GIFT OF v O y Harry Eeal Torrey. Pp. 205-221; 16 text- figures. 11. (XXXI) "^o1e on Geotropism in Corywiorpha, by Harry Beal Torrey. Pp. 223-224; 1 text-figure. Nos 10 and 11 in one cover. August, 1910 20 12. The Cyclostomatous Bryozoa of the West Coast of North America, by Alice Robertson. Pp. 225-284; plates 18-25. December, 1910 60 13. Significance of White Markings in Birds of the Order Passeriformes, by Henry Chester Tracy. Pp. 285-312. December, 1910 25 14. (XXXIII) Third Report on the Copepoda of the San Diego Region, by Calvin Olin Esterly. Pp. 313-352; plates 26-32. February, 1911 40 15. The Genus Gyrocotyle, and Its Significance for Problems of Cestode Structure and Phylogeny, by Edna Earl Watson. Pp. 353-468; plates 33-48. June, 1911 - 1 - Index, pp. 469-478. * Roman numbers indicate sequence of the Contributions from the Laboratory of the Marine Biological Association of San Diego. UNIVERSITY OF CALIFORNIA PUBLICATIONS IN ZOOLOGY Vol. 11, No. 14, pp. 377-5 10, pis. 2 1-24, 5 text figs. February 27, 1914 A DETERMINATION OF THE ECONOMIC STATUS OF THE WESTERN MEADOW- LARK (8 TURN ELL A NEGLECT A) IN CALIFORNIA BY HAROLD CHILD BRYANT CONTENTS PAGE Preface 378 Introduction 382 History of methods in economic ornithology 389 Investigation of the economic status of the western meadowlark in California 395 A comparison of methods in economic ornithology 397 The western meadowlark (Sturnella neglecta) 400 Field investigation 401 Abundance 402 Nesting habits 404 Depredations 404 Experiments on captive birds 409 Amount of food required 410 Time of digestion 412 Examination of the stomach contents 413 Collection and preservation of material 413 Material 415 Examination of stomach contents 416 Identification of stomach contents 420 . Food of the western meadowlark in California 420 Vegetable food 420 Animal food 426 Inorganic matter 436 Principal articles of diet 437 378 University of California Publications in Zoology [VOL. 11 PAGE Examination of feces 437 Quantity of food 440 Capacity for good or evil as evidenced by the number of birds taking different kinds of food 444 Pood of nestlings 447 Variation of kind of food 448 Variation in food according to time of year 450 Variation in food habits according to locality 454 Influence of age and sex on quantity of food 455 Combination of field and laboratory work 456 The relation of birds to insect outbreaks 456 Verdict of ranchers as to the value of the western meadowlark 462 A determination of the economic status of the western meadow- lark in California 466 Suggestions for the protection of crops 474 Recommendations as to legislation 475 Some interesting side-lights on the investigation 477 Parasitism 477 Malformation 477 Albinism 478 Incubation and moult 478 The effect of systematic destruction on the numbers of meadow- larks 478 Natural death-rate 479 Do protective adaptations of insects protect them from the at- tack of birds? 480 Availability as a factor in the kind and quantity of food of birds.. 487 Solved and unsolved problems in economic ornithology 488 Summary 490 Bibliography 494 Explanation of plates 504 PREFACE The impetus given in late years to the study of the relations of birds to agriculture in the United States is traceable to the extensive work of the United States Bureau of Biological Survey. From the organization of this department of the United States Department of Agriculture, July, 1885, to December 31, 1911, members of the Biological Survey have prepared and published one hundred and thirty-one documents relating wholly or in part 1914] Bryant: Economic Status of the Western Meadowlark 379 to the food of birds. Notes on the economic status of over four hundred species of native birds and of over fifty species of foreign birds are to be found in these publications. In many cases extended studies have been made of the food of birds by the examination of stomach contents. In no other country has economic ornithology been accorded the attention it has received at the hands of the United States Department of Agriculture. In a newly settled state like California, where large tracts of land are being brought under cultivation, disturbances of the natural order of bird life arise in two ways. First, the natural food supply of birds is destroyed through cultivation. Second, a new source of food is often supplied by cultivated crops. Hence birds become of great economic importance. The variety of conditions to be found in so large a state as California makes a study of these economic relations of birds complex and difficult. In spite of the need, therefore, of a knowledge of the value of birds, comparatively little work along this line has been done in this state. Until recently two bulletins entitled "Birds of California in Relation to the Fruit Industry," by F. E. L. Beal, published by the U. S. Bureau of Biological Survey, and a few scattered notes in ornithological literature afforded the only pub- lished material on the economic relations of birds in California. Complaints of the depredations of birds in this state have been numerous. The injury to fruit caused by the linnet (Car- podacus mexicanus frontalis) is so great that this bird is branded as a pest by the fruit-grower. In recent years grain-growers have complained of damage to sprouting grain caused by western meadowlarks (Sturnella neglect a). These birds have been ac- cused, and rightly so, of boring down beside the sprout with their long bills and pulling off the kernel of grain. Ranchers have maintained that in some cases whole fields of grain have had to be reseeded because of the loss occasioned by these birds. In fact, there has developed so much sentiment against the meadowlark that there has been a persistent attempt made at each legislative session to take protection from the bird. As the western meadowlark is a bird defended by many because of its insectivorous habits, the agitation following the complaints has afforded an exceptional opportunity to deter- 380 University of California Publications in Zoology [VOL. 11 mine scientifically the economic value of this bird by a thorough investigation. Such an investigation has been made possible through the patronage of the California State Fish and Game Commission, which established a research fellowship in the De- partment of Zoology of the University of California. In Jan- uary, 1911, I was appointed Fellow in Applied Zoology on the Fish and Game Commission Foundation in the University of California, with instructions to carry on an investigation into the relation of certain California birds to agriculture. The com- mission assumed the expense and through its deputies furnished the material for stomach examination. The office work and the laboratory work have been carried on in the Zoological Depart- ment of the University of California. The investigation has, therefore, been conducted through the co-operation of the State Fish and Game Commission with the University of California. To the men past and present who have preceded me in this line of work and who have furnished the world with the under- lying facts which have established the science of economic orni- thology I wish to give due credit, for without their contributions this work must necessarily have been far more elemental. The helpful criticisms and valuable suggestions of Professor Charles A. Kofoid of the University of California, under whose direction the work has been done, have inspired and assisted me in the task. To Dr. Joseph Grinnell, Director of the Museum of Verte- brate Zoology in the University of California, who has often given me of his time to discuss certain features of the work, I owe much. The help also of Mr. E. R. Ong as laboratory assist- ant, and of Professor C. W. Woodworth, Dr. E. C. Van Dyke, Dr. F. E. Blaisdell, Mr. John Bridwell, Mr. W. L. McAtee, Miss Anna M. Lute and others in the identification of insects and weed seeds has lightened the burden and facilitated this part of the investigation. Although collections have been made of a number of birds about which complaint has been received (western robin, bicolored red-wing, Brewer blackbird, horned lark, western mourning dove, and roadrunner), yet, because it was the special object of attack, effort has been concentrated on determining the eco- nomic status of the western meadowlark. 1914] Bryant: Economic Status of the Western Meadoivlark 381 The investigation has consisted primarily of field investiga- tion, experimentation, and stomach examination. A large part of the field work has been carried on at Lathrop, San Joaquin County, California, a place admirably suited for the work in hand. Duties on the agricultural and horticultural demonstra- tion train, which toured the state in 1911 and 1912 under the auspices of the Department of Agriculture of the University of California and the Southern Pacific Railway Company, have afforded additional opportunity to study conditions from one end of the state to the other. Economic ornithology is a new science and has hardly pro- gressed further than the stage of preliminary interest and study. As a result practically all of the work attempted thus far has been of the extensive rather than of the intensive sort, and has been made up largely of a study of the food of birds. In this investigation the attempt has been made to improve on past methods and, by determining the food of birds taken in the same locality each month, or twice each month, to furnish reliable evidence as to their food throughout the whole year. A study of the bird in the field, its depredations, and its life-history, has also been made in order that all available evidence might be obtained. Considerable difficulty has been experienced in that there has been, and now is, a difference of opinion as to the criteria to be used in the determination of the economic status of a bird. The ideas which have been advanced in the past, and even those of the present day, appear to be unsatisfactory, or at least un- trustworthy. It seemed, therefore, that a review of past methods, with the addition of such new ones as appeared to be valuable, might prove not only interesting but of considerable value to future workers in the field. A similar lack of information re- garding methods of stomach examination has been evident. A detailed account of the method used in this investigation, there- fore, seems justified. The service which birds render to agriculture has doubtless been overemphasized. On the other hand, the position taken by some that birds are of no value as insect and weed-seed de- stroyers hardly seems justified by the facts. If there be a mis- 382 University of California Publications in Zoology [VOL. 11 conception as to the utility of birds, it is high time we sought to destroy it and to establish truth in its stead. The economic relations of birds must necessarily become more and more important. As they do so, the extensive study of the past few years will give way to the intensive study necessary to solve the greater problems of the future. Probably no one thing will play a greater part in the conservation of wild life than will this intensive study. Eecognition of the economic value of a single bird will stimulate interest in the protection of all. This "work is published, therefore, in the hope not only that the facts and data here presented may be of general interest and of value to future workers in economic ornithology, but that it may be a factor in promoting the conservation of wild life in California, a state which still possesses enough of its original fauna to make its conservation important and eminently desirable. INTRODUCTION Doubtless if our knowledge were not so limited we might be able to find a use for most living things. As it is, we designate animals as useful, neutral, or injurious because of their effect on ourselves or our interests. A thorough study of the inter- relations of such animals often reverses our original decisions regarding them. Not many years ago insects as a class were called injurious because some of them destroyed certain crops. Today only a part of the insects are considered destructive, and we are yearly finding that others are of neutral or beneficial character. Not many years ago birds were looked upon either as pests or as marks for the gunner. Today most of them are looked upon as valuable assets of the agriculturist. As science in the past has slowly lifted us to a plane where we study the complex interrelations instead of the single and obvious ones, so in the future we may expect that more and more each form of life will be found to fill a particular niche in its environment better than any contiguous form. 1914] Bryant: Economic Status of the Western Meadowlark 383 And yet, viewed from the utilitarian standpoint, there is a certain value in classifying organisms as injurious or beneficial. The danger lies not in the classification itself, but in the risk attendant upon a judgment hastily made or one based on circum- stantial or partial evidence. Being the dominant form of life on the earth, it is only natural that we measure the usefulness of things by their immediate effect on ourselves or our interests rather than on the whole complex of nature. An intimate knowledge of the use of wild life is indispensable to sane conservation. Anything known to be useful may justly demand protection. Anything known to be of no utility in nature may justly be accorded destruction. Ignorance has caused the waste characteristic of the past. Knowledge must prevent waste in the future. What may not seem to be of use today may be of great importance tomorrow. It appears that the economic value of wild life seldom becomes evident until the form becomes extinct, or at least diminished in numbers. It was only a few years ago that fish were so abundant that no attention whatever was paid to their life-histories. Today the study of ichthyology, including fish-hatching, is a necessity, in order that the supply of this kind of food may continue to be available. Not many years ago people believed that there was an inexhaustible supply of game. Today strict game laws and the most careful conser- vation alone prevent the extinction of many forms. Fifty years ago the farmer in the east may have lost some grain and corn from the depredations of birds, but he either planted an extra acre or two to make up for the loss or took it as a matter of course. At the present time, however, when we find not only much of the available land under cultivation, but even that cut up into small tracts and men attempting to earn a living on ten or twenty acres instead of on eighty or a hundred, the depredations of birds are more noticeable. The loss of a sack of grain is hardly noticeable in a large field, but let the same amount be lost in a two- or three-acre field and the loss becomes relatively important and very apparent. It is only natural, therefore, that at the present time complaints against birds are more frequent and more insistent. 384 University of California Publications in Zoology [VOL. 11 Depredations probably increase also as the natural food sup- ply of birds is destroyed and they have to rely on the products of civilization. The change of food caused by change in environ- ment sometimes causes an increase in the number of birds of a species, and thus increases the extent of the depredations. Ap- parently there are a number of birds in California which have directly profited by the change of environment and are increasing in numbers. The linnet, western meadowlark, and mockingbird undoubtedly belong to this class. As the crops change, and consequently the food supply, we may even expect that in the future the food habits of birds will change. Hence a knowledge of the food habits of birds at the present time may be of far greater value in the future, when such data are needed for comparison. Whether it is best to destroy certain birds because of their depredations, or to preserve them because of their value as insect or weed-seed destroyers, has become a real problem. There is not a farmer who is not at some time of the year confronted with this problem. In a newly settled country the question as to the value of certain birds is often of grave importance. If nature were not so closely woven together we might easily solve the problem by simply exterminating those birds which cause damage. In the early days this was tried. It is experience that has taught us the danger attendant on the indiscriminate ex- termination of any form of life. The problem stated, the next thing to be considered is : How shall it be solved? Observation has proved an unsafe method of determining the true value of a bird. Mere sentiment fails to convince a large number of the class of people deeply interested from actual contact with the problem. The method which has proved the most dependable is a thorough scientific investigation. The problem is complex. It involves a knowledge of the life- history of birds, insects, and plants, a conception of the inter- action of organisms, and an appreciation of the accompanying ecological relations. A scientific investigation as a means of determining the status of a bird presents just as great possibilities as this method has in other fields. In preventive medicine we see the results of 1914] Bryant: Economic Status of the Western Meadowlark 385 scientific method. The efficiency of the forest service can be attributed to the same methods. If, then, this type of investi- gation can help us to conserve our health and our forests, it should also lead to the best method of conserving our native birds. Professor Charles S. Minot has denned the method of science as "the art of making durable, trustworthy records of natural phenomena. ' ' He goes on to say : ' ' The method of science is not special or peculiar to it, but only a perfected application of our human resources of observation and reflection to use the words of von Baer, the great embryologist. To secure relia- bility the method of science is, first, to record, everything with which it deals, the phenomena themselves and the inferences of the individual investigators, and to record both truly; second, to verify and correlate the personal knowledges until they acquire impersonal validity, which means, in other words, that the con- clusions approximate so closely to the absolute truth that we can be safety guided by them." These statements justify the use of scientific method for any modern problem and especially for the problem in hand. "But putting aside economic and utilitarian considerations, there is to some of us a greater stimulus to solve the problems of nature. With the birds, and the insects and plants upon which they feed, we share a common heritage, and the more we learn of the life of these, our fellow-workers, the nearer we ap- proach solution of the great riddle of the Universe, the mysterious law-abiding scheme of Nature. The book of knowledge to which we add some iota is marred with mystery, superstition and error, but each proved fact cleanses its pages. 