 266 seen), 10 (3 taken, 3 seen), 13 (1 taken, 2 seen), 14 (4), 15 (9), 16 (1), 20 (3), 21 (16), 23 (1), 26 (3), 29 (5) ; August 2 (2 taken). Ithomia agnosia agnosia Hewitson. Species Range: Venezuela, Colombia and northern Peru. Subspecies Range: Venezuela. Field Characters : This clearwing was quite indistinguishable from seven or eight other species. Number: Total recorded, 23. Taken, 23. Sex: Both sexes taken. Date : April 30 to September 3. Frequency: Solitary flyers and low, just clearing the undergrowth. Record : Only 4 migrants were taken at the Pass in 1948, whereas eighteen were taken at kilometers 20 and 21 in 1946. Dates cor- responding as to months, I have included the latter as probable migrants. 1946— April 30 (1) ; May 15 (1), 24 (1), 26 (1), 28 (1) ; June 4 (2), 7 (1), 14 (1), 18 (1), 23 (1), 25 (1), 26 (2) ; July 2 (3), 17 (1); September 3 (1). 1948— May 26 (1) ; July 6 (1), 26 (2 taken). Ithomia iphianassa iphianassa Doubleday and Hewitson. Field Name: Common Small Ithomiid. Species Range: Venezuela. Subspecies Range: Venezuela. Field Characters : The small size combined with transparent patches and bright colors make this very abundant species easy to identify in flight. Number: Total recorded, 1,403. Taken, 205. Many thousands must have passed un- recorded. Sex : Both sexes taken. Date: April 27 to September 8. Frequency : This species was so abundant that on some days a continuous stream was passing. No condensed flocks but the dom- inant butterfly on many days. The low counts at the Pass in 1946 due to infrequent observation. Flight: Moderately low and never very rapid. Record: 1946— May 28 (1), 29 (1) ; July 3 (2), 17 (1); September 7 (4), 8 (3). In addition 109 males and 37 females were taken by Mr. Fleming at kilometer 20 and lower in 1946. 1948 — -Records are so numer- ous that I condense as follows: April 8 (1 taken, 8 seen), 28 (7 seen); May (on 17 days, 33 taken, 98 seen) ; June (on 30 days, 97 taken, 175 seen) ; July (on 28 days, 75 taken, 925 seen). Mileria cymothoe (Hewitson). Field Name: Small Clearwing. Species Range: Venezuela and Colombia. Field Characters: Small and almost com- pletely transparent, indistinguishable in life from 5 or 6 other species. Number: Total recorded, 12. Taken, 12. Sex: Both sexes taken. Date: May 5 to July 21. Record: 1948 — May 5 (1 male), 6 (1 male), 23 (1 male), 26 (5 females), 29 (1 male) ; June 6 (male), 28 (female) ; July 21 (female) . Oleria victorine graziella (Oberthiir). Species Range: Central America, Colom- bia, Venezuela, Ecuador, Bolivia and upper Amazons. Subspecies Range: Venezuela and Colom- bia. Field Characters : A small clearwing with strong black borders, indistinguishable in life from six other species. Number: Total recorded, 7. Taken, 7. Sex: Both sexes taken. Date: April 8 to July 10. Record: 1946— April 8 (2), 13 (1), 16 (2) ; July 10 (1). 1948— May 29 (1). Oleria makrena makrena (Hewitson). Species Range: Colombia, Venezuela and Ecuador. Subspecies Range : Colombia and northern Venezuela. Field Characters : A clearwing with wings tinted with whitish. Only once could I be sure of it before capture; in the sunlight resting on a leaf. Number: Total migrants, 7. Taken, 7. Sex: Both sexes taken. 62 Zoologica: New York Zoological Society [35: 3 Date: June 6 to July 21. Record: At lower levels in 1946 Mr. Flem- ing took 51 males and 21 females. These may very likely have been early migrants but we had no definite proof. 1948 — June 6 (1), 15 (1), 16 (1); July 5 (1), 6 (2 seen alighted), 8 (1), 21 (1). Oleria phemonoe phemonoe (Doubleday and Hewitson). Species Range: Colombia, Venezuela and Brazil. Subspecies Range: Venezuela. Field Characters : A rare clearwing indis- tinguishable in life from related species. Number: Total recorded, 11. Taken, 11. Sex: Both sexes taken. Date : May 29 to July 23. Record: 1946 — April 8 (2, km. 21); May 29 (2 at Pass) ; June 20 (female, km. 27) ; July 5 (male, km. 27), 28 (female, km. 27). 1948— May 27 (1 at Pass) ; July 8 (1), 23 (2). Aeria eurimedia agna Godman and Salvin. Field Name: Six-lemon-striped Ithomiid. Species Range: Central America, Colom- bia, Ecuador, Venezuela, Guianas and Ama- zonia. Subspecies Range: Central America, Col- ombia, Venezuela and Trinidad. Field Characters : The six elongate lemon spots surrounded by spots make this small species easy to identify even in flight. Number: Total recorded, 56. Taken, 29. Sex: Both sexes taken. Date: April 7 to September 7. Record: 1946 — Twenty specimens taken at or near kilometer 20, undoubtedly mi- grants; dates as follows: April 7, 8, 13, 30, May 28, June 10, 16, 30, July 3, 17, 25, 28, August 13, September 7. 1948 — May 25 (2 taken, 16 seen), 26 (1) ; June 6 (2), 9 (1), 22 (1 taken, 11 seen) ; July 5 (1), 8 (1). Bircenna euchyehma Felder. Species Range: Colombia and Venezuela. Field Characters : A medium sized, trans- lucent, dusky, red-brown species, indistin- guishable in life from Dircenna jemina. Number: A single specimen, 481392, taken July 26, 1948, migrating through Pass in company with ithomiids of other species. Dircenna jemina (Geyer). Species Range: Venezuela and Colombia. Field Characters: A medium, smoky- brown, translucent species recognizable in life only when it has alighted close by. Number: Total recorded, 52. Taken, 18. Sex: Both sexes taken. Date: March 15 to July 21. Frequency : Taken and seen singly, except on March 15, April 27 and May 13, when 12, 16 and 6 individuals were seen resting close together just beyond reach. Record: 1946— May 3 (1), 13 (1); June 26 (1), 29 (1), 30 (1) ; 1948— March 15 (1 taken, 12 seen), April 27 (2 taken, 16 seen), 29 (1): '(ay 5 (1), 6 (1), 13 (1 taken, 6 seen) ; June 17 (1) ; July 8 (1), 13 (1), 16 (2), 21 (1). Ceratinia tutia tutia (Hewitson). Species Range: Central America, Colom- bia and Venezuela. Subspecies Range: Venezuela and Colom- bia. Field Characters: A small, smoky-brown, translucent species not distinguishable from Hypoleria and Godyris. Number: Total recorded, 237. Taken, 19. Sex: Both sexes taken. Date: May 26 to August 7. Frequency: Seen and taken singly except on August 8, 1946, when a cloud of this spe- cies came up to the Pass, after heavy rain, and alighted near by. I took 3 and counted 218. Record: 1946 — August 8 (3 taken, 218 seen), 12 (1). (Fourteen males and 2 fe- males taken at kilometer 20 or lower, not counted as migrants). 1948 — May 26 (2); June 6 (1), 7 (1), 9 (2), 12 (1), 15 (1), 16 (1), 24 (1), 28 (2) ; July 8 (1), 13 (1), 21 (2). Episcada hymenaea (Prittwitz). Field Name: Small Brown-veined Clear- wing. Species Range: Widely distributed in South America. Field Characters: A very small, brown- veined, patternless clearwing, not distin- guishable in life from Mileria or Pteronymia aletta. Number: Total recorded, 44. Taken, 44. Sex: Both sexes taken. Date: May 21 to July 26. Breeding Note: Female No. 48592, taken May 25, contained many fully formed eggs, with surface sculpture, including vertical grooves and finer markings apparent. Record: 1946 — May 28 (1); June 26 (male, km. 20), 29 (2 males, km. 20). 1948 — May 21 (3), 23 (3), 24 (3), 25 (10), 26 (7), 29 (3) ; June 6 (4), 17 (1), 18 (1) ; July 10 (1), 15 (1), 21 (2), 26 (1). Episcada sylpha Haensch. Species Range: Venezuela. Field Characters: Indistinguishable in life from Episcada hymenaea, but very rare. Ntimber: Two specimens only, 1946, June 29, male; and 1948, May 30, male. Hypoleria ocalea (Doubleday and Hewitson). Species Range: Colombia, Venezuela and Trinidad. Field Characters : Small, half smoky, half transparent; not distinguishable in life from other similar species. Number: Total recorded, 4. Taken, 4. Date: May 26. Record: 1948 — May 26 (4 taken). Pteronymia adina Hewitson. Species Range: Venezuela. Field Characters: A small clearwing, in- distinguishable from others in life. 1950] Beebe: Migration of Butterflies at Rancho Grande, Venezuela 63 Number: Total recorded, 4. Taken, 4. Sexes : Both sexes taken. Date: May 21 to July 20. Record: (2 males and 11 females taken in March by Mr. Fleming at kilometer 20 and lower. Owing to early date not counted as migrants). 1948 — May 21 (1), 25 (1) ; June 6 (1) ; July 20 (1). Pteronymia aletta (Hewitson). Species Range: Venezuela. Field Characters : A very small, pale smoky clearwing, hardly to be distinguished in life from several others. Number: Total recorded, 7, Taken, 7. Sex: Both sexes taken. Date: May 24 to July 15. Record: 1946 — June 7 (1), 23 (1 taken, km. 20)i 1948— May 24 (1), 25 (2), 29 (1) ; July 15 (1). Pteronymia asopo (C. and R. Felder). Species Range: Northern Colombia and Venezuela. Field Characters : Small clearwing with large white splash. Not identifiable in life. Number: Total recorded, 11. Taken, 11. Sex: Both sexes taken. Date: April 20 to July 17. Record: 1946 — April 20 (2 taken) ; June 29 (2 taken). 1948 — June 17 (3 taken), 18 (1), 22 (1), 24 (1) ; July 10 (1 taken, km. 20). Pteronymia beebei Fox and Fox. Species Range: Northwestern Venezuela. Field Characters: Medium clearwing. Not distinguishable in life from several other species. Number: Total recorded, 14. Taken, 14. (All taken at Pass). Sex: Both sexes taken. Date: May 25 to August 5. Record: 1946— August 5 (1 taken at Pass). Thirty-seven males and 7 females, including type of species, taken by Mr. Fleming at kilometers 18 to 21. 1948 — May 25 (1), 26 (3), 29 (1); June 6 (1); July 8 (1) , 15 (2), 16 (1) , 21 (1) , 26 (1) ; August 2 (1). Pteronymia nubivaga Fox and Fox. Species Range: Northwestern Venezuela. Field Characters: Small clearwing, indis- tinguishable in life from several other species. Number: Total recorded, 14. Taken, 14. Sex: Males only taken. Date: April 14 to July 8. Record: 1946 — -April 14 to July 8 (12 males, taken by Mr. Fleming around km. 20). 1948 — May 9 (2 taken at Pass). Pteronymia veia (Hewitson). Field Name: Yellow-spotted Clearwing. Species Range: Venezuela. Field Characters: Small clearwing with bright yellow spot. Number: Total recorded, 44. Taken, 44. Sex: Both sexes taken. Date : May 6 to August 2. Frequency : Usually single, but now and then several in association. As with all clear- wings they fly low, slowly, just above or more usually threading through underbrush. Record: 1946 — May 28 (1 taken at Pass). 41 males and 11 females taken by Mr. Flem- ing around kilometer 20 and lower, in 1946. Not counted as migrants, in spite of circum- stantial evidence. Extreme dates March 8 to July 15. 1948— May 6 (1), 9 (1), 21 (1), 24 (1), 25 (1), 26 (1), 29 (1), 31 (1) ; June 6 (5), 15 (1), 17 (2), 18 (1), 22 (3), 27 (2) ; July 5 (1), 7 (1), 8 (2), 13 (1), 14 (1), 20 (1), 21 (3), 23 (1), 26 (3), 28 (1) ; August 2 (1) . Godyris kedema kedema (Hewitson). Species Range: Venezuela and Colombia. Subspecies Range : Venezuela. Field Characters: Medium, pale smoky, half transparent. Often flies so slowly and low that identification is possible. Number: Total recorded, 21. Taken, 21. Sex : Both sexes taken. Date : May 21 to August 2. Record: 1946 — June 7 (1 taken at Pass). Seventeen males and 14 females collected by Mr. Fleming at low altitudes to the south of Pass, around kilometer 20, in 1946. Extreme dates were March 13 and August 4. 1948 — May 21 (1), 25 (1), 29 (2), 31 (1) ; June 6 (1), 9 (1), 28 (1) ; July 13 (1), 15 (5), 17 (1), 21 (2), 26 (3) ; August 2 (1). Pseudocode fimna (Hewitson). Species Range: Colombia and Venezuela. Field Characters: Small clearwing, with much white on forewing. Not distinguish- able in life from several others. Number: Total recorded, 7. Taken 7. Sex: Both sexes taken: Date: May 6 to July 15: Record: 1946 — None taken at Pass. At kilometer 20, Mr. Fleming collected 16 males and 2 females, the extreme dates being April 8 and July 15. 1948— May 6 (1), 24 (1), 25 (2), 29 (1) ; June 24 (1) ; July 15 (1). Hymenitis andromica andromica (Hewitson). Species Range: From Guatemala south through Colombia, Venezuela, Ecuador, Peru and Bolivia. Subspecies Range: Venezuela and Colom- bia. Field Characters: Medium clearwing with considerable white on forewing. Indistin- guishable from others. Number: Total recorded 21. Taken 21. Sex: Both sexes taken: Date : May 24 to August 2. Record: 1946 — July 3 (1 taken); August 1 (1 taken at Pass from flock of 19, probably the same species). At kilometer 20 and low- er, in 1946, Mr. Fleming collected 66 males and 43 females, doubtless migrants, but not counted as such. 1948 — May 24 (2), 25 (4), 29 (1) ; June 6 (2), 9 (1), 22 (1), 28 (1) ; July 8 (1), 13 (1), 14 (1), 15 (1), 16 (1), 24 (1), 26 (2) ; August 2 (1). 64 Zoologica: New York Zoological Society [35: 3 Hymenitis dercetis (Doubleday and Hewitson). Species Range : Venezuela and Colombia. Field Characters : Medium clearwing, bright yellow splash. Indistinguishable from several others. Number : Total migrants reported, 31. Taken, 31. Sex: Both sexes taken. Date : April 30 to September 5. Record: 1946 — May 28 (1 taken at Pass), September 5 (1 taken). At kilometer 27 and lower, Mr. Fleming collected 72 males and 12 females. 1948 — April 30 (1); May 1 (2), 9 (2), 24 (2), 25 (1), 26 (2), 29 (1), 31 (1) ; June 6 (2), 11 (1), 15 (2), 17 (1), 24 (1), 27 (1), 28 (1); July 3 (1), 10 (1 taken, many probably of this species, seen), 15 (3), 16 (1), 17 (1), 21 (1). Probable Migrants But Not Taken at Pass. Athesis clearista clearista (Doubleday and Hewitson). Field Name: Semi-transparent Clear- wing. Species Range: Colombia and Venezuela. Subspecies Range : Venezuela. Field Characters: A medium, orange- veined clearwing, very similar to several other species. Record: Two male specimens only taken, and neither actually at the Pass. Two males, 46106, taken February, at Red Bridge, less than one kilometer from the Pass, but in the dry season. Hyalyris coeno coeno (Doubleday and Hewitson). Species Range: Nicaragua, Colombia, Peru and Venezuela. Subspecies Range: Venezuela. Field Characters: Indistinguishable in life from Hyalyris cana. Record: Not taken actually at Pass. In April, 1946, 3 males, 1 female, taken at kilometer 27, north of Pass, feeding on flow- ers. CalUthomia agrippina alpho (C. and R. Felder). Species Range: Panama, Colombia and Venezuela. Subspecies Range: Eastern Colombia and Venezuela. Field Characters : A smoky clearwing not distinguishable in life from several other species. Record: No specimens taken at Pass and only one near by but lower. A female, at kilometer 21, June 23, 1946. Migration of Acraeidae. The members of this small family seem always to appear at the Pass on days of in- tensive migration, and in life they are so similar to the smaller ithomiids, as well as to many heliconids, that they were caught and enveloped without special attention. Later, when their true nature was discov- ered in the laboratory, it was too late to re- call flight characters or to be certain of how many others had passed unobserved. Acthote anteas (Doubleday and Hewitson), forms anteas (Doubl. and Hew.) and straminosa Jordan. Field Name: Striated-hindwing Ithomiid- mimic. Species Range: Guatemala to Colombia, Venezuela, Trinidad and Tobago. Field Characters : Inconspicuous in flight, closely resembling the smaller opaque itho- miids, and even more exactly the heliconid Heliconius vibilia vialis. Number: Total recorded, 32. Taken, 32. Sex: Both sexes taken. Date: April 29 to July 26. Record: 1946— July 3 (1 taken). 1948 — April 29 (1 taken), 30 (2) ; May 4 (1), 21 (2, 48546), 23 (3); June 6 (1), 27 (1, 48966) ; July 3 (1, 481541), 6 (1), 13 (1), 15 (7), 16 (1), 21 (2), 26 (2). Actlnote hylonome Doubleday. Field Name: Small White-banded Black. Species Range: Venezuela and Colombia. Field Characters : Small, black, with wide oblique band across each forewing. So re- sembles other small butterflies and even day-flying moths, that only once were in- dividuals noticed, other than those cap- tured. Number: Total recorded, 20. Taken, 12. Date: April 27 to July 16. Record: 1948— April 27 (2, 48378, 48398; 8 others seen) ; May 9 (1), 23 (2, 48573), 25 (1), 26 (2); June 6 (1); July 3 (1), 8 (1), 16 (1). Migration of Heliconidae. Of all groups of butterflies the heliconids are nearly the most difficult in which to designate typical patterns or colors. In this respect the eighteen species and subspecies of heliconid migrants through Portachuelo Pass seem to have evolved in an effort to resemble species in other groups, to offer themselves as models or mimics, or at least to have developed intensely independent and individual forms. This, combined with the habit of low flight and slow flapping with much leisurely glid- ing, makes sight identification in most spe- cies a very easy matter. A key based on pattern and color was of the greatest help in day-by-day recording of numbers, and for what it may be worth to others I reproduce it here. It was of course intended only for my own use with the mi- grants at Rancho Grande. Heliconia Sight Identification Pattern and Color Key. Fritillary-like (northern genera; Agrau- lis, Argynnis, etc.) Silvery Fritillary — Agraulis vanillae Dark Silvery Fritillary — Dione juno 1950] Beebe: Migration of Butterflies at Rancho Grande, Venezuela 65 Large Orange Fritillary — Dryas iulia Dwarf Orange Fritillary— Heliconius aliphera Yellow-banded, Red-hindwing Solid-red-hindwing — H. procula H. clysonimus Striated-red-hindwing — H. doris (form, transiens ) Yellow-banded, Blue-hindwing Large, Blue-hindwing- — H. doris (form, doris) Small, Blue-hindwing— H. sara Light-striped Black 4-white-striped Black— H. antiochus antiochus 4-yellow-striped Black — H. antiochus aranea Zebra-striped Black — H. charithonius Red-banded Black— H. melpomene Acraeid-mimic — H. vibilia vialis Ithomiid-mimic — H. isabella dynastes H. anderida holcopliorus H. anderida semiphorus H. eucoma metalilis Green-spotted — Philaethria dido Agraulis vanillae vanillae (Linnaeus). Field Name : Southern Fritillary. Species Range : New York to Chile and Ar- gentina. Subspecies Range : Panama, northern South America, Amazonia and Lesser An- tilles. Field Characters : The same species as our northern silver spangled fritillary, but sev- eral minor charactei’s provide it with a sub- specific name, characters impossible to tell on the wing. All specimens taken were char- acteristic, however. The absence of the dark hue of the underwings easily distinguishes it from Dione juno. Number : Total recorded, 188. Taken, 40. Sex: Both sexes taken. Date : May 9 to July 29. Frequency : Usually singly or in twos. On June 22 and 24 great numbers were observed high in air, hundreds of which must have escaped our counts. Condition: A minority only were observed to be torn and faded. Record: 1946— May 28 (1), 25 (1). 