ALBERT 
 LIB 
 
 CORNELL UNIVERSilll 
 
 V 
 
ALBERT R. MANN 
 LIBRARY 
 
 AT 
 
 CORNELL UNIVERSITY 
 
Natural History 
 of the Honeybee 
 
 Second Edition 
 
>mt ((atfap of Agriculture 
 
 At djontcll Uhitteraity 
 
 Jtfcaca, &. f. 
 
 Stbrari} 
 
Natural History of the Honeybee 
 
 or 
 
 Are Bees Reflex Machines? 
 
 By 
 
 H. v. Buttel-Reepen, Ph. D, 
 
 Translated by Mary H. Geisler 
 
 Second Edition 
 
 THE A. I ROOT COMPANY 
 Medina, Ohio, U S. A. 
 1917 
 
A translation of ' ' Sind die Bienen 
 Renex-machinen?" by Dr. H. v. Buttel- 
 Reepen. This paper first appeared in the 
 11 Biologischen Centralblatt, " Vol. XX., 
 1900, and was reprinted, with the addition 
 of a table of contents and index, in the 
 same year by Arthur Georgi, of Leipsig. 
 Footnotes in this English edition which 
 are marked " E. F. P." are by Dr. E. F. 
 Phillips, United States Department of Ag- 
 riculture. 
 
TABLE OF CONTENTS. 
 
 Introduction l 
 
 The Hive Odor and the Reactions Resulting from it 4 
 
 Modification of Reactions toward the Hive Odor 5 
 
 The Swarming-out of a Queenless Colony 5 
 
 Intensifying the Reaction 6 
 
 Overcoming the Reactions toward Hive Odor 6 
 
 The Odor of the Queen 7 
 
 The Brood Odor 9 
 
 The Indifferent Odor of Young Bees 9 
 
 Failure of the Hive-odor Reactions in Queens and Drones 10 
 
 Abnormal Hive Odor 10 
 
 The Means of Communication in Bees 12 
 
 Investigations with Colonies from which the Queens are Taken 12 
 
 Behavior of a Queenless Swarm 14 
 
 Disregard of a Queen in Open Air 14 
 
 Hearing Capacity and Sensations of Sound Present 15 
 
 Experiments on Swarms 15 
 
 The Infecting Influence of the Swarm-tone 16 
 
 The Enticing Note of Bees 16 
 
 The ' ' Teeting ' [ and ' ' Quahking " of a Queen 17 
 
 The Queen 'a Tone of Fear 17 
 
 Memory of Place in Bees 19 
 
 The ' ' Paths ' ' of Bees and their Direction 19 
 
 Disappearance of the Memory for Location through Narcotization 23 
 
 The Box Experiment 23 
 
 The Loss of Memory for Location through Swarm Dizziness, etc 27 
 
 Associations of Impressions 28 
 
 Memory for the Feeding-place in the Hive 28 
 
 Conduct of Bees in the Buckwheat Season 29 
 
 Are Bees Attracted by the Color of Flowers or by the Nectar? 29 
 
 Place Perception in the Queen 29 
 
 Memory for Locality in Scouting Bees 30 
 
 The Eyes of Bees 32 
 
 The Flight of Orientation 32 
 
 The Finding of the Hive through the Senses of Sight and Smell 33 
 
 Bethe 's Tree Experiment 34 
 
 Special Capacity for Orientation in Bees 36 
 
 The Influence of Weather Conditions on the Sense of Sight, therefore on the 
 
 Ability to Orient 37 
 
 Influence of Color on Bees 38 
 
 Some Further Contributions to the Natural History 40 
 
 The Flight of Bees into a Room 40 
 
 The Behavior of Bobbing Bees 41 
 
 The Origin of Hostile Conduct 40 
 
 The Vanishing of Instincts with the Decrease in the Strength of a Colony 41 
 
 Reactions toward Flight 41 
 
 The Formation of the Honey-comb 42 
 
 The Play Instinct in Bees 43 
 
 Conclusions 44 
 
 BlBLIOCxRAPHY 46 
 
INTRODUCTION. 
 
 The following observations are submitted after fifteen years' study, and are further to 
 be used for a General Biology of the Honey-bee; but this larger work can not be completed 
 until later, because of the heavy demands on my time at present. It seems desirable to 
 publish this much of the investigation now, however, and I wish specially to consider the 
 most vigorously debated question of the day, that of the psychical faculties of social insects, 
 because of Bethe 's * interesting study of this question with regard to ants and bees. I shall 
 take up bees only, as there are many scientific observations on ants 2 and as especially the 
 great myrmicologists Aug. Forel 3 and Wasmann 2 have already overthrown many of Bethe 's 
 conclusions. 
 
 That I do not accept Wasmann *s definition of Instinct will be apparent from what 
 follows. I refer to the discussion of instincts in my " Stammesgeschichtlichen Entstehung 
 der Bienenstaates, Leipzig, 1903, as well as to Forel 'a excellent paper, "Gehirn und Seele, " 
 5. u. 6. Aufl., Bonn, 1899, pp. 34 and following. 
 
 As early as 1872, Dohrn 4 recognized that it would benefit science if more biological 5 
 investigation were carried on, and the results made useful practically. It seems to me that 
 the biological knowledge concerning Apis mellifica 6 which has been gained by practical 
 bee-keeping has scarcely entered scientific literature, and, strangely enough, the results 
 are little regarded; it has not passed over into the flesh and blood of science. In proof 
 
 1 Albrecht Bethe, Durfen wir Ameisen und Bienen psychische Qualitaten zuschreiben ? Arch. f. die 
 ges. Physiologie, Bd. 79, 1898. (Also appears as a separate with other paging.) 
 
 2 Wasmann, Die psychischen Fahigkeiten der Ameisen. Stuttgart, Erwin Negele, 1899. 
 
 3 Forel, Fourmis de la Suisse. Nouveaux memoires; de la societe Helvetique. Zurich, 1874. Expe- 
 riences et remarques critiques sur les sensations des Insectes. Rivista di Scienze Biologiche. Como, 
 1900, 1901. The Psychical Faculties of Ants and some Other Insects. Smithsonian Report for 1903, 
 pp. 587-599. Washington, 1904, etc. 
 
 Lubbock. Ants, Bees, and Wasps. International Scientific Series. New York, 1883 ; German trans- 
 lation, Leipzig, 1883. 
 
 Janet. Etudes sur les Fourmis, les Guepes, et les Abeilles. Limoges, 1897, etc. 
 
 Emery. Die Entstehung und Ausbildung des Arbeiterstandes bei den Ameisen, Ueber Entstehung 
 des Soziallebens bei Hymenopteren. Biol. Centralbl., Bd. XVI., 1894. Le Polymorphisme des Fourmis 
 et 1' alimentaire. Compt. rend. III. Congr. internat. d. Zoo>l. Leyden, 1896, etc. 
 
 Wm. Morton Wheeler. The Compound and Mixe4 Nests of American Ants. American Naturalist, 
 Vol. XXXV., 1901. Some Remarks on Temporary Social Parasitism and the Phylogeny of Slavery among 
 Ants. Biolog. Centralbl., 1905. Ethological Observations on an American Ant. Journal fur Psychologie 
 und Neurologic Leipzig, 1903, etc. K. Escherich. Die Ameise. Braunschweig, 1906. 
 
 Bethe, 1. c. «. 
 
 4 A. Dohrn. Der gegenwartige Stand der Zoologie, etc.: XXX. Bd. Preuss. Jahrb., 1872. 
 
 5 The German word Bioldgie refers particularly to the study of life-history and habits rather than to 
 all phases of animal and plant life — the sense in which it is used in English. — E. F. P. 
 
 c Apis mellifica L., or Apis mellifera L. ? As is well known, the rule of priority begins as far back 
 as the tenth edition of Linnaeus' "Systema Naturae," in 1758. In that year we find the honey-bee 
 desigated by Linnaeus as Apis mellifer^. Three years later he named it Apis mellifica, probably because 
 he recognized that the name first given was erroneous, for the bee does not carry in honey but nectar, 
 making the honey in the hive. It is, therefore, not a carrier of honey (mellifera) , but a maker of 
 honey (mellifica) . Be that as it may, the first author himself undertook the correction after a fhort time. 
 Then developed an overwhelmingly rich literature concerning Apis mellifica (not meUifera), which is now 
 inexhaustible, even to the specialists, if the non-scientific literature on bees be included. The bibliography 
 of the late Mr. Edward Drory, of Berlin, relative to the. honey-bee alone, embraces, for example, more 
 than 2500 works. In the catalogue "Elenchus Librorum de Apium Cultura," Bibliographia Universale de 
 Apicultura raccolta per Augusto Keller, 2300 works on mellifica are cited. 
 
 Under the circumstances Friese (as writer on Apidae for the Tierreich), and I have thought that, 
 in spite of the rule of priority, we would not be justified in re-establishing the name "melMfera*," which was 
 recognized as incorrect, and shortly rejected by the first author. This is the result of manure delibera- 
 tion at various times during many years. Not only were the reasons just mentioned arguments for 
 retaining the name adopted 150 years ago, but also the knowledge that its establishment proves how 
 powerless the regulations of the priority rule itself are in definite cases. They are powerless in spite 
 of the rules for the "Tierreich," in spite of Delia-Torre's "Catalogue Hymenopterorum," and even in 
 spite of Friese and othera of the originators of the priority rules for each case. For example, for the 
 genus of solitary bees "Anthrophora," the name "Podalirius" was introduced, or at least according to 
 the new system an attempt was made to introduce it. But here the case was like that of melUfica — the 
 first author himself, for certain valid reasons, changed the name after a short time from. "Podalirius" 
 to "Anthrophora." Thus the name Podalirius did not prevail. It has not succeeded in becoming estab- 
 lished in the lapse of years, and there is not the slightest probability that it will change in the future. 
 I qxiote this discussion from my work which has just appeared, "Apistica. Beitrage zur Systematik Biologie, 
 sowie zur geschichtlichen "und geographischen Verbreitung der Honigbiene (Apis meUifica L.), ihrer 
 Varietaten und der ubrigen Apis-Arten." Mitth. d. Kgl. Zoolog. Museums in Berlin, 1906. 
 
2 Natural History of the Honeybee. 
 
 of this there are the vague, defective assertions which are found in the newest editions of 
 scientific works. Thus in a well-known text-book of zoology we read that a calony of 
 bees contains about 10,000 workers. Such a small colony as that, 1 kilo of bees (5009 
 generally counted to a half -kilo) is not capable of developing under ordinary circumstances, 
 nor of Irving over the winter. If a colony of medium strength is considered normal, then 
 a normal colony contains at least 20,000 to 30,000 bees. The following statement also is 
 wrong: "If a young queen emerges from her cell, then the former queen leaves the 
 hive with part of the colony (first swarm) to found a new colony." Normally the swarm 
 has issued^ by the time the cells containing young queens are sealed. Since' it tfikes 16 
 or 17 days for the complete development of a queen, and the cell iar closed on the ninth 
 day, the hive is without an emerged queen for seven or eight days after the issue 
 of the first swarm. The after-swarm, therefore, normally issues seldom before the ninth 
 day after the issue of the first swarm. In Beehhold's Lexikon der Naturwissenschaften, 
 1894, there are similar incorrect statements. Among other things, a vigorous colony is 
 said to contain, at most, 30,000 workers. But since strong icolonies can sustain swarms of 
 three or four kilos (27,000 to 36,000 bees), it follows that the whole number of inmates 
 may amount to 60,000 or 75,000 and more, for usually a little more than half remain in the 
 hive. Only 9000 swarming bees are counted to the kilo, because the honey-sacs of all are 
 filled. Claus, 8 in a paper which contains many other errors, doubts that bees are able 
 to hear. No zoologist who has done any experimental bee-keeping can have the least 
 doubt that bees have an excellent sense of hearing, since observations yield him hundreds 
 of proofs. The man who is not familiar with biological facts might recognize nothing of 
 the kind with certainty, for up to the present the organ of hearing has not been discovered. 
 
 It appears, however, that 1 , the work of Otto Schenk indicates an advance in this direc- 
 tion (Schenk: Die antennalen Hautsinnes-organe einiger Lepidopteren und Hymenopteren, 
 Zool. Jahrb., 1902). In this work on antennae the presence of organs is demonstrated, 
 which, with more probability than has been possible heretofore, may signify organs of 
 hearing. Again, the work of Vitus Graber, * ' Die Insekten, ' ' Miinchen, 1879, in picturing 
 the social life of insects, shows a series of errors and incorrect assertions so far as bees 
 are concerned (Part II., 2d half, pp. 232-248). A refutation would take up too much 
 space, and would lead me too far; but I might mention, for example, that Graber declares 
 that the queen normally flies about inside the hive, because she "can not always pass per 
 pedes through the vertical paths inside" (1. c, p. 88). Further, Wundt (Vorlesungen uber 
 die Menschen- und Tierseele, Leipsig, 1892), buildjs numerous far-reaching theories upon 
 an incorrect biological basis. I shall speak of this in my l i Conclusion ' ' to this paper. The 
 foregoing list of errors is not very complete, but let the matter end here. 
 
 While the founders 9 of our present knowledge in this sphere depended largely upon 
 the results gained in practice, we see now and then an almost complete ignorance of the 
 rich material which distinguished bee-keepers have laid down in the literature of practical 
 bee-keeping. 10 
 
 As to terminology, I shall not use Bethe's phrase, "psychic qualities" (Psychische 
 Qualitaten), since it can be interpreted in various ways. Bethe gives it a definite significance. 
 
 8 Claus, Der Bienenstaat. Part 179 of the collection of scientific papers published by Virchow und 
 Holtzendorf. Hamburg, 1873. 
 
 9 Francois Huber, Nouvelles Observations sur les Abeilles. Geneva, 1814; German by G. Kleine, 
 Einbeck, 1856; English editions in 1823 and 1841. This investigator, blind since his twentieth year, could 
 not have carried on his observations without the help of his unusually efficient bee-keeper Burnens. 
 
 v. Siebold Wahre Parthenogenesis bei Schmetterlingen und Bienen, Leipzig, 1856, etc. 
 
 Leuckart, Zur Kenntnis des Generationswechsels und der Parthenogenesis bei Insekten. 1858, etc 
 Surely the histological findings of these last-named investigators with regard to the presence of spermato- 
 zoa in bee eggs can no longer be esteemed as free from objections. They doubtless depend upon. an{ 
 illusion, for, as I have shown before (Der Befruchtiungsvorgange im Bieneaiei, Bienen w. CentraJblatt 
 Nr. 16, v. 15 August, 1899; further in Aus den Wundern des Bienenstaates, ibid, 1900), according to 
 my investigations, the spermatozoa undergo the transformation into sperm nuclei in about 15 to 20 
 minutes (cf. Weismann, Vortrage uber Deszendenz Theorie, II. Aufi., 1904, p. 250, I. Bd.) ; while v. 
 Siebold says he has seen living spermatozoa in eggs after 12, 15, and 22 hours. I believe, therefore, 
 that in Blochmann's paper (Ueter die Zahl der Richtungskorper bei befruchteten, und unbefruchteten 
 Bieneneiern, Morphol, Jahrb., 15 Bd., p. 85-96, 1889), and in Paulcke's investigations (Zur Frage der 
 parthenog. Entstehung der Drohnen (Apis mellif.), XVI Bd., pp. 474-476, Anat. Anzeiger v. 5 October, 
 Jena, 1899, Vorlaufige Mitteilung),, we have the first safe histological corroborations of the theory of 
 parthenogenesis in the honey-bee. In the meantime Petrunkewitsch has gwren full confirmation (cf. 
 Petrunkewitsch, Alex., Natural and Artificial Parthenogenesis, Amer. Naturalist, Vol. 39, 1905). 
 