'Facts,' says Laing, 'are the spokes of the ladder by which we climb from earth to heaven.' " (See Coward, 1912.) The labor, time and cost of such an investigation as this is amply justified by the results to be expected. A knowledge of the real economic status of a bird means dollars in the pocket of the rancher, for the destruction of any bird which causes serious damage, or the preservation of any bird that is a benefit, has a direct bearing on the size of the crop raised. Nor is the value to the rancher the only value to be considered, for, as will 386 University of California Publications in Zoology [^OL. 11 be seen, a bird has a certain value to society that cannot be reckoned in dollars and cents. Some entomologists, seeing in insecticides the only successful control measure against insects, are inclined to minimize the value of such a natural control as birds. True it is that birds apparently do not prevent or entirely control insect outbreaks, for insects continue to ravage crops, no matter what the bird population. However, if a certain number of insects cause a certain amount of damage, it must follow that a diminution of the number of insects causing damage must cause some dimin- ution of the damage done even if it be not proportional. We are justified in saying, therefore, that the fact that birds destroy great numbers of injurious insects shows them to be important agents in contributing to the safety not only of crops, but of all vegetation. Judging from the great numbers of insects destroyed at the time of an insect outbreak, we can safely infer that birds may be instrumental in preventing the appearance of insects in abnormal numbers, by helping to keep the numbers near the normal, which we approximate by the phrase, "the balance of nature. ' ' In this practical age almost everything is viewed from the standpoint of dollars and cents. Hence it is desirable that we study the economic value of birds. There is danger, however, in so doing, for such studies may tend to minimize to a certain degree a value which cannot be expressed in dollars and cents. To say that a meadowlark is worth so many dollars to the rancher each year may obscure its esthetic value. The strongest opponents of the theory that birds are bene- ficial often emphasize the esthetic value. The following is a quotation from one of these opponents (Baskett, 1910) : "Make their song, beauty, grace and interesting habits a part of our culture and their preservation part of our ethics, but do not try to foist them on the farmer as an economic asset, for he knows better in many cases. If the soldier can make better marches under the martial influence of the 'spirit-stirring drum' and * ear-piercing fife, ' so can the farmer gather inspiration from the activity and cheerfulness of birds." 1914] Bryant: Economic Status of the Western Meadoivlark 387 One need only point to the place which birds take in art and literature to prove their esthetic value. The inspiration for some of the finest paintings, poems and other pieces of literature has come from a knowledge of bird life. That exhilaration and inward joy awakened with an acquaintanceship with birds has a real value. What mental pictures stay longer with us than do those gained first hand from nature? Those things which make the world more beautiful make it more fit to live in. Few birds there are that have a greater esthetic value than does the western meadowlark. One reason for this is that it is a conspicuous bird and therefore known to every one. Its song has been pronounced far sweeter than that of its eastern relative. Its plumage and general habits add to its attractiveness. A bird associated with the fields and plains, it adds great interest to the general loneliness and monotony of our great treeless areas. What person traveling along a lonely country road has not been cheered by the bird which stands bobbing on many a fence post and telegraph pole and continually pours forth its "Eh heu wheel'iky, wheel 'iky, wheel'iky"? (For the musical notation of the song of the western meadowlark see Allen, 1881.) The educational value of birds has more utilitarian aspects. Birds teach a code of ethics exceeded only by that of man him- self. The fidelity of parents to each other and to their helpless young and the industry, cleanliness, grace, and cheerfulness exemplified by them add much to the finer ideals of life. The educational and esthetic points of view can even be considered economically. To many, this type of presentation detracts instead of adds. Nevertheless, it is true that the esthetic and educational value of birds has its economic relations. Many a summer resort is chosen because of the abundance of birds in the vicinity, and many a summer vacationist is influenced in his choice of destination by the presence of birds in the vicinity. The value of suburban property is enhanced by the presence of birds. That many a business man has been attracted to certain suburban localities because of the presence of the meadowlark and its song is self-evident. Carrying this point of view to an extreme has often antag- onized certain classes, and herein lies a danger. Yet the facts 388 University of California Publications in Zoology [VOL. 1.1 here presented can easily be verified. A determination of the status of a bird must include a study of the bird from every point of view. To many city folk the esthetic and educational value of a bird is the more important, for they never see it from any other point of view. Perhaps the rancher is an extremist on one side and the city resident on the other. A modification of the views on both sides is very desirable to a sane appreciation of the value of birds in general and the western meadowlark in particular. The advance made in investigations of the economic relations of birds since Professor Aughey (1878) studied the relation of birds to the locust ravages in Nebraska up to the present, when government experts give the whole of their time to such inquiries, demonstrates the growth of the science of economic ornithology. A study of this advance, however, shows that only a beginning has been made. Although we know in general the food habits of our common birds, yet conditions vary so greatly that we cannot definitely predict the food in any given locality. The work thus far has afforded us a general survey of the food habits of birds and in some few instances has given us definite knowl- edge as to the usual food of certain birds. The thing that eco- nomic ornithology has not afforded us as yet is a detailed study of the food of a particular bird in a given locality throughout the whole year. The importance, then, of a thorough knowledge of the eco- nomic relations of a bird in addition to its life-history is evident. It has been left to one of the new sciences, economic ornithology, to tell us of these economic relations and to explain the real status of birds. The agitation coincident with the establishment of this science had made known at least five facts : 1. Birds are very largely insectivorous, and as a result are important in keeping the numbers of insects in check. 2. The amount of food required by birds is enormous. 3. Birds often considered injurious are really beneficial, and vice versa. 4. Birds change their food habits and feed on the kind of food most easily obtained. 1914] Bryant: Economic Status of the Western Meadowlark 389 5. Birds are very important in preserving that balance of nature most suited to the interests of man, and their place can- not be filled by any other class of living things. These facts are now familiar. They have furnished a basis for a sane protection, have demonstrated the intricacy of the interactions of organisms, and have helped develop the economic view of birds. Although economic ornithology is fundamentally the study of birds from the standpoint of dollars and cents, and, therefore, includes their use as food, as cage birds, etc., yet emphasis has rightly been placed on the study of the food of birds. As a result, economic ornithology is most often used in a restricted sense and has reference to the study of the food of birds. Great activity is evidenced in this line of work at the present. Not a month passes that there is not some important contribution to economic ornithology, and there is scarcely an entomological re- port that does not mention the value of birds as insect destroyers. To appreciate the work of the present, however, there must needs be some knowledge of the work of the past. A brief historical review of the subject, with emphasis on the methods used, will furnish this needed information. HISTORY OF METHODS IN ECONOMIC ORNITHOLOGY We need only to examine in detail the progress of our sciences to be convinced that there is such a thing as evolution. As we interpret their progress step by step, and thus survey their gen- eral trend, it would seem that the development has been of the orthogenetic type. The biological sciences have been a little slower than others in their development, but they are now taking front rank. One of the most marked tendencies to be noted in history is that of a change from the period when biologists drew conclusions from facts gained from observation only to the present period when more intensive study and experimental evidence are demanded. The period of time previous to 1850 may be termed the prim- itive period, for during this time we find only an occasional mention of the food habits of birds, the entire time of the workers 390 University of California Publications in Zoology [VOL. 11 in this field having been given over to classification and habit notes. The following period (1850-1865) marks the time when the specialized science of economic ornithology was founded. It was a period of interest and agitation. Before the conclusion of this period the modern methods of investigation were introduced into America by Jenks (1860). The period since 1865 is best considered the modern period a time when the attempt to reach truth is backed by experimental evidence and the work becomes intensive rather than general. (See Locy, 1908.) To the first period belong such men as Catesby, Edwards, Forster, Latham, Bartram, Hearne, and Barton men who took an active interest in natural history and enriched ornithological literature with what observational facts they were able to glean. In the latter part of this first epoch there began a marked tend- ency to gain more than superficial facts by observation, and so in the writings of Wilson, Audubon, and Baird we find mention of the food of birds. (See Palmer, 1899.) It was not till 1860 that Jenks (1860) applied scientific method to the study of the food of birds. Previous to this time there had been considerable agitation concerning the value of birds, and many papers dealing with the question appeared in agricultural journals. Le Baron's "Observations of the Birds of Illinois Interesting to the Agriculturist" is a good type. Other writers at this time were "Walford, Holmes, Kirkpatrick, Dodge, Allen, Elliott, and Samuels. The hour was ripe, there- fore, for economic ornithology really to take a place among the sciences as distinct from ornithology itself. The work carried on by Jenks (1860), Treadwell (1859), Aughey (1878), and Forbes (1880, 1882, 1883, 1903) gave the science its real foun- dation, and inaugurated the modern methods now well exemplified and used by the United States Bureau of Biological Survey. At about the same time we find the science getting a start in Europe. Jenks (1860), when introducing the method of stomach examination into the United States, followed Prevost's (1858) method. M. Florent Prevost was evidently the pioneer in Europe. His paper, "A Memoir on the Alimentary Regimen of Birds," presented to the Imperial Zoological Society of Paris in the year 1858, and translated by Jenks in 1859, should still 1914] Bryant: Economic Status of the WesternMeadowlark 391 be considered a classic. In it he presents an original method of stomach examination and draws some very sane conclusions from the results of thirty years' work. It is interesting to note that this early worker had a vision of the very method which is advocated at the present time, but which has seldom been followed in detail. He says : "It ap- peared that it would be of interest to gather, at different periods of the year, the stomachs of all birds which it might be possible to procure, to examine the contents, to note down the exact results of this examination, with the date of the observation, and to preserve these pieces in order to form, in time, a collection by means of which one can in the future verify each of the regis- tered facts. ' ' He goes on to point out what he had accomplished in thirty years' work and the methods which he used in pre- serving stomach contents. These methods were: drying and mounting on cards, drying and preserving in a vial, and pre- serving in alcohol. His method of examination appears to have been thorough, for he suggests that an "attentive examination," in many cases, made "fragments such as antennae, jaws, lips with their feelers, feet, and often entire heads" give the means of determining the family, genus, and, in some cases, even the species. Following this is a discussion of the results of the work. The tables used are described thus : "To this end I have drawn up a uniform table for all the species of birds ; each copy of this table concerns a species whose name figures at the head. It represents a series of columns, of which each bears the title of an alimentary regi- men ; it is in these columns, and conformably to their title, that I have inscribed both the date of the observation and the indi- cation of the objects found in the stomachs. In fine, each of these tables contains a sufficient number of lines to register observations made during twelve months of the year, and at five different dates in each month." Early workers in America have often failed to consider the food of a bird for the whole year. They have also failed to take into consideration the fact suggested by Prevost in the following words: "The studies which I have pursued after the method indicated above will establish the fact that the same species of 392 University of California Publications in Zoology [VOL. 11 bird changes its food according to the age and season of the year." Also that "the moment when certain insects inundate a country with individuals without number, . . . coetaneously, this very abundance seems to invite a crowd of different species of animals to feed upon them." It was not till 1882-1885, nearly twenty-five years later, that Forbes clearly pointed out these interrelations between birds and insects. Prevost concludes: "I am in the course of proving that birds are in general much more useful than injurious to our crops, and that even in respect to the greatest part of the graniv- orous species the evil which is done to us at certain times is largely compensated by the destruction of insects which they accomplish at other times. It is important, then, that we do not destroy these species, but only divert them from the crops when they injure them. Their destruction would permit, with- out counterbalance, the development of many species of insects more fatal still to agriculture. The study of the alimentary regimen has furnished me also some information which I believe useful in comprehending the reunions, the separations, and peri- odical emigrations which are observed so commonly among birds." (See Prevost, 