1948 — May 9 (2 seen), 13 (2), 15 (1 taken, 4 seen), 23 (2); June 6 (13), 17 (2), 22 (many high, 19 counted), 24 (many high, 64 counted) ; July 2 (2), 5 (1), 6 (11 seen), 8 (2 taken, 17 seen), 10 (1), 13 (1, km. 15), 14 (1), 15 (3), 16 (4), 17 (2, km. 30; 15 seen), 19 (2), 21 (1), 29 (14 seen). Dione juno (Cramer). Field Name: Dark Silvery Fritillary. Species Range: Central America to Brazil. Field Characters : Distinguished from Dryas iulia above by broader, darker wings and smaller size, and from Agraulis vanillae below by dark setting of silver spangles. Number: Total recorded, 194. Taken, 19. Sex: Both sexes taken. Date: May 25 to September 9. Frequency: Decidedly gregarious, but in moderately sized flocks. Condition : Many worn and tattered speci- mens, many perfectly fresh. Record: 1946 — -September 9 (1 taken). 1948 — May 25 (1 taken, 8 seen) ; June 17 (2 taken, i4 seen) ; July 1 (1 taken, 12 seen), 5 (1 taken, 10 seen), 9 (48 seen high), 10 (1 taken, 19 seen), 11 (2 seen, km. 31), 14 (1 taken, 3 seen), 15 (1, km. 15), 16 (1), 17 (8 seen), 19 (2 taken, 9 seen), 22 (1 taken, km. 35, 28 seen), 29 (21 seen). Dryas julia (Fab.). Field Name: Large Orange Fritillary. Species Range: Trinidad, Guiana and Brazil. Not before recorded from Venezuela. Field Ch aracters : A large, narrow-winged orange fritillary, to be confused only, and that hardly, with the silver-spangled Dione juno. Number: Total recorded, 969. Taken, 6. Sex : Both sexes taken. Date: April 15 to September 7. Frequency : On July 18, September 1 and 7, 1946, seven-eighths of all observed speci- mens of this species passed through the Pass. In the following year only flocks of moderate size were seen. Condition: In many of the flocks the fresh and the worn and faded individuals seemed about equal in number. Record: 1946 — May 28 (1); July 18 (450 + seen), September 1 (321 counted), 7 (95 + counted). 1948 — April 15 (1), 30 ( 4 seen) ; May 4 (3 seen) ; June 28 (1 taken, 22 seen) ; July 3 (1 taken, 28 seen), 10 (2 seen, km. 20), 13 (1 taken, km. 15), 17 (3 seen), 21 (4 seen), 24 (2, km. 31), 26 (42 seen) , 29 (16 seen) . Heliconius aliphera aliphera (Godart). Field Name: Dwarf Orange Fritillary. Species Range: Mexico to Paraguay. Subspecies Range: Colombia and Peru to Trinidad and Paraguay. Not before reported from Venezuela. Field Characters : A diminutive copy of the preceding species, Dryas julia. Number: Total recorded, 2,313. Taken, 35. Sex: Both sexes taken. Date : May 28 to September 9. Frequency: As with Dryas julia, great numbers were seen in early September, more than half of all observed on September 1, 5 and 7. Hundreds, if not thousands, must have passed unobserved. The majority of this host were faded and torn. Several taken gave out a strong scent of witch hazel, much like that of Heliconius melpomene. Record: 1946 — -May 28 (3 taken) ; July 3 (1, 46725), 8 (1, 46752), 30 (200 seen); August 12 (411 seen) ; September 1 (285 in one hour, all worn), 5 (1 taken, 461134; 463 seen, worn), 7 (700 + seen), 9 (1, 461162). 1948— July 14 (1, 481168), 15 (3 taken, 63 seen), 16 (1), 17 (6 seen), 20 (2), 21 (8 taken, 150 + seen), 24 (1 taken, strong witch hazel scent), 26 (1), 31 (1). 66 Zoological New York Zoological Society [35: 3 Heficoniiis pr ocula (Doubleday and Westwood). Field Name: Red-hindwing Yellow-band- ed. Species Range : Venezuela and Colombia. Field Characters: With the yellow fore- wing spots and the solid red hindwing band this species is not distinguishable in flight from H. clysonimus clysonimus. Number: Total recorded, 867. Taken, 22. Sex: Both sexes taken. Date: April 29 to September 8. Frequency: A third species in which large numbers appeared in late July and again in early September. At other times they usually were seen singly. Condition: Almost all of those seen were in fresh, unfaded condition. Record: 1946 — July 18 (600 + seen), Sep- tember 8 (3 taken, 461176; 213 seen). 1948 — April 29 (1) ; May 26 (1) ; June 6 (6) ; July 6 (1 taken, 6 seen), 7 (1, 481021), 14 (27), 15 (2), 17 (1, km. 30), 18 (1), 27 (2 taken, 26 seen) . Heliconius vibllia vlalls (Stichel). Field Name : Acraeid Mimic. Species Range: Guatemala to South Brazil and Peru. Subspecies Range: Guatemala to Colom- bia. Field Characters : Hindwing, black-veined striations on orange; with only forewing double (instead of single) band of yellow spots to distinguish it from acraeian spe- cies. Number: Total recorded, 43. Taken, 8. Sex: Both sexes taken. Date : April 29 to September 8. Condition: All specimens fairly fresh. Record: 1946 — September 8 (1, 461143a; 13 seen). 1948 — April 29 (1, 481442) ; May 23 (1 taken, 48545; 3 seen), 26 (1), 31 (1) ; June 28 (1 taken from flock of 10) ; July 3 (1 taken, 3 seen), 6 (1 taken, 7 seen). Heliconius isabella dynastes Felder. Species Range: Panama to central Brazil and Bolivia. Subspecies Range: Venezuela. Field Characters: So exact a mimic of typical ithomiid pattern that it is indistin- guishable in life from several species, espe- cially Mechanitis doryssus veritabilis, as well as the heliconid H. anderida holco- phorus. Number: Total recorded, 7. Taken, 7. Date: June 17 to July 26. Condition: All were in good condition. Record: 1946 — July 3 (1, 46725, taken at Pass; 2, 461023, taken at km. 20). 1948— June 17 (1 taken at Pass). July 16 (2 tak- en), 26 (1 taken). Heliconius anfioc has antiochus (Linnaeus) , form alba Riffarth. Field Name: Four-white-striped Black. Species Range: Colombia to south Brazil and Peru, Subspecies Range: Venezuela to south Brazil and Peru. Field Characters: This black species with its four, forewing, oblique white stripes is unmistakable, except for the rare aranea subspecies in which the stripes are yellow. Number: Total recorded, 228. Taken, 39. Sex: Both sexes taken. Date: April 29 to July 27. Frequency: Decidedly gregarious, occa- sionally singly. Condition: Mostly fresh-appearing. Record: 1946 — May 28 (4 taken, 48 seen) . 1948— April 29 (1, 48445), 30 (1) ; May 9 (2 seen), 24 (1, 48583), 28 (1); June 6 (5), 10 (2 taken, 27 seen), 15 (2), 16 (3 taken, 14 seen), 24 (1 taken, 48897; 12 seen), 28 (2 taken, 66 seen); July 5 (3 seen), 9 (3, km. 31), 13 (1 taken, km. 15), 14 (2 taken, 4 seen), 15 (1), 17 (6 seen, km. 30), 19 (2 at Pass, 3 km. 31), 22 (7 seen, km. 35) , 23 (pair taken, km. 15) , male with strong odor), 27 (1 at Pass, 1 km. 16). Heliconius antiochus aranea (Fab.). Field Name : Four-yellow-striped Black. Species Range: Colombia to south Brazil and Peru. Subspecies Range: Colombia, Venezuela, Ecuador, Guiana and Amazonia. Field Characters : This rare species is not to be distinguished from H. antiochus antio- chus on the wing. Number: Total recorded, 5. Taken, 2. Sex: Both sexes taken. Date: July 8 to 23. Record: 1948 — July 8 (1 taken, 481116), 10 (3 alighted, seen close), 23 (1, 481543). Heliconius anderida holcophorus Staudinger. Species Range: Honduras to Colombia and Venezuela. Subspecies Range: Colombia and Vene- zuela. Field Characters: A mimic with typical ithomiid pattern and coloring. Indistin- guishable from several species, especially close to Tithorea harmonia furia. Number: A single specimen, a male, taken, May 21, 1948, at Pass. In somewhat worn condition. Heliconius anderida semlphorus Staudinger. Species Range : Honduras to Colombia and Venezuela. Subspecies Range: Colombia and Vene- zuela. Field Characters: A large size ithomiid mimic, but with basal half of all four wings rich orange. Indistinguishable in life but closest to Olyras crathis crathis. Number: Only a single specimen, a male, taken, July 15, 1948, at Pass. Heliconius ch arithonius (Linnaeus). Field Name: Zebra-striped Black. Species Range: North Carolina to Brazil and Peru. Field Characters: Unmistakable in its 1950] Beebe: Migration of Butterflies at Rancho Grande, Venezuela 67 pattern of yellow and black bands across all four wings. Number: Total recorded, 6. Taken, 3. Sex : Both sexes taken. Date: May 24 to July 21. Record: 1948 — May 24 (1, 48584) ; June 6 (3 seen) ; July 21 (2). Heliconius dysonimus dysonimus Latr. Field Name: Solid Red-hindwing. Species Range: Costa Rica to Venezuela and Ecuador. Subspecies Range: Venezuela, Colombia and Ecuador. Field Characters: Indistinguishable from H. procula, otherwise quite distinct. Number: Total recorded, 90. Taken, 17. Sex: Both sexes taken. Date: May 5 to September 8. Frequency: Usually singly. Only twice seen in any number. Record: 1946 — May 5 (1 taken) ; Septem- ber 8 (1, 481177; 39 seen) ; 1948— May 11 (2 seen), 21 (1), 23 (2) ; June 6 (2 taken, 12 seen), 10 (15 seen), 18 (1 seen); July 2 (1), 3 (4 seen), 13 (1), 15 (1, km. 15), 17 (1), 21 (1), 23 (2 at Pass, 2 on trail), 25 (1). Heliconius doris (Linnaeus), form transiens Staudinger. Field Name: Striated-red-hindwing. Species Range: Colombia and Venezuela to Brazil and Bolivia. Form Range: Mexico to Colombia and Venezuela. Field Characters: When flying slowly close at hand or alighted, easy to distinguish from H. procula and H. chjsonimus. Number: Total recorded, 77. Taken, 8. Sex: Both sexes taken. Date: April 29 to September 8. Record: 1946 — September 8 (1 taken, 461175; 10 seen). 1948— April 29 (1, 48453) ; J.une 22 (6 seen) ; July 8 (4 taken, 22 seen), 9 (4 seen), 19 (1 taken, 27 seen), 21 (1, 481540). form doris (Linnaeus). Field Name: Large Blue-hindwing. Form Range: Mexico, Colombia and Vene- zuela to Bolivia and Brazil. Field Characters: Identical with form transiens except for color of hindwing. Of this remarkable mutation-like form only three individuals were observed, although it was striking in appearance and easily iden- tified in its slow flight. Number: Total recorded, 3. Taken, 1. Date: July 8 and 10. Record: 1948 — July 8 (1 taken, 481116), 10 (2 seen distinctly). Heliconius eucoma metalilis Butler. Field Name: Ithomiid-mimic. Species Range : Panama to Peru and Ama- zon Valley. Subspecies Range ; Colombia, Venezuela and Curasao. Field Characters: An ithomiid mimic, in- d'st'nguishable in flight from several spe- cies, as well as from the heliconid H. anther ' da holcophorus. Number: Total recorded, 36. Taken, 29. Sex: Both sexes taken. Date: April 8 to July 26. Record: 1946 — April 8 (1) ; June 10 (1), 16 (1) ; July 1 (1), 10 (1), 14 (1), 18 (1), 25 (1). 1948— May 2 (1, 481022), 23 (1, 48573), 28 (1) ; June 27 (1, 48966) ; July 2 (1, 481021), 3 (2 taken), 6 (1 taken from group of 4, all alighted), 11 (1, 481148, km. 30), 13 (2 taken), 14 (2 taken), 15 (2 taken from flock of 6), 16 (1), 18 (1, 481248), 26 (3 taken) . Heliconius melpome ne (Linnaeus), form melpomene (Linnaeus). Field Name: Red-banded Black Heliconid. Species Range: Northern South America to Peru and central Brazil. Field Characters : A common migrant and resident at Rancho Grande. Slow flyer, usu- ally through underbrush and along trails, easily caught with fingers, always giving out a strong scent of witch hazel. In pattern and color very close to the female pierid Perente charops meridana, and to the nym- phalid Adelplia lara lara. In both cases the wings are broader and the flight nervous and swift in comparison with the heliconid. The three species are sometimes seen asso- ciated. Number: Total recorded, 2,566. Taken, 42. Thousands were not counted on days of in- tensive migration of other species. Sex: Both sexes taken. Date: April 29 to September 7. Note: Never saw them hurrying; often alighting on leaves or blossoms; occasionally courting and mating. Record: 1946— May 22 (1), 28 (2), 29 (1 taken, 6 seen) ; August 3 (213 flying slowly through Pass) ; September 7 (6 seen, 461145; first in a month). 1948 — April 29 (1) ; May 23 (1 taken, 30 seen), 25 (1 taken, 8 seen), 26 (1, 6 seen), 31 (1 taken, 27 seen); June 6 (2 taken, 30 seen), 7 (1, 6 seen), 9 (1, 61 seen), 10 (509 counted and many missed ; 4 and 5 males hovering over single female. Many alighted on my sleeves), 11 (18 in twenty minutes), 15 (16 seen), 16 (17 seen), 17 (117 seen), 18 (48 singly, 68 in groups), 19 (2 at 8.30 A.M.), 21 (28 seen), 22 (53 seen), 28 (27 seen, one with nymphalid mimic, flying around each other), 29 (33 seen) ; July 3 (1 taken, 66 seen), 5 (243 singly or not more than 6 together), 6 (2 taken, 64 seen), 8 (1 soaked in rain, 108 seen), 9 (29 at km. 31 headed for Pass), 10 (66 seen), 11 (4 seen, km. 15), 13 (130 seen, 3 at km. 15), 17 (8 feeding on lantana, km. 30), 21 (368 seen), 29 (1 taken, 66 seen) . Heliconius s ara s ora (Fab.). Field Name: Small Blue-hindwing. Species Range: Panama to south Bi'azil and Bolivia. 68 Zoologica: New York Zoological Society [35: 3: 1950] Subspecies Range : Panama, Colombia and Venezuela. Field Characters-. Only to be confused with the very rare H. doris, form doris, and that hardly, for the latter is a full third larger, although one individual sara is un- usually large. Number-. Total recorded, 3. Taken, 3. Date: April 29 to July 25. Record: 1946— May 29 (1 taken). 1948 — April 29 (1, 48445) ; July 25 (1, 481369). Philaetria dido ( Clerck) . Field Name : Green-spotted Heliconia. Species Range: Honduras to Peru, Bolivia and southern Brazil. Field C haracters : Only to be confused with the nymphalid Victorina stelenes, from which it can be distinguished by much nar- rower, more elongated wings, and by a slow, wavering flight. Number: Total recorded, 625. Taken, 4. Sex: Both sexes taken. Date : March 15 to September 8. Frequency: Decidedly gregarious, five large flocks being recorded. Record: 1946 — May 26 (13 passing km. 30), 28 (98 seen), 31 (1 alighted and oriented itself to sun) ; July 8 (196 seen) ; September 7 (125 passing slowly but steadily), 8 (250 +, bucking high wind). 1948 — March 15 (12 seen) ; April 15 (16 seen), 29 (1, 48444), 29 (1 taken) ; June 6 (2 taken) ; July 15 (1 taken), 21 (1 taken). Beebe: Home Life of the Bat Falcon 69 4. Home Life of the Bat Falcon, Falco alhigularis alhigularis Daudin.1 William Beebe. Director, Department of Tropical Research, New York Zooloyical Society. [This is one of a series of papers resulting from the 45th, 46th and 47th Expeditions of the Department of Tropical Research of the New York Zoological Society, made during 1945, 1946 and 1948, under the direction of Dr. William Beebe, with headquarters at Rancho Grande in the National Park of Aragua, Venezuela. The expeditions were made possible through the gen- erous cooperation of the National Government of Venezuela and of the Creole Petroleum Cor- poration. [The characteristics of the research area are in brief as follows: Rancho Grande is located in north-central Venezuela (10° 21' N. Lat., 67° 41' W. Long.), 80 kilometers west of Caracas, at an elevation of 1,100 meters in the undis- turbed montane rain forest which covers this part of the Caribbean range of the Andes. The migration flyway of Portachuelo Pass, which is also the watershed between the Caribbean and Lake Valencia, is 200 meters from Rancho Grande. Adjacent ecological zones include sea- sonal forest, savanna, thorn woodland, cactus scrub, the fresh-water lake of Valencia and various marine littoral zones. The Rancho Grande area is generally subtropical, being uni- formly cool and damp throughout the year be- cause of the prevalence of the mountain cloud cap. The dry season extends from January into April. The average humidity during the expedi- tions, including parts of both wet and dry sea- sons, was 94.4% ; the average temperature dur- ing the same period was 18° C. ; the average annual rainfall over a five-year period was 174 cm. The flora is marked by an abundance of mosses, ferns and epiphytes of many kinds, as well as a few gigantic trees. For further details see Beebe and Crane, Zooloyica, Vol. 32, No. 5, 1947. Unless otherwise stated the organisms discussed in the present paper were observed in the montane cloud forest zone, within a radius of one kilometer of Rancho Grande.] Introduction. This is a record of observations, during a period of one hundred and sixty-four days, of the lives of a pair of bat falcons, from February 20 to August 1 inclusive, in 1948. Most of the watching was done through open windows from within our laboratory at Rancho Grande, the perches of the parent birds and their nest being in full view, at a distance of about one hundred meters. Meth- ods included the naked eye, a 7 X 30 hand binoculars, and Zeiss giant binoculars of 12, 20 and 40 powers. This last was of course mounted on a tripod, and except when in use 1 Contribution No. 872, Department of Tropical Research, New York Zoological Society. elsewhere was constantly in place focussed on the perches or the nest of the falcons, so that instant contact could be made at any desired moment by myself or any member of my staff. With this battery of glasses it was possible to see every detail of color, pat- tern and plumage of both falcons and their victims, as well as such minutiae as the coil- ing and uncoiling of the tongue of a still living morpho butterfly held in the hawk’s talons. During two preceding years we had become proficient in sight identification of almost every bii’d, mammal and insect which en- tered into the diet of these falcons, and within arm’s reach in the laboratory was a collection of skins of the birds themselves, loaned by Dr. William H. Phelps. This col- lection included all the species of small and medium birds which might possibly occur in the vicinity of Rancho Grande. Falco alhigularis, as a species, has an ex- tensive distribution, covering 48 degrees of latitude, from central Mexico, south through- out South America, reaching as far as south- ern Brazil and northern Argentina at about 27 degrees South Latitude. Two subspecies have been described. The bat falcon is fairly common through- out much of its range, and is accounted a bird of tropical lowlands, rather than of subtropical elevations. Brief accounts of nests, eggs and food have been published, the latter based chiefly on stomach contents. The principal contribution of the present account is the data afforded by the opportu- nity for detailed identification of the diet of this pair of falcons throughout more than five and a half months, and the recording of the rise, development and in some cases de- cline of various instincts in the young birds. Previous Records. In accord with the reputed limiting of these falcons to areas of low elevation are my notes of their presence at sea-level, at