 10 v. Siebold (1. c, p. 57) expressly refers to the findings of bee-keepers as "the most important work." 
 
Xatural History of the Honeybee. •> 
 
 comprehending under it everything acquired in the life of an individual, and also every 
 capacity for sensation and learning; in short, everything that transcends unperceived reflex 
 activity. 11 He considers ants and bees as mere reflex; machines: "It seems to me that 
 these insects have no senses, have no ability to make experiences and modify by it their 
 actions; that all stimuli remain below the threshold of perceptible sensations and percep- 
 tions, and that they execute, in a purely mechanical way, all the apparently reasoned 
 actions " (Bethe, 1. c, p. 98). Whether this view is warranted, we shall see further on. 
 
 The terms " reflex " and "instinct, 7 ' I shall use in the significance Bethe gives to the 
 word ' ' reflex. ' ' Instinct is complicated reflex. 12 He lets pass as reflex the inherent faculties ; 
 the course of nervous processes is determined by inherited instincts. That acquired in the 
 life of the individual shows the opposite; here the course of the nervous processes indicates 
 experience, memory, learning, capability for association, etc. 13 I shall restrict myself here 
 to these short statements, but shall add some psychological views in the concluding chapter. 
 
 I shall first discuss the "Hive Odor" and reactions resulting from it, then give my 
 experiments and opinions on the capacity for intelligence in bees, and end with my con- 
 clusions concerning the homing instinct in bees, or the ability to find the hive, etc. 
 
 13 Bethe drops this expression in '-Beer, Bethe, und J. v. Uexkull, Vorschlage zu einer objectivirenden 
 Nomenklatur. Biol. Centralbl., 19 Bd., 1899, Nr. 15, p. 517 (also in Centralblatt fur Physiologie, 1899, 
 Nr. 6) ; and further in Bethe's " Noch einmal uber die psyehischen Qualitaten der Ameisen," Arehiv. f. d. 
 ges. Physiologie, Bd. 79, 1900, p. 45. 
 
 12 H. E. Ziegler, Ueber den Begriff des Instinkts. Verhandl. d. deutsch. zoolog. Gesellsch., 1892. 
 
 - 3 H. E. Ziegler, Theoretisches zur Tierpsvchologie und vergleichenden Neurophysiologie. Biol. Centralbl., 
 Bd. XX., Nr. 1, 1900. 
 
THE HIVE ODOR AND THE REACTIONS RESULTING FROM IT. 
 
 In the first section of "Investigations on Bees, " Bethe (1. c.) treats the question as to 
 how bees recognize the hive, and concludes that the recognition results solely from an 
 odorous chemical compound (Geruehsstoff) which he calls the "hive substance" (Nest- 
 stoff). 14 He avoids the term "hive odor," because he thinks bees have no sense of smell. 
 I use the expressive term "hive odor" (colony odor, exhalation odor) under which I com- 
 prehend only the existence o-f gaseous substances disseminated in a colony, the presence 
 of which is detected by the bees ' sense of smell. 
 
 I believe that the following odors are present in a colony of bees: 
 
 1. The Individual odor. It can be easily 'demonstrated that the queen odor (see p. 7) 
 varies with different individuals, and on the same ground .(germinal variation), an indi- 
 vidual odor should be assigned to the workers. 
 
 2. All offspring of one mother (queen) have a common inherited family odor in addi- 
 tion to the individual odors, belonging only to the progeny of one queen. 
 
 3. The brood and chyle odor (p. 9). 
 
 4. The drone odor (p. 10). 
 
 5. The wax odor. Since the wax is a granular secretion, an exuded product, it may be 
 safely taken for granted that, considered apart 1 from the specific odor of wax, the indi- 
 vidual odors of the wax-generators adhere to the honey-comb. Accordingly the wax 
 structures of different colonies have different odors. 
 
 6. The honey odor. That the honey of each colony (mixed with a secretion of the 
 salivary glands) has its specific odor is readily seen from the old practice of bee-keepers 
 to which Bethe also ^alludes. If a queen be daubed with honey from ai queenless colony, 
 she will be accepted readily by that colony when inserted. 
 
 7. The hive odor (exhalation odor, colony odor). The hive odor is composed normally 
 by a mixture of the preceding odors, or of some of them. Single bees, therefore, besides 
 their individual odors, possess the family odor and especially the common adhering hive 
 odor, which forms the dominant factor in the various actions toward hive mates and 
 hive strangers — that is, in mutual recognition between bees. 
 
 If a strange queen is put into a queenless colony in a cage, a confinement of twenty- 
 four hours is usually sufficient for the queen tk) have received the ' ' hive substance, ' ' as 
 Bethe calls it; that is, she has become "scented." 15 The external adherence of the colony 
 odor therefore suffices to make the bees permanently friendly with the foreign queen. If 
 a swarm is made of the bees hanging in front of several hives, 16 which in strong colonies 
 often form great clusters (beards), by scooping them up in a ladle, there will probably 
 be bees from ten, twenty, or thirty different hives. If they are put into a hive and given 
 a queen, with the observance of proper precautions, these bees from, let us say, thirty differ- 
 ent hives, form a peaceful colony in a few hours, which adjusts itself in its new dwelling 
 and takes up the ordinary tasks of the day. Here we have thirty family odors, and about 
 
 14 According to Bethe we must consider the existence of this "hive substance" as a 'family odor." 
 "I believe that these family odors, common to all the members of one family, and differing slightly from 
 those of other families of the same species, play an important part in the life-history of the social hymenop- 
 tera. This family difference is due to the varying proportions of the constituent odors" (1. c, p. 3*1). 
 This "hive substance" must be inherited, as Jager likewise maintains (ZeitschrSft fur wissensch. Zoologie, 
 Bd. 27, 1876, p. 327). 1 might mention here that I find no proof in Jager's paper that he concludes the 
 "exhalation odor" of a colony to be solely a "f amity odor" as Bethe does, and still less proof that he 
 considers this common odor to be "inherited," because it is a mixture of many thousands of inherited indi- 
 vidual odors. 
 
 Bethe includes, under the term "hive substance," two separate substances, one giving the family odor 
 and the other causing the "various reactions toward hive mates and hive strangers." That this conception 
 will not cover all cases, I believe I can demonstrate in the course of my paper. 
 
 35 Further on I shall show that in many cases the bees very likely became familiar with the odor of the 
 queen, whir-h is a mixture of the queen odor and the hive odor. 
 
 16 G. Dathe, Lehrbuch der Bienenzucht, 5 Aufl., Bensheim, 1892; Bienenzeitung, 7 Jahrg., No. 19. 
 
Natural History of the Honeybee. 5 
 
 thirty to forty thousand individual odors united into a special hive odor, peculiar to this 
 colony alone; merely by the scooping together. The proof of this can be found in the 
 fact that a queen placed in such a colony can be set free frequently after only twelve to 
 twenty-four hours; she has taken up the hive odor. 
 
 We see, therefore, in this case, a colony without a specific family odor conducting 
 itself exactly like colonies which are made up of the offspring of one mother. In each 
 case the hive odor is formed by the union and mixing of individual odors. 
 
 Now the question arises: Is this hive odor inherited? No, surely not, for the hive 
 odor, therefore the common odor, that each individual can take up in an external way is 
 something purely exogenous or acquired. 
 
 The power of the hive odor seems to me, therefore, to lie not in the inherent family 
 odor nor in the inherited individual odors, but in the exogenous mixture of the two. 
 What Bethe (1. c, p. 43) alleges for compound nests (Schuttelnester), and communicates 
 in his experiments on bees (p. 71), is not a contradiction of this. He says that bees 
 that had been transferred to another hive did attack almost not at all or with only slight 
 enmity their sisters remaining in the mother hive because the common family odor les- 
 sened the hostile reaction. This proves merely that the hive odor differs from the family 
 odor. If the latter were the predominant factor, bees of the same family would always 
 be friendly with each other, which is not the case. 
 
 Further, I can not regard this, the only experiment which Bethe employed (1. c, p. 71), 
 as conclusive, since very many circumstances which have nothing to do with the hive odor 
 affect the friendly or hostile attitude of bees, be they sisters or strangers. The forage, 
 weather, time of year, strength of the colonies, quantity of provisions, etc., all have an 
 influence, as is well known to those who have studied the peculiarities of bees for many 
 years. In my experience, estrangement seems to take place between separated hive mates 
 generally much more quickly than Bethe states, yet I think this is on the whole a second- 
 ary matter which does not touch upon the principal point. 
 
 MODIFICATION OF REACTIONS TOWARD THE HIVE ODOR. 
 It is contested by Bethe that the " various reactions" are based upon hive-mate and 
 hive-stranger modifications. 17 It seems to me that it is hardly possible to grant such a 
 far-reaching conclusion when Bethels investigations give no striking proof that reactions 
 toward the hive odor are incapable of modification. We learn nothing from the illus- 
 tration g^iven above. In the critical examination of robbing between colonies, we find that 
 a great number of factors enter, which can be discovered only after years of keen obser- 
 vation and fortunate circumstances. In the manual of bee instruction which I helped to 
 publish 18 (see p. 9), I recommended also fumigation of the hive of the robbing and not of 
 the robbed colony, but that is no proof that the hive stimulus can not be modified. Bethe 
 believes that no psychic elements enter into the actions of a colony and apparently the 
 short time he could devote to bees led him to this view. I submit the following investiga- 
 tions on this question. 
 
 THE S WARMING-OUT OF A QUEEN LESS COLONY. 
 If two hives are placed close together, and the queen and brood are removed from 
 one, it sometimes happens that the entire colony, from which every possibility of rearing 
 a queen has been removed, will enter the queen-right 19 (weiselrichtigen) colony, humming 
 
 17 "It has teen thought possille to make the individuals of a hive 'recognize' each other better by 
 fumigating the colony with some strong-smelling substance — Camphor, naphthaline, baldrian, for instance. 
 If this be done to a colony exposed to frequent plundering it is thought that the tees of this hive will 
 more readily detect the robbers to whom the scent does not adhere. Were this correct, then it would 
 prove that the various reactions toward hive mates and hive strangers could be artificially modified. 
 
 "I believe that I can assert positively that such treatment does not increase the reaction toward 
 strange bees in the least, but only that the bees of all strange colonies react more vigorously toward the 
 individuals thus scented. (Therefore if it is desired to defend a colony from robbing, it is the robbing 
 and not the robbed which must be fumigated.) 
 
 "We see, therefore, that here, as in ants, the various reactions toward hive mates and strangers bring 
 us back to a simple chemical reflex" (Bethe, 1. c, p. 71). 
 
 18 G. Dathe, Lehrbuch der Bienenzucht, 5 Aufi., published bv R. Dathe and H. Keepen (v. Buttel- 
 Reepen), Bensheim, 1892, p. 181. 
 
 19 There is no word in common use among English-speaking bee-keepers to indicate the normal con- 
 dition of a colony in the possession of a queen, either mated or i:nmated. It eeems desirable, therefore, to 
 translate the German term literally, although " queen'right " is an undenraHe term. It has already been 
 used by some American writers. — E. F. P. 
 
6 Natural History of the Honeybee. 
 
 "joyfully." These bees seldom sting, and are received in a friendly manner, although the 
 normal queen-right colony should react hostilely, if it followed a chemical reflex inca- 
 pable o£ modification. What causes this swarming-out of the queenless colony we shall 
 see further on. 
 
 INTENSIFYING THE REACTION. 
 
 If a colony tolerates robbing without attempting to suppress it, a stimulating food 
 should be given it, such as fermenting buckwheat honey 20 several years old or a mixture 
 of honey and branny, and the hive should be shaken to arouse the anger of the bees,, etc. 
 Then a better defense is made, the irritability of the colony is increased, and robbers are 
 better recognized and repulsed. 
 
 The "courage" and the "attention" of the colony are increased, and therefore we 
 have intensified reactions toward hive strangers. 21 
 
 OVERCOMING THE REACTIONS TOWARD HIVE ODOR. 
 
 It is interesting that this powerful reaction toward hive odor, though generally act- 
 ing hostilely, can be subdued and turned in other directions. Thus bees* which blunder 
 into the wrong hive, as they return from the fields with filled honey-sacs, are seldom 
 attacked. 
 
 The behavior of such "begging" bees is most peculiar. With the abdomen dragging 
 on the flight-board, the proboscis far extended and dealing out the honey to her tor- 
 menters, she is trying to insinuate herself into the hive in order to deposit her honey, 
 and in this she often succeeds. (Here and further on I use anthropomorphic terms for 
 better demonstration. It is understood that many of these actions rest upon reflexes 
 and instincts, but it make's the case clearer to use such terms, as "to insinuate" for ex- 
 ample.) 
 
 The old practice of transposing hives is based on the fact that strange bees, laden 
 with honey, are received in a friendly way. If a weak colony needs improving, it may 
 simply be put in the place of a very strong one, the two hives changing places. This 
 transfer will be successful only on days in which there is a good honey-flow and all flying 
 bees return heavily laden. If the hives have the same outward appearance, and (most 
 important) have their entrances at the same height, almost all bees will fly in as usual. 
 Thus the weak colony will be strengthened from the strong one, and stinging seldom occurs. 
 
 Also, if one wishes to prevent swarming, the flying bees may be drawn off in the 
 same way. I can not at this point go into the many modifications of this practice, and 
 must refer to the manuals of instruction. 
 
 If a strong colony stands next to a weak one during a rich honey-flow, and succeeds 
 in filling its own chamber to overflowing, it happens now and then that the strong colony, 
 having no further room, will help to fill the hive of the weak one yi the most peaceful way. 
 So. we see that of two colonies which should react hostilely toward each other, either one 
 or both may be put under peculiar conditions in order to handle both in the same way with 
 a peaceful reaction between them. For example, if two colonies are sprinkled with some 
 strong-smelling liquid, or powdered with meal, they may be united without risk. 
 
 If bees are stupefied with saltpeter, chloroform, ether, or puff-ball, the same thing 
 can be done with no danger of mutual stinging, but at the same time the earlier orientation 
 is completely lost. When the bees recover from their stupor they no longer recognize their 
 own colony, and can be united to other colonies at will. What conclusions are to be 
 drawn from this capacity for learning, etc., we shall see later on. 
 