1858, translation by J. W. P. Jenks.) Since the work of Prevost, economic ornithology has grown rapidly. Germany has probably been most active in the work. Hawks and owls have received the most attention throughout Europe, probably for two reasons. They have been most widely attacked because of their size, and their value is most apparent upon investigation. A number of societies and institutions scat- tered over the continent are actively engaged in studying the economic status of birds. Chief of these are the Kaiserliche Anstalt fur Land- und Forstwirtshaft zu Berlin, Ornithologische Gesellschaft in Bayern, Paris Museum of Natural History, and the Koniglich Ungarische Ornithologische Centrale. The names of Berlepsch, Key, Ouster, Rorig (1903), and Hollrung (1906) have become well known as workers in this field in Germany. Rorig is the one man who has attempted a computation of the comparative amounts of food by a weight method. Csiki (1909) and Greschik (1910, 1911) have been the principal workers in Hungary. Their researches have been mainly confined to the birds of prey. 1914] Bryant: Economic Status of the Western Meadoivlark 393 A committee appointed by the British Association for the Advancement of Science is now investigating the feeding habits of British birds by a study of the contents of the crops and gizzards of both adults and nestlings, and by collation of obser- vational evidence, with the object of obtaining precise knowledge of the economic status of many of the commoner birds affecting rural science. Data as to the environmental conditions under which the bird was feeding and the available food supply are obtained with each specimen. All data obtained from the stomach examination are tabulated, and the weight of the bird and the condition and weight of the gizzard contents are recorded. The United States Department of Agriculture has carried on the most extensive work in economic ornithology ever attempted by one institution. Study along this line was begun in 1885. Since that time over sixty thousand stomachs of birds have been examined, and the results, with the addition of data collected in the field, have been published in more than one hundred and thirty bulletins. Other investigations have also been conducted. The most extensive work has been done in Illinois by the pioneer economic ornithologist, Professor S. A. Forbes. Massachusetts, Wisconsin, and Pennsylvania have also carried on investigations, the work being done by Forbush (1908), King (1883), and Warren (1888) respectively. Practically every state has been supplied with some literature on the subject by the state university or the agricultural experiment station. At the third ornithological congress at Paris in 1900, the section of economic ornithology and bird preservation reported in favor of urging all countries and even their governments to take up seriously the subject of the utility or harmfulness of birds as being of the greatest economic importance. It was urged that "inquiries should be instituted on regular business lines, that migratory and non-migratory species alike should be observed during every month of the year and for several years in succession, that the contents of their stomachs should be care- fully noted, and lists prepared of their action towards the farmer's crops." The amount of careful work along these lines which has been done since this time is very encouraging. The last few years 394 University of California Publications in Zoology [VOL. 11 have seen a number of the leading magazines take up the subject of the economic value of bird life, and with illustrated articles they have brought to the attention of many the value of birds to the farmer. Largely because of their depredations, the demand for an intimate knowledge of the food of birds has become very pressing, and yet it is interesting to note that attention to the economic side of ornithology was not aroused by the depredations of birds, but by the depredations of insects. A comparison of the methods used up to the present will clearly show the progress which has been made in the science of economic ornithology. Such a comparison is afforded by the following sequence of methods used in determining the economic value of birds: Sequence of methods used in determining the economic value of birds : 1. Observational notes on the food of birds. (Wilson, 1808- 1814; Audubon, 1827-1838.) 2. Critical observational study of the food of birds. (Le Baron, 1855; Holmes, 1857; Weed, 1903.) 3. Examination of the stomach contents of birds. (Jenks, (King, 1883.) 4. Experimental feeding of captive birds. (Treadwell, 1859.) 5. Observation plus stomach examination. (Aughey, 1878; Judd, 1902.) 6. Observation plus stomach examination plus experimenta- tion. (Forbes, 1903 ; U. S. Biological Survey.) Similar progress can be noted in the methods used in de- termining the food of birds. Their sequence has been as follows : 1. Investigation of food with no reference to time or locality. (King, 1883.) 2. Investigation of food at time and locality of depredations. (Forbes, 1903; Wilcox, 1892; Aughey, 1878; Bryant, 1911, 1912d.) 3. Investigation of food according to the month, regardless of exact locality. (Jenks, I860; Beal, 1907, 1910.) 19M] Bryant: Economic Status of the Western Meadowlark 395 4. Investigation of food according to the month in the same locality. (Forbes, 1903; Bryant, 1912a.) 5. Investigation of food according to the month in the same locality, with a comparison with many different localities. (See p. 454.) The determination of the economic status of birds has like- wise progressed. The sequence of the criteria used has been as follows : 1. Inferential evidence. 2. Circumstantial evidence. 3. Number of injurious insects eaten. 4. Proportion of percentage volume of injurious, neutral, and beneficial insects and seeds destroyed. 5. Contrast of all harm vs. all good, including knowledge as to life-history. From these comparisons it can be seen that great progress has been made. To infer that a bird is injurious simply because it is seen in a grain field or orchard, or to brand it as injurious because of circumstantial evidence in the form of grain or fruit found in the stomach, are obsolete methods today. Furthermore, we recognize at the present time that a bird may eat some bene- ficial insects and still be a valuable bird. Nothing less than a knowledge of the food for the whole year, combined with a knowledge of the life-history of the bird concerned, allowing a balance of all the benefits conferred with all the damage done, meets the requirements of the present. INVESTIGATION OF THE ECONOMIC STATUS OF THE WESTERN MEADOWLARK IN CALIFORNIA Interest centered around the meadowlark for some time pre- vious to the institution of an investigation. A rather dormant complaint against the depredations of the meadowlark in sprout- ing grain fields was brought to a head in a bill (no. 229) intro- duced by Assemblyman Stuckenbruck of San Joaquin County 396 University of California Publications in -Zoology [VOL. 11 into the State Legislature on January 11, 1909. The bill, which proposed to amend section 637 of the penal code of California, passed through the committee, but was refused passage, the vote standing 32 to 28. On the motion to reconsider, the bill was again brought to a vote and passed with a vote of 41 to 28. The Committee on Fish and Game of the Senate reported favorably on the bill, but it was refused passage on a vote of 17 to 12. In 1911 Assemblyman Stuckenbruck, at the request of his constituents, introduced a similar bill with the proviso that in the counties of Tehama, Butte, Sutter, Sacramento, Yolo, Colusa, Glenn, San Joaquin, Stanislaus, Tulare, and Kings the meadow- lark be not included among the birds protected by the act, hoping thus to allay the opposition met from other parts of the state at the former session of the legislature. This bill, being referred to the Committee on Fish and Game, was returned to the Assem- bly with a majority report in favor of its passage and a minority against its passage. It failed of passage on March 20. Continued complaints from the farmers and fruit growers of the state have been made to the State Fish and Game Com- mission regarding the losses to crops caused by the depredations of birds. The commission has been repeatedly urged to take strong measures to avert the damage done. The usual measure urged is that the particular bird in question should be placed on the unprotected list. On the other hand, many scientists and others interested in birds have pointed out the fact that birds confer a great benefit in keeping down the number of injurious insects and weed seeds, and thus they fill a niche in the economy of nature most suited to mankind which is not and can not be filled by any other form of life. Experience has shown that many belonging to the first class have based their complaints on circumstantial or partial evidence or on evidence not sufficiently reasoned out. Furthermore, these complaints have brought out the fact that really very little is known of the food habits of birds of California. Certain it is that a knowledge of the food habits of a bird is necessary to a determination of its economic status. As a result, therefore, the commission thought it wise that legislation should be based on scientific investigation as to the value of birds, and not on circumstantial evidence. Conse- 1914] Bryant: Economic Status of the Western Meadowlark 397 quently an investigation into the relations of the birds of the state to agricultural and other interests was instituted. The institution of the investigation was largely due to the interest and energy of Mr. John P. Babcock, Chief Deputy of the California State Fish and Game Commission, 1910-11, and Professor Charles A. Kofoid of the Department of Zoology of the University of California. A COMPARISON OF METHODS IN ECONOMIC ORNITHOLOGY The attempt to show the amount of the different kinds of food contained in the stomachs of birds has led to the use of two distinct methods, both of which must be considered valuable, and both of which approximate the end sought. A method introduced by King (1883), and later used by Newstead (1908), gives the total number of birds taking the different kinds of food compared with the total number of stomachs examined. The second method, employed by the United States Biological Survey, depends entirely upon the comparative volume of the different kinds of food found in the stomach, calculated in per cent of total volume and averaged. A third method, in which actual counts of the insects found are made, has been used in a few instances (Mason and Lefroy, 1912; Fisher, 1893). Workers in this field in Great Britain have used the numerical system almost entirely, depending for a criterion upon the number of birds taking a certain kind of food. Since all of these methods appear to furnish certain infor- mation not furnished by the others, a combination of all three methods has been used in this investigation. Dependence is laid on the first method for an idea of the percentage of birds of a species feeding on a particular insect, on the second for an idea of the comparative amounts of the different kinds of food taken by individuals and by the species, and on the third for an idea of the actual numbers of the different elements of food. The counting of weed seeds and insects found in the stomach of a bird is difficult and fruitful of error. Yet the fact that smaller 398 University of California Publications in Zoology [VOL. 11 numbers are always counted than truly exist, owing to the com- minuted condition of some, makes it evident that no exaggeration is possible here. Consequently it affords dependable evidence as to the numbers of weed seeds, insects, etc., taken by a bird. On the other hand, the percentage-of-volume method can be depended upon only to furnish an idea of the comparative quantities of the different kinds of food. Personal error in estimating has to be allowed for in this method, for whereas a certain insect might be to the eye of one person ten per cent of the volume, it might represent fifteen per cent to the eye of another. The furnishing of complete data as to the bird whose stomach is examined (date, locality, kind of field, collector, etc.) should afford information, first, as to the variation in the amount of food taken by birds during the day, month, and year, and second, the food preference of birds in a given locality and in different localities. The record of the exact time of day, the month, and year when the bird was collected furnishes the basis for the first, the record of the habitat, as, for example, the kind of field, orchard, or vineyard, the basis for the second. An attempt has also been made to improve the method used in determining the economic status of a bird. As has already been pointed out, the economic status of a bird was originally determined by inference. A bird in the grain field must be eating grain and therefore is injurious. Experience has taught that such reasoning is fraught with error. And further exper- ience has taught us that even though a bird may cause consid- erable damage, yet because of its usefulness as a weed-seed destroyer, as an insect destroyer, or as a bird important in keep- ing the balance in nature most suited to man, it may be more beneficial than harmful. At one time the total good accom- plished by a bird was held to inhere in the number of injurious insects it destroyed. Today, although we still retain this idea, we see a little further and conclude that a bird may be beneficial because it destroys insects (almost all insects being potentially destructive), and not because it chooses a particular class of insects arbitrarily classified as harmful by man. An attempt has been made to arrive at the average volume of food taken by the meadowlark, by determining the volume 1914] Bryant: Economic Status of the Western Meadowlark 399 of food contained in a large number of stomachs in cubic centi- meters and taking an average volume. This allowed the record- ing of each stomach as being of average volume, over the average, or below the average. Identification of the various insects and weed seeds found in the stomachs has been difficult. Help from the experts of the United States Department of Agriculture, from the Department of Entomology of the University of California, and from others has facilitated greatly the identification. Certain field work, embracing studies of the abundance of birds, the depredations of birds, nesting habits, the relation of the birds to insect out- breaks, the kind, amount, and availability of food, and the time of digestion, has afforded needed supplementary information. In addition to the importance of this investigation to agri- cultural interests, it has been fruitful of valuable data from the standpoint of science. Although the investigation has been car- ried on primarily to furnish practical information as to the exact relation of the western meadowlark to agriculture and horticul- ture, yet no pains have been spared to collect data of purely scientific interest. The importance of a knowledge of life-histories has been emphasized only of late. Information as to the food of any form of life constitutes one of the most far-reaching phases of its life-history. As Forbes (1903) pointed out: "Since the struggle for existence is chiefly a struggle for sub- sistence, a careful comparative account of the food of various competing species and genera, at different places and seasons and at all ages of the individual, such as has not heretofore been made for any class of animals, cannot fail to throw much light upon the details, causes and effects of .this struggle. The flexi- bility of the food habits of the widely ranging species, the direct effects of normal departures from the usual average of food elements upon the origin of variations, and the general reactions of birds upon their organic environment, are examples of subjects upon which light should be thrown by this investigation." In this investigation the difference in food habits of the nestling and adult has been clearly demonstrated by the exami- nation of a large number of specimens. The difference in the kind and amount of food taken by the two sexes is made available 400 University of California Publications in Zoology [VOL. 11 for the first time. Field investigation has been fruitful of an increased knowledge of the general habits, nesting habits, abun- dance, depredations, and distribution of the western meadowlark. The handling of so large a number of specimens taken from all parts of the state and during each month of the year has furnished information as to variation, albinism, parasitism, and malformation. Critical evidence as to the value of certain so- called protective adaptations of insects has also been afforded. The investigation of the relation of birds to insect outbreaks has emphasized their importance at such times and furnished critical evidence as to the interrelations of these organisms. THE WESTERN MEADOWLARK (STURNELLA NEGLECT A) In spite of its name, the western meadowlark is not a true lark, but belongs to the family Icteridae along with the black- birds and orioles. It is easily recognized