Kartabo, British Guiana. In 1920 I found this species living in dense jungle in pairs, always perched in the tops of the highest trees. The adults were feeding chiefly on honey-creepers, both Cyanerpes caeruleus caeruleus and Chlorophanes spiza spiza. Three of each of these species were found in an equal number of stomachs. Rufous- throated birds of the year were insect eaters 70 Zoologica: New York Zoological Society [35: 4 for the most part, large and medium butter- flies, giant grasshoppers, together with small flying hemiptera and coieoptera, forming the larger proportion of their diet. At Rancho Grande I have notes of only two occurrences of these birds, other than the pair under consideration. On July 6, 1945, at nine in the morning, I was seated near the gate of Rancho Grande compound, watching a pair of tanagers in a distant tree, when a sharp ke-ke-ke-ke drew my attention, and directed it to a bat falcon in the top of a lofty candelo tree, about one hundred meters away. This was the first record for this locality. The bird looked straight at me as I shifted the barrels of the big binoculars, then shifted its gaze upward and I followed. High in a air, a flock of what to the naked eye might have been dragonflies, resolved into the en- tire company of Rancho Grande blue and white swallows, Pygochelidon cyanoleuca cy- anoleuca. They were completely absorbed in the small hawk, but circling at a safe dis- tance. Occasionally one or two reckless birds would swoop down and rush past in high gear, twittering loudly, but attracting not the slightest attenion. Every feather on the body of the hawk was frequently raised, shaken and settled, as the bird preened itself. Once a small plume fell out and drifted slowly downward, watched by the owner with complete concentration. Most of the time it stood quietly, the head moving quickly from side to side, bobbing up and down in falcon fashion. During the ensuing hour it called three times, a con- tinued rattling, a shrill, high ke-ke-ke-ke, five to twelve times, very fast. Once I heard a variation whew-ke-ke-ke. Twenty minutes after first discovery the bird rose, called loudly, circled the tree once and started due north. Every swallow vanished, but in less than a minute the hawk returned to a perch a few feet way and equally high. As it flew over every bird sound ceased in the sur- rounding jungle. A tree crashed near by and there came a loud hum from a plane over Valencia, but the hawk paid no attention. When a fly alighted near its toes, it was watched closely until it took to flight. At ten o’clock the hawk left again, this foray, after the conventional wasp-like cir- cle, heading it eastward over the Rancho building. This time it was attended by a streaming comet-tail of worried swallows at judicious distances, all twittering as the hawk flew over their nesting crevices. Again the hawk returned but hardly had it resumed its perch when six more bat falcons circled over, high up, and with loud chittering my bird joined them, fell into their circling pat- tern, and all disappeared northward, directly through Portachuelo Pass. They were ap- parently migrants only from lowland to low- land through the flyway. About a month later, on August 4, a single falcon drifted slowly up Limon valley toward the Pass, but soon disappeared into the thin neblina. The following is a selection of day by day notes from my journal, excluding all unim- portant matter which would gain nothing by repetition, and omitting a number of event- less captures of duplicate species prey. In the first section, mention of individual prey is by species only, this being in ac- cordance with sight identification. In the final systematic list, the subspecies are added, it being understood that these are based on geographical expectancies. Chronology. Adult falcons first seen, and mating Nest discovered Eggs estimated laid Incubation estimated Hatching estimated Week-old chicks first seen April 11 Last visible down gone May 5 Young male left nest May 10 Young female left nest May 13 February 20 March 30 March 1 to 5 March 5 to April 5 April 5 Bat Falcon Diary. February 20, 1948j Heard a hawk calling at 7 :30 A.M. and later saw a bat falcon with something in its claws alight in candelo tree No. 3. The large glasses showed the prey to be a male eastern swallow-tanager, Tersina viridis. The blue plumage filled the air as the hawk plucked out beakfuls of feathers, when suddenly thei’e entered my binocular field a larger bird. With an upward swoop the new arrival snatched the prey, and flew off to the top-most twig of candelo No. 2. She — for that proved ultimately to be her sex — tore away at the swallow-tanager while the male watched, and picked at the shreds left in his claws. Half an hour later the male flew straight across to the other tree, mated with the fe- male intermittently for fifteen seconds, and then back to his own perch. Both birds left a few minutes before nine. (Note: Directly south of Rancho Grande laboratory, about one hundred meters dis- tant, were three very tall candelo trees, Gy- r anther a caribensis. No. 1, close to the road, fell in the rainy season of 1947. Of the re- maining pair, No. 2 was to become the fa- vorite perch of the female falcon, while No. 3, a few meters to the west, was the male’s perch more than thirty meters above the ground, all in full view through the open windows of the laboratory. Both trees were on the northern rim of the steep Limon val- ley, facing and to the south of Rancho Grande. This valley led downward toward Maracay and Lake Valencia, and in the other direction ended at Portachuelo Pass.) February 22 and on succeeding four days: Male arrived in early morning and left im- mediately. February 27: Both birds on their respec- tive perches at 6 o’clock, the male with a male American redstart, Setophaga ruticilla, which he plucked and ate. At 8 the male went to the perch of the female and chittered fre- quently. 1950] Beebe : Home Life of the Bat Falcon 71 February 29: Birds at 6:30. Later the male brought a female euphonia, either Tanagra xanthogaster or laniirostris. When plucked and partly devoured it was taken by the female and carried away. March 1: Both birds at 6:30. Male made two flights far down Limon valley and finally returned with a white-breasted martin, Progne chalybea, a bird abundant about Maracay and Turiamo but unkown at Rancho Grande except as a migrant through the pass in late July. The female took the bird at once. Its long wings interfered with her landing and she had to circle and make a second upward swoop to her perch. March 2 to 13: During our absence the birds were reported almost daily, the male carrying prey, but no definite identifications were made. March 14 : Both birds on their perches at 8:30. They were soaking wet after the night’s heavy rain, indicating an exposed sleeping perch. Long and thorough preening, and sun bathing with fluffed-out plumage. The male made a fifteen-minute trip and at 10 o’clock returned with a small bird. As he revolved once or twice on his perch I made out glimpses of the pattern and finally iden- tified a male blackpoll warbler, Dendroica striata. The air was filled with feathers as he pulled and tossed them out of his beak. Remiges and rectrices were with difficulty jerked out, but the warbler was completely plucked before the falcon began to feed. From the beginning of the defeathering at 10:05, twenty minutes elapsed before the end of the meal. There remained a head and wingless half-devoured skeleton body with a single leg. He clutched this cadaver in one lowered and extended foot, and at 10:30 yawned widely twice, and shut both eyes for five minutes. At 10:40 he ate some more and dropped what was left of skull and skeleton. I am impressed with his lack of plucking and tearing power compared with that of his larger mate. He sat quietly until 12:15, made three short flights, capturing a honey-creeper, Chlorophanes spiza, and then sat in the heavy rain until his plumage was again soaked. Left at 1 P.M. and the female fol- lowed a few minutes later. She had spent the entire morning sitting motionless on her perch. At 3 o’clock both birds reappeared, the male with a freshly caught female redstart, which the female falcon took from him, plucked and devoured. Even in the midst of violent plucking, both birds, at times, would suddenly stop and watch the path of a slowly falling feather until it was out of sight in the foliage far below. March 15: Male perched at 6:25 with a male thick-billed euphonia, Tanagra lanii- rostris, in his claws. In ten minutes the bird was cleanly plucked and then eaten. A bit of blood and a feather fastened on the upper curve of the beak and would not be shaken off. Although the stub on which he perched was only about an inch across, and a rather strong wind was blowing, yet the falcon gripped his prey in one foot, lifted the other and slowly and deliberately scratched and cleaned his mandible. The female made a belated appearance and a futile attempt to snatch the remains of a bird from her mate. She then perched and kept up a violent ke-ke-ke at her mate, at the top of her voice, revolving the while and picking at the wooden splinters of her perch. She left soon afterward but returned just in time to snatch a honey-creeper, Coereba flaveola, from her mate. The sunlight was unusually brilliant and through the forty powers lenses the bird might have been perching on my wrist. Part- ly exposed blood sheaths were distinct and, as I watched, a flat feather-fly slid sideways from beneath the plumage and vanished be- low the scapulars. When watching the bird uttering its call, I became aware that the first beak and throat effort began definitely before my ear caught the first ke, while no visible effort accompanied the final ke. I then timed threes kes with a stop watch and found their duration summed up as exactly a second. This verified a long forgotten fact that sound travels about three hundred and thirty meters a second. So my estimate of the hundred meters from eye to bird seemed reasonably accurate. When the hawks were very excited or hungry, the acceleration of Ice-ke-lces put all this estimate of trisyllabic tempo to naught. March 16: Male around at 6:45 with an- other male thick-billed euphonia. The female falcon circled for a few minutes, then dived out of sight. After plucking the tanager the male also vanished among the foliage. When one or both birds appear there is a very evi- dent subsidence of sound in the neighbor- hood and no small birds are visible, but this local alarm soon passes. Birds from blue tanagers up in size, caciques, toucans, para- keets, pay no attention. At 3:30 the male brought a short-tailed swift, Chaetura brach- yura. March 21 : After our four days of absence I saw the male falcon this morning on his perch at 6:38 with prey. This was plucked beyond specific recognition, except that the short, thick beak and a few green feathers indicated another female euphonia. A high wind, thirty miles per hour through the Pass, made perching on one foot most diffi- cult. As I watched, the bird was blown bod- ily backward; he swung out over the valley and dived out of sight still carrying the bare carcass. March 22: Male at 6 A.M. Away until 9:30 when he reappeared with small bird prey but dived into the foliage too swiftly for further identification. On perch ten min- utes later, for an hour. He makes a perfect weather vane, and today, with variable winds, faced in turn, north, north by east and east. March 23: Male at 5:35 A.M. Returned at 6:15 with blue tanager, Thraupis virens, thereby betraying the apparent fearlessness 72 Zoologica: New York Zoological Society [35: 4 of this species in the hawk’s presence. Long periods of watching the horizon and unus- ual plucking used up an hour in prepara- tion, and it was not until 7:10 that the tan- ager was denuded of feathers. It was then carried down and away. Two pairs of blue tanagers have their nests in crevices of Rancho Grande’s outer walls, together with the blue and white swal- lows. This afternoon one tanager was miss- ing from the pair in the eastern wing so I presume it was the victim. This is the first time I have evidence of prey captured in the vicinity of the laboratory. Within a week a new tanager mate joined the single bird. March 24: Male at 6:05. Two trips with- out prey. The female came at 7:10, preened for a time and both left, flying northeast over the laboratory. This is the first appear- ance of the female in a week. March 25: Male on duty at 6 A.M. Left and returned shortly with a dark bundle of prey which the female took at once. In- stead of perching and making a complete meal, she soon dived and swooped up among the mid-foliage of the male’s candelo. The big glasses showed her in the dense shade, hopping along a large horizontal branch cov- ered with moss and small orchids, and hold- ing a good-sized brown bat in her beak. The skinny fingers of one wing were moving slightly, and she put her burden down, bit it several times, then wiped her beak. Next she picked it up, carried it a short distance farther along a sloping limb and pushed it down hard, nudging with her beak again and again, between two leaves of a radiating monstera. The rounded head and face and the upright ears of the commonest bat of Rancho Grande were plainly visible, Lon- choglossa caudifera, the culvert bat. She flew away, but at 3:10 in the afternoon returned, salvaged her bat, tore at it a few minutes and carried it away. March 26: The male arrived at 5:30 and at 6 : 15 returned with a still living male black- headed seedeater, Sporophila nigricollis, plucked it, dived and returned with empty talons. Before ten o’clock, in the course of a short flight he caught a female of the same species of seedeater which he ate leisurely. After the meal he faced the sun and I again used the 40-power lenses. I was struck with the separate muscular control of adjacent but pigmentally separated plumage areas. The rufous mid- and lower belly and flanks were often flat and tightly compressed, while the cross-barred breast, upper and lower sides were fluffed out, distinctly raised above the level of the rufous, blowing about in the breeze, wholly unlike the firmly-held rufous. The male remained on his perch all morn- ing and at 2:05 P.M. turned his back to the sun, spread wings and tail to widest extent, fluffed out his plumage and for fifteen min- utes seemed half asleep and making the most of the rare undiluted sunshine. He then preened thoroughly and left. March 27: Male arrived at 5:30. Quiet until 6:10 when he left his perch and flew around the near side of the candelo; then made three complete circles, gaining height rapidly until he was only a speck, and van- ished. In less than three minutes he was back on the perch with a bird. I rushed to the glasses just in time to see the female appear from somewhere, fly swiftly to him, turn on her side, grasp the bird and tear it from his claws while going at full speed. Through the glasses I saw the wings of the prey stretch wide, and put them down as a swift’s. I followed her to her own perch. She began tearing the bird to pieces and I soon identified the blue and white as one of our Rancho Grande swallows, Pygochelidon cy- anoleuca. The feathers began to fly and she ate for some time and then shifted to a perch a few feet higher on the same dead limb. At 6:30 she dropped like a plummet straight down and disappeared in the lower foliage of her mate’s gigantic candelo. Search as I might, I could not locate her. At 7 :30 the male returned with an uniden- tifiable bird, mostly plucked before he brought it to his perch. The female later took it and carried it away. As she passed, a smoky flycatcher which I was watching van- ished at full speed into the nearest leaves. The male soon flew southeast toward the first Limon ridge, between Rancho Grande and Lake Valencia. Suddenly it dropped faster than I thought any bird could ever fall. At 8 : 05 the female appeared and chittered excitedly. At 9:00 the male reappeared with what seemed to my eyes a parula warbler. When I checked up on range, I realized that the victim of the hawk was the closely related pitiayumi warbler, Compsothlypis pitiayumi. This is the Venezuelan representative of our northern parula, and is an inhabitant of lower, more tropical levels than Rancho Grande. The hawk plucked the small bird and ate bones and all, even swallowing several rectrices. The female watched from her perch a meter above him. After he finished his meal she flew and he took her place. When they are near to one another the difference of nine and twelve inches in size is very evi- dent. Her creamy white collar is paler than his. At 12:30 the male returned with a dead rufous-winged flycatcher, Myiozetetes cay- ennensis. The female started toward him but he would not relinquish his prey, flew heavily out of sight with it, returning at 3 P.M. March 28: Male at 6 A.M. At 7 he leapt a foot up and let himself fall straight down. Far down, below the tops of the lesser trees, he passed through a shrieking flock of ara- tinga parrakeets, then swept up on the op- posite side and back to the female’s perch. The parrakeets might have been stationary in mid-air as far as speed was concerned. They were too big and too well-armed for even him to tackle. An hour later he appeared with a second rufous-winged flycatcher. March 29: At 10:05 the male came to his perch and the moment I reached the glasses I saw he had an unusual bird. A white rump 1950] Beebe: Home Life of the Bat Falcon 73 was distinct and I knew it for a swallow, and rough-winged came to mind. Then, little by little, I saw every detail. The falcon plucked slowly and turned the prey all the way around. The tail was emarginate, the rump white, the primaries brownish, and as he began on the secondaries I saw that they and their coverts were edged with white. The head was gone but the body plumage was partly white and partly shining green, un- questionably a white-winged swallow, Irido- procne albiventer. This is a lowland bird, new to Rancho Grande but recorded by Wet- more from El Sombrero. Later in the year we were to see several hundred migrating through the Pass. The treatment of this prey by the hawk was most interesting and I watched the whole process. For twenty minutes he plucked, in a high wind, working with ter- rific energy on wing feathers, now and then resting by plucking a few beakfuls