 20 Dathe, 1. c, p. 179. 
 
 21 The Luneburger bee-keepers sometimes employ a particular method for putting a stop to robbing. 
 If the honey-flow is strong, so that the powerful odor of honey issues from the hive-entrance, la general 
 robbing occasionally goes on ; but the robbers are hardly noticed by the bees of the hive, apparently the 
 strong odor of honey covering the strange odor of the robbers, or the certain, quiet entrance deceiving the 
 inmates of the robbed colony. In order to call out a stronger reaction, the bee-keepers turn over the 
 skeps so that the wider opening of the skep is turned to the front. One would think that, since the whole 
 honey-comb is now open to the air, robbing could be effected more easily; but this is not the case, for the 
 powerful odor of honey is now easily "dissipated," and the robbers, because of the change in the position 
 of the entrance, are uncertain, and hesitate. Thus the attention of the colony is drawn, to the robbing, the 
 fortification is well protected, and every attack is beaten off (cf. G. Lehzen, Hauptstucke der Luneburger 
 Bienenzucht, 1900, Hannover). 
 
Natural History of the Honeybee. 7 
 
 Incidentally, I might mention here a curious aberration of instinct. The literature 
 ©f bee economics records some cases in which bees showed unaccountable hostile reaction 
 toward their own hive mates. Thus it has been observed sometimes that a colony will 
 sting a great number of its own flying bees as if they were intruders. This behavior has 
 been noticed for a long time, and it finally did lead to the destruction of the colony. Possibly 
 here -there is a degeneration of that instinct which causes colonies to place bees at the 
 entrance to control the flying in. Perhaps differences of odor come into account, or perhaps 
 both stimuli act together. 
 
 It is probable, too, that we may be dealing with defective observations, although errors 
 of instinct in bees are not at all uncommon, and frequently in the spring it can be proven 
 that bees, in an inexplicable mistake, even attack their own queen and ball her. The colony 
 is then in wild confusion, and between the combs or on the bottom-board is the poor queen 
 in the middle of a cluster of bees; she often comes out of the tumult a cripple. Another 
 ease of failure in instinct is as follows: As is well known, bees now and then build queen- 
 cells over drone eggs. 22 If a queen-cell is of more than ordinary length it is tolerably sure 
 to contain a drone larva. 23 The too invigorating royal jelly ** seems to be unwholesome 
 for the larva; it falls out from the food because of the abnormal size of the cell, and that 
 is the cause of their always lengthening the cell. Usually the drone dies. 
 
 THE ODOR OF THE QUEEN. 
 
 The individual odor of the queen is doubtless in many cases a very small part of the 
 odor of the hive, but often may form the dominant factor. The exhalation of the queen 
 is so intense that it can be perceived by man. It is very characteristic and adherent, being 
 somewhat like the odor of thyme. If a queen is crushed on a board, the bees of her colony 
 smell for several days around the place where she was killed. If the 1 bees are allowed 
 to run over the board, they gather there; and, lifting the abdomens, fan their wings in a 
 peculiar way. 
 
 It often happens that after-swarms, also swarms with young queens, fly together and 
 unite into a powerful swarm-cluster. The bees in this cluster do not attack each other, 
 in spite of the varied hive odors. The ' ' swarm-dizziness ' , extinguishes the reactions 
 toward the foreign hive odor, just as it also almost destroys the sense of orientation, so 
 that the impulse to seek the parent colony is lost, 25 at least under normal circumstances. 
 The swarming bees, instead, remain in the home they have taken up; the field bees, which 
 some days before, or immediately before the swarming, have been bringing honey, pollen, 
 and water to the parent colony in the usual way, will, a few hours later, after they have 
 become oriented, bring their burdens into the new hive. Under the proper conditions this 
 can be placed adjoining the old one. The memory of the old birthplace has completely 
 disappeared. I shall speak later on of an exception to this. 
 
 If it be wished to separate these united swarms, the whole cluster may be put into a 
 large box containing as many twigs of a tree as there are swarms. Over night the colonies 
 separate of themselves, each hanging on a twig. 
 
 It is safe to take for granted that a purely mechanical separation takes place here, 
 and, evidently, according to the various hive odors. I nevertheless believe that a still more 
 powerfully determinative stimulus enters — the odor of the queen. Each colony congre- 
 gates around its queen; 26 and if the queen is taken away from one swarm it will unite 
 with another possessing a queen, in spite of the hostile hive odor. 
 
 22 O. von Roth, Ueber abnorme Zustande im Bienenvolk. Berichte der Naturf. Ges. Freiburg i. Br., 8 
 Bd., 1894. 
 
 23 There is also a prevalent idea that such cells are not as rough as normal cells containing queens; 
 some books on bee-keeping even go so far as to try to figure the difference. — E. F. P. 
 
 24 Concerning the varied chemical composition of pabulum for the three kinds of bees, see A. von 
 Planta. "Uber den Futtersaft der Bienen," and "Nochmals uber den Futtersaft der Bienen," Schweiz. 
 Bienenzeitung, 1888-'9 ; further, A. von Planjta, Zeitschrift f. phy. Chemie von Hoppe-Sevler, 1888, Bd. 12, 
 Heft 4, pp. 327-354, and also Bd. 13, Heft 6, pp. 552-561; further, Dathe 1. c, p. 24. 
 
 25 Bethe, in dealing with the "psychical qualities," has not considered either the act of swarming, which 
 is an all-important factor, or the individual odor of the queen. "We shall see presently what interesting 
 bearings swarming has on this question. 
 
 26 Some may take exception to this statement, perhaps, and believe that this gathering together is due 
 to other instincts — the sex instinct or the instinct to swarm which is characteristic of bees; but it must be 
 kept in mind that the bees separate into single colonies only if the queens of the various colonies are present, 
 manifesting their presence by the scent which exhales from them. 
 
8 Natural History of the Honeybee. 
 
 It may be thought that here the family odor, the possession of all the offspring of 
 one mother, enters its claim; but I notice that the swarms may be united ingeniously, even 
 if the queen does not originally belong to the colony in question, but comes from a queen- 
 cell taken from a strange colony. In addition, we continually have to do with unfertilized 
 queens in normal after-swarms, therefore we can not speak of the offspring of one mother 
 at all. The queen of an after-swarm is the sister of the workers, if I may so express myself. 
 
 It is the queen odor familiar to the bees which acts with the hive odor (to be sure, 
 the reaction toward the latter seems to disappear during the swarming), and perhaps 
 holds the community together, but the queen odor is the dominant factor. This may be 
 seen from the first experiment (p. 5) also, to which I shall now return. The bees of 
 the queenless colony scent the queen in the neighborhood hive, and, paying no attention to 
 the foreign hive odor (toward which, under ordinary circumstances, they would react 
 sharply), they go over into the enemy's camp ''humming joyfully. " Very probably sound 
 perceptions also come into consideration — that is, the reaction toward the humming of the 
 " queen-right ' ' colony (see later). 27 
 
 If the odor of the queen is so powerful,; it is clear that this individual odor alone 
 determines the special character of the hive substance, especially when this is particularly 
 strong. This would happen in the spring during the increased sexual activity 2S which is 
 demonstrated by the enormous number of eggs laid (2000 to 3500 and more in twenty-four 
 hours). That it is not always the determining', factor, follows from the fact that there 
 remains a hive substance capable of causing reaction, even if the queen is taken out and 
 kept at a distance. 
 
 At such a season, during the strongest breeding period, the bees care for the queen 
 with special zeal. It may not be the very ' ' sympathetic ' ' odor M alone which causes them 
 at this time to surround her in a close cluster, for her increased need of nourishment 3 ° 
 increases the constant care of the bees around her. Nevertheless, the singular pleasure that 
 bees take in the strong mint-like exhalation from the queen is shown in the attachment 
 of the ' ' court, ' ' this affection being demonstrated by the single ' ' courtiers, ' ' who from time 
 to time lick the abdomen or thorax most zealously, humming continually in a characteristic 
 " contented' * way. 31 
 
 The bees wish to sniff the odor, so to speak. A queen in an observation have can be 
 seen constantly surrounded by a circle of brood nurses who continually turn the head to 
 her. If she advances slowly, the bees yield the: way, moving backward. From time to 
 time the "caressing," licking, and feeding go on. It can be easily understood how this 
 attitude has often been interpreted as the expression of a peculiar reverence toward the 
 ' '- ruler of the community. ' ' The ' ' respectful ' ' walking backward, the ' ' tender ' J licking, the 
 irreproachable service (as the queen never leaves the hive, she is forced to deposit her 
 faeces in the hive, and the ' l courtiers ' ' immediately clean away all traces) closely approach 
 anthropomorphic appearances. But perhaps it might be proven that the individuality of the 
 queen is the dominant factor. A foreign queen with the same strong odor of mint would 
 be stung in spite of the pleasure in the smell. 
 
 In autumn and winter, also early in the spring, the bees concern themselves relatively 
 very little with the queen, and still less with a young virgin. The lessened exhalation (it is 
 as yet very weak) may explain this phenomenon in part. 
 
 It seems to me that if the odor of the queen is extremely penetrating and adherent, 32 
 
 27 In this there might be found an apparent contradiction to the "disregard of the queen" p. 14). 
 The finding of the queen in the next hive is not by scenting from hive to hive, for the' distance is too 
 great, The agitated queenless bees run searching over the front wall of their hive and also over the 
 front wall of the one standing next. In this way some come to the entrance of the neighboring hive, 
 where the odor of the queen and the sound of contented humming are issuing forth. At once they begin 
 to lift the abdomens, and fan with their wings. Their mates near by take up the humming, and soon the 
 whole colony enters the queen-right hive in order, with lifted abdomens and fanning 1 wings. 
 
 28 Jaeger, Ueber die Bedeutung des Geschmacks- und Geruchsstoffes. Zeitschrift f . wissensch. Zoologie 
 Bd. 27, p. 327, 1876. ' ' 
 
 29 In order to keep a swarm in the hive, the hive may be rubbed with thyme. 
 
 30 As is well known, the queen can eat only honey independently; but the probosces of the workers are 
 necessary to feed her with nitrogenous food. 
 
 31 I might say again that anthropomorphic designations are chosen for purposes of clearer demon 
 stration. 
 
 S2 I took the queen from a strong colony, put her in a cage, and after a few minutes removed her 
 Fifteen minutes later I placed the empty cage upon the flightboard of the colon v concerned. Imme- 
 diately the bees scented the odor, and alighted, f uttering upon the cage, which thev had complete'- ignored 
 before (see also p. 7). 
 
Xatural History of the Honeybee. ' 9 
 
 as it doubtless is, the view (mentioned also by Bethe) that the queen takes on the hive 
 odor of the queenless colony in which she is placed in a cage, is not correct in all cases, 
 for the colony is "scented" by the strong odor of the queen, which is distinctly perceptible 
 to our sense of smell. Very rightly, therefore, O. von Rath' 3 says that "the workers 
 first accustom themselves to the odor of the queen.' > The stronger, then, the exhalation 
 from the queen, the more easily must be her acceptance, or scenting of the colony proceed; 
 and consequently the mutual friendliness also. I find from many investigations that a 
 mated queen heavily laden with eggs is more easily accepted than an unfertilized one. The 
 following rules 34 of bee-keeping harmonize with this statement: 1. "An unfertilized queen 
 is not accepted by many colonies if formerly they have* had a fertilized one." 2. "The 
 older a queen is, the more readily is she accepted. " 3. " Weak colonies accept queens more 
 easily than strong ones." 4. "Queens of the same kind are accepted with less difficulty 
 than those of different varieties; as for example, Italians, Caucasians, Cyprians, etc." Here 
 in every case the different odor of each variety plays its part. 
 
 It may seem obvious that it is easier to demonstrate this scenting, whether it be on only 
 cne side or mutual, with a weak colony, but here other instincts come into consideration 
 too, as we shall see further on. 
 
 THE BROOD ODOR. 
 
 It is highly probable that it is not only the odor of the queen at all times which 
 causes a queenless colony to enter a hostile camp (see p. 5), but during the monthsi of 
 greatest increase, from March to June or July, a powerful constituent! of the hive odor 
 is the odor of the brood, a warm characteristic vapor exhaling from the thousands of 
 brood-cells. The chemical processes going on at this time in the cells for the development 
 of the larvae are of so intense a nature that) a temperature of 28 to 32° C. is maintained 
 in the brood-chamber. On spring days this exhalation from the brood, which smells like 
 freshly baked bread, may be perceived some distance away in the direction of the wind. 
 Very probably this strong odor sets free the same reaction in queenless and broodless 
 colonies as the queen odor does; but as such colonies always go over to the queen-right 
 colony, even if there is no brood at all or only a minimum amount, so in spite of this 
 vapor from brood and pabulum (the larva? swim in food), it is very likely the queen odor 
 and the humming of the queen-right colony which are of first importance. 35 
 
 Bat, on the other hand, the attachment of the bees to the brood is very strong, as 
 may be seen from the fact that an unruly swarm, which has already withdrawn from one 
 unaccepted hive, will certainly remain if a frame of brood is hung in it. Again, swarms 
 can often be enticed from an undesirable place for capturing them (the middle of a 
 hedge, for example), by means of a frame of brood. 36 
 
 THE INDIFFERENT ODOR OF YOUNG BEES. 
 Young bees, particularly those just emerging, have an apparently faint and indifferent 
 odor, and they are, therefore, not attacked in a strange colony; the same thing is true 
 of young queens. In cutting out queen-cells, it happens repeatedly that the young queens 
 which are entirely or nearly "ripe" free themselves from their compartments. I: such 
 a queen is immediately allowed to run through the entrance of a queenless colony, the 
 queening is usually successful. This depends, perhaps, upon the fact that, as described, 
 the individual odor is not yet developed, analogous to the similar indifferent "infant odor," 
 and that the common reaction-loosing hive. odor has adhered but little so far. It is curious 
 that there are colonies which will not allow themselves to be requeened;"' 7 then all artifices 
 are in vain. 
 
 ■'■ 3 O von Roth, Uel.er abnorme Zustande im Bienenvolk, Beriehte der Naturf Ges Freiburg i Br 
 8. Brl . 1894. - " ' M 
 
 ™ Dath-e, 1. c, p. 211. 
 
 '■" The "constancy" of 1 ees to their queen, which is always the dominant instinct, manifests itself 
 ma Mai\irg' colcny Ly the fait that the queen is always the last to die; she is fed to the end l,v the 
 dying bees. In order to confirm this, I put a queen with a few bees and very little food into a box 
 covered with wire gauze. After forty-eight hours the bees were very weak; two days later only four 
 were still alive; the next day but one was living, while the queen was apparently as vigorous as ever 
 The last surviving worker lay on its side unable to crawl; but when the hungry queen' approached demanding 
 food, it tried feebly to join its proboscis to that of the queen in a vain attempt to give her food Fina'ly 
 the queen turned away; and when I looked again, half an hour later, the last worker was dead but the 
 qreen shoved no sign of weakness. I then put her back in the, colony. 
 
 • T! Dathe, 1. c, p. 225 and 230. 
 
 r7 Bienenwirtsch. Centralblatt, Jahrg. 28, Heft 19, p. 298, 1892. 
 