by its medium size, gray- and brown-streaked back, brilliant yellow throat, black V-shaped collar, and its conspicuous white outer rectrices. The meadowlark is widely distributed over North America. The eastern meadowlark (Sturnella magna magna) differs from the one found in the west in size, color, and song. The western meadowlark is slightly larger than the eastern bird, is paler in color, and has a much richer song. For these reasons the western form is considered a distinct species and is called the western meadowlark (Sturnella neglecta). It is found from Wisconsin, Illinois, Iowa, Texas, etc., west to the Pacific Coast, and from central and western Mexico to British Columbia and western Canada. It is to be found through- out the State of California from sea level to 7000 feet elevation in the mountains. The western meadowlark is resident throughout the year. A slight altitudinal migration perhaps takes place, governed largely by the available food supply, but usually the bird is to be found in the same general locality throughout the year. 1914] Bryant: Economic Status of the Western Meadowlark 401 The western meadowlark is a conspicuous bird of treeless areas and a frequenter of meadow, pasture, or uncultivated grass land. Although a poor flyer when compared to some birds, the meadowlark, with its peculiar hovering flight, is possessed of a method of locomotion sufficient for its needs. Its mode of life necessitates but a small amount of flying. During the fall and winter months meadowlarks gather in flocks of five to fifty or more. During the spring, however, they are seen singly or in pairs. Of a nervous temperament, they are wary and do not often allow of close approach. Both the male and female are good singers. Their cheerful and varied song is sometimes given from mid-air, but more often from a fence post, shrub, or clod. The western meadowlark appears to be one of the few birds which is profiting by the increased cultivation of land. Alfalfa furnishes particularly good food and cover for the bird, and grain fields are often chosen for a home. With the furnishing of still more good food and cover, combined with the destruction of some of its enemies, this bird may be expected still further to increase in numbers. The western meadowlark feeds almost exclusively on the ground. It seldom perches in a tree of any kind. The early morning hours are spent in obtaining food, whereas the middle of the day is usually spent quietly hiding in the grass. The food, composed largely of insects, grain, and weed seeds, is procured not only from the top of the ground, but also by probing beneath the soil and by searching under clods, manure, etc. Alfalfa and grain fields appear to be the favorite feeding grounds of these birds in cultivated districts. FIELD INVESTIGATION Next to the knowledge of the food of a bird in determining its economic status is a study of the bird at the scene of action, or, in other words, a knowledge of the habits of the bird. Evi- dence along this line can be afforded only by field investigation. Over a month's time was spent at Lathrop, San Joaquin County, California, studying the abundance, feeding habits, nest- 402 University of California Publications in Zoology [VOL. 11 ing habits, depredations, etc., of the western meadowlark. As has been stated, this particular locality was chosen because it afforded not only an abundance of birds, but also a favorable proportion of cultivated and uncultivated land, thus allowing a study of food preference. The field work carried on can be grouped under three heads : studies of the abundance, of the habits, and of the depredations. Studies of the relations of birds to insects and to insect outbreaks have also been included in the field investigation, but will be discussed in another place. (See p. 456.) ABUNDANCE OF THE WESTERN MEADOWLARK Several findings in connection with the field work have tended to minimize somewhat the depredations of the meadowlark. Per- haps one of the most important is the preference which the bird shows for uncultivated land. Censuses have absolutely demon- strated that, during hours of feeding, western meadowlarks are more abundant in pasture land than in cultivated fields. Evi- dence as to the abundance of the western meadowlark is also afforded by the censuses. The following are censuses taken in the vicinity of Lathrop, San Joaquin County, on a two and one-half hour drive : Uncultivated Cultivated ( Pasture ) ( Grain and alfalfa ) February 28, 1912 158 65 May 31, 1912 69 27 These two censuses covered practically the same length of time and the same territory. As the birds were seen in flocks in February, it is only natural that more birds were recorded. May being in the nesting season, the birds were then more widely scattered and not so easily seen. It is apparent that the meadow- lark prefers uncultivated land even at the time of feeding. A similar census taken at Acampo, San Joaquin County, during an hour's walk (2-3 P.M.), resulted as follows: Number Orchards, vineyards, and pasture 25 Grain fields ... 2 1914] Bryant: Economic Status of the Western Meadowlark 403 Meadowlarks were more abundant in the vicinity of Lathrop than in the vicinity of Acampo. There is very little pasture land in the vicinity of Acampo. This, if not the main reason, is one of the important ones which account for this contrast in abun- dance. Another contrast in abundance can be noted from the following censuses, two taken at Los Banos, Merced County, July 11, 12, 1912, and the other at Merced, Merced County, July 17, 1912. Those at Los Banos were taken while walking less than five miles, and occupied about four and three hours respectively. The one at Merced was taken while driving about fourteen miles, and occupied the time between 1 :30 and 5 o'clock. Meadow- Average Locality Date Time larks seen per acre Los Banos July 11, 1912 3-4:30 36 1.0 Los Banos July 12, 1912 2-5:15 67 1.0 Merced July 17, 1912 1:30:5 23 .1 It is evident, therefore, that the abundance of this species of bird is largely affected by locality. Probably in the last analysis food supply is the important factor. This brings us to the question : Can the western meadowlark obtain its natural food in cultivated fields as easily as in the uncultivated? If we con- sider insects as vegetable feeders, then we should expect to find the best insect supply where plant growth was most luxuriant. The cultivation of land destroys much of the natural plant growth, and therefore must diminish the food supply of the insects enough to vary the abundance. Grasshoppers, cutworms, and wireworms can usually be found more abundant in grassy pasture land than in orchards or grain fields. Hence it is a natural consequence that we find meadowlarks frequenting un- tilled land more often than tilled land. The censuses taken also, demonstrated the fact that meadow- larks were found in the pasture land in greater abundance during the middle of the day than in the morning hours of feeding. During the hotter periods of the day these birds hide in the grass. Open fields are seldom chosen at this time of the day. 404 University of California Publications in Zoology [VOL. 11 NESTING HABITS The nesting season of the western meadowlark lasts from March to August. The nest is a well-concealed one, built of dry grasses usually in grass, alfalfa, or grain fields, in a de- pression in the ground. A canopy of dry grass stems usually arches the top of the nest and a runway two to five feet long leads to the nest. Ofttimes this runway is the only clue to the location of the nest. The female bird does most of the work of incubation and feeding of the young, while the male acts as a guard. Eggs are usually five and are white, variously marked with brown, purple, and lavender spots and lines. Work in the spring of 1911 and 1912 substantiated the fact that western meadowlarks usually nest twice each year. The first nesting usually occurs in April and May and the second in July and August. Probably on an average not more than three young are in a brood, although the number of eggs laid is usually five. Second nestings examined usually show an incom- plete set of eggs. A preference for pasture land for nesting sites was shown, at least eighty per cent of the nests found being so situated. The time of incubation was found to be twelve to fourteen days. The young stay in the nest but a short time, eight to ten days. Nestlings are exposed to many enemies, such as skunks, weasels, rats, and hawks, and the number of broods successfully reared is less than that of most other birds. That over ten per cent of the nests in most localities are destroyed by predacious animals and birds seems a very conservative estimate. These facts have an important bearing on the economic rela- tions of the meadowlark. Proximity of breeding grounds to cultivated crops naturally has an influence on the amount of damage done. The rate of reproduction influences the amount of damage, owing to the number of individuals to be expected in any locality. DEPREDATIONS OF THE WESTERN MEADOWLARK No small part of the field work has consisted in investigations of the damage caused by the western meadowlark. In most cases the field work has been supplemented with stomach examinations. 1914] Bryant: Economic Status of the Western Meadowlark 405 The principal complaint lodged against the western meadow- lark has been that this bird destroys sprouting grain. The field investigations have proved that this complaint has a real foun- dation. An interesting side-light on this habit is afforded by a paragraph from Coues's (1874) "Birds of the Northwest": "In April, before pairing, hundreds used to frequent daily the parade ground of Fort Randall, where, as the grass was yet scarcely sprouted, good opportunity was offered of observing their characteristic habit one not so generally known as it should be, since it is related to the peculiar shape of the bill. The birds may be seen scattered all over the ground, busily tugging at something; and on walking over the scene of their operations, the ground, newly softened by the spring thaw, is seen to be riddled with thousands of little holes, which the birds make in search of food. These holes are quite smooth not a turning over of the surface of the ground, but a clean boring, like that made by sinking in the end of a light walking stick; just as if the birds inserted the bill and then worked it about until the hole was of sufficient size. Whether they bored at random, or were guided by some sense in finding their prey, and what par- ticular objects they were searching for, I did not ascertain ; but the habit was so fixed and so continually persevered in as to attract general attention." This habit of boring into the ground to obtain sprouting seeds and possibly insects is therefore a habit of old standing, and is not one recently developed. A careful investigation of a sprouting grain field where meadowlarks are abundant will demonstrate to any one that the western meadowlark pulls sprouting grain. At times the drill row is followed for distances of four to six feet and apparently every sprouted kernel is pulled up (pi. 21, fig. 1). With its long awl-like bill, the meadowlark bores down beside the sprout, grips the kernel and pulls it up. The kernel is occasionally eaten, but more often it is simply crushed in the bill to obtain the milk and then dropped (pi. 21, fig. 2). Consequently stomach examination cannot be relied upon to furnish accurate evidence as to the total amount of grain thus destroyed. Certain fields examined have given evidence that the deeper furrows made by drills w r ere most frequented by the birds. 406 University of California Publications in Zoology [VOL. 11 Whether this was due to the better cover or to the ease with which the sprouting grain could be obtained it is impossible to state. Where meadowlarks are very numerous and the field of grain small and isolated, considerable damage results. In some cases such fields have had to be resown. Since the loss of a small patch of grain means far more to the small farmer than does a larger amount of grain to the large rancher, the bird's depre- dations here are important. The investigations have shown, however, that fields apparently badly damaged by meadowlarks when the grain was sprouting yielded the usual crops at harvest time. This can be accounted for in this way: Jhe birds can succeed in pulling the grain for only a short period of time after it appears above the ground. By the time the second and third leaf appear the plant is well enough rooted so that the loss of the kernel, even if it should be removed, would not injure the plant. Consequently the apparent devastation is largely mini- mized by the further sprouting of other kernels and the successful survival of the sprouts of many of the kernels removed. A certain amount of thinning may at times indeed be desirable. The lack of uniformity in the thinning accomplished by meadow- larks is, however, an argument against them. Broadcasted grain, unless harrowed in very deeply, suffers more than drilled grain. It is the universal verdict of the grain ranchers of the state that deeply drilled grain suffers less than the shallowly drilled or the broadcasted grain. Experiments have shown that the greatest yield comes from drilled grain. The following table is from University of California Publi- cations, Agricultural Experiment Station, Bulletin 211: KELATIVE KESULTS FROM DRILLED vs. BROADCAST SEEDING UPON THE YIELD OF GRAIN Average of 22 trials Barley Wheat Drilled 70.80 34.85 Broadcast 64.43 31.60 6.37 bu. 3.25 bu. Percentage increase 9.9 10.3 Money value $3.18 $3.12 1914] Bryant: Economic Status of the Western Meadowlark 407 The proper depth for grain to be planted in sandy soil is three to four inches. If all grain were planted at this depth very little damage would be possible, for meadowlarks are unable to bore more than two and one-half inches at most. Holes meas- ured average about an inch and a half in depth. Margins of fields bordering pasture land usually suffer most. Ofttimes a noticeable difference in the amount of grain growing along the edges of fields can be attributed to the work of meadow- larks. Owing to the concentration of large numbers of meadowlarks on a single field, fields of grain planted early suffer most from the depredations of these birds. Grain sown late in the year suffers much less, for. insects become available in small numbers and there is less concentration of damage, due to the larger amount of available food. Meadowlarks are more able to obtain grain planted in sandy soil. Their ability to bore deeply into soil after kernels of grain varies directly with the hardness of the soil. Hard, dry, adobe soil precludes attack. Sandy soil, especially after being softened by a rain, is easily manipulated to advantage. An apparent preference of the birds for oats has been shown not only by field investigation, but by the complaints of the ranchers also. Probably its availability is a greater factor than any preference shown by the bird. That less damage is possible to wheat and barley because the kernel is more easily removed without damage to the plant is one theory proposed. Certain it is that there is a difference in the damage to adjoining fields of oats and barley. Beyond an occasional instance of meadowlarks pulling sprout- ing garden seeds, the only other complaint of importance is that they destroy melons by boring holes in them. Most of the com- plaint has come from the San Joaquin Valley, and especially from Delano and Pixley, Kern County. Melon growers, although admitting that meadowlarks cause considerable damage to melons, have been unable to demonstrate the actual damage in the field. Opinion is divided as to whether they cause any damage. Some prominent growers affirm they are never troubled. Others com- plain of a considerable loss. For instance, the names of two 408 University of California Publications in Zoology [VOL. 11 melon growers in the vicinity of Dinuba, Tulare County, were handed in as those of men who were greatly troubled. One re- turned a verdict of "not guilty," and the other reported that the damage was not very great. All of the growers report that as soon as there are broken melons in the field the birds cease to be troublesome. Apparently the depredations of meadowlarks on melons have been exaggerated. There is evidence to support the view that meadowlarks bore into the melons to obtain water. Whether or not they are at- tracted by the sweet taste we cannot say. The placing of water in a field as an experiment would doubtless confirm or disprove this view. An occasional complaint that meadowlarks are injurious to grapes has been received. Inquiry in grape-growing sections of the state has led to the conclusion that such damage is negligible. A number of birds are destructive to grapes, chief of which are the oriole and grosbeak. Both of these birds are well known as fruit eaters. The meadowlark, on the other hand, seldom turns its attention to fruit of any kind. No damage to grapes caused by the meadowlark has been noted in the field. Investigations of the damage caused by meadowlarks has led to the following conclusions: 1. The western meadowlark is destructive in sprouting grain fields, because of its habit of drilling down beside the sprout and pulling up the kernel. The amount of damage done is de- pendent on the particular location, the abundance of the birds, the character of the soil, the time of year, depth and method of planting, and the kind of grain. The damage to oats is greatest, wheat suffers considerable damage, whereas barley suffers but little. Broadcasted grain suffers more than drilled, because not being sowed so deeply it is more easily obtained by meadowlarks. The birds often follow the drill row and pull almost every kernel. Occasionally, where meadowlarks are very numerous and the quantity of grain small, fields have had to be resown. The real amount of damage done has evidently been overestimated, for fields apparently badly damaged have given the average yield later in the year. After the second and third leaf appears on the grain, the bird can do little damage. This fact reduces the 1914] Bryant: Economic Status of the Western Meadoivlark 409 duration of their depredations to less than two weeks, and conse- quently minimizes the amount of destruction possible. Deep planting and drilling as against broadcasting are important as measures for protecting crops. 