of body plumage. The wing got tougher as the larger primaries were reached. The thumb plumes refused to be uprooted as did one or two of the secondaries. The entire wing was torn loose at last and swallowed, remaining feath- ers and all, with great difficulty. The feathers stuck out of the beak for a time, until by strong effort they went down. The rectrices came out easily, and finally there remained the body, clean and intact. He had nibbled at the top of the breast but practically noth- ing was eaten but the entire wings. The small legs were torn off and dropped. At 10:30 the female appeared, circling high up, and chittering. The male watched but did not answer. The prey was held in the left foot which rested on the heel. At 10:34 the female dropped, swerved inward, seized the body and left with a single motion with- out touching the perch or her mate. She went direct to her perch on the nearer candelo, and with ferocity ate the entire body, bones and all. The intestines came first and then she tore the body structure apart and little by little swallowed it. After a meal both sexes occasionally in- dulge in a curious continuation of the feed- ing motions. They tug at projecting splinters of the stub and even at adjacent leaves. At last the beak is wiped and the bird settles down. Her plumage on this occasion needed much arrangement as if she had been brood- ing. There was not the slightest doubt that the male prepared the prey for her. I suspected that he might take it away to the nest, until she appeared and took it. I have often seen him pluck and eat a bird wholly without the nicety and neatness of plucking and leaving the body intact when it seems intended for his mate. March 30: At 5:45 the male perched with a small tanager, Chlorospingus ophthalmi- cus, swooped down into the foliage of his tree, and at 6:15 returned with the bird plucked and dewinged as before, but the legs intact. He watched and waited for forty-five minutes holding it out in his left foot, with heel on perch. He would gaze down into the tree below him for minutes at a time. At 7 the female came, swooped up, took the bird and ate it. He wiped his beak again and again, and picked each toe clean. The first few times I watched this performance the male appeared reluctant to give up his prey. Now he prepares it and literally holds it out as far as possible from him. It appears as if the instinct of share your catch was incom- pletely developed at first, but now has achieved a definite routine. Every time the female takes food from her mate she returns to her own perch, calls, and before she be- gins to tear or feed, she evacuates. From 8 :30 until noon I was at Portachuelo Pass watching a heavy insect migration, and in my absence Jocelyn Crane took over, watching the hawks from the laboratory. She was fortunate enough to discover the nest of the falcons, which for the past month has been in full view from the laboratory windows. Her notes are as follows; 9:50 A.M. Was photographing the male falcon on his perch with a telephoto lens when he vanished. Five minutes later he re- turned with a freshly caught bat, looking like the same species which he killed five days ago. He plucked casually at the fur but ate nothing. Saw two flat bat-flies sail away from the fur or wings, both together. At 10 the female zoomed by, making a perfect pickup. She banked back to her tree, to her regular perch while the male disappeared. She ripped off the skin and wings, eating both, one after the other. At 10:14 she plummeted, then curved over to the trunk of the right candelo, the perch- ing tree of the male. She vanished by the time I had refocussed the giant glasses in the general area, and then I suddenly found her in her nest. She still had the bat and was scuffling around in the shallow, apparently empty hollow, appearing and disappearing, occasionally mumbling the bat. Finally she squatted, both she and the bat sinking almost out of sight. 2:15 P.M. (Beebe takes over). Hawk call- ing. The head of the female was visible in the nest, no attention being paid to the loud ke-ke-ke-ke of her mate. The latter was on his perch with a second bat, decidedly larger and the general orange color made it un- doubtedly Cliilonycteris rubiginosa. He skinned the bat, tore off one wing and swal- lowed it. Called again and then dived. I fol- lowed with the glasses in time to see him reach the nest, swooping up from beneath, and deliver the bat, the female reaching out her beak to take it from him. He perched on the rim for a split second, took off and rested quietly on his perch for an hour. The Nest. Candelo number 3, one hundred meters from our laboratory, was 30 to 35 meters high, and the nest, only now discovered, was about half way up, a large but shallow cavity in the trunk. Many years ago, perhaps in the 74 Zoologica: New York Zoological Society [35: 4 course of a great storm, one of the largest branches was wrenched away and fell to earth. This left a gaping wound, which sap and wood set to work to heal. Out-flaring, wavering lips of bark began to curl over the rim, then rot or water or some agent inter- fered and the still exposed heart wood soft- ened and flaked away. This in no way inter- fered with continued growth of the enormous bole, but left a semi-hollow, shallow but well cupped, which in the year 1948 was destined to provide a home for our pair of bat falcons. The setting of the arboreal eyrie was very striking. Three sides of the two-by-one-foot opening were exposed, lichen-covered. Down the fourth side a mass of air-plants cascaded from an enormous burst of foliage, twenty feet above. Within a few feet of the hawks’ home were many kinds of lush growths which at the first hint of rains put forth new shoots, leaves and blossoms; orchids flow- ered, as did bromeliads, jungle arums and tropical mistletoe. Four long, dangling air- roots hung at one side with their beginnings so high up that they pendulumed slowly in the slightest breeze, and when high winds and neblina swirled through the Pass and around the candelos, the vegetable ratlines flapped and thrashed like wrecked rigging. March 31: At 5:30 the male brought a male black-headed seedeater, and after pluck- ing, gave it to female at nest. April 1: Male with prey at 10:30. Mr. Fleming reports that this was a large sphinx moth, Pholus obliquus. The hawk ate the in- sect, discarding the wings. April 2 : Male flew with bat to nest with- out plucking or dewinging it. Later came with a gnatcatcher, Polioptila plumbea, which must have come all the way from Maracay. April 4 : Male on perch at 6 with bat. With- out dismembering it, he called and the female came and took it to the nest. She fussed about with it for a few minutes, on the rim and inside the nest, then took flight, circled once and alighted on a branch half-way be- tween the nest and the perch of the male. Here she laid it down, shifted it, and finally pushed her way into the leaves of a large bromeliad, and backed out without the bat. Flew back to the nest. This is the second instance of caching unwanted food, bats in both cases. (Note: from April 5 to 9 no notes were made owing to our absence on a trip to the llanos.) April 10: Hawk heard at 7 A.M. Fog so dense all day that no trace of candelos, hawks or the nest was visible. April 11: Hawk on perch at 6:30 with a remarkable yellow grosbeak in his talons. It is Pheucticus chrysopeplus, and with its huge beak and heavy body, the streaked back and the white-banded dark wings it gives an appearance of size and weight above that of its captor. I would have given much to see the attack. My only record at Rancho Grande is of a pair feeding in a high tree. The female hawk was on the nest, squat- ting low. At 7:15 I heard a chattering and saw the female feeding three young birds. They were good-sized, about a week old and clad in creamy white down. At this distance they had the general appearance and size of recently hatched domestic chicks. They were fed with small bits of meat which the parent tore from the body of the grosbeak. One got less than the others. After five minutes, she stood on the rim chattering, then flew to her perch and evacuated. The babies pushed each other about. One fought to get on top, finally succeeded, turned tail toward the open rim and shot a shower of white lime drops far out, on and over the edge. The female is now marked by slight lime stains on her longest primaries and outer edges of x-ectrices, so even aside from her superior size and regular perch she is fur- ther distinguishable from her mate. The deep shade and the distance of the nest from our windows had kept the secret well hidden from us. To the naked eye the nest and the bird on the rim were barely dis- tinguishable when we knew exactly where to look. April 12: At 10:15 the male bat falcon swung up to his perch in the top of the ean- delo tree with a swift. Through the 20-power glasses I could see every detail and in every respect of the cephalic pale color it seemed to be the rarest of all Rancho Grande swifts, Cypseloides cryptus. There was no trace of the supraloral white spots of cherriei, and the area around the base of the beak showed the pale grizzled appearance so apparent in both of the specimens taken in 1946. If the bird had been in my hand I could not have had a better view of every detail. It was slowly and thoroughly plucked and as the female did not appear, the male proceeded to eat the eighth known individual of this swift. At 12:15 the male was on his perch again and the female was feeding chicks. Subse- quent onrush of fog hid everything. April 13: At 9:50 male arrived with a chestnut and gold tanager, Tangara arthus, plucked and dewinged it and called. Female came at once, carried it to her perch, quickly ate head and fore body and carried the rest to the nest. Could see only two young birds. Their eyes seem to have more pigment around them. The parent left after feeding, and yelling at the top of her voice, flew to perch. April 14: While still dusk, at 5:30 I saw a large swallow-like bird dash past my bed- room window and leaned out in time to see the male falcon swoop back close beneath me and snatch a small bat from the air. The bat seemed to be standing still, so swift was the hawk. The legs went out and the mammal was seized with almost no slowing up in the space of a couple of wing beats. The hawk flapped in a half circle and then effortlessly glided with sheer impetus up to his perch. The fog prevented seeing any details of plucking but something of the kind went on, then the watching female swooped from her 1950] Beebe : Home Life of the Bat Falcon •75 perch, snatched in her turn, and took the prey to her young. A half hour later when I entered the laboratory she was just reaching the nest with a second bat, this one twice as large. She pulled it apart and ate it her- self. When finished, she came to the nest rim, and for five minutes never ceased call- ing. The strong adverse wind kept the sound inaudible until I cupped my hands and con- centrated. More angles are now visible on the chicks and the down is less dense. Before leaving she cleaned feet and toes thoroughly, one by one. April 15: A bat caught early, the female taking it as it was, from the male at 6 o’clock. At 7:50 male caught a hummingbird. Fe- male snatched it at once, pinched the back of its neck, swung it in her beak for a while, then to nest and fed the chicks. She was through and calling for more all within three minutes. The bird was iridescent purplish- blue on head and breast, green elsewhere, tail bronze and the beak medium long and curved, a male blue-headed hummingbird, C hrysuronia oenone. At 10:05 the male brought a blue-black grassquit, a male, Volatinia jacarina. He must have gone far down the Limon valley before finding this lowland species. It was fed to the young. The nestlings now crane their necks up when the parents call. A second blue-headed hummingbird ar- rived with the male at 1:05, and this time the female took and ate it all, feathers and bones. April 16: Male came at 7:10 A.M. and brought a Tangara arthus and before he had a chance to pluck, the female had it on a branch above the nest, plucked it and tore oft the long primaries with no trouble. She always appears much stronger than the male. She fed it to the young, half plucked. The third, very small, chick only got a bit now and then, and was trampled on again and again by the other two. I thought at first he was a male and the others females, but it is apparent that he is the weakling of the brood. At 9:30 the male brought another culvert bat, and the female took it straight to the nest, tore it apart and fed fur and all. It was heart-breaking to see how the two stronger stood and pushed and gaped at the expense of the smallest. His beak would come up and as a bit of flesh was offered in the parent’s beak the larger ones would snatch it and fairly trample on their weaker broth- er. He has not lost his egg-tooth, and is hardly half the size of the others. The mo- ment she finishes feeding, she steps on the edge of the nest, calls at the top of her syrinx, cleans her toes and claws and flies to her lofty perch. All small birds in sight leave at headlong speed but soon return. Their fear seems deep-seated but evanescant. Also the male seems hai'dly ever to hunt within sight of perch or nest. At 10:30 male caught a red-headed green tanager, Tangara gyrola, and it was fed, body feathers and meat, to the young. Her instinct ends with pulling off bits of meat and pushing them into the first available beak. By sheer accident, now and then, when the larger chicks are half choking on a beak- ful of feathers, the little one gets a piece. At noon another bat appeared, with a repeat performance as of 9:30 this morning. After the rain the rim of the nest is quite free from lime, and the trajectory of the birds is now much higher and stronger, the fine but visible spray of white drops arching high over the nest rim. At 1 :20 another blue-headed humming- bird was caught, and required no dressing or plucking befoi'e feeding. April 17: At 5:20 heard the ke-ke-ke and reached the window just in time to see the male dive at a bat close to the laboratory. The bat swung inward with such force that it struck the glass but not enough to stun it. The hawk went on with no stopping or slackening of speed, up to his perch. At 8 :25 he returned with a half-dead giant white-collared swift, Streptoprocne zonaris, almost as large as himself. The female took it to her perch, and there furiously plucked wing and tail feathers and about half the plumage. There is no doubt about identifica- tion, the long wings, almost as long as the hawk’s, the black plumage with the white collar and maximum size of the more com- mon species. The female hawk tore off and ate the wings. This species appeared twice more in the diet of the falcons. In the nest the littlest hawk was not visible. At 9 : 50 a partly living white-winged swal- low was captured and brought back. The fe- male took it at a higher speed than I have seen before, tearing past, reaching out with all eight claws, and snatching it without effort from the out-stretched foot of the male. Chick number one was still too full of giant swift to open his beak, but number two was ravenous. The female daintily tore off a clean bit of meat, turned her head side- ways and pushed it into the open beak. If by chance he got it wedged crosswise she nudged the side of his beak, loosened the bit of food and rammed it down. Finally she ate the re- mainder, feathers and all, and then hovered the brood. No sign of Dopey. I fear he has succumbed to the cruelty of an unvarying instinct; bringing about the extinction of the unfit. It is significant the number of migrants the hawk is taking, as if their direct flight southward through the Pass and perhaps the slowing up against the usual updraft of wind gave him an extra shade of opportunity. At 11:50 male came with bird, female took it, flew to her favorite horizontal, secondary perch above the nest, plucked and ate most of the wings and some bony parts. Then to nest where Number One got four-fifths of the food, only at the end when he was stuffed allowing his larger, Number Two, nest mate to get a small share. No signs of the weak- ling, Number Three. The prey was a black- faced oriole, Icterus chrysaetus, not new to 76 Zoologic a: New York Zoological Society [35: 4 Rancho Grande, but very common at lower levels. At 12:40 the male caught a stripe-headed warbler, Basileuterus tristriatus. The male plucked and ate wings, and gave it to the female, beak from beak. The two chicks fed greedily although their crops were bulging. April 18: 11:15. The first prey was caught in dense neblina. The female took it to some hidden perch before we could identify it. No signs of Number Three. The smaller of the remaining two gets the major share. The parent never lets any food stick on the chick’s beak, but takes time and care to remove it. When she stands on the rim she systemat- ically cleans her legs and feet, but they still remain blood-stained. The grip which these birds have in their toes when perching is remarkable. Often in a high wind, they will draw up one leg and keep perfect balance and poise, looking around nonchalantly, with only four toes for support. A dog-faced bat brought at 2:30 and fed to young. Later a blue tanager came close to the nest, climbing up the lianas and searching the tangle of leaves, then coming within two feet of the young hawks, and leaving without haste just before the fe- male arrived. This shows a lack of localiza- tion of the source of danger, as the tanager was nesting in a hole of Rancho Grande, a hundred meters away. April 19 : A bat brought at 5 :30. An excel- lent series of views showed it to be the same species as we took in Rancho Grande the night before. It was a white-lined bat, small, with a number of white lines extending over the fur, around the head and down the mid- back, Uroderma bilobatum. We could plainly see the yellowish base of the ear, a character which I had written on the label of the one we took and skinned. At 2 : 15 the female was on the nest, facing out, close to the rim with a chick on each side, when a twelve-inch anolis lizard, the green and black banded Anolis squamulosus , climbed up the trunk. It made a sudden rush, up and over the rim, and the hawk with a quick movement reached forward and seized it by the head. A single pinch and it was drawn in, torn apart and fed. The chick which got the tail had a difficult time, for that organ had become detached, and began a frantic twisting and turning on its own, wrapping about the face and head of the small falcon. Finally the parent had to help poke and untwist, before it vanished, still wriggling. April 22: (On our return from Caracas). Female took a culvert bat from her mate at 6:10 and fed it to the young. These have grown appreciatively. Although still en- tirely covered with grayish-white down, the wing feathers project about an inch. The size difference is pronounced, probably sex- ual, the smaller seeming stronger or more active than the other. The faces are light leaden blue. They are alert, watching every bird or other creature which comes within view, now and then lifting flabby, heavy wings and flapping them as much as possible in the confines of the nest. They have, from the first, showed the vertical head bobbing of the species, as they change the direction of concentration. The whole morning was solid fog, and both adult birds sat close to- gether on the female’s tree. Later a pair of blue tanagers perched and searched for in- sects almost at the entrance of the nest, with both chicks looking at them. A hawk in flight would send these tanagers tearing for cover; a perching one would hold their attention and suspicion, but they seem to see nothing wrong about chicks and nest. The young now call like the parents but weaker, and have lost their baby cheeping. 