10 Natural History of the Honeybee. 
 
 However, in the winter months, when the bodily functions which determine the inten- 
 sity of the individual odor are for the most part quiet or weakened, the strength of the 
 individual odor decreases. In consequence of this we have prompt friendliness with 
 strangers, since the lessened odor of the hive excites only a weak reaction. Therefore a 
 union of colonies can be undertaken in early spring without observing the precautions 
 otherwise necessary. * 
 
 I noticed one day a change in two hives standing close to each other, which was totally 
 unintelligible to me. Colony " A, ' ' in which a vigorous increase was to be expected, showed 
 a constant diminishing of numbers, while colony "B M strengthened in a surprisingly short 
 time. By good fortune colony "A" was of the native brown variety, and colony "R, " a 
 yellow Italian hybrid. This color distinction of the varieties, which has helped to solve 
 so many mysteries of the domestic economy of bees, brought the explanation in this case. 
 I was soon sure that colony "B" was taking up the young; from the other hive, and I 
 found out how after long observation. A passage through possible crevices could not take 
 place, nor could the crossing be from one alighting-board to the other; therefore only 
 the flying bees had to be taken! into consideration.* I noticed after some time that the 
 young Italian bees always took their flight of orientation earlier than their neighbors. 
 Now, when a thick cloud of these young bees, humming loudly, would be flying in front 
 of colony "B, ' the native eolony would gradually begin to send out bees for orientation, 
 but its mass of humming bees was always considerably smaller. This was because many 
 bees, attracted by the loud buzzing, immediately plunged into the neighboring tumult, there 
 oriented themselves, and accordingly entered colony "B" thereafter. There they were 
 accepted on account of their indifferent odor, and chiefly, perhaps, because of their entirely 
 "harmless" conduct. 
 
 FAILURE" OF THE HIVE-ODOR REACTIONS IN 7 QUEENS AND DRONES. 
 
 It is of interest that the hive odor of a strange colony causes no reaction at all in a 
 queen. Queens never react either peacefully or hostilely toward strangers or toward bees 
 belonging to the hive. They demand nourishment from every bee, and they maintain them- 
 selves even in the most hostile colony as long as it is queenless. Even the * ' angrily buzzing ' ' 
 bees which besiege the queen-ea'ge in a solid mass, and which try to bite and sting the 
 queen through the wire cloth, put the required f ood «into the extended proboscis of the 
 queen. In this manner a queenless colony will often feed ten or twenty confined queens; 
 but if one should accidentally free herself and be accepted by the colony, then the bees 
 will let the rest starve. 
 
 The queen recognizes as an enemy only her ' l rival, ' ' even if reared in the same colony 
 (daughter or sister), and who, therefore, must have the same family or hive odor. If 
 two queens come upon each other, only one will remain on the battlefield. 88 
 
 If the queen is pleasing to every bee in every colony, the same thing may be said of 
 the drones, who are extremely cosmopolitan, and who loaf about from hive to hive, and 
 in consequence, apparently, of their specific odor, they are received peacefully everywhere, 
 provided, of course, that the killing of the drones has not yet begun. At no time do they 
 display the smallest response of any kind toward other bees, except when they accomplish 
 the object of their existence in the mating -flight. 
 
 ABNORMAL HIVE ODOR. 
 It is worthy of note that drone-producing (fertile worker) colonies, that is, colonies 
 in which the workers take to egg-laying because a queen is lacking, and on account of 
 
 38 The queen is normally the absolute "monarch" in the colony; but in spite of that we not seldom 
 find cases where there are two egg-laying oueens. Here we have the successor encroaching upon, the 
 old decrepit queen before the latter dies. "But under such circumstances there are always two brood- 
 nests — the queens do not come together. The following observation stands alone, and is the more 
 remarkable because it has to do, not only with two queens, but with two of different varieties. "Since 
 I had the opportunity," writes one Mr. Breuer, in the Rheinischen Bienenzeitung, "to obtain a pureV 
 mated Carniolan queen, I took out the old queen on July 17th, and put in the Carniolan. She was 
 accepted without delay and immediately began laying. Another queen was positively not present in 
 the colony. The brood developed cmickly, but I kept noticing among the young bees 'Germans as well 
 as Carniolans. When I revisited the hive I found upon the same frame, hardly five centimeters apart, 
 two magnificent queens, peacefully together — one German, the other Carniolan. The brood-nest was 
 not divided, but jnst as normal as if the eggs had all Teen laid bv cne queen." (See reference in 
 Bienenw. Centralblatt, No. 22, 1899. Hannover.) 
 
Natural History of the Honeybee. 11 
 
 lack of brood, are not only difficult to requeen, but also equally difficult to unite with 
 queen-right colonies. This is doubtless in consequence of the peculiar hive odor called 
 forth by the presence of so many egg-layers whose number increases the longer this 
 abnormal condition lasts. According to the investigations of Dohnhoff, almost all the 
 bees finally lay eggs without conducting themselves differently from the usual non-laying 
 bees. 39 A true queen odor does not seem to develop, and I have observed that drone- 
 laying queens are never rendered the " homage" which a normal queen receives. As long 
 as she is unfertilized, she seems to be unnoticed by the inmates of the hive; but as soon 
 as she begins laying eggs she has around her constantly a ring of "courtiers" (see p. 7). 
 Other colonies very frequently may be united without special precautionary measures, but 
 not so with fertile-worker colonies, which can be joined successfully only by the appli- 
 cation of very special precautions. 40 We therefore have here an abnormal hive odor of a 
 peculiar kind/ 1 • r 
 
 In any case it is evident from the foregoing that the hive odor is exceedingly com- 
 plicated — much more so than would appear from Bethe's account; and the idea of a 
 simple chemical substance and a chemical reflect, incapable of modification, is not enough 
 to clear up the proceedings which are involved. 
 
 89 Bienenfceftiung, XIII. Jahrg., No. 20. In passing, I might submit the following : It has been 
 shown experimentally that, as in the circumstances cited!, the workers (normally sterile) take up egg- 
 laying. I shall here mention only one such investigation, and shall not take up the various anthropo- 
 morphic explanations which are usually given. There is a wide-spread view among bee-keepers that the 
 larvae of workers which are located nearest to the queen-cell are fed occasionally on royal jelly by 
 mistake; on account of this exceptional nourishment, it is believed that a better development of the 
 ovaries may take place. This view is held also by v. Siebold and Huber. (The ovaries of workers 
 normally consist of about twenty to thirty egg-tubes, whereas those of the queen contain about four 
 hundred. Parallel with and partly qualified by this view is the incorrect idea that there is onlv one 
 or a few egg-laying bees in a di one-producing colony; but if one realizes that almost all workers in 
 such a colony lay eggs, this view is already weakened. In my opinion we here have to do with, the 
 same reflexes which induce a colony to produce a young queen during the lifetime of an old weakened 
 queen that is laying eggs in insufficient numbers. What "prudence" and "deliberation" on the part 
 of the bees this suggests to the ordinary observer! Again the same reflexes impel a colonv to erect 
 queen-cells if the queen is kept confined for a long time (see p. 14.) It is, I believe, for the greater 
 part an unsatisfied instinct for feeding. In the first case we have the pabulum, which is produced in 
 great quantity, and which is not reaching] its natural destiny, acting to produce a hyper nutrition of the 
 bees, and consequently the stimulation of organs which normally are not stimulated at all. 
 
 40 Dathe, 1. c, p. 161. 
 
 41 I shall not consider further the special abnormal hive odors generated bv disease (dvsenterv, foul 
 brood, etc.), nor through the irritability over the lack of the queen. 
 
THE MEANS OF COMMUNICATION IN BEES 
 
 According to Bethe, there is not '/the slightest doubt" that bees recognize each other 
 or hive strangers only by odor (chemical substance), and that no special means, neither 
 a " sound" nor a "definite movement of the antenna?/' comes into question as possibilities 
 for communication. 42 
 
 Long-continued and careful observations, however, yield many data which do not 
 coincide with this view. 
 
 ' INVESTIGATIONS WITH COLONIES FROM WHICH THE QUBENS ARE TAKEN. 
 
 If the queen is taken from a very strong colony of 50,000 to 60,000 bees or more, the 
 loss is first noticed sometimes after an hour, sometimes after many hours. This is par- 1 
 ticularly the case if the removal is made during a rich honey-flow when the bees are 
 busily occupied with bringing in and storing the nectar. A striking change then suddenly 
 takes place, the comfortable humming gives w T ay to a louder, long-drawn-out, lamenting 
 buzz. 4,0 The guards at the entrance and those providing the ventilation become uneasy; 
 excited bees come out of the hive and run over it as if seeking something; single bees fly 
 away quickly, then back; the whole character of the colony is changed — not only in outward 
 behavior but also in inner disposition. They are -very irritable, and inclined to sting. 
 I have sometimes noticed the queenlessness of colonies, which usually were very gentle, 
 by the increased desire to sting. Such notice is, of course, possible only to one who is 
 used to working without veil or gloves. 
 
 If the door 44 of a queenless colony is opened, the same agitation is seen in the interior, 
 and smoke blown in only increases the buzzing. This excitement over the loss of a 
 queen often appears very soon after her removal, particularly when there is no forage or 
 if the colony is weak. 
 
 The difficult question now arises, how bees notice the absence of the queen and how 
 they communicate this loss. Is it the sudden absence of her odor? Hardly, at least not 
 in all cases, for we have seen in a previous paragraph that the odor of the queen is 
 exceedingly adherent, and therefore the walls, as well as the bees, must be impregnated 
 with it. But the intensity is gradually becoming less. However, this odor is variable 
 in the customary course of events, 4 "' and so no uneasiness of any kind arises on this account. 
 Furthermore, how does it happen that the bees xiHhlmli/ become agitated after the queen 
 has been taken out sometimes for an hour or more? 
 
 If Bethe is right, that the means of communication depend upon the outflow of some 
 chemical, then the above observations would prove at least that the odor of the queen 
 is a very dominant one in the colony. The queen then does not, as Bethe declares, take 
 on the hive substance, but just the opposite happens; the queen influences the hive odor 
 considerably, or at least a mutual scenting takes' place. On the other hand, if we agree 
 with Bethe in his statement that the queen takes up the hive substance of the colony, 
 the means of communication must depend, not upon reflexes of smell alone, because the 
 colony odor would not be influenced by the removal of the queen. 
 
 42 Bethe, 1. c, p. 70. 
 
 53 That "agitated bees" constantly "buzz" is a misleading expression of Bethe's. Particularly can 
 the humming with raised abdomens and fanning wings upon the flight-board l.«e sharply distinguished 
 by the trained ear from what is usually called the ''buzzing" of bees, by the pitch . 
 
 44 Reference is here made to the German style of hive with a door in the rear. The American style 
 of hive opening at the top is little used in that country. — E. F. P. 
 
 45 If the queen gets into the honey-chamber, which is generally separated from the brood-chamber 
 beneath by a strong piece of board with slits in for the passage of workers, and if she there continues 
 to lay eggs, it is clear that her odor must soon be tost to the bees beneath. This is especially true if 
 there is an entrance into the honey-chamber. Under these circumstances, however, excitement at the 
 queen's absence never breaks out; at most the "isolated" bees after a few days occasionally build queen- 
 cells, but more often not. 
 
Natural History of the Honeybee. 13 
 
 If we were to ask bee-keepers their opinion in this matter, they would probably answer 
 that the young bees which care for the feeding of the brood and the queen "miss" the 
 latter after a longer or shorter period, and begin to seek her diligently, communicating 
 their anxiety to the whole colony until the " conviction ' ' that the queen is gone comes 
 into consciousness, when the comfortable humming changes to a loud buzz. 
 
 But since the young bees, as all bee-keepers know, can be taken away very well with 
 the queen it naturally follows that they take no very important part in this matter. Then 
 the old bees will begin the loud buzzing. 
 
 One can, however, ingeniously gather a colony of old workers and place within it a 
 queen enclosed in a cage. The excited colony is quieted as if by magic. The hostile bees 
 next to the cage begin to list their abdomens, fanning their wings, 46 with a characteristic 
 humming. This hum is taken up by other bees, and suddenly there is a peaceful quiet. 
 
 Before we formulate conclusions from the foregoing let me communicate further 
 observations. 
 
 The following experience is interesting, and proves that the feeding of the queen as 
 one of the chief phenomena of life needs not to be taken into consideration in the breaking- 
 out of the agitation over the loss of a queen, apart from the odor. 
 
 In bringing in a first swarm, I crushed the queen by accident. Since I could not care 
 for it on that day nor the next, and I did not have another queen at my disposal to sub- 
 stitute, I knew that the swarm would certainly fly out again and join the mother colony. 
 To prevent this, I arranged the following experiment: I fastened the dead queen with 
 a needle to a piece of cork, and hung it in a cluster of bees. The colony remained 
 quiet, and I knew later on that it had felt entirely queen-right because the comb which 
 it made showed no drone-cells, and the dead queen was surrounded by licking bees. 47 How 
 long a dead queen can supply the place of a living one I am not able to tell. 
 
 However, even a dead queen may be dispensed with. In order to do this, I put a 
 weak colony into swarming condition — that is, took away all combs so that the agitation 
 could be more quickly observed. The queen was put into a cage. The next day I took 
 the cage out of the colony, and very soon I noticed the typical signs of queenlessness. 
 When the excitement seemed to have reached its height I opened the glass door and held 
 the cage from which the queen had been quickly taken in the midst of the bees. Imme- 
 diately it was surrounded by many ( * joyful ' ' bees with lifted abdomens, fanning with their 
 wings, and the buzzing ceased. Here we have conclusive proof that the odor of the 
 queen is enough to satisfy all, the instincts which appear unsatisfied during the excitement 
 over the absence of the queen. 
 
 The conclusion may easily be drawn that, if the weak odor adhering to the queen- 
 cage causes a cessation of the agitation, and certainly it is the odor alone, the disappear- 
 ance of weakening of the same causes the outbreak of the uneasiness. But this deduction 
 does not seem conclusive* to me; for, as I have said above, the\ odor does not disappear 
 with the removal of the queen, but is, however, weakened; and if the colony is vigorous, 
 and their instincts are diverted by a rich honey-flow, the bees notice the loss only after 
 some time has passed. But in the case of the sudden outbreak of uneasiness over the 
 absence of the queen, we have to do with an intensified instinct, for which the shadowy 
 memories in actual requirement are sufficient for the needs of the case, if I may so 
 express it. Thus bees in the spring, impelled by necessity in a region where there is little 
 pollen, will gather the dust from thrashing, coal dust, or brick dust, instead of the lacking 
 pollen. Once I observed bees collecting even sawdust. It is just as little permissible to 
 draw from this the conclusion that brick dust is equivalent to pollen, because bees react 
 in the same way toward both, as it is to Conclude that it is the missing odor alone which 
 
 46 In the original, the author characterizes this peculiar humming as "sterzeln." This he describes 
 in a footnote as follows: 
 
 "By 'sterzeln' is meant that characteristic raising of the abdomen, accompanied by a slow whirring 
 of the wings, which indicates joy. Therefore it is seen in the above case, or when they discover the 
 hive after going astray, but never on finding a honey supply, even if very hungry. This is similar to what 
 I might call the attitude of fright. If the finger is approached to the entrance of a hive, single bees 
 guarding the passage are seen to hasten toward the hostile object, raising their abdomens in threatening 
 manner,°and they remain in rhis position. 
 
 17 A queenless colony v. ill erect .drone-cells if it builds at all. 
 
14 Natural History of the Honeybee. 
 
 causes agitation in a stock, because the bees are satisfied with the remembered odor in 
 place of the real queen. We shall have better proof further on. 
 
 BEHAVIOR OF A QUEENLESS SWARM. 
 