2. Damage to other cereals, such as corn and maize, and to fruit is negligible. 3. Investigation of complaints that meadowlarks are destruc- tive to melons has shown that damage caused in this way has been exaggerated. Melon growers, although claiming that the birds cause considerable damage, have often been unable to demonstrate the actual damage in the field. 4. Censuses have demonstrated that the western meadowlark prefers grass land to cultivated land, nearly forty per cent more birds being found in the former. EXPERIMENTATION ON CAPTIVE BIRDS Experimentation on captive birds as a means of determining food preference has been suggested by Forbes (1903) and Judd (1901). No doubt such experimentation furnishes considerable evidence as to the food preference of the bird if carried on with proper controls. Thus far this sort of experimentation has not furnished dependable generalization as to what the bird would have taken under natural conditions (McAtee, 1912). This does not mean that better devised and controlled experiments would not furnish dependable evidence. The difficulty of keeping in cages birds with the temperament of the western meadowlark and the difficulty of procuring for them proper food has prevented the use of feeding experiments in this investigation. In order properly to estimate the quantity of food consumed daily it has been necessary to determine the time of digestion. It was in this determination, and in the determination of the quantity of food, that experiments on captive birds became of value. There are four methods of determining the quantity of food required by young birds. First, the quantity of food carried 410 University of California Publications in Zoology IT L - U to the young by the parents may be observed ; second, the stomach contents may be examined and the quantity estimated; third, experimental feeding of caged birds may be used; and fourth, the quantity of food may be determined by a daily weighing of nestling birds and of their excreta. The first method, owing to the difficulty of observing in the field the feeding of the young of so shy a bird as the western meadowlark, has been largely neglected in the interest of the other more practical methods. However, some observations as to the number of trips to the nest made with food have been made. In one instance a female western meadowlark carried food to the nest three times in twenty minutes (6:15-6:45 A.M.). As the presence of the observer caused some nervousness on the part of the parent birds, this cannot be considered the normal rate. AMOUNT OF FOOD REQUIRED BY WESTERN MEADOWLARKS Stomachs of nestling western meadowlarks examined con- tained as high as two grams of insect food. Maxima of seven large cutworms, of twelve grasshoppers (three-quarters of an inch in length), and of eight beetles have been found in the stomachs of nestlings. One stomach contained twenty-four ants and parts of a ground beetle. The volume of nestling stomachs and of their capacity in terms of the common elements of food follows : Volume of average cutworm 5 c.c. Volume of average ground beetle 3 Volume of average grasshopper 75 Volume of average stomach of $ western meadowlark 3.00 Volume of average stomach of $ western meadowlark 2.50 Capacity of average <$ stomach in cutworms 6 Capacity of average $ stomach in cutworms 5 Capacity of average stomach in ground beetles 10 Capacity of average $ stomach in ground beetles 8 Capacity of average $ stomach in grasshoppers 4 Capacity of average $ stomach in grasshoppers 3 A nestling western meadowlark after obtaining no food for three hours was fed twenty-eight small grasshoppers (one-half inch in length), equal in volume to about three cubic centimeters. Another one was fed four grasshoppers (one inch in length), twelve small grasshoppers (one-half inch in length), one robber 1914] Bryant : Economic Status of the Western Meadowlark 411 fly, one beetle, and five ants. A third one was fed thirty grains of wheat inside of ten minutes. Weighings of nestling birds demonstrated the fact that they gained very nearly seven grams (0.6 ounce) in weight daily. Solid excreta averages 0.48 gram. The weight of excreta voided ' during twenty-four hours must be near 3.6 grams, thus making the weight of food required daily over ten grams. This com- putation does not take into account the weight of excretory products given off through the skin or the weight of carbon dioxide given off through respiration. The ratios of solid, liquid, and gaseous excreta are not known. The tabular results of weighings follow: WEIGHINGS OF NESTLING MEADOWLAKKS Average weight of 2 western Time of weighing meadowlark nestlings Nest No. 1 Nest No. 2^ .75 oz. 1.25 1.50 1.75 .75 oz. .50 oz. 2.9 1.00 .75 2.0 2.25 1.50 1.25 1.75 1.50 Weight of egg ready to hatch 135 oz. Weight of day-old nestling 25 Weight of eight-day-old nestling 2.50 Weight of average adult 4.00 Although more experiments are necessary to establish the exact gain in weight of nestling birds, yet these experiments have furnished evidence as to the enormous quantities of food con- sumed by nestling birds. When one considers that there is a gain of about three ounces (93.3 grams) in weight inside of two weeks, and that this added weight must be calculated by the weight of food consumed minus the waste thrown off in the various forms of excreta and expired air, the quantity of food necessary is evident. Not only is enough food needed to maintain energy, but an additional amount to maintain weight increment is demanded during the period of growth. Probably each young Date May 26 Nest No. 1 6:00 a.m. Nest No. 2 May 27 8:30 a.m. May 28 8:40 a.m. May 29 May 30 May 31 7:15 a.m. 7:45 a.m. 4: 25 p.m. 5:45 p.m. 5:00 p.m. June 1 June 3 7: 30 a.m. 6:30 a.m. 6:15 a.m. 412 University of California Publications in Zoology [VOL. 11 western meadowlark consumes something like five ounces (155.5 grams) of food during the time it remains in the nest. This weight in grasshoppers would mean 311 individuals and in cut- worms 415 individuals. The time of digestion of the western meadowlark was deter- mined by feeding captive juvenile birds and examining the con- dition of the food at intervals after feeding. The following table gives a summary of the results of these experiments : TABULAR EESULTS OF EXPERIMENTS ON THE TIME OF DIGESTION Time Time between Exper- without feeding and Food given Condition of food iment food killing on examination 1 3% hrs. 2 hrs. 4 large grasshoppers Finely comminuted (Camnula pellucida) and largely digested 12 small grasshoppers 1 robber fly 1 beetle (Coniontis sp.) 5 ants 2 5% 3% 20 grasshoppers About one-quarter of (Camnula pellucida) volume remained; all 10 ants soft parts digested 3 3^ 2 30 kernels wheat 15 kernels left undigested; hulls still undigested 4 3% 2 1 May beetle Only hard parts (Ligyrus gib~bosus} left in stomach 1 weevil (Bhygopsis sp.) (heads, wing-covers 12 grasshoppers knee joints, etc.) (Camnula pellucida) 5 4 5? 28 grasshoppers Stomach empty From these data it can be safely concluded that insects are digested in two to four hours and that the stomach is completely emptied every four hours. Beetles and ants, owing to the chi- tinous parts, remain longer in the stomach than do grasshoppers. Cutworms doubtless are digested much more rapidly. Grain is more difficult to digest than insects and remains in the stomach longer (four to five hours). These results compare very favorably with the results of similar experiments by other investigators. Experiments carried on by Treadwell (1859), Forbush (1907), and Weed and Dear- born (1903) have demonstrated that birds have a very rapid 1914] Bryant: Economic Status of the Western Meadoivlark 413 digestion, most of them requiring from one to four hours only to digest a meal. All evidence of food had disappeared from the excreta of a crow within two and one-half hours after feeding. Professor Treadwell showed that juvenile robins digested a meal every two to four hours. All evidence, therefore, points to the fact that four hours can be considered a sufficient period of time to assure the diges- tion of the stomach contents of a western meadowlark. The contents of the stomach at the time the bird is killed must have been taken within four hours previous to the collection of the bird. The daily consumption must, therefore, be considered about three times the capacity of the bird's stomach. As the birds start each day with an empty stomach and with the addi- tional stimulus of hunger, the greater amount consumed during early morning hours compensates for the smaller amount taken during the middle of the day. EXAMINATION OF THE STOMACH CONTENTS COLLECTION AND PRESERVATION OF MATERIAL Birds in sufficient numbers to furnish reliable data, collected every two weeks during a year, and from over twenty different localities in the state, have been made available through the co- operation of the deputies of the Fish and Game Commission. Each bird was tagged with data as to date, time of day, locality, kind of field or orchard, and collector. Dependable data re- garding abundance of food were not available as a general rule. The birds were then preserved in formalin. On the arrival of shipments at the laboratory the stomach (gizzard) was removed and data as to the species and sex of the different birds added. The tag bearing complete data was then wrapped with the stomach in a small cloth, and preserved in ten per cent formalin until microscopically examined. In determining the localities where collections have been made an attempt was made to select well-settled parts of the state representative of the different agricultural sections. The fol- lowing instructions were sent to each deputy: 414 University of California Publications in Zoology [VOL. 11 INSTRUCTIONS TO COLLECT BIRDS FOR SCIENTIFIC INVESTI- GATION OF THEIR RELATION TO AGRICULTURE DEAR SIR: In order to obtain material for our field investigation of the relations of game and other birds to agriculture, we propose to collect specimens of all the field and orchard birds in the State other than quail, ducks, geese, and crows during the first and third week of each month. We want especially one-half dozen each of meadowlarks, robins, and blackbirds collected during these weeks in grain fields and the same number in vineyards or orchards in sections where there are both. We want, also, two doves from the same section for each of the two weeks mentioned. For the present we will confine our attention to meadowlarks, horned-larks, robins, and blackbirds. Where practicable obtain specimens of each species of your district from the same field or fields of a like character near by, because we wish to show just what they feed upon throughout the year. As each bird is killed fill out one of the string record tags furnished you, being careful to note the hour, date, character of field, orchard or vineyard, the location and the name of the owners and the nearest post- office. Then securely fasten a tag to each bird. In writing upon the tags use plain lead pencils only, as the preserving fluid will destroy ink and indelible pencil marks. After filling out the tag, attach it to the bird and then record in a notebook devoted to this work a similar record to the one on the tag. Upon returning from the field proceed to preserve the specimens as follows: From any drug store purchase a 40 per cent solution of formalin or formaldehyde. Then place in an ordinary two-quart fruit jar one-half pint of the 40 per cent formalin and then fill up that jar with clear water and mark it "Jar No. 1." Then place your specimens in another jar (or jars) and fill it with the liquid from "Jar No. 1," and seal the top to prevent evaporation. Don't place too many specimens in a jar; give them room enough to become thoroughly saturated. Keep the specimens in the solution, for at least a week. See that the tag of each specimen is uninjured. At the end of each month take your specimens from the solution and wrap in a cloth wet from the solution in "Jar No. 1" and place in a thin cracker box or a tight wooden box and ship as hereafter directed, together with a copy of the record from your notebook. The solution from which you remove your specimens can be used several times, supplying the necessary additional solution from ' ' Jar No. 1." Recember that birds have no value unless each is securely tagged with a record of the hour, date, etc.; that our work depends absolutely upon the accuracy and reliability of the record. Send separate bills covering your purchase of shot, formaline, jars, traveling and other expenses con- nected with this work to this office. Do not include them on your regular bills. The necessary permit to take these birds throughout the year is fur- nished you herewith. 1914] Bryant: Economic Status of the Western Meadowlark 415 Inform your local press and others interested just what you are doing and the object we have in making this collection, to wit, investigating the relations of our birds to agriculture, so that there may be no mis- understanding. Finding that it worked a hardship on the deputies to spend certain days of each month collecting birds, the original instruc- tions were modified as follows: Our order concerning the collection of non-game birds, for scientific investigation, is hereby modified to permit collecting at convenient times instead of during the first and third weeks of each month. The collection, however, must be distributed evenly so that nearly the same number of specimens are secured in each bi-weekly period. Under this order we believe the deputies themselves can, while on other duty, take all the specimens needed for our work and thereby greatly reduce expense. In consequence we direct that all special collectors be dismissed and their permits and collecting material taken over by the regular deputies responsible for their engagement. MATERIAL The accompanying map (fig. A) shows the localities in which collections have been made. The localities from which complete series, that is, birds collected each month of the year, were made, and those localities from which incomplete series were obtained, are both indicated. The attempt was made to have a minimum of six specimens collected each month. In several instances collections of a dozen birds each month were obtained. In order that the work of 1911 might be verified, collections were continued at Live Oak, Sutter County ; Sacramento, Sacra- mento County ; Newman, Stanislaus County ; and Salinas, Monte- rey County, during 1912. This afforded a comparison of the food in two successive years, and has acted as a check on the results obtained the first year. A total of 2070 stomachs of western meadowlarks has been available for examination, of which number nearly two thousand have been examined and the results tabulated. The largest col- lection available, composed of one hundred and seventy-five birds, was from Hanford, Kings County. The largest number available taken in a single month was twenty-four. 416 University of California Publications in Zoology [VOL. 11 s 2 31 r^ Z sppg snoau ^ LO 10 UJOO UI uuaq ui ui ! sptau -euiA ui UtBj CO(MQOC5 COCOC B. g 1914] Bryant: Economic Status of the Western Meadowlark 417 sppg snoau UJOO UI oc CO 00 il ^H oo 00 flyr U7 H r- 1 1C : co H 1 3 usaq ui N Conclu jers taken suap.iBS ui c ^ ! rH I 1 sppg oc oc ) ) ir t> CO co cc ) 05 00 s M w H 9jn^sBd ui c<- 5 IT I ) ir i I 05 IT r ) < rH STOMA spas.? -auiA ui t> r~ t> ^ T- : os i ^ (M !