3:12 P.M. Male caught a blue and white swallow, dewinged and beheaded it, eating these parts, and called intermittently for ten minutes. At 3:30 the female, who had been on a perch below him, taking no notice, finally came ambling on foot along the branch separating them and took the prey from his beak with hers. Flew off, circled and returned to her regular perch. For ten minutes she kept her foot on it, yelling now and then. At last she cached it in a bromeliad above the nest and returned to her perch. Throughout this time the young were gorged, sunk far down in the nest, their nictitating membranes pulled far over. April 23: At 5:45 the male was on the female’s perch with a small bat which she soon took and fed. At 6:30 he had a female seedeater. At 7 :30 the male brought and female took a golden siskin, Sicalis flaveola. In the skinning process all the loveliness of the greenish-yellow, orange and bright clear yellow came into view. This is a lowland bird. She fed it after removing only the large wing and tail feathers. A Tanagra musica, a black- throated euphonia, followed within a half hour. I was uncertain about the black throat and wished I could see it, whereupon the hawk held the entire bird up in her raised foot, and, with me, gazed steadily at the throat and lower parts ! Again and again we should never have been quite sure of species or sex were it not for the deliberateness of the preliminary examination, and the fact that the wing "feathers were the first to be removed, leaving in full view all the other portions of the plumage. At 8:55 a turkey vulture swooped low over the road and nest and the female flapped high, dived full speed and struck with talons, spang on the great bird’s back. The vulture left hurriedly. At 2 : 15 a bat was brought and fed, and soon afterwards a black-headed seedeater was caught. These birds are lowland species but are fairly common on upland intrusions, living and nesting in the small patches of grassy open clearing along the winding road. April 24 : Several series of outcries proved false alarms, male and female sitting quietly, while yelling with all their might. We missed the early feeding. Mr. Fleming was walking along the road several kilometers down hill when a head-and-wingless swift fell at his feet, the hawk passing on out of sight. This 1950] Beebe: Home Life of the Bat Falcon 77 circumstantial evidence makes it possible to add another species to the list of prey, a chestnut-collared swift, Chaetura rutila. Two more individuals were later included in the diet. April 25: Bat brought at 6:30. At 7:10 the male came yelling with prey. Female dived, swooped in a large circle and took the bird. Sat quietly, now and then holding up the bird and looking at it. All this fuss concerned only a small, female, curved-bill green hum- mingbird, Chalybura buffonii. The length and curve of the beak, the general size, the light gray under parts, were as distinct as if in the hand instead of one hundred meters away. The female plucked head and wings and then took the trouble to swoop to the nest and offer a chick the half beakful which remained. Both parents perched for some time. Then the male aroused and went through his reg- ular routine. He shook his head, did a single circle pattering dance on his stub, and con- centrated. He bobbed in all directions but chiefly in the direction of the valley to the south. Three times he bobbed, each time four or five ducks of the whole head. He then shot off, flapped rapidly, evidently lost sight of what had attracted his attention, and re- turned. When he returned empty-taloned he never called. At 8:10 both birds left in a hurry and overtook a big black eagle, Urubi- tinga urubitinga, both in turn striking it feet first. The great black bird ducked and dodged, but three times the falcons rose and fell upon it, until it dived for the shelter of the trees in the valley jungle. Later, the male caught a magnificent black, gold and tur- quoise mountain tanager, Compsocoma flavi- nucha, and ate it himself. April 26 : A bat and a black-spotted green tanager, Tangara chrysophrys, comprised the morning’s larder. Both birds left at 11:15. April 27: At 5:45 the female had a large frog in her claws which the male had just brought. Glimpses made it certain that it was a zipper-backed frog, Gastrotheca ovi- fera. She carried it to the nest but the chicks did not feed while T was watching. T was sorry for this lack of verification of whether this amphibian was edible or not. An hour later she brought a plucked and evidently stiff corpse of a small bird. It had no ap- pearance of recent killing and I can only sur- mise that it was a cached prey which she had hidden some time before, if not the after- noon of the preceding day. April 28 : At 6 : 15 the male brought a chest- nut-collared swift which the female took, plucked, ate most of the head and then fed to the nestlings. At 7:15 the male returned and the female met him, calling loudly, and took a. hummingbird, Phaethornis augvsti. Plucked carefully it made onlv about two bites, one for each chick. After she has plucked a bird, large or small, the female may swoop down to the nest at once, or sit for many minutes, waiting until immediately after an evacuation. The young have grown rapidly and be- come dull grayish-white, owing to the grow- ing feathers. They continually preen all over, and even one another. Each has a dark cap on the center of the crown, and the legs and feet are strong and bright yellow. All of the major pterylae are now distinctly visible through the ragged down. The bobbing, almost from the first week, is continuous. The head is turned, the glance fixed in a certain, definite direction and in- stantly there is a vertical bob or two of the whole head. Then the head is turned quickly toward one or the other side, and another series of jerks takes place. It is as difficult to account for as the foreleg patting or pad- dling of the lizard Cnemidophorus , or the slight withdrawing of the head and refixing of concentration on the same object as is customary in owls. All shifts of vision in the falcons take place in quick sideway turns. April 29: A bat was brought at 6:15. An- other sortie resulted in a male Tanagra xan- thogaster. At 9:15 the female had vanished, the chicks were fast asleep, and the male was on his high perch. Eight short-tailed swifts rushed past, close to him, but neither species paid the slightest attention. A perching hawk and the same bird high in air above are two very diffei’ent factors in safety or danger, in relation to possible prey. At 11 : 15 the male brought a white-breasted martin, Progne chalybea. This is only a casual migrant, so the hawk must have brought this exceedingly heavy bird from far down the valley toward Maracay. 2:45. Note by Miss Crane: The hawk brought two hummingbirds of the same spe- cies within twenty minutes, the female tak- ing both. They were both male racket-tails, Ochreatus underwoodii, breeding along the ridge at Portachuelo Pass. All four rackets were uninjured when they were brought, be- fore plucking. April 30 : The male came at 7 and the fe- male left. For ten minutes he continued his dance. He half raised his wings, then whittled his beak; turned a little and repeated. This continued until he had made two complete circles, sharpening and scraping on first one, then the other side of the perfectly clean mandibles, eyes closed, and rubbing with all his might. It seems to be some strange nerv- ous preceding of hunting. At 7 :45 he left and in a short time re- turned with a rough-winged swallow, Stel- gidopteryx ruficollis. It seemed a bad day for this species for a few minutes after ten he caught another. May 1 : The down on the chicks is now re- duced to narrow lines of pale gray, the rufous collar and breast are distinct. First one, then the other waves the wings frantically, each in turn squatting out of reach of the other’s waving pinions. There is no doubht about the pair being brother and sister, judging from relative size. At 8:30 another white-breasted martin was provided. Again and again I watch the male set a straight course down Lirnon valley and keep on until quite out of 78 Zoologica: New York Zoological Society [35: 4 sight. This must bring it into the lowland, tropical, martin territory. A bat was brought back at 10.30. The two most common species of bats which have formed a considerable portion of the food are both culvert bats, by which I mean that they roost during the day by preference singly or in clusters under the numerous culverts which guide various streams and brooks across the road. Twice I have seen the male falcon dive at these places, once passing through. At neither time did he pick a bat off the walls, but this cir- cumstantial evidence plus the late hours in the day when bats are captured indicate that this source of manna is frequently used. In early morning I have already told of seeing the hawk take bats from mid-air in front of the laboratory. In this case the bats were snatching moths from the window glass. May 2 : Saw one of the chicks tearing up a good-sized, almost unplucked prey in the nest. There is less and less of careful prep- aration of the food on the part of the parents, the change nicely geared to the increasing ability of the young hawks to pluck and tear for themselves. The male clings more and more to the lateral sloping rim at one side and above the floor. May 3: Female arrived at 5:45, called and did her food dance, revolving twice on her perch. She looked for the male, the only time she fails to bob, and he soon arrived with a vireo, Vireo olivaceus. At 6:28, with the female sitting quietly and the chicks with full crops, he appeared and called, this time with a male violet and yellow siskin, Spinus psaltria. The male ate most of it. At 10:15 she fed one chick with a racket-tailed hum- mingbird, a male. She left; both chicks edged toward the rim, yawned in concert and went to sleep. May 4: At 5:45 the female arrived and went through a frantic hunger dance, revolv- ing and tearing at the branch stub beneath her feet. Male brought a culvert bat which was fed almost without plucking. Twenty minutes later a hummingbird was brought, and the female took it and held it in her beak for five minutes. It was then shifted to the left foot. She evacuated, gazed at her prey, and with almost a single motion swallowed it whole. It was one of the smallest of Rancho Grande hummingbirds, a female short- beaked Chaetocercus jourclanii. A skin on my laboratory table measures only 69 milli- meters. The young now exercise regularly and alternately, one climbing up the left side, giving room for wing exercise of the other. Five minutes later they will usually have changed places. They seem to find titbits on the nest floor or at least do a lot of search- ing. The prey is torn or plucked less and less each day. When the parent does tear the prey apart, the separated piece is snatched from her beak at once. Late in the morning the male brought a male racket-tailed humming- bird. May 5: There is little to choose in size between each parent sex and the correspond- ing nestling. The throat and three-fourths collar are rich rufous, justifying the old synonym rufigularis. The corresponding parts of the adults are creamy white. When the young wings are spread, the blood sheaths are plainly visible, but the tail seems almost full grown. Saw the smaller, the male, tentatively put his foot on the rim, lean for- ward and look down, then hastily draw back and scuttle behind his sister. A dog-faced bat and a blue-and-white swallow were early morning food. Again and again I notice that when the female is watching for the return of her mate, she never bobs, only turns her head sideways and up and down, scanning every bit of sky. But when looking for prey or danger or down at the nest both she and the male bob continuously, and the chicks never stop. At 11:14 the male brought a bird, tore off the head and one leg and devoured them and the female then carried it to the nest. The bird was a female American redstart. She hesitated a moment and then left the rim of the nest, carrying the prey. At 1 P.M. she retuimed with the same beheaded red- start, dropped it in the nest and watched the chick on the floor and the female clinging to the side. When one of them began to tear at the bird she left. May 6: Hummer caught at 10. Female pulled out a few feathers and left it at nest. It was a male Anthracothorax nigricollis. Both young birds now come occasionally to the rim and look down for a long time, then hustle back. The male flopped from his eighteen-inch upper lateral perch down to the floor, knocking over his sister in the process. Within a half hour’s time the male brought a white-lined bat and a Tanagra laniirostris. The latter he ate himself. May 8: Bat taken from male at 5:25 and fed. At 7:40 he brought a diminutive hum- mingbird, Chlorostilbon aliciae, tore off head and ate it, removed a few wing feathers and swallowed all the rest. The young take turns now sitting on the rim and flapping. At 10:15 the young male was tearing at something in the nest, and lifted it. It was a male black-headed seedeater, Sporophila nigricollis. None of the wing feathers or rectrices had been removed. At 4:20 Miss Crane called out that the male had prey. I reached the glasses in time to see a large, flat wing twist and fall, giving a brilliant blue heliographic flash in the sun- light. It was a fore wing of the only Rancho Grande morpho butterfly, Morpho peleides corydon. As I watched, the second forewing fell, and then the two hindwings dropped into the leaves of an airplant below. The male watched them spiral down, and then ate the body of the insect. This was unexpected, and another proof of the swiftness and accu- racy of the hunting power of this falcon. May 9: An almost untouched blue-and- white swallow was brought to the nest at 8 :20 and the nestlings tore it apart by them- selves and devoured it. At 8:30 the female called from her perch and took another bird 1950] Beebe: Home Life of the Bat Falcon 79 from her mate and ate it all. It was a female bronze-tailed hummingbird, Aglaiocercus kingi. The sun was out strongly and with the 40-powers I examined the colors. There is not the least hint of white in the collars of the young falcons; the belly and flanks are cinnamon-rufous, and the breast collar is apricot buff. This color extends almost around the neck, a half inch at the nape representing the break in what would other- wise be a complete ring. At 8 :43 the female brought a mouse, prob- ably the common jungle mouse, Heteromys anomalus. This she tore apart and fed with as much care as she showed in the first week of their lives. This time the young male got the first five pieces, then retired with bulg- ing crop, and his sister filled up. The head was torn off and swallowed by the parent, whereupon she left. She simply leans for- ward over the rim, drops with closed wings, and in about three of her lengths the wings open and she is in the full impetus of flight. It looks so easy to me, and perhaps to the youngsters, who follow to the rim, flap, look, bob and turn back. At 2 P.M. she brought a gray-breasted swift, Chaetura cinereiven- tris, after plucking, and the nestlings did their own tearing apart. As I watched, I distinctly saw the mass of small ants when the crop and stomach were torn apart, and even several large marble-like abdomens of Atta queens, which were eaten by the small hawks. May 10: This proved to be the fateful day, marking the break-up of the nesting of the bat falcons. At dawn I came down to the laboratory and started watch. At 5:30 the female came to her perch, and the male brought a short-tailed swift. It was carried to the nest and both birds fed on it. At 6 the male caught another swift which was taken to the nest where the female picked it almost clean. Had continued perfect views of it; a white-spotted swift, Cypse- loides cherriei. This was one of the nine birds recorded at Rancho Grande, making in all twelve known of the entire species. On his arrival the male falcon held the bird in his talons for several minutes, with the dangling head in full view. There was hardly any chin white, but very large and fluffed-out eye spots. As happens sometimes in over-anticipated crises, the actual event came and passed al- most unnoticed. I left the glasses a few mo- ments and when I returned I saw, without emotion, the female leave the nest. When too late, I realized that it was the rufous- collared male nestling who had dropped from the rim, the remaining bird being the con- servative sister. I swiveled up to the candelo perches in time to see the youngster make a crash landing — a messy five-point landing with outspread wings, tail and legs— in a clump of airplants. Slowly collecting his limbs and their feathers, he took off again, fluttered waveringly to his mother, upset her, and clung with agonizing