 As a first swarm was issuing I caught the queen at the entrance of the hive. Xow 
 instead, as happens in most cases, of the swarm returning to the hive after the vainly 
 searching circling, the bees hung to a branch united in the well-known cluster. Since no 
 alarm was shown, I took it for granted that a second queen had swarmed with them; but 
 a prompt examination of the hive revealed no queen-cells from which a young queen 
 could have emerged. The swarm was perfectly quiet for over half an hour, then sud- 
 denly broke up and went back to the hive — a certain proof that it was queenless. In this 
 case, then, there was a considerable time between the removal of the queen and the 
 breaking-out of the uneasiness over her absence. It was evidently not the disappearance 
 of the odor of the queen that caused the discontent, for the odor was lacking from the 
 beginning. Queenless swarms rarely take place; and if they do, they do not settle and 
 disband again; they miss the queen immediately. Coming out into the open air, the odor 
 of the queen seems to be of little value. This is evident from the fact that if one keeps 
 a caged queen in a circling swarm, it has tol remain there for a long time before she is 
 scented by the bees, if she becomes scented at all. But if a bee once places itself upon 
 the cage, "joyously" lifting the abdomen, and humming, then it is not long before the 
 rest are attracted by the sound of the "satisfied" hum. I might, incidentally, mention that 
 there is a so-called ' ' diamond rule ' ' among bee-keepers. This rule is that the queen is to 
 be enclosed in a cage for a short time before and during a rich honey-flow in order to 
 hinder her from laying eggs. The "bees then will have little brood to care for; the con- 
 sumption will be diminished, and the honey store should increase. This, however, fre- 
 quently does not take place, because the normal condition of the colony is disturbed. The 
 colony often believes itself queenless and builds queen-cells in spite of the imprisoned 
 queen in the middle of the colony; but this phenomenon does not appear in all colonies. 49 
 Apparently there is here, as has been demonstrated before, an unsatisfied feeding instinct. 
 The nurses can not distribute the richly prepared larval food, and so the impulse is 
 generated to remove the abnormal condition by building queen-cells. I can imagine this 
 as an example of a clear reflex. There is no need whatever for "reasoning" of any kind. 
 
 I shall now return to the very vigorous colony mentioned before. If the queen is 
 taken away from such a colony, which compactly filled the brood-chamber and honey- 
 chamber in a large hive, the signs of uneasiness over the absence of the queen will go on 
 as described. When the colony" is in greatest excitement, a cage containing a queen r.^ay 
 be pushed into the honey-chamber in the upper part of the hive which opens from behind, 
 and then the condition of the bees at the entrance at the opposite end of the hive under- 
 neath in the brood-chamber may be observed. Almost at once a change in the behavior 
 of the uneasy bees is apparent; the buzzing dies out in the hive, and the bees, searching 
 around the entrance, enter with lifted abdomens and fanning wings. 
 
 The explanation can here certainly not be the influence of the odor, since the odor of 
 the queen could not penetrate in an instant to the entrance which is separated from the 
 honey-chamber by the whole of the brood-chamber. If the extraordinary penetrating odor 
 of female insects be cited, as, for example, that of the moths (sphinx, etc.), then I would 
 reply by referring to the disregard of an imprisoned queen in a swarm. 
 
 DISREGARD OF A QUEEN IN OPEN AIR. 
 
 In order to demonstrate this question still further I hung a cage containing a queen 
 on a stick and stuck this into the ground so that the cage was at the same height as the 
 hive entrance, about thirty-five centimeters to one side of the flight-board. None of the 
 
 48 It is cunou,s that queen-cells are also established if the queen is decrepit. If, therefore queen-cells 
 are found, and at such a time either strongly defective brood or drone brood in worker-cells (on account of 
 the exhaustion of the supply of sperm-cells), then one can be sure that the old queen will shortly disappear 
 from the hive. Perhaps the same instinct is active here as in the case of an imprisoned crueen (see 
 footnote, p. 11). ■ \- 
 
Natural History of the Honeybee. 15 
 
 numerous bees flying in and out scented the queen, which remained entirely unnoticed 
 while the colony continued agitated. 
 
 HEARING CAPACITY AND SENSATIONS OF SOUND. 
 
 If we now conclude from the foregoing observations in what way the discontent 
 because of the absence of the queen, as well as the content because of her presence, is 
 communicated, and what the means of communication are,. we must conclude that the 
 odor is not the only factor. We have been able, on the one hand, to determine with rea- 
 sonable certainty that, if the odor were sufficient to inform the colony of the presence or 
 absence of the queen, no other special kind of communication need be used; on the other 
 hand, we saw indications of discontent (for example, the building of queen-cells) among 
 bees when a queen was present in a cage. So we see signs of peace or agitation in cases 
 where the influence of odor seems completely excluded. In all observations, when the 
 colony notices the absence of the queen there is always a change in the usual characteristic 
 sound in the hive, the bustle of the colony, if I may so express it. 
 
 There is, therefore, not the slightest doubt in my mind that bees communicate with 
 each other by sound. The tone of " peace' ' attracts hive mates or quiets them; the louder 
 buzzing excites them; it disappears if the queen is given back. At the same time the 
 whole character of the colony changes, 49 the queenless irritable bees become quiet and 
 peaceful, and again take up their work, which was laid aside during the excitement. We 
 must admit the possibility of communication between bees by sound, therefore of hearing 
 capacity and sound sensations. 50 Each bee has the instinct to join in the tone of discon- 
 tent if it hears it; therefore if the absence of the queen is noticed; by one bee the agitation 
 is very quickly propagated throughout the colony. x 
 
 How the first notice of the lack of a queen takes place is naturally, as before stated, 
 very difficult to determine — perhaps there are several possibilities. Often the absence of 
 the queen odor may bring about the result; it may be the absence of the characteristic 
 humming described, which the bees occupied with the queen produce. Further, it is 
 possible that the queen herself makes sounds, the absence of which is noticed. I have 
 never heard such a humming, but it is not impossible that such tones, not perceptible to 
 the human ear, exist. 
 
 EXPERIMENTS ON SWARMS. 
 The following observations will show to what a great extent the means of communi- 
 cation between bees depend upon sound perceptions. The loud buzzing of swarms is 
 readily distinguished from the usual hum by anybody who has been occupied with bees 
 for any length of time. The swarm presses out from the hive in impetuous haste in the 
 peculiar swarm dance; it surges and whirls in " bacchanal delight," as if the bees were 
 really drunk with "joy." The "swarm dizziness, 77 as bee-keepers say, has seized it. In 
 this dizziness it forgets everything connected with the old dwelling, and bees even forget 
 to sting. 
 
 "Swarming bees do not sting" is an old bee-keepers' saying. From this we get the 
 old fable that bees recognize the bee-master, because, generally, outsiders approach a 
 bee-hive only to see the interesting spectacle of a swarm, and then notice the bee-keeper, 
 often quite unprotected, quiet and composed, stand in the middle of the tumult without 
 being stung. 
 
 The "Deutsche Bienenf reund, " 1894, reports the following: "A boy about ten years 
 old was standing near a hive, bare-headed and shirtsleeved, when a swarm issued. After 
 flying here and there, the queen alighted on the boy 's head, and thousands of bees quickly 
 
 46 Careful observation will show that every colony has its particular character, which is determined 
 ln P art by. tne strength of the colony. On the other hand, we often see what are apparently great 
 differences in colonies of like strength. One colony is gentle; another constantly desines to sting. One 
 defends its hive entrance carefully; another almost not at all, although it ia a strong colony. One flies 
 constantly earlier than its neighbor, etc. Lubbock and Herman Muller have shown that individual bees 
 vary also. (Herm. Muller, " Versuche uber Farbenliebhaberei der Honigbiene," Kosmos, Jahrg. 6. Lubbock, 
 ' Ants, Bees, and Wasps," German translation published in Leipzig, 1883, English edition in the Inter- 
 national Scientific Series, New York, 1883.) 
 
 50 In the attraction of a queenless colony into the hive of a queen-right one, then, perhaps, in the 
 first place the hearing capacity comes into question (see p. 5). the sound of the contented humming acts 
 as a powerful stimulus and causes the migration. 
 
16 Natural History of the Honeybee. 
 
 followed. The boy's father, recognizing immediately the state of affairs, called to him 
 hastily (he had often looked on during the catching of a swarm), 'Do not move, Hans. 
 Shut your mouth and eyes, and breathe through your nose. I will sprinkle and capture 
 them! ' The boy obeyed, the father poured water over his head, bent it forward, and 
 with a feather stroked the whole swarm into a basket underneath. The boy had not a 
 single sting. " 
 
 After my own experiences, I consider this tale entirely trustworthy. 
 
 In 1893, i i Studers ill. Schweizer. Bienenf reund, , ' showed the picture of a young bee- 
 keeper who had been photographed with a swarm which hung down from his hand. In 
 the issue of the swarm he caught the queen with his fingers, thus causing the swarm to 
 surround his hand. When the photograph was completed, three-quarters of an hour had 
 elapsed, which was endured very quietly, the arm being supported' with a stick. Head 
 and hands were not stung. Similar pictures have been published in "Gleanings in Bee 
 Culture ' ' during the last few years. 
 
 It is not in contradiction to the above to say that most disasters are caused by swarming 
 bees; for nervous blows or accidental crushing irritates the swarming bees; and first one 
 sting, then hundreds follow, excited by the strong odor of the poison. 
 
 Shortly before the issuing of the swarm, single bees are seen coming out with uneasy 
 motion; they press through the bees around the entrance, which often cling closely together 
 like a long " beard ' '^hanging from the flight-board. It can hardly be doubted that sounds 
 of some kind perhaps serve here for communication — sounds lost to the human ear in 
 the general hum; 51 at least there is no more plausible explanation for the peculiar result, 
 for suddenly the ' ' beard ' ' loosens, the bees enter the hive quickly, assail the honey stores, 
 and fill their honey-sacs. All the others desiring to swarm do the same thing, and sud- 
 denly the swarm breaks forth. Those just returning from the fields heavily laden with 
 pollen are involved in the tumult, infected with the sound of swarming, and fly with 
 the swarm. 
 
 THE INFECTING INFLUENCE OF THE SWARM TONE. 
 
 That the sound made in swarming, which is given only in flight, is infectious is. well 
 known; for now and then it happens that the neighboring colony, though not nearly ready 
 to swarm, will follow the swarm-tone and swing into the air. If two swarms issue at 
 the same time, they mutually attract each other and unite. 
 
 That we have hereto do with a chemical reflex, as is thought by Bethe, who, strangely 
 enough, does not take swarming into consideration, seems improbable to me. If the rich 
 possibilities for sound communication between bees are borne in mind, it is evident that 
 this capacity must have some object, and that it is not practicable to ignore these data. 
 
 THE ENTICING NOTE OF BEES. 
 
 If a swarm is shaken out on a sheet spread over the grass, and a hive never before 
 used is placed, for instance, on the north side of the sheet, it will remain unnoticed if it 
 has not come in direct contact with the swarm. But if a handful is scooped up and 
 thrown at the entrance, they will immediately draw near with specially loud humming, 
 part standing outside, however, on the flight-board with lifted abdomens and wings fan- 
 ning. All the other bees near by turn, if they have been facing away from the hive, and 
 follow the alluring sound, lifting their abdomens and fanning with their wings. More 
 keep following, and a broad band march into the dwelling. If now the hive is taken away 
 and placed on the south side, the bees quickly march on further north, but for only a short 
 time. Then the increase of the attracting note from the south causes a gradual turning 
 of the stream, which enters the hive little by little. It may happen that a little cluster 
 of bees advances firmly northward, although many mates turn toward the south. The 
 alluring tone does not seem to penetrate further; they heard it coming from the north 
 and follow the memory. There seems to be no other explanation for this course. In my 
 opinion the fact that not only a single bee but plenty of them, a small cluster, follows this 
 
 51 Movements of the antennae should certainly be considered, since the play of the antennae is always 
 lively, and often a mutual touching takes place. 
 
Xatural History of the Honeybee. 17 
 
 memory is a proof of the impulse for imitation which is undoubtedly present in bees for it 
 is not likely that every bee in the small cluster has this memory but perhaps only a few of 
 them. The others follow only on account of this impulse for imitation. If the first bees 
 remain true to the direction, then a great number will turn quickly with them. 
 
 It must really seem strange, to any one familiar with the natural history of the bee, 
 that there can be any doubt as to the possibility of communication through sound, when 
 the whole life of a bee is one continuous humming, if I may so express it. Can this 
 "audible speaking ' ' have no purpose? Only the dead bee is quiet. 51 
 
 If the previous assertions are not convincing, we may find proof of their capacity for 
 hearing in the following: 
 
 THE "TEETING" AND "QTJAHKING" OF THE QUEEN. 56 
 
 When the first swarm has issued with the old queen, normally nine or ten days pass 
 before the after-swarm follows. One or two days before the swarming, a strange concert 
 may be heard in the hive on a quiet evening at a distance of a couple of steps. It is the 
 ' * teeting ' ' and ' ' quahking ' y of the young queen. As is well known, the old queen, before 
 her exit, lays eggs in the queen-cells at intervals every day or so; the consequence is, 
 that the young queens do not all emerge at once. 
 
 The first queen to emerge falls upon the other queen-cells in order to kill her rivals. 
 If the colony wishes to swarm, however, the workers prevent this destruction, and the 
 queen begins to "feet" in "jealousy. " She presses her head against the comb, as I have 
 observed, and sounds a clear, ringing, long-continued "teet, teet, ' ' apparently using the 
 stigmata on the thorax. Immediately the most mature of the queens still in the cells 
 answers with a short deep ' ' quahk, quahk. ' ' Thus this interchanging goes on for hours 
 or days with shorter or longer interruptions. If rainy weather prevents the issuing of 
 the swarm, and more queens mature, they i ' quahk ' ' likewise. They ' ' dare ' ' not emerge as 
 long as the "teeting" goes on; and since they need nourishment they cut a little slit with 
 their mandibles, through which they stick their probosces, and so are fed by the workers. 
 
 It would lead me too far to go into these interesting circumstances. 
 
 If we can prove from the above an undoubted capacity for hearing, we can also 
 from the following experiment: 
 
 THE QUEEN'S TONE OF FEAR. 
 
 If a strange queen is put into a queenless colony by merely allowing her to run in 
 upon a comb, then the bees nearest her fall upon her and bite her legs or ride upon her 
 to sting her. The powerful queen runs quickly away from her persecutors, but she is 
 continually seized by others. Now in her "fear" she utters loud cries which throw the 
 whole colony into excitement. One could here conjecture the influence of odor; but a 
 strange queen in a cage does not cause such excitement, especially not if the colony is queen- 
 less. It is doubtless the tones of fear which stir up the whole colony. We have here again 
 an unmistakable reaction toward sound-sensations. 
 