> BB g a spaBqojo ui c o- 5 r- 9 * ^ C^ o 1 (M rH l> te ) CO CO SUITABK pa^uasaj V } Ci T 5 C^ H 1C i : 5 r- > C T- ) 'T| 1 O- 1 cc 9 > t> T 1 O 5 CO O5 C*" t- G 3 r- 5 H o ^H -daa sqjuora c oo co | o g a o K >, EH b 6 g o * 418 University of California Publications in Zoology [VOL. 11 LEGEND O Lesstfian six montkly collections vl7 Sixtoten monthly collections I Complete series f< '* c or one year omplete series for two years o^- ^ Fig. A. Map of California showing localities in which collections of western meadowlarks have been made for the purpose of stomach exami- nation. A complete series is a minimum of six birds collected each month in a year. Table I, pages 416-417, gives a summary of the material available. EXAMINATION OF STOMACH CONTENTS On removing the stomach (gizzard) and tag from the cloth, the stomach was carefully cut open, the incision being made with a scissors along the longest axis and through the muscles, starting at the cardiac opening, and the contents emptied upon a glass 1914] Bryant: Economic Status of the Western Meadoidark 419 plate. Great care was taken to see that every bit of the contents was scraped from the walls of the stomach. A Zeiss binocular was used in examining and determining the material found. Where possible, counts were made of all vegetable and animal elements of food. The amount of mineral matter used as grinding material was computed by a calculation in per cent of the com- parative volume. The comparative volume of each kind of food was calculated in per cent of total volume contained in the stomach examined. In the estimation of apparent volume there is always a personal error. However, since this method is depended upon to furnish evidence as to the comparative amounts and not actual amounts, the personal error is largely distributed in the averages. Parts of insects and weed seeds used in identification were wrapped separately in small pieces of paper to prevent their mixing with the rest of the stomach contents. Where possible, the heads of insects were used as a safe cri- terion of the number eaten. In many cases dependence was necessarily placed on an enumeration of the mandibles. In the case of grasshoppers the mandibles are probably retained in the stomach longer than the soft parts, but experiment has shown that the stomach is completely emptied in four hours, so that it is necessary only to give a long enough period of digestion to make such an enumeration dependable. The fact that the mandibles of grasshoppers may be found along the entire length of the digestive tract also supports the view that this criterion is trustworthy. Beetles, bugs, bees, and ants were readily counted, because the heads and thoraces of these insects remain long undigested. Partly digested grain and weed seeds were computed in per- centage volume, but in addition the undigested kernels and seeds were counted. Owing to the finely comminuted condition of the food to be found in the intestines, that found in the stomach alone was used as evidence of the food taken. The stomach alone gives the best unit of volume. The consideration of the food to be found in the intestines could at best but show evidence as to food for a longer time previous to the death of the bird. 420 University of California Publications in Zoology [VOL. 11 In all cases the stomach contents were preserved in vials, so that verification of the results will be possible at any time. As a rule, the contents have been preserved by drying, but where certain animal matter such as larvae was present the material has been preserved in seventy per cent alcohol. After each examination the kind and quantity of food was recorded on a stomach blank. Complete data were recorded on larger blanks. Summaries giving the results of the examinations of the different collections were made in the form of tables. IDENTIFICATION OF STOMACH CONTENTS Collections of insects and seeds for comparison were most helpful in identifying the different insects and seeds. Insects and seeds, if in good condition, can be determined at least to the genus by this method. Floating out the wings of certain insects in water and the mounting of other parts in some clearing fluid are methods which have had to be resorted to occasionally. In the examination of a large series of stomachs it is nearly always possible to obtain a fairly good specimen of an insect which is commonly taken as food, for some bird is usually found which has taken such an insect just before being killed. FOOD OF THE WESTERN MEADOWLARK IN CALIFORNIA The food of the western meadowlark is made up of both vegetable and animal matter. The vegetable food is largely com- posed of grain and seeds. The animal food is made up largely of insects. The accompanying diagram (fig. B) shows the rela- tive amounts of the different kinds of food taken during the year. A discussion of each kind of food is followed by a statement of the "amount destroyed" and the "economic importance." VEGETABLE FOOD Grain Grain forms the largest percentage (seventy-five per cent) of the vegetable food for the year and makes up thirty and eight- tenths per cent of the total food for the year. But a small part (one per cent) of that found in the stomachs has been sprouted. 1914] Bryant: Economic Status of the Western Meadowlark 421 This was therefore in all probability pulled from the seeded fields. Oats is the grain most often taken. It is not only preferred, but is the most available. Much (about seventy per cent) of that found in the stomachs is wild oats (Avena fatua). This oat is so mixed with the tame varieties that part of it must be consid- ered a loss to the rancher, for it makes good feed. Nearly half PROPORTIONS OF DIFFERENT KINDS OF FOOD FOR THE YEAR ANIMAL FOOD VEGETABLE FOOD 63.3% 36.7% Fig. B. Diagram showing relative amounts of different kinds of food taken during the year by western meadowlarks. 422 University of California Publications in Zoology [VOL. H of all the grain taken by the birds examined was consumed during the three winter months November, December, and January. It is apparent, therefore, that availability, lack of insect food, and possibly the sprouting condition of the grain are responsible for this. Owing probably to the feeding habits of the bird, mixtures of different grains are seldom found. The stomachs usually contained one kind of grain only. Barley, because of its greater availability, is more often taken than wheat. The barbs are seldom found in the stomach. One hundred and fifty-five out of one thousand and nine hundred birds had eaten barley. Wheat is taken less often, probably because it is less available. One would suppose that a grain without the hull would be the more palatable. Field corn (Zea mays) was taken by only seven birds out of one thousand nine hundred and twenty, while ten had eaten white milo maize (Andropogon sorghum) and other varieties of Egyptian corn. Quantity destroyed. During the months of December and January the meadowlark feeds largely on grain. Grain forms nearly thirty-one per cent of the food for the year. Stomachs are often found entirely filled with oats, barley, or wheat. As many as thirty kernels, with enough hulls to account for as many more, have been found in a single stomach (pi. 22, fig. 3). Less than one per cent of the grain found in the stomachs has been sprouted grain. Practically no grain is found in the stomachs during the months of March and July. In that grain is more slowly digested than insects, a smaller volume of grain is prob- ably consumed daily. Economic importance. For its destruction of sprouting grain, the western meadowlark justly deserves criticism. In small fields, where the birds are numerous, losses are great. The facts which tend to minimize the damage done are as follows: Much of the grain found in the stomachs is wild oats. Since cultivated oats always contains more or less wild oats in California, it can- not be said that injury does not result from the destruction of the latter. However, much of that eaten must be taken in places where no injury results. All of the grain taken in the months of August, September, and October must be considered 1914] Bryant: Economic Status of the Western Meadoiclark 423 waste grain and of little economic importance. Meadowlarks are not known to attack heads of grain. Whatever field grain is taken is picked up from the ground. Damage done to sprouting grain can result only during a limited period of time (two weeks). After the second leaf appears no damage can result. Weed Seed Weed seed evidently forms the principal part of the vegetable diet of this bird where or when grain is not available. The seeds of such weed pests as tarweed, mustard, tumbleweed, Napa thistle, pigweed, amaranth, canary grass, Johnson grass, foxtail, and sunflower are consumed in large quantities. The seeds of such forage plants as the burr-clover and filaree are commonly eaten. The seeds of filaree (Erodium cicutarium) form the largest per- centage of the weed seeds taken as food. A stomach has seldom been found completely filled with weed seed, for some sort of grain, especially wild oats, is nearly always available with the weed seeds. Nevertheless during the late fall weed seeds make up a considerable part (twenty per cent) of the diet. Grass has been occasionally found in the stomachs. Whether or not it was taken intentionally it is impossible to state. Small sprouts from sprouting grain and sprouting seeds have been found in some instances. As a rule the seeds appear to be sep- arated from the large sprouts when eaten. Small pieces of straw and other vegetable fiber found in the stomachs can be classified as rubbish picked up with the food. There has been a slight complaint that meadowlarks damage sugar beets by feeding on the sprouting seeds. Mr. F. J. McCoy, assistant manager of the Union Sugar Company, Betteravia, California, says on this point: "I have noticed meadowlarks in early spring in our beet fields, but noticed they were feeding on insects. ' ' Stomach examination has failed to disclose any beet seeds. The stomachs of birds collected in beet fields have been found filled with insects. Quantity destroyed. The maximum consumption * of weed seed occurs in October, when nearly one-fourth of the food is made up of this kind of food. Weed seed amounts to five and 424 University of California Publications in Zoology ["VOL. 11 three-tenths per cent of the food for the year. Over 150 seeds of filaree (Erodium sp.) have been taken from a single stomach. Tarweed, pigweed, tumbleweed, mustard, turkey mullein, Napa thistle, Johnson grass, canary grass, foxtail, sunflower, burr- clover, and nightshade seeds have been found in numbers ranging up to fifty. In some few instances stomachs have been found entirely filled with weed seeds. Western meadowlarks appear to feed upon weed seeds to a considerable extent during the time in which they are available. Most weed seeds do not mature until late summer and fall. After plowing begins they are no longer available in cultivated districts, except along fence rows and in uncultivated fields. Economic importance. The destruction of weed seeds must be considered of value to the agriculturist. Weeds even in small numbers take a toll in the grain field. The destruction of weed seeds accomplished by western meadowlarks must help to limit, in some measure, the number of weeds which grow in fields and fence rows the following year. Meadowlarks feeding in grain fields must destroy weed seed that would not otherwise be de- stroyed. Their habit of feeding on sprouting seeds increases their efficiency as weed-seed destroyers. Seeds eaten are digested. In no case have undigested seeds been found in excrement. Fruit No vegetable matter found in the stomachs has been identified as fruit. Grape seeds have been found in a number of cases and there is no doubt that western meadowlarks eat grapes to a slight extent. No serious complaint as to their depredations in this direction has been received. The stomachs of practically all of the birds collected in vineyards have been filled with insects, mostly beetles. A systematic list of the grain and weed seeds found in the stomachs follows : Grain Oats Barley Wheat Field corn (Zea mays) Sorghum (Andropogon sorghum subsp.) 1914] Bryant: Economic Status of the Western Meadowlark 425 Fruit Grape seeds (Vitis sp.) Weed seed Gramineae (Grass family) Andropogon sorghum halepensis. Johnson grass Panicum sp. Panic grass Chaetochloa glauca. Bristly foxtail Chaetochloa viridis. Foxtail Phalaris minor. Small canary grass Avena fatua. Wild oats Bromus sp. Brome grass Lolium temulentum. Darnel Hordeum sp. Barley grass Cyperaceae (Sedge family) Carex sp. Sedge Polygonaceae (Buckwheat family) Rumex sp. Dock Polygonum sp. Knotweed Chenopodiaceae (Goosefoot family) Chenopodium sp. Goosefoot pigweed Amarantaceae (Amaranth family) Amaranthus graecizans. Tumbleweed Amaranthus sp. Amaranth Portulacaceae (Purslane family) Lewisia sp. Bitter root Eanunculaceae (Buttercup family) Ranunculus sp. Buttercup Cruciferae (Mustard family) Brassica nigra. Black mustard Brassica sp. Mustard Leguminosae (Pea family) Medicago hispida. Burr clover Medicago arabica. Spotted medick Melilotus indica. Yellow melilot Melilotus sp. Sweet clover Trifolium sp. Clover Lupinus sp. Lupine Geraniaceae (Geranium family) Erodium cicutarium. Red-stem filaree Erodium sp. Filaree Euphorbiaceae (Spurge family) Eremocarpus setigerus. Turkey mullein Malvaceae (Mallow family) Sida hederacea. Alkali mallow Onagraceae (Evening Primrose family) Oenothera ovata. Golden eggs Primulaceae (Primrose family) Anagallis arvensis. Pimpernel 426 University of California Publications in Zoology [VOL. 11 Boraginaceae (Borage family) Amsinckia intermedia. Amsinckia Amsinckia sp. Amsinckia Cynoglossum sp. Hound's tongue Eubiaceae (Madder family) Galium sp. Bedstraw Solanaceae (Nightshade family) Solanum sp. Nightshade Compositae (Sunflower family) Lactuca scariola. Prickly lettuce Centaurea melitensis. Napa thistle Centaurea solstitialis. Barnaby's thistle Carduus sp. Thistle Hemizonia sp. Tarweed Helianthus annuus. Common sunflower Iva axillaris. Kagweed Economic importance of vegetable food. The destruction of sprouting grain means a loss of dollars and cents to the rancher. This loss is minimized somewhat by the limited time during which injury is possible and the possibility of protective measures. A much smaller loss can be attributed to the destruction of grain picked up in newly sown fields. Grain on or near the surface of the ground in seeded fields is of doubtful value, as it cannot be depended upon to furnish a strong, healthy plant. Much of the wild oats and some of the tame oats must be considered waste grain or uncultivated grain. Its destruction, in spite of its utility as feed, cannot be considered a direct loss in money value. Practically all the other seeds destroyed are the seeds of weed pests. The destruction of the seeds of certain forage plants such as filaree could be considered a detriment if they were destroyed in large enough quantities to make any difference in the amount of forage available. This same plant is considered a weed in many places. Any destruction of the seeds of thistles, sunflowers, Johnson grass and like weeds must be considered a benefit. ANIMAL FOOD Coleoptera (Beetles) The most constant article of diet of the meadowlark consists of beetles. The habitat of the bird would forecast this fact. Ground-beetles (Carabidae, Tenebrionidae) , click-beetles (Elate- 1914] Bryant: Economic Status of the WesternMeadoivlark 427 ridae), and weevils (Rhynchitidae, Calandridae, Otiorhynchidae) form the largest per cent of this kind of food. Representatives of practically every family of the Coleoptera, however, have been found in the stomachs. A svstematic list of the beetles identified follows : Cincindelidae Cincindela sp. Carabidae Calosoma sp. Amara californica Dej. Amara conflata Lee. Calathus ruficollis Dej. Platinus bicolor Lee. (?) Staphylinidae Staphylinus tarsalis Mann. Creophilus villosus Silphidae Silpha ramosa Say Dermestidae Dermestes sp. Erotylidae Languria sp. Histeridae Saprinus fimbriatus Lee. Buprestidae Elateridae Cardiophorus tenebrosus Lee. Anchastus einereipennis Mann. Drasterius livans Lee. Drasterius sp. Megapenthes aterrimus Horn Dolopius lateralis Melanotus variolatus Lee. Limonius infuscatus Mots. Limonius canus Lee. Limonius californicus Mann. Lampyridae Telephorus censors Lee. Malachidae Collops marginellus Lee. Scarabaeidae Aphodius subaeneus Lee. Aphodius granarius Linn. Aphodius rugifrons Horn. Hoplia sp. Cotalpa ursina Horn Ligyrus gibbosus De Greer Harpalus pennsylvanicus Dej. Agonostoreus maculatus Lee. Pterostichus sp. Bladyeellus rupestris Say Anisodactylus dilatatus Dej. Anisodactylus sp. Dytiscidae Agabus lugens Lee. Chrysomelidae Glyptoscelis albidus Lee. (?) Gastroidea sp. Diabrotica soror Lee. Disonycha sp. Chaetocnema sp. Microrhopala melsheimeri Cr. Tenebrionidae Blapstinus gregalus Casey Blapstinus rufipes Casey Blapstinus elongatus Casey Coniontis subpubescens Lee. Coniontis viatica Esch. Eulabis pubescens Lee. Eurymetopon sp. Eleodes sp. Meloidae Otiorhynchidae hnigopsis effracta Lee. Rhigopsis sp. Curculionidae Sitones californicus Fah. Sitones sordidus Lee. Lixus perferatus Lee. Cleonus virgatus Lee. Centrocleonus near angularis Lee. Baris cuneipennis Casey Baris sp. Calandridae Sphenophorus vomerinus Lee. Sphenophorus simplex Lee. Sphenophorus sp. 428 University of California Publications in Zoology [VOL. 11 Quantity destroyed. Beetles are taken every month of the year, and form 21.3 per cent of the total food. Stomachs have often been found filled with nothing but beetles. From twenty to fifty have been found in a single stomach. Wireworms, the larvae of click-beetles (Elateridae), are taken in large numbers where they are available. As they are less often seen above ground than cutworms, it is only natural that they do not form nearly so large a percentage of the food for the year. The adult click-beetles are also taken (see pi. 24, fig. 8) . Economic importance. Wireworms are injurious to the roots of plants. Damage by them to garden truck and pasture land is of common occurrence. The ability of the meadowlark to probe into and remove from the soil such insects increases its value as an insect destroyer. The destruction of wireworms must be considered a benefit of considerable importance, especially in meadow and pasture land. By far the greater number of beetles taken as food are the common ground beetles (Carabidae). These beetles are often classified as beneficial insects, because they are supposed to feed on other injurious insects. Certain ones are predacious and are known to feed on fly and beetle larvae in California. Of the food habits of others little is definitely known, and we are justified in speaking of them as neutral, for they do practically no harm and are not known to do any particular good. Tiger-beetles (Cincindelidae) and carrion-beetles (Staphylinidae), eaten to some extent, must be numbered among the beneficial beetles de- stroyed. The meadowlark, however, does feed upon many injur- ious beetles, chief of which are click-beetles (Elateridae), pina- cate beetles (Eleodes sp.), leaf -beetles (Chrysomelidae), snont- beetles (Otiorhynchidae, Curculionidae), and weevils (Calan- dridae). Among the leaf-beetles is numbered the destructive California flower-beetle (Diabrotica soror). This and other members of the family constitute some of the worst beetle enemies of our crops. Snout-beetles (curculios) and weevils (Spheno- phorus sp.) are well-known pests of fruit and grain. The con- tinual destruction of large numbers of these injurious beetles must be considered a decided benefit. 