flapping to her perch, and finally made it. He now did three things — looked at distant Lake Valen- cia on the horizon, chittered ke-lce-ke-lce, and bobbed. Life had really begun. [NOTE: In order to effect ease of recognition and clarity of distinc- tion, which are the principal objects of nomenclature, from now on I shall use the following terms : adult male falcon — Male; adult female falcon — Female ; young male falcon — Bob; young female falcon — Nod. Q. E. D.] At 6:30 the female swooped back and forth around the perch to which Bob was still glued. She then dived with great speed be- yond her candelo down the valley and a few seconds later came fluttering up with a bird, the first we had ever seen her catch since the beginning of the nesting. She held it for a time, then half plucked it, ate the head and went to her favorite branch, pushed it down among bromeliad leaves and returned to her perch. The bird was the warbler, Basileu- terus culicivorus. Both adults now made swooping circles around the candelos. At 6:40 she returned to her prey, retrieved it and carried it to the nest where Nod disposed of it. From 7 : 03 we watched the remaining nest- ling. It was a period of great emotion, the strain of the pull out into the great world, versus the safety of the nest. The bird would balance on the outer rim, with wings half raised, teetering forward on her toes, almost go, then turn back. Then would ensue a period of violent flapping, in as wide an arc as the confines of the nest permitted. She would climb to the top of the side wall and pretend to swoop on some prey, a bit of left-over food on the floor, and finally while the fine frenzy lasted, again step to the nest edge, duck and bob, and gradually subside into a yelling mound, crouched on the bot- tom. The trigger was not quite set. Mean- while both parents and brother were out of sight high overhead, with no encouraging calls or even casual visit, or any show-hows. At 7 :40 the male swooped southeast toward Guamitis and returned in three minutes with a hummer, Chlorostilbon aliciae, which he ate complete, head, bill, all. Off he went and at 7 :46 brought another hummer which the female took. This was Chalybura buffonii. At this moment the young bird, Bob, flew across the gulf to near the top of his nesting tree, not far below where the male was perched on his favorite topmost stub. The female brought the re- mains of the hummer to him. He had landed in a mess of smallish dead twigs and slipped and slid, trying to change the bird from beak to claws. But it was too much and the prey slipped and fell far down through the leaves, probably to the ground. He then found a small smooth area, surrounded by small bromeliads and some tiny-flowered orchids. He had a grand time climbing up and down, into the bromeliads, pecking at 80 Zoologica : New York Zoological Society [35 : 4 lichens. A circling fly occupied all his atten- tion and he almost twisted his head off trying to follow it. Then he preened. Meantime the male left twice, flying far out over Limon Valley, and dropping like lightning to the tree-tops, but returned with- out prey. This was at 8:30, and again he went and failed to kill. At 8:55 a red-headed green tanager, Tangara gyrola, hopped about just behind the young hawk, arousing not much more interest than had the fly. But when the male returned, the tanager fled with every bit of wing power. The young bird now discovered a distant turkey vul- ture, and squatted and bobbed as if he was about to take to wing and attack the strange creature. At 9 : 12 the vulture approached and the female drove it headlong away. At 10:55 the male came with a blue-and- white swallow and the female took it to Bob, on his broad, horizontal branch. He ate about half and accidentally dropped the rest. When the female came near to watch the eating, Bob turned his back and chittered. Then she went to a slightly higher branch and Bob followed, foot over foot, parrotlike. He came very close and yelled for more food. Twice she moved and he followed. At 11:10 she swooped down, took a quick look at the nest and Nod and left, leaving a furor of flapping and pseudo-prepai'ation for flight, with no result. Nod seems afflicted with incurable hypsophobia. Noon, no change; everybody preening. At 12:10 Bob got up to the top perch of the male, and took up his stand no more than an inch or two away. There he remained, staring at his parent and yelling unceasingly. Parent, from my human view point, looked embarrassed and nonplussed, and gazed everywhere but at his offspring. It was amus- ing and at the same time very interesting that Bob, who had never seen or been fed by his father, should choose to go not to the more accessible female’s perch, but labori- ously clamber up to the male. At 12 :30, practically pushed off his perch, father fled, followed awkwardly but persis- tently by Bob, the latter circling twice with his father in mid-air and then collapsing in his mother’s tree. Again Bob tried to approach his male parent, but father was fed up and refused to budge or pay atten- tion to his clamoring issue. Bob at last flopped to his more horizontal, comfortable branch where he had eaten his first meal. At 1:50 Bob flew to his mother, knocked her off and alighted. From now on neblina prevented all observation. Late in the day I saw Bob fly into the heart of his mother’s tree. May 11: 5:15 Bob yelling like mad. 5:30 male brought bird, female plucked it thoroughly and fed to Bob who was still in the heart of the tree. The prey was a Coereba flaveola. Bob later flew out to a branch between those of his parents. At 6:15 the female flew to nest, went in and looked at Nod, stayed a moment and left. Five minutes later returned with partly plucked calliste, the black-spotted Tangara chrysophrys, and Nod fed on it. At 6:30 Bob flew down near nest, then into it. He took the remains of the calliste from Nod, and cowered over it in a corner, tore it apart and devoured what was left. Then he climbed up the slanting side of the nest, and before he flew, soiled Nod’s plu- mage with a shower of lime. He had expe- rienced only twenty-four hours of freedom, yet in this short time he had lost the mutual, or at least not unfriendly sharing of food, as well as all directive instincts of nest san- itation. 7:15 male brought hummingbird, female took it and ate it all, a small Chlorostilbon canivetii. The color characters showed dis- tinctly. At 7:25 the female herself caught a Rancho Grande blue-and-white swallow, brought it to nest and both birds fed. At 7 :35 male caught a Chalybura buffonii, took it to his perch and ate it ail. It came from near a flowering tree, some distance down the road, greatly haunted by birds of this family. It may be that the abundant supply of hum- mers was taken as they fed from jungle trees. 7 :40 both young in nest preening. Adults circled trees several times, calling loudly. At 8:05 Bob out into the thin air and alighted near the male. 8:20. Male left and flew out over Limon Valley with his swift-beating flight, a very rapid beating of wings, followed by a space of slower beats, then repeated. I followed him with No. 7’s. Before long I saw him waver and dodge; then four birds of his own size swooped at him, two or three times. Once he turned sideways and I could see his out-stretched claws. He seemed to turn back and I lost him, but the four birds flew toward me and passed north directly over Rancho Grande, a quartet of white-collared giant swifts. Not very long ago he had brought one of these birds to the female. Apparently they are not afraid of him when on a level ; they can probably outfly and out dodge horizontally, but when he has eleva- tion, with the lightning-like speed of his plunge, it would seem that no flying creature can escape. He was back on his perch within the minute. 8:45. Bob returned to nest and ten min- utes later left again. He returned twice more to the nest. Actions of all obscured by neb- lina. 4:00 P.M. Male caught, female plucked and fed a Tangara arthns. Both young in nest. Bob seized it and hovered over it in a corner, and after the female left, every bit that Nod got was only after a struggle and a severe tussle. All tolerance and friend- ly sharing were gone after a few hours of the wude world. His whole psychology was changed. Yet sufficient independence had not been acquired to prevent a return to the nest. For an hour or two in the afternoon Bob perched on the topmost stub usually occupied by the female. 1950] Beebe : Home Life of the Bat Falcon 81 May 12: 5:15. In the dusk Bob called loudly. 5:25. In dim light, male brought a cul- vert bat, female ate some, then carried it down to the nest to Nod. 6:00. Bob now appeared, damp and di- shevelled, having apparently spent the night in the open. He flew to the long oblique bough in female tree, and ascended it by fits and starts, half like a woodpecker, half like parrot, then flew, pushed off the female and on to the highest stub of the tree. Male made several forays. When he re- turned empty-clawed he did his frantic dance of sharpening his beak, revolving several times on top of his narrow perch. Then he brought a short-tailed swift, ate half and the female took it and ate the re- maining half. Bob yelled continuously. He can fly well in wide circles but makes ter- rible landings, especially crashing in thick foliage. He soon stopped calling, so must have had a meal while I was away from the glasses. 8:50. Second bat brought, male to female to Nod in nest. 9:02. Female caught big cicada and ate it. I had heard the loud zzzzz of a “six o’clock bee” stop suddenly in mid-roar, and it may have been this individual. Wings not eaten. 9:40. Very small hummer with a short, almost straight beak; another Chlorostilbon canivetii. Male to female. She plucked all long feathers, flew to Bob ten feet away and thrust it at him. He spent much time yelling and looking in all directions before he ate it. 11:06. Male caught hummingbird, gave to female who plucked it and carried it to Nod in nest. Male Chrysuronia oenone. 4:30 P.M. In gaps of swirling neblina saw both young in nest. May 13 : Early this morning we heard an uproar of chittering and at 6 : 00 through the glasses we saw the nest was empty. Nod had flown at last. Later we saw her high up near the female’s regular perch. 10:05. Male brought a mockingbird, Mimus gilvus, to his perch, plucked off a few tail feathers and the female took it and vanished in the heart of a mass of airplants in her tree. The hunter must have gone beyond the limits of the National Park, even to Limon, to get this lowland species. Neblina shut down on fur- ther observations. May 14: No chance of naming birds brought to youngsters. Rain and neblina opaque. At 6:10 this evening, when sky cleared for a time, saw Nod back in nest. May 15: Took glasses to bedroom, and in morning saw female come to her perch ex- actly at 5:30 A.M. Nod left the nest and alighted below female five minutes later. Then the old and young males arrived. The parent made six dives in the thin neblina before he caught a Chlorospingus ophthal- micus. Gave it to female who gave it to Nod, who ate for a short time, then Bob took it and finished it. The difference between the ages are the pale throat and collar of the adults and the yellow of the old ones instead of bluish cere. Twice the young flew against the old birds and dislodged them, yelling for food. Once the young followed the old male in flight, but when the parent wished to lose the youngster, he side-slipped and turned like a streak, leaving his offspring floundering in mid-air. On the other hand, the most interesting thing of this morning was the occasional brushing off of their perch of the young birds by the parents, a very evident intention to get them on the wing, again and again. At 7 :30 the male caught a female Anthra- cothorax nigricollis, followed by the regular routine of female plucking, eating head and giving the rest to Nod. As she was eating it I saw two birds shoot past at great speed, and the next glimpse showed the male falcon in pursuit of giant swifts. A moment later the falcon returned empty-taloned. Hori- zontally, there is no doubt the swift can out- fly the hawk. At 8:10 there was a quick triple play of a tiny iridescent green hummingbird, Chloro- stilbon canivetii, one of the emeralds, hardly more than two and a half inches overall. After its arrival, it passed in a matter of seconds from male to female to Bob, who flew off and made a crash landing among leaves but clung to his morsel. When he climbed into the open, having transferred the bird to his beak, he stood, chittered to the world and watched the horizon for three minutes before he partly plucked and de- voured the little creature. At 11 :30 the female flew to her perch with a small, compact bundle which turned out to be the white-throated head of a Rancho Grande swallow. She must have plucked the bird, decapitated it, and given the rest to her young. At noon I happened to look at the nest and there was Nod with the rest of the swallow, working on it by herself. May 16: Took glasses to room. Female flew to perch at 5:20 A.M., followed almost at once by the other three, Nod coming from nest. All flew about calling and perching now and then, then both settled on high branches and the parents vanished. At 5:35 female took a small bird from male. Bob flew up and clung partly to the female’s perch. Only after a tussle would the female give up the bird, but finally Bob got it. It was Chry- suronia oenone. Soon the male brought a short-tailed swift, when neblina settled down and stopped observation. At 1 P.M. the female herself caught a male V olatinia jacarina and after plucking and eating the head, gave it to Bob. He did not immediately eat it and his mother tried to get it back from him, whereupon he devoured it hastily. May 19: (After two days in Caracas). Missed early catches but at 7 A.M. the female arrived with male Sporophila nigricollis, was attacked by Bob but shook him off in mid-air in no uncertain way and ate her bird in peace. After removing most of the feathers she ate every part, including head, wings, 82 Zoolofjica: New York Zoological Society [35: 4 legs and feet, then for some time she per- formed her picking dance on the stub. At 7:30 female, Bob and Nod close to- gether at the top of the candelo. The young are greenish-blue around the eyes and on the cere where the parents are bright yellow. They have not learned yet to face up wind, so their plumage often blows the wrong way. Also their stance is uncertain, and they still creep and crawl along a branch with help of waving wings. In flight they have good control and are strong, but alighting is still weak, and I have seen no attempt to learn to hunt. The male brought a hummingbird at 8:20, a Chrysuronia oenone, and ate it. This seems to be the regular early morning routine, a bird to each young, or perhaps a bat, or a second bird, then one each eaten by the adults. There is often a second morning’s period of activity, but little hunting in the middle of the day; a final season of hunting activity comes in the afternoon. At 10 a vulture soared low over the candelo tree and the two falcons took after him at once, and alternately hit him again and again, driving him dodging into the valley. The youngsters followed behind their par- ents, screaming lustily. This day after day watching impresses me with the total lack of play or any alleviation of their routine. They arrive in early morning, watch for a time; dash off, kill, feed their young and themselves, perhaps repeat, and when all are fed they sit quietly for hours, forever look- ing about and preening, seldom sleeping. In late afternoon the activities are repeated, and sometimes before dark they vanish, go- ing somewhere to sleep. At 4:30 all four falcons were, for the first time, within the same field of view of the 20-powers. At 4:45 the male made two un- successful forays. The young got very im- patient and bothered the female. Then both, on separate perches, began tearing at the surrounding leaves, making the bits of old airplant flower stalks and dead leaves fly, while Nod ripped up bits of loose bark and lichens. It was an interesting imitation of the adults. May 20 : After the first terrific rainstorm of the rainy season, at 12 :30 saw Bob in nest eating the remains of a bat. He left and all four did much circling and chasing one an- other. At 4 o’clock all four joined in a re- newed season of flight, diving at one another, and playfully striking with out-reaching talons. I was wrong in the matter of play practice. At 4:30 female caught a white- breasted martin. Yesterday I saw about forty of these birds passing down Limon Valley. May 25: Twice the young birds have been seen to drop prey which had been given them or taken from the parents. The first time it was a blue-and-white swallow, which dropped and flew off apparently unhurt. The second time a female euphonia dropped head- long when the young lost its grip. Early morning and especially in late afternoon, just before the birds disappear for the night, the aerial evolutions continue. Several times I have seen the female, and once the male as well, literally brush or knock the young birds off their perch, and the young then pursue their parents at full speed. The favorite circling and pursuit area is near the Pass and down over the caciques’ tree, the latter seeming to pay no attention. May 27: Male on perch at 5:15, others soon after. Nod swooped down for a brief visit to her nest, perching on the rim for five minutes. A bat at 6:10 from male to female to Bob, a small species with small, rounded ears. She had trouble with the wings. A bird at 6 :55, male to female to Nod, who took it to a special perch of her own. This was a jacamar, Galbula ruficauda, the first of its kind. The wing and tail feathers came out easily, but she could do nothing with the head or long beak and ate only what she could detach. At 7 :15 male dined alone on a Adelomyias melanogenys, the first of this species to enter into the falcon’s diet. May 29: Late afternoon notes: 4:45, both adults left for the night. At 5 both young flew in large circles, over Rancho Grande down south and back, occasionally closing wings and making a little swoop. Returned, perched close together, and yelled in concert. Bob vanished. Nod returned from a short flight with something in her claws, which she dropped in mid-air, a yellow