 The attempt might be made to weaken these/ assertions, so far as they apply to the 
 existence of hearing capacity, by pointing out that no one has yet succeeded in showing 
 in insects a reaction toward sounds artificially produced. Then I would call to notice 
 that recently an American has obtained undoubted reactions toward the notes of tuning- 
 forks in two members of the Formicidae 56 (Lasius American us and Formica nitidiventris) 
 in the two Myrmicidse (Crematogaster lineolata, and a kind of Aphoenogaster). The 
 well-known myrmecologist William Morton Wheeler has meanwhile published in an 
 important paper (Ethological Observations on an American Ant, Journal fur Psychologie 
 und Neurologie, Bd. II., Heft 1 u. 2, Berlin, 1903), his investigations on hearing in ants. 
 He writes as follows: 
 
 " Stridulation, at least among the Myrmicidse, Ponerida?, and Dcrylinae is an impor- 
 
 M 1. Ernst Haeckel, Die Weltrateel, Bonn, 1899, p. 145. 
 
 54 2. Now and then the colonies settle down in their winter sleep into a condition of complete quiet, 
 but usually it is accompanied by a hardly audible murmur. 
 
 55 It is, of course, very difficult to represent these sounds in words ; but the ones here given are often 
 used in English to represent them. No bee-keeper will have any difficulty in knowing what is meant. The 
 author represents them in German by the words "Tuten" and "Quaken." — E F. P. 
 
 5(5 Science (The Sense of Hearing in Ants), No. 5, Vol. X., No. 256, 1889. See also my Conclusions. 
 
18 Natural History of the Honeybee. 
 
 tant means of communication, which Bethe has completely ignored, and even Forel and 
 other Myrmecologists have failed to appreciate. It readily explains the rapid congrega- 
 tion of ants (Myrmicinae) on any particle of food which one of their number may have 
 found, for the excitement of finding food almost invariably causes an ant to stridulate 
 and thus attract other ants in the vicinity. It also explains the rapid spread of a desire 
 to defend the colony when the nest is disturbed. This is especially noticeable in, species 
 of Pheidole, Myrmica, and Pogonomyrmex. It is the secret of being able in a short time 
 to catch ants like Pogonomyrmex molefaciens in great numbers by simply burying a 
 wide-mouthed bottle up to its neck in the mound of the nest. An ant approaches and 
 falls into the bottle. It endeavors to get out, and, failing, begins to stridulate. This afc 
 once attracts other ants which hurry over the brim and forthwith swell the stridulatory 
 chorus till it is audible even to the human ear.' More ants are attracted, and soon the 
 bottle is filled. If it be corked and shaken for the purpose of still further exciting its 
 contents, and then held over another Pogonomyrmex colony whose members are peacefully 
 sauntering about on the dome of their nest, the wildest excitement will suddenly prevail, 
 as if there had been a call to arms or — to dinner. Even more remarkable is the stridula- 
 tion in a colony of Atta fervens, the Texan leaf-cutting ant. Here the different ants, 
 from the huge females through the males, large soldiers and diminishing casts of workers 
 to the tiny minims, present a sliding scale of audibility. The rasping stridulat^on of the 
 queen can be heard when the insect is held a foot or more from the ear; to be audible 
 the male and soldier must be held somewhat closer, the largest workers still closer; whereas 
 the smaller workers and minims, though stridulating, as may be seen from the movements 
 of the gaster on the postpetiole, are quite inaudible to the human ear. It is not at all 
 improbable that all this differentiation in pitch, correlated as it i3 with a differentiation 
 in the size and functions of the various members of the colony, is a very important factor 
 in the co-operation of these insects, and of ants in general. The contact-odor sense, impor- 
 tant as it undoubtedly is, must obviously have its limitations in the dark subterranean 
 cavities in which the ants spend so much of their time, especially when the nests are 
 very extensive like those of Atta. Under such conditions stridulation and hearing must 
 be of great service in maintaining the integrity of the colony and^ its excavations. The 
 fact that as yet no unquestionable auditory organs have been discovered in ants is of 
 secondary importance when it can be so easily shown that these insects really respond 
 in an adaptive manner to the sounds produced by other members of the colony." 
 
 I know very well that Miss Adele M. Fielde and George H. Parker are of the opinion 
 (The Keactions of Ants to Material Vibrations, Proc. Acad. Nat. Sc, Philadelphia, Sept., 
 1904) that "it is misleading to ascribe or deny hearing to ants; they are very sensitive 
 to the vibrations of solids, not to those of air; their reactions could be as appropriately 
 described as resulting from touch as from hearing." But the above investigations on 
 bees have been neglected by these authors, and I am still of the opinion that a sensitive- 
 ness to the vibrations of air can not be denied so far as bees are concerned, and the 
 above experiments of Wheeler seem to show the same in ants. 
 
MEMORY OF PLACE IN BEES. 
 
 In general, we should not set up a new unknown 
 force to explain natural phenomena until it has been 
 proven that they can not be explained by the known 
 forces.— Aug. Weismann, p. 539. 
 
 According to Bethe, bees are "led back to the hive by a force entirely unknown to us. 
 This force does not adhere to the hive itself, and it does not lead bees back to the hive 
 itself, but to the place in space which the hive usually occupies. It does not act at 
 boundless distances. It is an old experience of bee-keepers that they can take a colony 
 to another stand without fearing that the bees will return to the old place, if the new 
 spot is only more than six kilometers from the old. It follows, then, that this force acts 
 at most at a distance of six kilometers, since the impulse to return to the hive is the 
 strongest of all impulses in bees. But' I\ believe that the zone of action of the force is 
 not a circle with a radius of six kilometers, but of only between three and four kilometers. 
 If the circle had had a radius of six kilc meters, then the bees of the transported hive 
 would be back into the circle of action if they got more than half that distance near the 
 old position in foraging, and would have to return to the old place. But this only happens 
 if the old position is less than sis kilometers away from the new. We must, therefore, 
 accept something near three kilometers as the boundary for this circle of action for this 
 force. " (Bethe, 1. c, p. 89.) 57 
 
 I believe that the foregoing, and likewise in many other of his observations, Bethe 
 furnishes such excellent proof df the existence of memory for location that one can 
 hardly wish anything better. But Bethe supposes "a force unknown to us, " with which 
 we do not know what to do, which offers support for every investigation. Under this 
 head I must depend more strictly upon Bethe 's work in order at the same time to point 
 out the errors in the chapter, "How do Bees Find the Hive?" (Bethe, 1. c, p. 72.) 
 
 THE "PATHS" OF BEES AND THEIR DIRECTION. 
 
 If the bee-hives are placed on an open heath, with no elevations such as trees or bushes, 
 the particular kind of flight toward and away from the hive may be studied, undisturbed 
 by local conditions. In August, 1898, I had opportunity to prove this condition on a 
 moor in the neighborhood of Oldenburg, corroborating entirely my earlier experiences. 
 
 The colonies were established on the moor by the bee-keeper shortly before the buck- 
 wheat-flow. The hive-entrances faced the east. On the first day the flight was weak, for 
 as yet there was nothing to gather. The bees flew out irregularly in smaller or larger 
 circles in all directions. There was nothing to be seen yet of " a path. ' ' M Great buckwheat- 
 fields stretched toward the north as well as to the south. When the flowers opened, the 
 mode of flight changed. Part of the bees flew from a point close to the entrance, sharply 
 
 57 The "unknown force" must act at much greater distances under certain conditions, for 'other 
 observations show that bees have flown five, six, or even over seven kilometers further under extraor- 
 dinary circumstances. (See Bienenzeitung, X., No. 14; ditto III.. No. 9, Dzierzon; Le Rucher, Amiens, 
 1876,* IV., p. 30). In these cases there was no forage near at hand. According to Dzierzon, some bees 
 under these circumstances scented a large "hundred-acre" rape-seed field which lay far outside their 
 usual circle of flight. Roth, the* leader of the Baden school for bee-keepers, in Durlach, observed that 
 some of his bees returned in thirty minutes with full loads from a buckwheat-field six kilometers away. 
 Granting that Roth's bees flew to the buckwheat-field from the south, there is no reason for thinking 
 that bees from an apiary at the same distance toward the north had not hastened to this same nectar 
 supply. Now if, for experiment, a colony should be taken from the southern apiary to the northern 
 one, then they fly south to the buckwheat-field, and there, coming into the field of action, would again 
 return to the southern apiary. The "unknown force" would reach twelve kilometers in this very possible 
 case. If it be asked from what distance bees can find their way back, the answer can not be made in 
 kilometer measurements, for it depends upon whether the bees are in their, flight or orientation (see the 
 same) or in their search for food have flown to greater oi* less distances, Tipon the definite direction of 
 the forage and the general orientation. 
 
 58 "It is well known that, in front of the entrance to a strong colony, there is always a long dark 
 cloud formed by the bees constantly going back and forth." (Bethe, p. 75.) This Is Bethe' s "bee-path." 
 
20 Natural History of the Honeybee. 
 
 to the loft (north) and another part sharply to the right (south). In consequence of 
 this the paths were very short, the terminations (the hives stood close to each other) 
 coming together in a common wave. 59 It is apparent from this that the position of the 
 pasture determines the direction of the flight in and out. 
 
 When, later, the buckwheat-fields ceased blooming and the moor was in full bloom, 
 the same short, though perhaps a little lengthened, paths were seen; but the common wave 
 was higher and apparently somewhat larger, from the great numbers hastening in from 
 all sides. 
 
 "Long paths " of bees, which Bethe describes as normal, are found chiefly only in 
 the case of bees in gardens where trees, houses, etc., modify the flight. The view is 
 wrong (Bethe p. 80), that "it is a well-known fact to bee-masters that the steepness of 
 bee-paths varies with the weather." 60 It is an error further, that; "bee-paths always go 
 out from the hive in the same direction with slight fluctuation. This is almost always 
 east, southeast, or south. Not only for. this reason, but also because bees need sun to 
 be industrious, the bee-keepers place their hives toward the south or east." (Bethe, p. 81.) 
 This last assertion is erroneous. 61 
 
 It is clear from the above, even if Bethe had not stated it himself exactly, that there 
 was nothing in the north for his bees to gather, since they never flew north (see Bethe's 
 following conclusions). In that case the town (Strassburg) must spread out northward 
 from the place of Bethe 's investigations. Bethe supposes that the town remained "un- 
 known ' 1 to the bees. 
 
 The garden of the Physiological Institute in which my bee-hives stand lies close to 
 the town wall. The wall is covered with green, and behind it stretch great blooming 
 meadows which teem with bees. On the other side of the Institute lies the city. There 
 are very few and very small growing plants in the city; and, in spite of many searches, 
 I have only once seen bees in the inner part of the town, on two isolated sunflowers. At 
 all events it must be admitted that only a few isolated individuals have been in the interior 
 of the city at any time in their lives, particularly as my bees always fly toward the south 
 to the meadows." (Bethe, p. 86.) 
 
 I must consider these assertions also as erroneous and not conclusive, for the handler 
 of sugar wares, honey-sellers, gingerbread bakeries, and housewives who leave sweet 
 preserves or jars of honey in the open windows well know they have to count upon much 
 unpleasantness from visiting bees. 62 There is not always something to be gathered from 
 the meadows (pause in the honey-flow, the cutting of the field) ; and, besides that, bees 
 in their first flight, which is devoted to orientation, orient themselves not only toward the 
 side of the pasture but naturally on all sides. 
 
 Bethe then let bees fly in the streets of the city, only within the short distance of 
 350, 400, and 650 meters from the hive, in spite of the "unknown force" which acts, 
 according to Bethe, within a distance of three kilometers. Because, when let go, these 
 bees found their way home just as well as those from the meadow, he decides that ' ' there 
 
 59 I noticed that hees of one and the same hive followed both directions. 
 60 1 admit that the French boy learns in his lesson : 
 
 "Quand les abeilles volent en haut 
 Nous aurons bientot de l'eau." 
 But this probably national idea is just as \Mong as the one that a hard winter is to. le expected if 
 bees propolize entrances strongly. In general, many statements to be found in the literature on bee 
 management should be taken with prudence. Safe judgment can be made only after a personal experience 
 of many years. 
 
 61 Dathe, Lehrbuch der Bienenzucht, 5 Aufl., p. 34, and following, Bensheim, 1892 : Berlepsch-Lehzen, 
 "Bienenzucht," Berlin, 1899 p. 28 and following; "Bienenzeitung," 2 Ausg., Nordlingen, 1861-'62, 
 2 Bd., p. 3 and following; v. Berlepsch, "Die Biene und die Bienenzucht," Muhlhausen, 1860, p. 219 
 and following; Dzierzon, Rationelle Bienenzucht, Brieg, 1861, p. 36, etc. 
 
 62 Because of this kind of trouble, bees are forbidden in the precincts of the city of Paris. The 
 Bw. Centralblatt> No. 19, 1899, announces: "In a small city a broker had stored comb from colonies 
 with foul brood in a room accessible to bees. The following year almost all the apiaries of the city 
 were infected," etc. During the dearth of nectar, specifically in Bethe's case, the flying in the town 
 where there was always something at which to nibble might have been more vigorous than to the fields 
 where nothing could be obtained. . Moreover, Bethe himself furnishes a good reason that bees flew 
 extraordinarily accurately through the city, for he asserts that he "had seen bees inside the city upon 
 two isolated sunflowers." How very inquiring the bees must have been to find a single flower! We 
 can be very sure that innumerable bees (even if not seen by Bethe) followed the same instinct which 
 drew these sunflower visitors into the city, for there are no recluses among bees, none which separate 
 themselves from the masses to follow a way of their own, and all possess the same* kind of instincts. 
 A single linden-tree in bloom in the city/ is enough to attract hundreds of thousands of bees. 
 
Natural History of the Honeybee. 21 
 
 can be no doubt at all that bees find their way back to the hive, not by means of memory 
 pictures' ' (Bethe, p. 89). 
 
 I cannot find in the foregoing the slightest foundation for this view of Bethe 's. If 
 he wishes to disprove the investigations of Bomanes, 63 then I think the attempt is unsuc- 
 cessful, for Bethe leaves to us the proving that the city has remained actually ' i unknown ' ' 
 to the bees. 04 
 
 But how is the mysterious behavior in the streets explained? We shall let Bethe 
 speak for himself: "All bees, if let fly, go upward in a corkscrew line, then suddenly 
 take a direction and" fly off in a straight line. This happens when bees are let fly in the 
 city streets almost always before they have reached the levels of the house roofs. It often 
 happens at a height of four to six meters above the street level, therefore generally before 
 they can have acquired a view of the neighborhood. Now, almost without exception, they 
 take the direction of the Institute, where the hives are. ' ' 
 
 Before I give the explanation for this, other parts of Bethe 's account continue: "Light 
 is the incentive to flight in these diurnal animals" (Bethe, p. 83) ; further, "Light regulates 
 flight." 
 
 I remember, too, in Herm. Miiller's investigations that he could' carry a bee in a 
 drinking-glass open beneath, the length of the garden without the bee flying out, for it 
 constantly pressed to the top of the glass toward the light. 65 
 
 I slipped a bee into a reagent glass and put it upon the window-sill so that the bottom 
 of the glass was toward the window. For eight hours the bee strove inside the glass in 
 vain efforts to reach the light. Then it died, although it would have been easy to crawl 
 out of the open tube and fly out of the open window. 
 