1914] Bryant: Economic Status of the Western Meadowlark 429 Orthoptera (Grasshoppers and Crickets) Grasshoppers and crickets form a large percentage of the meadowlark's food during the summer and fall, making up as high as eighty-five per cent of the food in August. The species of grasshopper taken is the one most available. Practically all of the species found in the state are doubtless represented in the stomachs. The common cricket (Gryllus) and the Jerusalem cricket (Stenopelmatus sp.) are common articles of diet. Katy- dids (Locustidae), being less common insects and having a different habitat, are not taken so often. A systematic list of the Orthoptera found in the stomachs follows : Gryllidae Arphia sp. Gryllus integer Scud. Dissosteira spurcata Saus. Gryllus pennsylvanicus Burm. Conozoa behrensi Saus. Locustidae Aeridiinae ( ?) Macrocentrum sp. Oedaleonotus enigma Scud. Conocephalus acutalus Scud. Melanoplus devastator Scud. Stenopelmatus irregularis Brun. Melanoplus differentialis Uhler Stenopelmatus sp. ( ?) Melanoplus unif ormis Scud. Acridiidae Fulgoridae Cedipodinae (?) Labia minor Camnula pellucida Scud. Quantity destroyed. Next to beetles, grasshoppers form the most important article of diet. Nearly fifteen per cent of the food for the year is made up of these insects. Parts of as many as twenty-six large grasshoppers (one inch or over in length) and fifty-eight small grasshoppers (one-half inch in length) have been found in a single stomach. The maximum amount of this food is taken in June, July, and August, the birds feeding almost exclusively on these insects during these months. The state over, some grasshoppers are taken by the meadowlarks every month in the year. The quantity taken closely parallels the abundance of these insects. Crickets as well as grasshoppers are relished by the meadow- lark. As many as fifteen pairs of mandibles have been taken from a single stomach, showing that at least fifteen common black 430 University of California Publications in Zoology [ VOL. 11 crickets (Gryllus pennsylvanicus) had been eaten within four hours by that particular bird. Five per cent of the food for the year is made up of crickets. Wood crickets, better known as Jerusalem crickets (Stenopelmatus sp.), being less abundant than the common cricket, are less often taken as food. Economic importance. Grasshoppers can be classed as injur- ious insects. The extent of their damage can be traced to their abundance rather than to the presence of any particular kind of grasshopper. The species which most often become abundant enough to cause serious losses in this state are the differential grasshopper (Melanoplus differentialis) , the pale-winged grass- hopper (Melanoplus uniformis), the devastating grasshopper (Melanoplus devastator), and the valley grasshopper (Oedaleo- notus enigma). All of these grasshoppers are destroyed in great numbers by the meadowlark. The more abundant these insects become, the more do these birds turn their attention to this kind of food. Where grasshoppers are abundant, meadowlarks have been found to average as high as fifty grasshoppers a day. (See Bryant, 1912d.) As a grasshopper destroyer, the meadowlark is unequaled by any other bird unless it be the blackbird, and then only because of greater numbers of blackbirds. As grasshopper outbreaks continue to ravage certain parts of the state each year, the meadowlark performs a service to agriculture that can hardly be overestimated, in that it helps to keep the insects down to normal numbers, so that losses do not result, and prevents greater losses by taking a greater toll at the time of an outbreak. Crickets are usually classed as injurious insects. The degree of injury, as with the grasshoppers, depends largely upon their abundance. Since the species of crickets fed upon by the mead- owlark feed almost entirely upon plants and are often destructive to grain, their destruction is to be desired by the rancher. This is especially the case with the Jerusalem cricket, which is very destructive to potatoes. Lepidoptera (Butterflies and Moths) The general law that birds do not eat butterflies to any great extent appears to hold good in the case of the western meadow- lark. (See p. 481). However, the following dependable obser- 1914] Bryant: Economic Status of the Western Meadowlark 431 vation made by Mr. John G. Tyler of Fresno, California, also furnishes evidence of the fact that butterflies are occasionally, at least, destroyed. "While strolling along the road east of this city the writer noticed a field of alfalfa that was infested with yellow butterflies. A nearer approach revealed the presence of several meadowlarks, and I was so fortunate as to see one of these birds seize a butterfly and make way with it. I am not prepared to say that the victim was actually swallowed, but it was certainly captured and killed." The larvae of butterflies and moths are common articles of diet. Cutworms and caterpillars form ten per cent of the food for the year, reaching a maximum in May and June, when they amount to nearly a third of the meadowlark's food. Even hairy caterpillars do not escape destruction. In one instance the larva of the mourning-cloak butterfly (Euvanessa antiopa) has been found in the stomach. Smaller hairy cater- pillars are of common occurrence. Both the larva and pupa of the sphinx moth have been taken from stomachs. Pupae do not form so important a part of the diet as do the larvae. The only Lepidoptera positively identified follow: LEPIDOPTERA (BUTTERFLIES AND MOTHS) Noctuidae Nymphalidae Peridromia sp. Eugonia calif ornica (Boisd.) (?) Euvanessa antiopa (Linn.) -(?) (?) Papilio sp. Sphingidae (?) Phlegethontius sp. Quantity destroyed. Cutworms and caterpillars form about twelve per cent of the food for the year. The maximum quantity is taken in the months of March and April, when almost half of the food taken is made up of these insects. Many of the stomachs contained as many as twenty large cutworms or caterpillars. One bird collected at Red Bluff, Tehama County, contained sixty- six cutworms and over thirty small beetles (pi. 23, fig. 5). 432 University of California Publications in Zoology [VOL. 11 Economic importance. Cutworms and army worms can be classed as two very important pests in California. Garden truck and even trees are sometimes defoliated when these insects become numerous. The depredations of the grape cutworm are only too well known in the state. Caterpillars are vegetable feeders and are always classed as injurious. The destruction of these pests in very large quantities by the western meadowlark must cause a direct saving to the rancher and fruit grower. Hemiptera (Bugs) Stink-bugs (Pentatomidae) appear to be relished in spite of their excretions, for they are taken in large numbers. Squash- bugs (Anasa sp.) have been found in only a few instances. Negro-bugs (Corimelaena) form the only other important Hem- iptera taken. Cicadas appear to be relished and often caught. Two stomachs have contained aphids (Aphis brassicae). The following Hemiptera have been identified : HEMIPTERA (Buos) Jassidae Coreidae ? Corizus sp. Aphidae Alydus pliosulus Aphis brassicae Linn. Anasa sp. Membracidae Pentatomidae Stictocephala franciscana Stal. Podisus pallens Stal. Cicadidae Podisus sp. Platypedia areolata Uhl. Eusehistus conspersus Uhl. Platypedia minor Uhl. Eusehistus servus Say Eeduviidae Corimelaenidae ? Corimelaena sp. Quantity destroyed. The commonest true bugs destroyed by western meadowlarks are stink-bugs (Pentatomidae), negro-bugs (Cormelaenidae), leafhoppers (Jassidae), and cicada flies (Cica- didae). They form nearly two per cent of the food for the year. As many as twenty stink-bugs have been taken from a single 1914] Bryant: Economic Status of the Western Meadowlark 433 stomach. Negro-bugs are not taken in such large numbers, nor are they so abundant. Two stomachs have been found almost filled with leafhoppers, and many others contained two to ten of these insects. Cicadas, near relatives of the eastern seventeen- year locust, are occasionally taken, probably as often as they are available. Bugs form over three per cent of the food for the year. Economic importance. Most stink-bugs are vegetable feeders and occasionally give trouble. Negro-bugs are troublesome on berries. In the destruction of these insects the western meadow- lark is also conferring a benefit. Leafhoppers are injurious to plants, because they secure their food by sucking the juice of the plant. Any destruction of leafhoppers, however small, is of value. The cicada in California is not abundant enough to be of economic importance. It lays its eggs in the sapwood of plants and trees. Since these insects, if they became abundant, would cause trouble as does the seventeen-year locust of the east, their destruction is to be looked upon with favor. Hymenoptera (Ants. Bees, and Wasps) Ants appear to be taken irrespective of size or kind, for they are to be found from the smallest to the largest. The common red and black ants (Messor, Pogonomyrmex) , field ants (Formica), and carpenter ants (Camponotus) are most abundant in the stomachs. Ichneumon flies are taken in considerable numbers. Bees and wasps form a less percentage of the food made up of Hymeno- ptera. In but one or two cases was a bee definitely identified as a honey bee (Apis mellifera). Solitary bees (Chrysis) and even bumblebees (Bombus calif ornicus) have been found. Cow- killers (Mutillidae) are occasionally eaten, although it has com- monly been supposed that they were well protected from attack by their sting, hairy covering, and warning coloration. For several reasons the Hymenoptera have been very difficult to identify. The finely comminuted condition of the insects has proved an almost insurmountable difficulty. The following: only have been identified: 434 University of California Publications in Zoology [VOL. 11 HYMENOPTERA (ANTS, BEES, ANb WASPS) Ichneumonidae Vespidae ? Polistes aurifer Sauss. Polistes minor Beauv. - ? Sphegidae 1 Ammophila sp. Mutillidae Formicidae Sphaerothalma californica Formacinae Sphaerothalma aureola Camponotus sp. Apidae Formica sp. ( ?) Apis mellif era Dolichoderinae Bombidae Tapinoma sessile Say Bombus californicus Smith Myrmicinae Chrysididae Messor andrei Mayr. Chrysis sp. Pogonomyrmex californicus Buckley Pogonomyrmex sp. Quantity destroyed. Bees and wasps form 3.6 per cent of the yearly food. In no case have ichneumon flies, which are valuable parasitic insects, been taken in numbers, five being the maximum found in a single stomach. Their rapid flight prob- ably prevents a greater toll being taken. Ants are often eaten in large quantities and form over two per cent of the food for the year. It is not an uncommon occurrence to find a stomach almost filled with ants. Over one hundred have been found in a single stomach. Economic importance. Most bees and wasps are considered beneficial insects. Ants are either injurious or neutral; few are beneficial. In the destruction of ichneumon flies the western meadowlark is destroying a valuable parasitic insect. The de- struction of bees and wasps must also be reckoned as a count against the bird. However, the small numbers destroyed mini- mize greatly the real and the possible damage done. The destruction of most kinds of ants makes little difference one way or the other, owing to their abundance and scavenger habits. Dipt era (Flies} A few members of the family Muscidae, a few flower-flies (Eristalis sp.) 'and crane-flies (Tipula sp.) and the pupae of syrphid flies (Syrphus) are the only representatives of the Diptera which have been found in the stomachs of western meadowlarks. The following have been identified: 1914] Bryant: Economic Status of the Western Meadowlark 435 DlPTERA Eristalis tenax Lucilia caesar L. Tachina sp. Musca sp. Syrphus sp. ? Tipula sp. Quantity destroyed. Flies do not constitute any important percentage (about one per cent) of the food for the year and when found have been in small numbers. A few green bottle- flies and other members of the same family are eaten, as are also flower-flies and drone-flies. Birds collected at El Toro, Orange County, during 1911 had eaten large numbers of the pupae of flower-flies (Syrphus sp.) . Crane-flies (Tipula sp.) are not taken as often as it would seem they would be from their abundance. Evidence is at hand, however, that western meadowlarks feed largely on the larvae when they become abundant. Mr. W. M. Hughes of Madera, Madera County, has made the following report: "When I visited the tract of land affected, I found myriads of blackbirds and thousands of meadowlarks on the ground making small holes into the ground at the roots of the plants and taking out the worm. Several hundred acres of fine crop was destroyed before the birds collected in numbers suffi- cient to destroy the pest. ' ' The outbreak referred to was in the vicinity of Minturn, Madera County, in 1909. Several hundred acres of barley were destroyed by crane-fly larvae at this time. Economic importance. In spite of the fact that the larvae feed upon decaying matter, most flies are considered pests be- cause some of them carry germs of disease. Green bottle-flies are disease carriers. The larvae of flower-flies feed upon plant lice and hence are considered beneficial. The larvae of crane- flies, on the other hand, are destructive to vegetation. The small numbers of flies taken and the fact that injurious as well as beneficial forms are eaten make the destruction of Diptera by meadowlarks of little consequence. Arachnida (Spiders) Quantity destroyed. Spiders and their egg-cases form less than one per cent of the food for the year. Most of the spiders taken are grass spiders (Agalenidae) and daddy-long-legs (Pha- 436 University of California Publications in Zoology [VOL. 11 langidae), the commonest of those found in pastures. In no instance have more than two spiders been taken from the same stomach. Economic importance. Spiders should be considered as doubt- fully beneficial or of neutral value to the agriculturist, in spite of the fact that they feed largely on insects. As a rule spiders are not abundant enough to be of great economic importance. Their destruction at the hands of the meadowlark is of no conse- quence, as the resulting effect on insect life is so small. Miscellaneous Animal Food Miscellaneous articles of diet form three and one-half per cent of the food for the year. The common sow-bug (Porcellio scaber) is the commonest crustacean found in stomachs. Two birds had eaten snails. Two birds from San Diego had each taken a scorpion. But few earthworms have been found in the stomachs. Centipedes (Scolopendra sp.) and millipedes (Julus) appear to form a constant part of the diet. They are evidently taken regularly where available. Two birds had eaten ant-lions (Myrmeleon sp.). Quantity destroyed. Centipedes, millipedes, scorpions, ant- lions, and sow-bugs may be considered oddities in the diet. Their slight availability may account in some measure for the small numbers taken by the western meadowlark. Economic importance. Of these miscellaneous elements in the diet, only millipedes and sow-bugs can be considered injurious. Centipedes are usually considered beneficial, scorpions injurious, and ant-lions of neutral value. None of these forms is taken in sufficient quantity to make their destruction either an injury or a benefit. Inorganic Matter Pebbles, used for grinding the food, make up the inorganic matter found in the stomachs. White and red pebbles, probably because of their conspicuousness, predominate. Pebbles appear to be necessary as an aid to the digestion of grain, but much less necessary for the digestion of insects. Pebbles are nearly always 1914] Bryant: Economic Status of the WesternMeadowlark 437 found with grain and weed seed, but seldom with insect food. Doubtless the chitinous parts of the insects largely take their place. Two nestlings and two adults contained parts of egg-shells. It is a well-known fact that birds often eat the broken shells after the young have hatched. In one instance, at least, the parent birds had fed the young on the shells. PRINCIPAL, ARTICLES OF DIET The kinds of food forming a definite part of the food of the western meadowlark for the year are as follows : VEGETABLE ANIMAL Grain 30.8% Coleoptera 21.3% Weed seed 5.3 Orthoptera 20.3 Miscellaneous .6 Lepidoptera 12.2 Hemiptera 1.7 Hymenoptera 5.6 Diptera .1 Arachnida .2 Miscellaneous insects 1.9 EXAMINATION OP FECES Other than stomach examination, the examination of feces would appear to give the best evidence as to the food of birds. That a considerable amount of knowledge concerning the food of meadowlarks can be obtained in this way is evidenced by the fact that the examination of feces of meadowlarks collected June 15, 1912, in the Berkeley Hills showed the following: Thorax of spider, heads of ants, mandibles and other parts of grass- hoppers, wing covers and mandibles of beetles and pubescence from wild oats. The examination of some feces collected from nestling birds at Lathrop, San Joaquin County, showed that these same hard parts passed through the digestive tract undigested. Conse- quently this affords a practical method of determining the kind of food taken. Its value as a means of determining the amount of food is much less, for only the more resistant parts can be found. 