butterfly, Phoebis eubule. It seems to have been her first catch. Later, adults appeared unexpectedly. Sat for a while, then they went for good. Bob flew into the heart of candelo number two, and Nod flew down for a night in the nest. May 30 : This morning got the clue to the beginning of hunt learning. They were both knocked off their perches and followed and struck repeatedly at one or the other of their parents. Then Bob circled and again and again dived and struck at some small insect, never getting it. Then repeated twice, and both youngsters did it. A large yellow leaf fell from the candelo and Nod swooped after it but missed. Yesterday she had been more successful with the pierid butterfly. This seems the transition from flight learning to the first attack. It would seem that the final finished lethal swoop must be many weeks away. May 31: All four at 5:30. Adults left al- most at once, young made a few circling flights. At 6:15 male returned with prey which Bob snatched and took behind a limb. Although I could not see it I knew it was a bat, for throughout the plucking in a fair wind, not a feather drifted away. Later Nod perched below her father and yelled con- tinuously until he left. Then she swooped and chased a swallow in front of Rancho Grande, missing it by a mile, more or less! Her first attempt on a real bird as far as my observa- tions went. June 5: (After return from Caracas). At 11 :54 three falcons in the same candelo, the male. Nod and Bob. Nod was holding a short- tailed swift and plucking it. Bob was watch- 1950] Beebe: Home Life of the Bat Falcon 83 ing with only casual interest so he must have already fed. Two primary feathers soon dropped and drifted away in the breeze. Bob made a feint after them and finally swooped and came back with both in his beak. He toyed with them, let them drop. Soon he repeated his capture with another feather and still another. In this way was skill attained. June 6: At 8:15 and again at 10 I was at the Pass, watching with amazement as flock after flock of sharp-tailed manakins, Chiro- xiphia lanceolata, went past overhead, from south to north, 8 to 26 in each group. These are rather solitary residents of lowland jungle and this was an unexpected minor migration. Between these times I was be- hind my glasses in the laboratory and at 9:15 I saw the female falcon holding and slowly plucking one of these beautiful birds. The tousled scarlet, blue and black was then given to Nod. In this case, as in many others, the falcons drew upon the regular or irreg- ular migration through the Pass. June 10: At 1 :30 a swallow-tailed kite ap- peared and the female falcon took after him, later joined by the male. They swooped with terrific speed, but the kite simply side-rolled, with effortless but lightning-like countering movements, and the falcons shot past and never hit. He rose a hundred feet and then returned, seeming to enjoy the conflict, giv- ing a sense of reserve strength and speed which put the utmost efforts of the past masters of the air — the falcons — to naught. Never have I seen such different, and such perfection of, aerial evolutions. While the kite seemed merely to be egging on the small birds, they were in deadly earnest, wasting much energy in loud chittering. When he de- sired, he rose in a spiral, losing only a few feet at each counter, and vanished through the Pass. The daily pursuit in mid-air of young and old falcons goes on steadily, morning and late afternoon for many minutes. It is def- initely a method of practice. June 11: After flight evolutions Nod was traced to a stub above the nest. She flew down and dug or scraped about on the floor for a long time. Male later brought a fork- tailed swift, Panyptila cayennensis. June 12: 8:15 Bob nowhere in sight. Nod quiet on top perch. Male brought bird, female began half-heartedly to pluck, held in it beak, then called loudly. The bird is a male golden siskin, Spinus psaltria. Female dived with it to nest, entered, waited, called loudly, stood on rim and left. She carried the prey up to a horizontal branch and cached it among some airplant leaves. June 13: At6 A.M. female arrived with bird and was mobbed by both young. Bob followed her to perch and almost knocked her off and got the prey. I followed him to his own perch, where he spent much time calling and bob- bing and looking around before he began plucking. I first saw the spiny tail, then the long, curved wings, then the head and knew it was another of the rare white-spotted swifts, Cypseloides cherriei, the ninth spec- imen seen by us at Rancho Grande and the twelfth known to ornithology. June 17: The young arrive before the old birds these mornings, and chitter till their parents appear. Then more calling until the birds start off hunting. Nod appeared with a long, thin object, and found she was carrying a good-sized stick, a half-inch through and four long. She circled again and again, dragging it with her, and finally tried to alight, and got it stuck with her own wing over a branch, finally having to let the wood go. But she retrieved it before it struck the earth in the best performance I have seen. Carrying it to a higher perch, she savagely bit off pieces of rotten chips. June 19: For two days Nod has been left alone by the other three birds, spending much of the time on a top branch calling. At 10:15 she was tearing apart a white-breasted swift, Aeronautes montivagus. Two hours before I had seen six of these birds rushing south through the Pass, and suppose that the falcon’s prey was one of these unusual mi- grants. Other individuals of this swift were taken on April 4 and June 19. June 22: The morning activities are now shaping into a new and final pattern. At 5:30 A.M. Nod arrived on her perch. For 45 minutes she called and did her hunger dance. At 6 : 15 the female swooped up, called, looked at her offspring and streaked for Limon Valley. Until 9 Nod waited and called. Then she flew down to the nest and lighted on the rim. This seems an unusual thing for a hawk to do. Forty-one days ago she left the nest and now she returned and the rolling neblina hid her from my view. Bob appeared at noon for a few minutes. Nod alone until she left at 6:15 P.M. June 23: Nod on perch at 6. The other three soon arrived and left. Nod did her dance, frantically picking at the lichens, patting with her feet and revolving on her stub. At 1 1 female returned with a humming- bird, a Chrysuronia oenone. Bob returned at noon, apparently well fed, for he paid no at- tention to his sister who, having disposed of the hummer, was now happy with a bat. June 26: Female brought a short-tailed swift. Nod met her in mid-air, took it and promptly dropped it. The female swooped but was too late to retrieve it before it reached the jungle foliage. Came back and chittered, with what sounded to my human mind appropriate sentiments, and left. When Bob came Nod chittered to him and begged as if from a parent. June 28: Nod at 5:30. Flew about and oc- casionally hesitated and shifted with out- stretched claws as if reaching for some in- visible prey. No parents, and soon neblina enveloped her. Just at dusk, as I was passing, I saw fresh lime on the nest rim. June 29: Nod flying in early morning in large circles, now and then striking at some- thing, but as far as I could see, getting noth- ing. On her next flight she slowed up abruptly and struck at what looked like a small hn+. She seized and took it to ner perch. As I 84 Zoologica : New York Zoological Society [35: 4 focussed, she tumbled off and perched on a lower branch. She held a full-sized male hercules beetle, Dynastes hercules. The elytra of the great insect seemed to have closed down over one of her toes, and she had trou- ble tearing it away. The elytra and the flying wings were then torn off and the body was eaten. June 30 : Nod flew up from the direction of the nest at 5:55. After I had finished break- fast I saw she had a small bat and the female falcon was just leaving for the valley. July 3: At 11 Nod left her perch and swooped down close to where I stood in the road, and clutched something in the top foliage of a tree. She emerged with a giant, red-winged grasshopper, Tropidacris dux, which fluttered as he was being carried. Fi- nally he freed himself, but Nod swooped and caught him fairly in mid-air, and ate all but the wings. The routine now is, Nod sleeps near or in the nest, and perches early. Bob and the parents appear later than before, sometimes coming for an hour around mid-day. The fe- male still feeds Nod but irregularly. She is usually alone in late afternoon and leaves about 6 P.M. July 5 : 9:15. Both young birds mobbed the female when she appeared, although she had no food. For most of the morning they hawked after insects. One caught a big yel- low butterfly, Phoebis eubule , and Nod flew after but missed two others. The birds are far from skilful but are eking out their re- duced rations with insects. July 8: At 10, fairly heavy neblina and chilly. The young birds hunted insects, once swooping over Rancho Grande and sending every swallow, old and young, into the rooms. When the old falcons are not around, the swallows seem to pay little attention to the young birds, very distinctly seeming to gauge their ineffectiveness in the air. Bob appeared and with Nod made twelve attacks on a lone yellow butterfly, but it flicked aside every time. As entomologists the young hawks use up a tremendous amount of energy for very small pickings. At 10 the female came to the tree with a chestnut-collared swift and Bob seized it. Be- fore it was devoured, neblina closed in and the rain came down in torrents. At 12:15 the weather had cleared and Nod was soon perched with a prize. She had caught herself a giant, white-striped nymphalid butterfly, Victorina epaphus. A few minutes before, we had seen four and twenty of these butter- flies flying south through the Pass. At her first peck the wings hung down in full view, then they fluttered away to earth, and the hawk ate every particle of the insect and picked at the stub splinters afterward. Later, when all four falcons were perched on their lofty perches, a rufous-tailed squirrel naively came and looked into the nest. He would probably have been perfectly safe ; much tbo large and heavy even for the valiant falcons. July 9: Male and female at 5:45 and both went through their dance. At 6 Nod caught a sphinx moth at the first attempt, and ate it. It was a green Xylophanes, species not recog- nied. At 7 :15 Bob dashed into a mass of tree- top leaves and emerged with an eighteen- inch slender greenish snake in his claws. Probably the common Herpetodryas cari- natus. He half lost the reptile, got a fresh hold, and when the serpent twisted upward lost it for good, and it fell still writhing into Limon Valley. At 12:30 Bob and Nod sailing around together. Bob dived at a passing in- i sect, caught it, and in mid-air transferred it from claws to beak. Saw its knobbed an- tennae and coiling tongue and then the wings of a papilio. Fleming looked and identified it as one of the tailless aristolochia group, Pa- pilio areas areas, a male. He had just taken one at kilometer 31, and I had seen it at the Pass this very morning. Bob pinched off one green and black forewing, which I watched as it fluttered slowly down to the very jungle floor. Then the scarlet and black hindwings dangled and showed every detail. She nibbled at the body and then discarded it, after all her work. This very likely reflects the sup- posedly inedible, bitter quality of the body juices of this group of butterflies. Both birds launched out and Nod returned with a sizeable green leaf, and did her dance, pretending to bite at the leaf as she held it in her claws. Rain started straight down and both took a real bath, spreading wings and tail to full extent and revolving on their tiny perches. It was the first real relaxation I have observed in the family. July 11: At 9:45 Nod suddenly swooped down to the nest and spent twenty minutes there, scuffling around the bottom, pecking and gazing from the edge at nothing in particular. The increased lime marks on the rim make it certain that she must sleep there. Today is only two days less than two full months since she left. July 12 : Much rain and neblina so I missed the hunting and feeding. At 8:45 Nod dived after and caught a yellow leaf and devoted much energy to its destruction. At noon, female came with bat, was mobbed by both young. All were absolutely soaked but Nod won out and would not give up the prey. July 14: At 10:30 we left the Pass and soon afterwards onrushing, low neblina sent the migrating butterflies high into the air and the young falcons had an exciting time. Bob dived twice at a dark butterfly which turned out to be a Victorina. Bob chased the butterfly and Nod chased Bob, but at last Bob found a safe perch and ate his prey. Nod went to the nest, climbed up on one side, then swooped out and caught a dark nymphalid and ate it quickly. A single glimpse of the under wings identified it as Marpesia coresia. Then out again and flushed a large caligo from a tree, Caligo eurilochus. The great owl butterfly rose swiftly and darted irregularly upward, but at her first down swoop Nod got it. She carried it to the nest and one huge wing fell to the bottom as she ate the insect. At 12:15 Bob got a small hummingbird 1950] Beebe: Home Life of the Bat Falcon 85 which his mother brought him, and devoured every bit. It was Chlorostilbon aliciae. I tried to locate Nod’s ke-ke-ke on the scale, and it proved to be the note B, two octaves above middle C, much higher than one would think. The long, dangling bunched masses of orchids almost overhanging the nest of the falcons have never fruited, although they have blossomed. This is doubtless due to their never having been visited by humming- birds, for obvious reasons, and therefore have never been fertilized. The orchid is Elleanthus species? (Department Tropical Research, Bot. Cat. No. 140 ). Similar plants and flowers on the neighboring trees have all fruited. July 18: For the first time in the history of the family, no falcon in sight all day, not even Nod. July 19: Nod on perch early. Bob at 6:15, both soon leaving. July 23: Bob on perch at 6, and saw Nod leave the nest to join him. July 27 : More and more the three hawks are away and Nod is alone, but occasionally fed by the female. She sleeps every night in the nest, and early this morning, after a long downpour in the night, I saw Nod on the rim and Bob behind her, both with dry plumage, so they must both have spent the night there. July 29 : At 4 P.M. a very severe electrical storm, and Nod appeared and flew to the nest where she looked out at the bad weather in perfect shelter. Either this behavior is a sign of weakness and inability to face the elements, or it may represent an advance on Bob, an ability to utilize shelter and thus husband energy. All depends on whether Bob has outgrown her in expert foraging for himself. At dark she was still crouched in the nest. July 31: Nod on perch alone in morning. At dusk Nod in the nest. August 1 : Early this morning Nod caught and brought to her perch a hummingbird, Chrysuronia oenone, and ate it. As far as I know this was her first vertebrate prey caught alive. At 10 A.M. we left Rancho Grande, so this is the last note made on the life history of Falco albigularis. Systematic List of Observed Prey and the Numbers of Individuals Captured by Falco albigularis albigularis. Species represent certain sight identifica- tions; subspecies are based on geographical expectancies. McammoSs — - 33 Heteromys anomalus (Jungle mouse) — 1 Carollia perspicillatum (Dog-faced bat) — 4 Chilonycteris rubiginosa (Large orange bat) — 3 Lonchoglossa caudifera (Common culvert bat) — 23 Uroderma bilobatum (White-lined bat) — 2 Birds — 163 Apodidae — 26 Streptoprocne zonaris albicincta — 3 Chaetura cinereiventris lawrencei — 1 Chaetura brachyura brachyura — 12 Chaetura rutila brunneitorques — 3 Cypseloides cryptus — 1 Cypseloides cherriei — 2 Aeronautes montivagus montivagus — 3 Panyptila cayennensis — 1 Trochilidae — 34 Phaethornis augusti augusti — 1 Anthracothorax nigricollis nigricollis — 2 (males) Chlorostilbon canivetii caribaeus — 3 Chlorostilbon aliciae — 3 , Chrysuronia oenone oenone — 9 Chalybura buffonii aeneicauda — 2 Adelomyias melanogenys aeneosticta — 1 Ochreatus underwoodii polystictus — 7 (6 males) Aglaiocercus kingi margarethae — 5 (4 males) Chaetocercus jourdanii rosae — 1 Galbulidae — 1 Galbula ruficauda ruficauda — 1 Pipridae — 1 Chiroxiphia lanceolata— 1 Tyrannidae — 6 Fluvicola pica pica — 3 Pyrocephalus rubinus saturatus — 1 Myiozetetes cayennensis rufipennis — 2 Hirundinidae — 17 Progne chalybea chalybea — 4 Stelgidopteryx ruficollis aequalis — 2 Pygochelidon cyanoleuca cyanoleuca — 7 Hirundo rustica erythrog aster — 2 Iridoprocne albiventer — 2 Mimidae — 1 Mimus gilvus melanopterus — 1 Sylviidae — 2 Polioptila plumbea plumbiceps — 2 Vireonidae — 1 Vireo olivaceus vividior — 1 Coerebidae — 5 Chlorophanes spiza spiza — 1 Coereba flaveola luteola — 4 Parulidae — 10 Compsothlypis pitiayumi — 1 Dendroica striata — 1 Setophaga ruticilla ruticilla — 5 Basileuterus tristriatus meridanus — ■ 1 Basileuterus culicivorus cabanisi — 2 Icteridae — 1 Icterus chrysaetus giraudii — 1 Tersinidae — 1 Tersina viridis occidentalis — 1 Thraupidae — 19 Tanagra musica intermedia — 2 Tanagra xanthogaster exsul — 1 Tanagra trinitatis — 1 Tanagra laniirostris crassirostris — 2 (males) 86 Zoologica: New York Zoological Society [35: 4: 1950] Tanagra females sp ? — 3 Tangara chrysophrys chrysophrys — 1 Tangara arthus arthus — 3 Tangara gyrola toddi — 2 Compsocoma flavinucha venezuelana — 1 Thraupis virens cana — 1 Chlorospingus ophthalmicus jacaueti — 2 Fringillidae — 12 Pheucticus chrysopeplus laubmanni — 1 Tiaris bicolor omissa — 1 Sporophila nigricollis nigricollis — 5 (males) Volatinia jacarina splendens — 2 (males) Spinus psaltria colombianus — 2 Sicalis jlaveola flaveola — 1 Unidentified birds — 26 Reptiles ■ — 2 Anolis squamulosus — 1 Herpetodryas carinatus — 1 Amphibia — 1 Gastrotheca ovifera — 1 insects — 19 Lepidoptera Marpesia coresia — 1 Phoebis eubule — 5 Victorina epaphus — 2 Papilio areas areas — 1 (male) Morpho peleides corydon — 1 Caligo eurilochus caesia — 1 Unidentified butterflies — 3 Pholus obliquus — 1 Xylophanes species — 1 Orthoptera Tropidacris dux — 1 Homoptera Cicada species — 1 Coleoptera Dynastes