 Now, if we remember that the city lies to the north of the seat of Bethe 's investi- 
 gations, we shall see that the sun must be in the direction of the Institute, and the 
 bees were let fly in "quiet, sunny weather" (Bethe, p. 87). In the darker streets, perhaps 
 unknown to them, they tried to orient themselves by mounting in circles, just as a 
 carrier pigeon does. 66 Then they fly instinctively toward the bright source of light (just 
 as in a room they fly unfailingly toward the bright window) until they become oriented 
 in familiar regions. "Light regulates flight" (Bethe, p. 83). 
 
 I, therefore, can not consider the "unknown force" and the conclusions based on it as 
 capable of proof. 
 
 I have held to Bethe 's assertions with regard to* these directions. From them I 
 understand that on one side of the Institute toward the soutfy are the meadows; and on 
 the other (therefore toward the north) is the city. Possibly Bethe did not liberate the 
 bees just in the center of the city but toward the edge, so that the Institute was shifted 
 from its southern position a little toward the west or east. Further investigations might 
 be necessary, perhaps, to prove that these bees did not return to the hive not through the 
 "unknown force." 
 
 We learn from Bethe (1. c, p. 87) that eight marked bees were let fly in the* city 
 
 63 Romanes, Nature, 1885, Vol. 32, p. 630, Homing Faculties of Hymenoptera. 
 
 64 That Bethe' s idea that bees had not been flying in the city is not at all convincing, follows from 
 something written by the editor of "Elsass-Lothring. Bienenzuehters." Karl ZwiUing, which reached 
 me subsequently. In my publication in the "Biologischen. Centralblatt" the same is given, therefore, in 
 a later passage. I quote the following: "* * * Outside of Strassburjg there are many apiaries close 
 to the wall, the bees of which never think of flying into* the city except in times when there i$ no 
 forage. Then they enter the candy factory of Mr. Pale, Tiergarten Strasse, and annoy the workers as 
 well as partake of the sweets. Even in the month of December, 1899, in the warm sunshine I observed 
 bees flying in the middle of the city on the Kleberplatz, where hundreds of pots of flowers were dis- 
 played for sale. There they gathered honey and pollen. Every year some swarms fly into Strassburg 
 and hang on the chestnut, linden, oak, and. - locust trees found in many parts of the city, and once a 
 swarm hung on the showcases of a glovestore near the cathedral. "When the trees bloom, bees fly 
 about them vigorously. In the interior of the city there are beautiful rows of chestnuts along the 111, 
 rows of lindens around the Kleberplatz, etc., all of which are sought out by the bees when they are in 
 bloom. The honey-venders and those who deal in sugar wares are not molested if they keep their wares 
 covered. As I live in the neighborhood of Strassburg (ten minutes by rail), and have been President 
 of the Bienenverein there for thirty years, I know the conditions exactly." 
 
 85 Herm. Muller, Versuche uber die Farbenliebhaberei der Honigbiene, Kosmos, Jahr. 6, p. 276, 1882. 
 66 H. E. Ziegler, Die Geschwindigkeit der Brieftauben, Zoolog. Jahrb., X. Bb., 1897, pp. 99, 278. 
 

 Distance of 
 
 
 400 meters. 
 
 1. 
 
 5 
 
 min 
 
 2. 
 
 7 
 
 min 
 
 3, 
 
 4. 10 
 
 min 
 
 5-8. ? 
 
 min 
 
 
 Distance of 
 
 
 650 meters. 
 
 1. 
 
 4:% min. 
 
 2. 
 
 5^4 min. 
 
 3. 
 
 5}4 min. 
 
 4. 
 
 7% min. 
 
 5. 
 
 9 min. 
 
 6, 
 
 7. IOI/2 min. 
 
 8. 
 
 I min. 
 
 22 Natural History of the Honeybee. 
 
 on a quiet sunny day at distances of 350, 400, and 650 meters from the hive. The result 
 is as follows, from an observation of twelve minutes at the hive entrance: 
 
 Distance of 
 350 meters. 
 
 1. iy 2 min. 
 
 2. 2% min. 
 ; j3. 2 1 / 4 min. 
 
 : 4. 2^4 min. 
 
 5. 2% min. 
 
 6. 3% min. 
 
 This time experiment alone proves that there can be no explanation for an "unknown 
 force" which "draws to the hive like a magnet;" for if it really existed these great 
 differences in time would be entirely inexplicable. Since there was no wind, and the 
 sun was shining, there were no hindrances to flight, and the bees ought to have reached 
 the hive-entrance in the shortest period and at the same time. Further, supposing the 
 existence of an unknown force which draws bees back to the hive directly, the length 
 of time for flight is in itself too long, because bees fly very quickly. A carrier pigeon 
 would undoubtedly cover the same distance in question under the same favorable con- 
 ditions in eighteen, twenty-four, and thirty-nine seconds, if we use as proof the calcula- 
 tions of Ziegler on the swiftness of carrier pigeons.* 7 Their flight is found to be often 
 as swift as 1000 meters per minute. If we now take the flight of bees to be half that, 
 500 meters per minute (a velocity which has been directly observed by Cowan es ) then 
 the ' ' unknown force ' 'should have led the bees back in thirty-six, forty-eight, and seventy- 
 eight seconds. If we double the time because the bees in the experiment were carrying 
 honey, 00 the time would be normally about one minute! twelve seconds, one minute 
 thirty-six seconds, and two minutes thirty-six seconds. Comparing this with Bethe 's 
 table we find a striking difference which is unintelligible if we accept thq idea of the 
 "unknown force," but this difference is easily explained if we consider that the bees 
 had to orient themselves with their eyes, and that they at first made errors in direction 
 so that they found the hive singly after three and one-half, ten, and ten and one-half 
 minutes had elapsed. 
 
 Of the nineteen bees which were let fly from a courtyard into the street, Bethe observed 
 that seventeen of them ("at about half the height of the houses," "before they flew as 
 high as the roof") "took the exact direction" for the hives at the Institute. That these 
 bees probably followed only light stimuli in taking this direction I have said before, 
 for that they really did not return directly to the hive is shown by the above table. 
 If all the bees, ' ' almost without exception, ' ' which were set free in an " unknown neigh- 
 borhood" under the conditions cited, took "exactly the right direction" they would 
 have come in very much more quickly. 70 I would here refer further to what I inci- 
 dentally work out from the box experiment (see p. 25). 
 
 The "unknown force" does not lead the bees back to the hive, but, Bethe suggests, 
 to the place where the hive stands or has usually stood. Is not this memory of location? 
 What does Bethe understand by the "unknown force"? We receive no explanation. 
 He thinks it is a power lying outside the bees which draws them to this place in 
 space as a magnet draws steel" (Bethe, p. 93). He^ has made no investigations to find 
 out whether this unknown force is connected at all with any organ of the bee. 
 
 But we shall attempt to get nearer to this mysterious power. 
 
 67 H. E. Ziegler, Die Geschwindigkeit der Brieftauben, Zoolog. Jahrb., X. Bd., 1897. 
 
 ^Thos. Wm. Cowan, The Honey-bee, its Natural History, Anatomy, and Physiology." London, 1890: 
 German edition, Braunschweig, 1891. 
 
 ""One can not take any bees at all for this experiment; but such as come into the hive laden must 
 be chosen, for otherwise we have no guarantee that the bees will come home by (he shortest wav and 
 not go foraging first." Bethe, 1. c, p. 87. 
 
 70 The assertion that beesf would fly directly to the hive, but that on the way they alighted at any 
 time, then flew again in the direct line, is in contradiction to the habits of tees. They flv many 
 kilometers without resting. 
 
Natural History of the Honeybee. 2H 
 
 DISAPPEARANCE OF THE MEMORY FOR LOCATION THROUGH NARCOTIZATION. 
 
 If bees are deafened with chloroform, ether, saltpeter, puff-ball, etc., the memory 
 for location entirely disappears. After waking from the stupor^ they may be taken from 
 the hive and they will not fly back to the place in space in which they are "accustomed 
 to find it." They no longer recognize their own home nor the place where it stands; 
 they have forgotten everything previously known. But an animal that can forget must 
 have possessed something to remember. The memory pictures are wiped away. We 
 see that the "unknown force" is identical with the memory for location which builds 
 up memory pictures. 71 
 
 It is necessary to mention that bees which have been stupefied become entirely normal 
 again — that is, they orient themselves with regard to their new home, and gather new 
 memory pictures by which they find once more the places for nectar and pollen as well as 
 their new hive. 
 
 Bees, therefore, have capacity for learning — a fact which Bethe disputes. 
 
 Let us look morel closely at the quintessence of Bethe 's investigations. Upon pages 
 81 and 89 he formulates the view about the "unknown force," the counterpart of which 
 is stated in this chapter (see p. 19). On page 94 we find, further conclusions in the 
 following words: "Yet I must repeat: Bees follow a force which: is entirely unknown 
 to us, and which causes them to return to the place in space from which they flew. 
 This place is usually the hive, but it need not necessarily be. The influence of this force 
 reaches over a circle of only a few kilometers." 
 
 But here we have no "repetition" of what has been said, but something that, on 
 closer examination, shows statements which are. contradictory among themselves, and 
 which contradict earlier assertions, if one takes into consideration the observations which 
 Bethe uses in formulating this conclusion (Bethe, p. 93). We shall now look somewhat 
 more closely at these observations. 
 
 THE BOX EXPERIMENT. 
 
 If the unknown force acts at a distance around the hive of only three or four kilo- 
 meters, as Bethe asserts, then all the bees of this colony which were set free within 
 this circle must return unresistingly to the place from which they flew which "draws 
 like a magnet." But this is not the case, as Bethe himself proves in the experiment 
 given on page 93. He says: "In my first experiment with letting bees fly from other 
 places I observed the following: I placed the box, in which the bees were transported, 
 on one of the large pieces of sandstone lying around in a stone-cutter 's yard and opened 
 the lid. The bees all flew out; and most of them, after a few circles ii^ the air, went 
 in the direction of the Institute. Two bees mounted to a height of about three meters, 
 made a few circles of four or five meters in diameter, and then alighted on the box. 
 I drove them away into the air again. They flew in large circles around it, and then 
 again lighted on the box. Now I took the box away and put it on another stone, having 
 driven the bees into the air once more. Both bees flew so high that I could no longer 
 see them; but a few seconds later they sank again and gradually flew around the place 
 where the box had stood." 
 
 We see, then, that a few bees were influenced by the unknown force, but others were 
 not. This changing condition Bethe can not and does not attempt to explain. 
 
 That the bees alighted on the stone where the box had been (which fact Bethe tells 
 with astonishment) is very easily understood, and furnishes further proof of the excellent 
 sense of locality in bees, for their admirable orientation with their eyes. Bethe 's objection, 
 "if they had been influenced by chemical or light reaction, then they would have flown 
 to the stone only two meters distant, and easily visible, on which the box was placed. 
 But they flew back to the place from which they had flown," signifies anthropomorphism 
 brought in in its highest potentiality. It is not possible to conclude for bees that they 
 would look for a box two meters away, which before had stood in the other place. Their 
 
 7i I might ask our ant-investigators to take up similar experiments with ants. It would be of great 
 interest to know whether a 'similar condition can be demonstrated. 
 
24 Natural History of the Honeybee. 
 
 memory of locality leads them with infallible certainty back to the place they had impressed 
 upon themselves by the circling orientation. 
 
 But why did these bees not fly back to the Institute if they possess such an infallible 
 sense of locality? For the simple reason that they were young bees which had not yet 
 developed their orientation by flying out, or perhaps old ones that had not taken their 
 orientation flight till to this spot. No one who is thoroughly familiar with the nature of 
 bees can have the slightest doubt about it. 72 
 
 But because of these experiments and others, Bethe was forced to take this position 
 in formulating hia final conclusions: "The unknown force causes the bees to return to 
 the place from which they flew," and, therefore, to modify his earlier declaration that 
 they return "to the place in space where they are accustomed to finding the hive." The 
 box experiment can not explain both the habitual return to a dwelling and a single return 
 to a place. The return to the place from which they flew takes place in those bees which 
 returned to the box and not in those which did not "return to the place from which they 
 flew" but turned toward the Institute. I must confess that I find no logic in these con- 
 tradictory statements. 
 
 Yet hear Bethe further: "I often repeated these experiments later. The further 
 away the bees were taken from the hive, the fewer returned to it; the more flew back to 
 the place from which they had flown. 73 I chose open fields for these experiments where 
 there were no large objects by which the bees might orient themselves optically. The 
 box was taken up immediately after the bees had flown out. I marked the place in the 
 grass accurately and stepped back a few steps. The bees returned with accuracy, and 
 made scarcely a mistake of more than a few decimeters. Many trod exactly upon the 
 spot. Often they stayed in the air for a minute, then returned. Once I noticed a bee 
 alight on some salvia in the meadow to take nectar there; then it flew away again and 
 then returned to the place from which it had flown. It was most startling to me when 
 I held the box open in the air until the bees had flown away and then stepped aside a few 
 steps. Four or six bees, after circling in the air, returned to the very spot where I had 
 held the box, making small circles at about the height of a man." 
 
 I can only remark here that, even if there were not objects in the meadow optically 
 discernible to human eyes by which they could orient themselves, there may have been 
 innumerable landmarks for bees. No one will doubt this who knows anything about the 
 wonderful certainty (far surpassing human ability) with which bees in swift flight pick 
 out their own homes among hundreds of bee-hives placed close together and of bewildering 
 similarity. And if Bethe could manage without marking accurately the place "in the 
 air ' ' (which I must say to my shame I did not succeed in doing in my control experiments, 
 in spite of eyes which are sharper than ordinary), then the bees might have been able 
 to do it too. In order to give the bees the least possible clue (the height of a man) I did 
 not stand but lay down and noted approximately the large circles of searching bees in 
 the air in the direction of the place from which they had flown. I can, therefore, find no 
 support in the experiment of Bethe for an unintelligible "unknown force." Moreover, 
 a few other control experiments gave such interesting and divergent results that I must 
 enlarge further upon them. 
 
 I let a few bees fly from a box placed on the short grass of a wide meadow, approxi- 
 mately five hundred meters from the apiary. I quickly stepped a few paces to one side, 
 greatly changing my position, as I did also in the following experiments. The bees 
 
 72 1 might point out in this connection that we see an analogous condition in carrier pigeons. In 
 Prof. H. E. Ziegler's writing already cited, we find the following abstract, Ueber die Orientierung der 
 Brieftauben: "After all I have read and heard of the flight of carrier pigeons, I am of the opinion 
 that their orientation depends upon memory alone, and that it is unnecessary to ascribe it to a mysterious 
 sense of direction." If pigeons are taken away to a place where they no longer have any point of 
 orientation they take up various directions and some make mistakes. The fanciers' society will not 
 undertake to send out pigeons from a distance for which they are unprepared, for in that way there is 
 constantly a greater or less loss of good pigeons. "The pigeon- fancier drills his pigeons in established 
 routes, and thus takes them by degrees to places further away, always in one direction." Also, pigeons 
 tan orient themselves in fog and darkness just as little as bees can. 
 
 73 This statement coincides excellently with, the foregoing statements against the existence of an 
 unknown force. The logical consequence from this assertion is that the "unknown force" compels bees 
 to fly in two different directions — to the hive and to the box ; the one here and the other there. This, 
 to be sure, can not be brought into harmony with what Bethe before considered the influence of the 
 unknown force, nor with the assertion that the "impulse to return to the hive is the strongest of all," etc. 
 