438 University of California Publications in Zoology [VOL. 11 pOOJ TH TH L^ rH O b- OS O rH GO rH GO b- CM CM IO CM CO tr.ir> T rH O* O rH* IO" O CO* 1O i-H OS* OS* CO CM* OS* TH CD TH TH COCOTHlOCOlOCOOSGOlOCMTHrH IOTHCO CO CO CO OS O CO rH O OS CM OS CM CO CO 00 IO b-- b- oo' os* os* 06 10' os" co 10" ao* o o co* t>* o 10* co 10' 10* COCOIOTHCOTHCO rHTHb-lOOOOSTHlOCDOS CM oo co co : : os : co TH co co b-; os CM oo co rH CM* ' rH j ! TH* j CM* rH rH CM* CO (epiuq0B.iy) oq CM TH : CM CM CM : : CM I-H o : iq r-i I-H iq sjgpidg rH : : iq os co CM co TH b-; : TH oo TH os iq to TH o TH * CO' j rH* CO* TH* " CO* IO* . b- O5_ O 1 IO CM b^ : CM OS CM CO IO b- TH CO CO ' CM* CO* 1O* i rH TH* TH i rH rH rH TH CM* CO* pUB S99^ \f CM TH iq : iq co O : rH co rH : I-H iq co co co b- "cOCMICMb-CM* rH rH lo'rHrH IO CM CO TH TH TH TH ! CD CO OS b- TH IO rH b- b- OS IO* CO CO IO' IO' CD OS* ! CO GO' CO TH OS* b-* TH CO* rH OS* i-H rH i I rH CMrHrHrH CO ICMOSO : i i iiooo ;rHio icoiq i c > * rH ! I I j CM* ! r-l b- OS_ CO CM rH CO O | O IO CO TH O : IO iq TH CM os* CM* 10* CM* oo' t^ oo' : CM' TH o CM' co' : t-* b-* CM* 10" rHrHCMCMCM CM rH CMTHCM oocooTHcocooo : :oscorHrH t> ; l ^* > ; r ~J ",_*,_" *co* :CM "0606 ICM'IO'IO'O rH rH rH O5 TH CM CO TH rH O CO IO CO O O TH IO b- IO OS CM CO* 00* CO* b-* 00* b-" CM* IO' b-^ O OS* TH rH* rH* TH GO OS IO' COCMCM CM rHCMCMCMCMrH CO b-COOOSCOrHCM JrHOSTHCOb-t^lOOOOrH " T-" TH TH O OS* ?O i b-* b-* CO* CM CM* GO rH b-^ CM' iq TH b- oq iq co_ oo o os oo CM TH co : cs TH co os o oo* cd 10* 10* co 10* 10* co co 10 TH os* : r-" co* CM* co COCMCOTHCMTHCMOSb-lOrHTH 'THCOCO Sq^UOUI OOOb-CO(MrHb-rHCMGOCOb- c C>'- 1914] Bryant: Economic Status of the Western Meadowlark 439 POOI t^ o CM CM 00 CM O rH >0 r i , (Baa^diuteH) ^ IO IO CO O5 10 : rH t. OO TH CO ^ w sSng i rH rH CO rH CO rt TH TH rH 1 ** SJBTItdjaiBO CD * IO O5 Tf< CO CO rH Th O CO OO co CM o puB sraaoA^iiQ ^ rH CO t^ TH ^ GO T | CM t- t- O5 LO CM' rH g 0allJ.dJ_ ' 1 O 8 LO l^. 05 TH CO co : -r 05 10 CM s s j9ddoqssBJ-) oo CD t>* H- CO 05 oo ; i^. CO rH t^ O TH z rH CM TH CM M fa rH CO CD (M t^ 05 05 : o CM TH t^ IO rH CM g T^^l . kj CO 05 00 t- CO ! ^ T 1 CM TH rH IO i CM rH I (Bja^doaioo) LO CO O 00 O CM O5 CO TH CD 1 I t~ CM O CO s S8J}33g O5 10 CO O5 rH 00 CO IO IO Tfi CM CD 00 CO CM CM CM CO O5 rH CO rH CM w >o E" 1 CD CO t- : LO LO rH 05 CO CO &4 p^da f^^^V ^ CQ TH 1 CO CO CO - CO TH rH t^ T 1 O5 00 CO CO oo cc rH i 1 O CM 1 TH O5 ( **$ JB8 A 05 05 05 05 05 O5 O5 05 05 05 05 O5 O5 O5 O 1 G rrt rH is E M O 43 c Z Si O | > s^ *G 1 Madera Fresno S "S W ^ 03 PQ ll Kern Ventura 1 ff & Riverside O 0) .H be ft : ~- Si W Imperial IVf isrplln n Is .I s 440 University of California Publications in Zoology L V L - n QUANTITY OF FOOD Three methods may be used in estimating the quantity of food in a bird's stomach. First, the articles found may be counted; second, they may be weighed; or third, they may be estimated by volume. The first method, although important as giving an idea of the bird's economic value by showing the number of injurious insects or seeds destroyed, fails to take into account the difference in size of the different articles, and does not show the relative amounts of each kind. The second method has been generally disregarded because of its impracticability. The third or vol- umetric method allows of a balance of the inequalities of size, and best portrays "the ratios each element bears to the others." By the numerical method, fifty ants would be placed against, say, six ground-beetles. A computation made by the percentage- by-volume method would doubtless show that these two kinds of food represented only three and twenty-six per cent, respectively, of the whole food. Hence the idea furnished in the first case (a ratio of 50 to 6) is a misleading one. Numbers of one insect cannot be balanced against the numbers of another insect. As each bird of the same species has a certain average stomach capacity, the ratio of each element to this average capacity gives the most accurate idea of the relative proportions of each kind of food. Although the first and last methods have been those most often used heretofore, the second presents certain advantages (e.g., mathematical accuracy) which should not be overlooked. A combination of the numerical and the percentage-by-volume method has been used in the present work. The number of birds taking the different kinds of food offers further evidence as to their capacity for good or ill. McAtee (1912) has pointed out that statements of numbers of individuals in stomachs has an interest in direct proportion to the bigness of the number. Believing this to be largely true, the maximum number of individuals found in the stomachs has here received emphasis. These maximum numbers should not be considered as averages. 1914] Bryant: Economic Status of the Western Meadowlark 441 It does not seem fair to compare the food of a species of bird of small size with that of one of large size without taking into account the bird's capacity. The degree of injury or benefit is largely dependent on the total amount consumed. In this investigation measurements in cubic centimeters of the stomach contents of a number of western meadowlarks have allowed the computation of the average stomach capacity. For male birds this average capacity is three cubic centimeters ; for female birds it is two and one-half cubic centimeters. The average capacity is two and three-fourths cubic centimeters. This method allows of some interesting computations of the capacity for good or harm of the western meadowlark. If there are twelve meadowlarks to the square mile, as there are in many places, these birds demand over one hundred cubic centimeters of food daily. This capacity, estimated in grain, means 2200 kernels, in grasshoppers 150 average-sized individuals, in cut- worms 125 individuals, in ground-beetles 300, in ants 2500. As digestion is constantly going on, much larger numbers are in reality taken, as is evidenced by a count of the insects in a stomach. These numbers, therefore, are not even a minimum. As pointed out (p. 412), the results of the experiments showed that meadowlarks must completely digest a meal inside of four hours. It was also found that grain takes longer to digest than do insects. Thus it can be seen that the food found in the stomach at any one time does not represent the total amount of food taken daily, but only about a third part of that consumed daily. Owing to the slower digestion of grain, the amount found in a stomach must represent more nearly one-half of the daily requirement. Taking an average capacity of two and three-quarters cubic centimeters and considering that each bird nils its stomach at least three times a day, one hundred western meadowlarks must consume near a liter, or about a quart of food each day. If the food be grain it can be seen that the amount of destruction is considerable ; if the food be insects it can be seen that meadow- larks take a large daily toll of insects. The same type of com- putation shows that a single western meadowlark must consume six pounds of food a year. 442 University of California Publications in Zoology [VOL. 11 Some idea of the average numbers of the common insects, grain, and weed seeds destroyed at each meal by western meadow- larks can be obtained from the following tables, which were computed from the results of stomach examination of birds col- lected in Sacramento and Stockton, California. The average for the day is three times that for each meal. NUMBER OF COMMON INSECTS, GRAIN AND WEED SEEDS DESTROYED BY WESTERN MEADOWLARKS COLLECTED AT SACRAMENTO, CALIFORNIA February to April, inclusive Grain 1 Weed seeds 61 ( Beetles ' 6 Cutworms and cater- Grass- pillars hoppers 8 Ants, bees, and Bugs wasps Spidei 2 71 3 3 6 5 4 5 1 10 4 12 1 6 8 7 5 8 3 1 9 2 2 5 16 10 7 11 3 5 3 12 . . 3 13 3 2 14 7 2 Totals 5 80 56 113 1 Av. i)er bird 3 6 4 8 9 10 11 Totals Grain 25 28 September to November, inclusive Weed seeds 60 61 155 65 37 3 200 93 80 180 15 949 Cutworm s Ants, and catei - Grass- bees, anc I Beetles pillars hoppers Bugs wasps Spiders 2 1 1 1 1 1 2 1 4 2 2 6 1 8 22 2 5 1 11 2 1 2 2 2 1 22 25 10 30 1 1 12 Av. per bird 2.5 86.2 2.2 2.6 .6 1914] Bryant: Economic Status of the Western Meadowlark 443 NUMBER OF INSECTS TAKEN BY WESTERN MEADOWLARKS AT STOCKTON, SAN JOAQUIN COUNTY, CALIFORNIA ii . s i i 1 1, i s . i i I 2 Date J I g J* 6 I ff 3 1 I 15 March-June, 1912 6 25 88 52 88 68 4 7 4 311 31 Av. per bird .4 1.6 5.8 3.4 5.8 4.5 .2 .4 .2 20 2.0 Av. per bird per day 1.2 4.8 17.4 10.2 17.4 13.5 .6 1.2 .6 60.0 6.0 Few people have any realization of the great quantities of insects consumed by birds. For instance, if we consider that there is an average of one meadowlark to every two acres of available land for cultivation (11,000,000 acres) in the Sacra- mento and San Joaquin valleys and that each pair of birds raises an average of four young, each one of which averages one ounce in weight while in the nest and consumes half its own weight of food each day, it takes over 343 y% tons of insect food each day to feed the young birds in the great valleys alone. The increased consumption of insect food due to nestling birds comes at a time when insects are most numerous, and so is instrumental in helping to prevent an undue increase of insects. As insects become injurious only when in maximum numbers, this increased con- sumption by birds is doubly important. A conservative estimate of the approximate amount of the different kinds of food consumed by the average meadowlark in California during a year is as follows: Grain 1% Ibs. Weed seed V 2 Insects 2% Total 6 The fact that the western meadowlark eats both animal and vegetable food is a point in its favor. If it were exclusively insectivorous the bird could not exist in such large numbers because of the lack of insect food during part of the year. The consequent destruction of insect life would therefore be much smaller. 444 University of California Publications in Zoology [VOL. 11 Certain species of birds when hungry will not only fill the stomach, but will continue eating until the gullet is also filled. This is often found to be the case with linnets and bicolored blackbirds. In no case, however, has the gullet of a western meadowlark been found well filled with food. At the most, the last insect taken before the bird was collected has been found in the lower part of the gullet. CAPACITY FOR GOOD OR EVIL AS EVIDENCED BY THE NUMBER OF BIRDS TAKING THE DIFFERENT ITEMS OF FOOD The percentage-volume method of estimating the proportion of the different kinds of food taken by a bird gives us the best idea of the relative importance of the different kinds of food in the diet of a given bird. However, the frequency of occur- rence of the different items in the food, shown by a statement of the number of birds taking each item, furnishes additional evidence as to the capacity of a species for good or ill. The number of birds taking a certain kind of insect food can be regarded as an approximate index of the availability of that kind of food, and to a much less extent as an index of food preference if we consider insects as being evenly distributed and birds as being but slightly influenced by psychological processes. A nearer approximation can be obtained by multiplying the number of birds by the number of insects taken. In such a computation the number of insects taken is considered, as well as the number of birds taking the different elements of food. By this method the index of availability of crickets at Live Oak, Sutter County, was four in 1911, whereas this index was 6902 at Hollister, San Benito County, in the same year. By the same method of calculation the index of availability of grasshoppers was 2162 at Live Oak and but 1541 at Hollister. In the first case crickets were 1670 times as available at Hollister as at Live Oak, and grasshoppers 1.4 times more available at Live Oak than at Hollister. The following table gives a comparison of availability of the commoner insects as evidenced by indices of availability. Preference is here classed as a factor in availability. 1914] Bryant: Economic Status of the Western Meadowlark 445 01 01 en Number of o o o o birds ^ ^ ^> ^> j 4 ^ 2 & > a g ^ t^ CD ^ CD CD O CS / H >-i 3 oo O5 to x i * 8 . i ^ : ^ rfx Wireworms ^ o Oats w 02 ft IN3 2 co co Bugs ^ ^j ^ (Hemiptera) j ^ 01 Barley g g g M Grain M i O GO ^ Cicadas i i o ?3 ^ i i_i . . ^ 01 Weed seeds J ^. CO 01 to Wheat bo Si > CO CO ^ h- ' tO o -^ ^ 05 to Ants O to to o ^ 5 o Beetles l*q QQ ^ 1 So ^ ees an( ^ co Corn oo J^ Sorghum 3 cr CD CD 1 tO M to wasps H ^z g g * ^Crickets 25 ^ to 01 r,. Beans S 1 72 __* ^j| ^. . H- ' H-i to to O CO P (Jl CO h^ O5 ^ oo 01 co o Grasshoppers ^ gg Weed seeds P^ o bo bi 5 V Butterflies ii ^ 8 r^ W >_. j_i co M Jerusalem -q ,_i o tc oo crickets i_5 co 05 p u pae Grape seeds 72 h- 1 CO Ol ^i o to o Cutworms and Tl CO to H3 O5 to caterpillars .^ w Flies r w Grass CO 5 ^1 00 01 oo H- o Bugs Cd 1 o O CO *" La ^ Fly larvae O 01 MX Beetles d LCl i_i E3 M Bees and es H (^7| to 2 o rf^ wasps ^ to Crane flies to g ^ -3 *. ^ ~^ N* W _ . . to co Crickets w C5 bo Ants ? ^ Spiders J^. ^ t i "^i W e3 Ol r^ 50 Spiders ^ M 01 Spider egg co cases o 2q Grasshoppers 01 C O2 a co J-> bs Miscellaneous ^ 05 g co Centipeds u, Jerusalem K.J crickets O =a ^ oo o rf^ Millipeds Ol o 01 O5 Animal ^, M food ^ . Sow bugs ^ ^ Cutworms and caterpillars W W 4- co Vegetable 00 co co food 446 University of California Publications in Zoology [VOL. 11 INDICES OF AVAILABILITY OF COMMON INSECTS EATEN BY WESTERN MEADOWLARKS No. of Cut- Grass- Bees and Locality birds Beetles worms hoppers Crickets Bugs wasps Ants Live Oak, gutter Co. 60 1,775 288 2,162 4 533 78 18 Hollister, San Benita Co. 60 4,307 7 1,492 6,680 197 265 35 The most available and the most popular food of the western meadowlark, if it may be judged by the frequency with which it has been found in the stomachs, is beetles. Seventy-five per cent of all the stomachs examined contained beetles (Coleoptera). Vegetable food in the form of oats is next in order of frequency, thirty-four per cent of the meadowlarks examined having taken oats as food. Grasshoppers, cutworms and caterpillars, and weed seeds were found in thirty per cent, twenty-six per cent, and twenty-five per cent respectively of the stomachs examined. The accompanying table gives a summary of the number of times each article of diet was taken and the percentage of each article in the diet of the species. Attention should be called to the fact that a very large percentage (seventy per cent) of the birds taking oats took wild oats (Avena fatua) instead of the tame varieties. Consequently not more than ten per cent of the birds examined had taken cultivated varieties of oats. Although the percentages showing the proportionate number of times each kind of food is taken to the number of birds exam- ined differs from the percentages showing the proportionate vol- ume of each kind of food, yet they parallel each other to a considerable extent. The accompanying table, giving only th^ principal articles of diet, shows this parallelism in the per- centages. IV. PERCENTAGE-VOLUME AND PERCENTAGE OF WESTERN MEADOWLARKS TAKING DIFFERENT ELEMENTS OF FOOD 1 II * ! 03 03 ,J3 CO Q, 11 I S I ii 35 6 & ~ 6 5 ""