hercules — 1 General Summary. Taking into consideration the many days away from observation, and the intermittent hours which we were able to devote to watch- ing, a conservative estimate of individual prey captured by the bat falcons during the five and a half months would be not less than 600 birds and bats. The total recorded is as follows: Mammals Species 5 Individuals 33 Birds 56 163 Reptiles 2 2 Amphibia 1 1 Insects 14 19 78 218 The proportion of essentially aerial forms is as follows: bats 4 species, 32 individuals; swifts 8 and 26; hummingbirds 10 and 34; swallows 5 and 17; lepidoptera 11 species and 16 individuals. This totals 38 and 125, which thus comprise respectively 50 per cent, of all the species, and 57 per cent, of all the individuals. These figures of essential flyers do not include such aerial-feeding birds as galbula and flycatchers. Another interesting fact is the large num- ber of lowland forms in the diet. Some of these may have been taken as they migrated through Portachuelo Pass, but the majority must have been transported several kilome- ters from the relatively low country about Limon and Maracay. There is no doubt but that the majority of the prey was captured at a considerable distance from the nest. Concerning avian prey alone, we can make the following distinctions, in regard to four phases of habitat: Species Specimens Aerial 23 87 Tree-tops 19 37 Open country 10 17 Jungle 4 5 A moment’s consideration shows the rea- sonableness of the respective divisions and their relative numbers. We are dealing with predators whose method of hunting is by keenness of vision, with stance on a lofty, exposed perch, and an ultimate dive from a still greater height, plummeting at full speed upon the victim. This explains the relative proportions of these subdivisions, the preponderance of aerial and tree-top species and numbers, the fewer number of open country, savanna and grass-inhabiting forms, and the dispropor- tionately small percentage of prey living in the heart of dense jungle. Pickford: Vampy romorpha of Bermuda Oceanographic Expeditions 87 5. The Vampyromorpha (Cephalopoda) of the Bermuda Oceanographic Expeditions.1 Grace E. Pickford. Bingham Oceanographic Laboratory, Yale University. (Text-figures 1-8). Introduction. It was the good fortune of the Bermuda Oceanographic Expeditions of the New York Zoological Society, under the direction of Dr. William Beebe, to capture no less than eighteen specimens of the rather rare, bathy- pelagic cephalopod, V ampyroteuthis infer- nalis Chun. The addition of this new material augments the number of recorded specimens to 95, 59 of which are from the Atlantic. Moreover, Dr. Beebe’s specimens, which are all larvae, constitute the largest series that has ever been taken at a single station, with- in so small and narrowly restricted a geo- graphical area. Furthermore they are the only specimens ever collected for which the depth of capture is rather accurately known, having been checked with the aid of a bathy- graph and not merely estimated from the length of the towline. Their capture in the Bermuda area extends the northern distribu- tion of this species in the western Atlantic by about 8° of latitude, the nearest previous records being those of the Pawnee and of the Atlantis, somewhat to the south of this region. The comparative rarity of this ani- mal is attested by the fact that it was taken by this expedition on only 18 occasions out of a total of more than 800 hauls at suitable depths. The collection provides a valuable opportunity for the confirmation, or rejec- tion, of certain conclusions drawn from pre- vious records and representing a total of 75 specimens taken from scattered stations in many different parts of the world. The pres- ent contribution is designed to lay special emphasis on this aspect of the problem. The conditions under which the collections were made have been described by Beebe (1931a) and were summarized as follows by Coe (1945) : “The collections were all made by Dr. William Beebe in a relatively minute portion of the deep subtropical Atlantic Ocean. This portion was a circular column of water eight miles in diameter, with its cen- ter located at 32° 12' N. Lat., 64° 36' W. Long., about nine miles southeast of Nonsuch Island, Bermuda. The nets were drawn hori- zontally across this area so as to collect sam- ples simultaneously at 200-meter intervals l Contribution No. 873, Department of Tropical Research, New York Zoological Society. from depths of about 1,000 to 2,000 meters. During the years 1929, 1930 and 1931 a total of 1,042 nets one meter in diameter were drawn at these depths in all directions across this eight-mile cylinder of water.” It may be useful, for the benefit of readers not already familiar with the story, to sum- marize some of the more interesting features in our knowledge of this remarkable animal. The biology and external morphology have been the subject of an extensive investiga- tion, the results of which are published in the Dana Reports (Pickford, 1946 and 1949a). The internal anatomy, which is still under in- vestigation, has been treated more briefly (Pickford, 1939b and 1940). Vampyroteu- this inf emails is an aberrant, but in many respects rather primitive and archaic di- branchiate cephalopod. It has ten pairs of arms, but the second pair is modified to form slender, sensory filaments which may be retracted into pockets on the aboral sur- face of the web. For this reason it cannot be placed either with the Decapoda, in which the fourth pair of arms are modified as ten- tacles, or with the Octopoda, which possess only eight pairs of arms. The order, or sub- order Vampyromorpha Robson (emend.), to which it is assigned, thus represents a third group of dibranchiate cephalopods related, by the form of the gladius, to the extinct Prototeuthoidea. The adult animal attains a mantle-length of 50 mm. in males, and perhaps 80 mm. in females. The head-width is nearly as great as mantle-length and the web is extensive (Text-fig. 1). The arms, like those of the Cirromorpha, bear a single row of suckers that alternate with pairs of cirri. A certain number of cirrus-pairs, the primary cirri, precede the first sucker on each arm. The ex- ternal surface of the body is jet black, except for the oral face of the web which is red- brown. Near the apex of the body there is a pair of paddle-shaped fins. Just behind the base of each fin there is a circular luminous organ which may be occluded by an eyelid; on the back of the neck there are two clusters of luminous nodules and almost the whole ex- ternal surface of the animal, except the oral face of the web, is decorated with minute, simple light organs. 88 Zoologica: New York Zoological Society [35 : 5 Text-figs. 1-7. Life history of Vampyroteuthis inf emails Chun. 1. Dorsal view of an adult male; the sensory filaments, representing the second pair of arms, are shown protruding from pockets in the B sectors of the web ; the remnant of the resorbed larval fin is indicated by a small pocket below the fin light organ. 2. The spherical, pelagic egg. 3. A post-embryo; dorsal view of the youngest known specimen showing one pair of larval fins, and the squid-like, anteriorly projecting mantle margin. 4. A typical stage 1 larva. 5. An early stage 2 larva, showing anterior fin rudiments in front of the fin light organs. 6. A typical four-finned, stage 3 larva. 7. A stage 4 larva showing the animal as it appears somewhat previous to the resorption of the larval fins. All drawings to the same scale. Drawings by Lisbeth Krause. 1950] Pick ford: Vampyromorpha of Bermuda Oceanographic Expeditions 89 The life history of Vampyroteuthis is most interesting since it undergoes what may be described as a double metamorphosis. The eggs (Pickford, 1949b) are about 3.86 mm. in diameter, spherical, and pelagic. The youngest larva ever captured (Text-fig. 3) has a mantle-length of 5.5 mm. and is pro- vided with a slender pair of larval fins that are set diagonally so that it appears probable that the animal must swim head downwards like other larval cephalopods. A most inter- esting feature of this youngest stage is the forwardly projecting, squid-like, dorsal man- tle-margin. In older larvae and in adults this becomes sunk within the enveloping, sub- cutaneous gelatinous tissue so that the ani- mal assumes a superficial resemblance to the Octopoda. The distinctive nature of the larval fin be- comes apparent in slightly older larvae (Text-fig. 4) in which the fin light organ is developed and is found to be situated in front of the fin base, not behind it as in adults. At a mantle-length of about 10 mm. one can find a minute anterior fin rudiment, situated in front of the fin light organ (Text-fig. 5). The appearance of this rudiment inaugurates the second larval stage. The future adult fin grows rapidly and becomes paddle-shaped, while the length of the larval fin increases but little if at all. When the two fins are subequal, at an average mantle-length of 18 mm., the animal is considered to be in the third, or typical four-finned larval stage (Text-fig. 6). As growth proceeds the an- terior fin becomes progressively larger and soon greatly exceeds the length of the larval fin; this is the fourth and last larval stage (Text-fig. 7). At a mantle-length of between 25 and 30 mm. the larval fin, whose length has scarcely changed since it became estab- lished in the 10 mm. larva, undergoes a sud- den resorption. This constitutes the defini- tive and final metamorphosis of the larva into the adult form (stage 5) and is accom- panied by many changes in the growth rela- tionships of the bodily parts. A minute ves- tige of the larval fin can be found in adult animals, in the form of a little pocket situ- ated just behind the fin light organ. Vampyroteuthis inf emails is found in the Deep Water of all tropical and subtropical oceans. It has rarely been taken at estimated depths of less than 1,000 meters and is cer- tainly absent from the upper 300 meters of the ocean. It is stenothermic and stenohaline and is commonly found just under the oxygen minimum, in water of density represented by CTt 27.6 to 27.8. The idea that it may be restricted to a constant density layer, which was suggested to me by Dr. E. F. Thompson, appears to be confirmed by a comparison of the vertical distribution in the different oceans. Specimens from the Indo-Pacific have usually been taken at greater depths than those from the Atlantic, in water that is colder and less saline but whose density is the same as that of the layer of maximum abundance in the Atlantic. As far as is known, the eggs are also restricted to this density layer. It seems to be quite certain that there is only one species of Vampyroteuthis. Previous descriptions by earlier investigators, who recognised as many as eleven species dis- tributed among eight supposedly different geneva, were based upon distorted and ab- normally preserved specimens, and upon ignorance of the life history. An exhaustive study of the external characters has shown that there are no differences of a specific or even of a subspecific nature between the populations of the three great oceans. Yet there is reason to believe that the Atlantic population constitutes a distinct race, sepa- rable from the Indo-Pacific group by the average number of pairs of primary cirri on the arms. Dr. Beebe’s collection of 18 speci- mens provides an admirable opportunity to examine this hypothesis which, it may be mentioned in anticipation, appears to receive adequate confirmation. It is a pleasure to acknowledge my thanks to Dr. William Beebe for the privilege of examining this important and interesting series of specimens, to Dr. Gordan A. Riley for assistance with the hydrographic data, to Miss Lisbeth Krause for the drawings of the life history (Text-figs. 1-7), and to Dr. Daniel Merriman, Director of the Bingham Oceanographic Laboratory, for his continued interest in my investigations. Seasonal Distribution, Vertical Distri- bution, AND HYDROGRAPHIC CORRELATIONS. The specimens captured by the Bermuda Oceanographic Expeditions are listed in Table I which gives the date, net type and depth. In the last column I have given the length of the towline, obtained by doubling the estimated depth. In regard to the latter, Dr. Beebe states (in litt.) that “The bathy- graph was used only occasionally to verify our estimates as to the depth of the trawling net. We found that at a speed of 2 knots and an angle of 30 degrees, the tests with the bathygraph agreed perfectly with our estimate of the depth as one-half of the amount of cable out.” It is most satisfactory to have this confirmation of the estimated depth of capture, enhancing, as it does, the validity of the hydrographic interpretations which, for all other expeditions, have been based upon arbitrary assumptions. Seasonal distribution. Previous investiga- tions have indicated that larval vampyro- morphs might be taken at all seasons, but the data were quite inadequate for a study of relative abundance. The Bermuda collec- tions were made during the summer months, from April to October, but a tabulation of the data (Table II) indicates a period of maximum abundance in this region during June and July. The depths are tabulated but provide no evidence of vertical migration and the total numbers at all depths may therefore be considered. Comparatively few hauls at suitable depths were made in April 90 Zoologica: New York Zoological Society [35: 5 TABLE I. List of Specimens. No.1 Net No.2 Date Net type Depth meters Towline meters3 78 331 VII.27.1929 1 metre 1,829 3,658 79 700 VI. 13. 1930 1 metre 1,463 2,926 80 807 VII. 16.1930 1 metre 1,463 2,926 81 800 VII. 15. 1930 1 metre 1,463 2,926 82 710 VI. 16. 1930 1 metre 1,280 2,560 83 668 VI.4.1930 1 metre 1,463 2,926 84 861 IX. 8. 1930 1 metre 1,280 2,560 85 328 VII.27.1929 1 metre 1,463 2,926 86 731 VI. 27. 1930 1 metre 1,463 2,926 87 785 VII. 7. 1930 1 metre 1,097 2,194 88 280 VII. 10. 1929 1 metre 1,280 2,560 89 25 IV. 15.1929 1 metre 1,463 2,926 90 788 VII. 7. 1930 1 metre 1,646 3,292 91 1229 VIII. 27. 1931 1 metre 1,463 2,926 92 724 VI. 25. 1930 1 metre 1,829 3,658 93 455 IX. 10.1929 1 metre 1,829 3,658 94 76 V. 8. 1929 1 metre 1,463 2,926 95 198 VI. 20. 1929 2 metre 1,829 3,658 1 Author’s register of known specimens; Nos. 1-77 are listed in the Dana Reports (Pickford, 1946 and 1949a). 2 Data from the published Station Lists (Beebe, 1931b, and 1932). "-Twice the estimated depth of capture (see text). and the paucity of Vampyroteuthis at this time is, therefore, of uncertain significance. The same is not true in May when a total of 114 hauls captured only one animal. In June and July, with 186 and 146 hauls re- spectively, a total of 13 specimens was taken. In August, as in May, an equally extensive series of hauls captured only one specimen. The data for September are even more signi- ficant; nearly twice as many nets were low- ered as in July and yet only two vampyro- morphs were taken. In October, as in April, too few hauls were made to permit inter- pretation of the negative results. Two interpretations are possible, either the larvae are more abundant in June and July, or they migrate into the Bermuda region at this season. The latter hypothesis receives support from two sources. Larval vampyromorphs have been taken from other parts of the Atlantic at all times of year and the otherwise rather inexplicable absence of even young adults from the Bermuda collec- tions might be explained along these lines since there is some evidence that the larvae range more widely than the adults. However, the absence of larger specimens might be due to the fact that most of the towings were made with one-meter nets. The rich collections of the Dana Expeditions were made, for the most part, with two- and three- meter nets. Vertical distribution. No specimens were taken at depths of less than 1,000 meters. This fact is significant since a total of 469 nets was lowered at depths ranging from 0-914 meters ; 172 of these were at 914 meters and if Vampyroteuthis occurred at this depth it seems px’obable that at least one or two specimens would have been taken. As a mat- ter of fact only one specimen was taken above 1,200 meters, the majority being be- tween 1,280 and 1,829 meters (Table II). The absence of specimens at depths of more than 2,000 meters cannot be interpreted on account of the small number of hauls, but in any case such towings must have been taken in close proximity to the bottom and the concensus of previous data suggests that Vampyroteuthis is not commonly taken with- in 250 meters of the bottom. In order to compare the vertical distri- TABLE II. Depth Summary of Data on Seasonal Distribution. N umber of specimens Total nets meters Apr. May June July Aug. Sept. Oct. by depth 1,097 _ 1 175 1,280 1 1 — 1 — 172 1,463 1 1 3 3 1 — — 174 1,643 — — 1 — — — 174 L829 — — 1 2 — 1 — 178 2,011-3,640 — — — — — — — 10 Total spec. 1 1 5 8 1 2 — Total nets by month 33 114 184 146 137 259 10 883 1950] Pick ford: Vampyromorpha of Bermuda Oceanographic Expeditions 91 TABLE III. Summary of Data on Vertical Distribution. Towline, in meters Group 1,000- 1,999 2,000- 2,999 3,000- 3,999 4,000 and over Bermuda — 13 5 — Previous Atlantic records 3 15 10 7 Atlantic Total 3 28 15 7 Indo-Pacific Total 1 5 12 14 bution of the Bermuda collection with that of previously recorded specimens it is con- venient to present the data in terms of the length of the towline (Table III). The re- sults show that the majority of specimens was taken with towlines of between 2,000 and 3,000 meters, in good agreement with previously published records from the At- lantic and in striking contrast to the Indo- Pacific in which the majority of specimens has been taken at greater depths. However, it must be remembered that, on account of the proximity of the bottom, few towings could be made with cable lengths of more than 4,000 meters and the absence of this species at greater depths, in suitable regions, is not therefore confirmed by the Bermuda collections. Hydrographic correlations. It will be seen from Table IV that the majority of speci- mens came from 1,500 meters, in water of temperature of about 4.2 °C., 35.0 %o salini- ty, and density 27.8 o-t. This information is in admirable agreement with previous esti- mates for Atlantic specimens, the majority having been taken in water ranging from 3.0 to 6.0 °C., 34.9 to 35.0 %o salinity, and