Natural History of the Honeybee. 2o 
 
 mounted ; slowly in widening circles, but keeping very low on the whole, at perhaps twice 
 the height of a man. After a minute one bee returned to the box; ir< one and a half 
 minutes, a second came. Then I drove both these off and took the box away. During 
 the next minute none returned to the place where the box had been, but I saw a number 
 zealously seeking, flying close to the grass in large circles, then flying a little higher. 
 Suddenly, for some unknown reason, numerous bees in a second box. under my coat buzzed 
 loudly and almost immediately the searching bees buzzed around me (I was standing 
 in the direction of the wind) so strikingly that my companions beside me exclaimed, 
 "They have heard the other bees.". Although I now replaced the box to its former posi- 
 tion it remained disregarded; but whenever we went in the near neighborhood the bees 
 followed us, and some alighted on my shoulders, hat, and coat. When I went further 
 away my companions were surrounded, although they had no bees with them. It is 
 noteworthy that the bees in the second box belonged to another colony. 
 
 I shall now describe the second experiment. The other box was held up high; the 
 inmates about thirty or forty in number circled in a small radius for a long time, then 
 gradually made larger circles. Some worked in a more up-and-down fashion, close to 
 the box, continually turning the head toward it, just as in? the first orientation flight from 
 the hive, about which I shall speak more in detail further on. About half a minute after 
 I took the box and drew back quickly, trying to impress accurately on my mind the spot 
 at the height where the box had been. Two bees appeared in a fev^ seconds, but in a 
 place a little to one side; and then, after making large circles, disappeared again. The 
 chief thing to be seen was the irregular search which they all carried on together after 
 a few minutes in ever narrowing circles, 74 apparently flying close over the ground/ about 
 two meters from the place where I had stood. The bees were no longer seeking the box, 
 but their home, the entrance of which (according "to my estimate) was just at the height 
 above the ground of the circles which they were making. I can give no other explanation 
 for this singular phenomenon. After the bees again wandered aimlessly for some time, 
 flying around us part of the time as described earlier, they followed us homeward for 
 about twenty paces, then turned back. That proportionately few flew back to the apiary 
 is explained by the fact that the bees shortly before had come from the moor, and in 
 the cool autumn weather could hardly have flown out, so that an orientation from the 
 apiary was not yet possible. The day of the experiment was sunny, with a moderate 
 southwest wind. 
 
 The -striking occurrence, that my companions and I were so thickly surrounded, can 
 be interpreted in so many ways that to explain it would lead us too far. 
 
 In the foregoing I see many proofs for the existence of memory for locality; but all 
 these box experiments are untrustworthy, since the investigations go on under entirely 
 abnormal ^ condition for the bees, and the results do .not harmonize with those gained 
 from the apiary. It seems very risky to me, therefore, to draw conclusions from them 
 for normal conditions, since we may easily be deceived. The time of year and the weather, 
 as well as the management; of the bees, will greatly modify the results of this kind of 
 experiments. If it is recognized how easily bees are influenced by various stimuli (e. g., 
 shaking, cooling, breathing on them, confinement, feeding, steange strong odors, etc.), 
 I say influenced, diverted, "confused," "subdued," "goaded on," or "calmed," then it 
 will be realized how difficult is the interpretation of such experiments and how cautiously 
 one must work on them. 
 
 If one ask Bethe why the unknown force acts at a distance of only three or four 
 kilometers, we receive no explanation from him: he does not even attempt to give one. 
 The explanation, however, is very simple. The mysterious force acts within the space in 
 which the bees have previously taken their flights of orientation, and acts only as far as 
 they have flown at any time and gathered memory, pictures. The proofs for this are 
 easy to bring forward. They show on the other hand that this unknown force so desig- 
 nated is identical with memory of location. 
 
 74 One sees from this that bees pay attention to each other, since they keep together and exhibit the 
 same behavior. Imitation instinct? Sense of hearing? 
 
26 Natural History of the Honeybee. 
 
 1. If young bees able to fly (brood-nurses), which have not yet had their flight of 
 orientation, are let fly not far from the apiary, none find their way back to the hive. 75 It 
 is also significant that first swarms, which are almost entirely made up of old bees, in 
 the loss of the queen quickly re-enter the mother colony, while an after-swarm, for the 
 most part composed of young bees, buzzes about for a long time, finally entering strange 
 colonies. 
 
 2. If old bees are let fly at a much greater distance they all find their way back. 
 
 3. If a colony is brought from a place more than seven kilometers away, and the 
 old bees are let fly only thirty or forty meters from their hive before they have been able 
 to make their flight of orientation, none find, their way back into the hive, supposing that 
 (in such a short distance) houses or trees intervene between it and the place from which 
 they were set free. 
 
 4. I had two colonies placed in the garden of the Zoological Institute in Jena for 
 the purpose of further investigation. At the end of the summer semester of 1899 they 
 were taken to the apiary of a bee-keeper about two thousand meters distant. Since the 
 colonies were not stupefied, it was presumed that many of the old flight bees would return 
 to the Institute apiary, and, for the refuge of these homeless ones, I placed a hive with 
 some empty frames exactly where their home had stood. Many hundreds returned, which, 
 in spite of complete freedom of flight, loitered around the empty hive uneasily for two 
 days. They were afterward chloroformed, and preserved in formaline, for purposes of 
 demonstration. Their memory of locality had led them back. 
 
 Naturally and easily, the varied conduct of bees in these four cases is explained if 
 we accept the theory of an orientation with the eyes^ through memory pictures, while the 
 unknown force entangles us in contradictions and inextricable mysteries. 76 
 
 It seems remarkable that Bethe attempts throughout to explain "how bees find their 
 way home, ' ' but never, how they find their way away from the hive. With regard to their 
 finding their way back to a place outside of the hive, he says only the following (Bethe, 
 1. c, p. 90): "The place where a honey supply has been found is again sought, I? think, 
 not because of memory of locality, but because of a reaction of the same force unknown 
 to us, etc." Bethe here refers to the observations of Lubbock (1. c.) and Forel 77 who put 
 down vessels containing honey, and observed that marked bees always returned to them. 
 
 Because of these assertions, Bethe's theory is still more complicated, and more difficult 
 to understand. We must not conclude, therefore, that "bees follow a force which causes 
 them to return to the place in space from which they flew, which place is generally,' but 
 not necessarily, the hive, ' ' for in an acceptance of this opinion the bees would have to 
 swing to and fro like a pendulum forced always between their dwelling and the honey 
 supply, whether there is any honey there or not. If they have flown from the hive or 
 from the place where the honey-receptacle was, the unknown force "impels" them to 
 return to both places. 78 
 
 75 As is well known, young bees fly for the first time about ten to fourteen days after they emerge. 
 During the first two weeks their existence is that of "house beds," "nurses," who perform all the house- 
 keeping and feed the larvae. This is, I might say, the firmly established management, from which there 
 is no deviation. Yet we can modify this activity substantially. For example, if we* form a colony of 
 bees which have just emerged, and give it a fertile queen, brood, and frames of honey, then we would 
 see that part of the young bees become "field bees" in five or six days, thus taking up the outside work 
 considerably earlier, even if everything* is present in the hive, which is necessary for the existence of the 
 colonv. 
 
 76 Bethe refers at different times to Fabre's investigations on Chalicodoma (Fabre, Souvenirs ento- 
 mologiques: Paris, 1879; Fabre, NouveaAix souvenirs entomol. ; Paris, 1882.). Yet as early as 1895 
 Weismann (Weismann, Wie sehen die Insekten? Deutsche Rundschau, 1895, pp. 434-452), had over- 
 thrown, in a very interesting paper, Fabre's view concerning the sense of direction. Weismann comes 
 to the following conclusions: "The only correct solution of the enigma of path-finding by Chalicodoma 
 is that the insects find their way back with their eyes." 
 
 77 Morel, Recueil zoologique Suisse, 1 Serie, T 4, 1886-88. 
 
 78 If an objection were here to be made, that very probably bees are compelled by the unknown 
 force to return to the place where honey or some stimulus is placed, just as long as the stimulus is 
 present, and that the action of the unknown force could cease with the disappearance of the stimulus, 
 such an exception might appear unauthorized for the following reasons: We should have two entirely 
 different kinds of unknown forces according to this view, since the one leading the bees back to the 
 hive, in fact, forces them to return to the place where the hive stands or no longer stands. Here the 
 stimulus, therefore, is acting in spite of the fact that the means of stimulation have been removed. Then 
 we should have a second force which vanishes with the disappearance of the means of stimulation. Some- 
 thing very different from Bethe's definition, which is not consistent, and decidedly not all conclusive, 
 occurs, for the observation may be made frequently that food or meal remains unnoticed, and is not 
 carried in, if Nature opens stores of honey and pollen (Dathe, 1. c, p. 176). Here the bees no longer 
 
Natural History of the Honeybee. 
 
 THE LOSS OF MEMORY FOR LOCALITY THROUGH SWARM DIZZINESS, ETC. 
 
 Through swarm dizziness, as well as through stupefaction (as before mentioned), 
 memory pictures are extinguished, or, at least, are without influence. 
 
 If a so-called artificial swarm is made by sweeping the bees from the frames of a 
 strong colony with their queen into a new hive placed upon a new stand, then all the 
 flying bees return to the original hive, and only the young bees remain in the new one 
 with the queen. In a genuine swarm, on the other hand, all the bees remain in the dwelling 
 of their choice. They have forgotten their old dwelling. But it is not a complete for- 
 getting, for, if a swarm becomes queenless within the first few days, then the bees 
 return to the mother colony— the memory for the old home is reawakened. The extinc- 
 tion of the memory for locality is not, therefore, as final as in narcotization, etc. The 
 old nerye-paths are maintained, but are no longer traveled over, because there is a diversion 
 into other nerve-paths; but if the stronger influences are removed by queenlessness (Weisel- 
 unruhe, see p. 12), then the old-trodden paths come into effect again and adjust the 
 direction of the stimuli in the former way; i. e., earlier memory pictures are reawakened 
 and the bees return to the mother colony. 
 
 An extinction of the memory of locality is brought about likewise by the apparently 
 narcotic influence of buckwheat honey 79 in the cases mentioned before. 
 
 Also bees kept in a dark room for many days, and those numbed with cold M appear 
 to lose their earlier memory for locality to a certain extent. Further, the throwing of 
 bees into water, the bathing of a colony, will cause the disappearance of the acquired 
 power of orientation. 81 A colony thus handled can be placed in a different position with- 
 out a return to the accustomed place of flight. Time has a substantial influence upon the 
 disappearance of memory pictures too. In approximately five or six weeks, or often 
 sooner, bees removed to a new position forget the influences of the old place. After this 
 length of time the hive can be changed back and put in any chosen position of the old 
 location without fear of the bees seeking the original spot. Memories disappear quickly 
 if new impressions obliterate the old. If bees stay in a hive, wintering for months, there- 
 fore receiving no new place impressions, the impressions which were received before the 
 wintering commenced remain. In very many cases it can be determined certainly whether 
 a transference shortly before the first spring flight can be undertaken without much loss 
 to the colony. If the first weather for flying is inauspicious, as is; generally the case, and 
 the temperature scarcely reaches the 7° to 8° C. in the shade necessary for flight, the 
 bees fly out, lingering for only a short flight with slow orientation, and execute the neces- 
 sary cleaning. They thereby impress upon themselves the new position. But if, as happens 
 now and then, after a long period of cold, a relatively very warm spring day breaks in, 
 the excitement in the hive is great; thousands press forth, and many hasten off for a longer 
 flight with only a hasty, careless orientation. Under such circumstances a greater or less 
 number, in coming back, return to the old place. 
 
 Francois Huber 82 reports that in the fall he had fed some honey to great numbers of 
 bees from a window; then the honey was taken away and the hives were kept closed all 
 
 return to the feeding-place, in spite of the fact that food is still present. Very significant for this view 
 is the following account by the well-known writer in apiculture, and editor of the Bienen Zeitung, Wilhelm 
 Vogel: "When I was a boy a beekeeper told my father, under the seal of secrecy, that a colony would 
 swarm in a few days if it were given daily a quart of honey. I immediately placed a dish of honey over 
 my strongest colony during the buckwheat-honey flow. After eight good days of forage the bees hadi not 
 touched the honey, although it was pure." (Bienen Zeitung, 1880, p. 10.) I might mention, likewise, that 
 the statements of Lubbock and Hermann Muller, about the pretended incapability of bees of not again 
 finding the placed honey will be explained through the mentioned characteristics of bees. The "slight 
 ability" of bees accepted by these investigators is, therefore, not proven through these investigations. In 
 a dearth of honey, on the contrary, the bee is very able to trace out places where extra-floral honey is to 
 be discovered. 
 
 79 If the buckwheat-honey flow is strong, then the bees seem to be unmistakably intoxicated, and they 
 go into the nearest hive-entrance with filled honey-sacs. The observation has been made that hives which 
 are passed over by bees from hives standing further back increase perceptibly in population and honey 
 at the cost of the hives behind" (Bw. Centralblatt, Nr. 3, p. 35, 1894) . 
 
 80 Deutscher Bienenfreund, 35 Jahrg. 1899, Nr. 4. 
 
 81 Francois Huber, 1. c. In men, after a violent illness, after a concussion of the brain, after stupefi- 
 cation, after poisoning with carbon dioxide, etc., a loss of memory and retrograde amnesia occurs (Aug. 
 Forel, Das Gedachtnis und seine Abnormitaten, p. 37, ff . ; Zurich, 1885; A. Goldscheider, Die Bedeutung 
 der Reize im Lichte der Neuronlehre, p. 28, ff . ; Leipzig, 1898). 
 
 82 Nouvelles Observations sur les Abeilles, 2d edition; 2 vol., Paris and Geneva, 1814; German by 
 Kleine, Einbeck, 1856; English editions in 1823 and 1841. 
 
^8 Natural History of the Honeybee. 
 
 winter. When they were again opened in the spring, the bees came again to the window, 
 in spite of the fact that there was no longer any honey there. 
 
 Bethe (1. c, p. 90) asserts as follows: "If bees cease to fly in autumn, then (as I 
 have proven myself) hives may be placed even inside a circle three kilometers in diameter, 
 each in a chosen spot. The bees do not return to the old location of the apiary when they 
 begin to fly in the spring, but from the first come to the hives in their new positions. ' ' 
 
 This experiment of Bethe 's is not conclusive and I would not recommend any investi- 
 gator or bee-keeper to try it in the way described, for in nearly all cases all the flying 
 bees will be lost. 
 
 If, following Bethe ? s suggestion, a colony is put in a different place in the fall when 
 "the bees have ceased flying" because of the cold weather, one can not tell whether, in 
 spite of the apparent approach of winter, in the next few days or in the first .three or four 
 weeks, a warm period with weather suitable for flight may not set in, during which the 
 bees still have the old position securely in memory. If this happens (and it frequently 
 does), then all the flying bees return to the old place and are lost. The next spring, of 
 course, no return to the old locality can take place, for all or nearly all that would recog- 
 nize the former position are already dead. 
 
 ASSOCIATION OF IMPRESSIONS. 
 
 During the dearth of honey, when bees are usually excessively fond of sweet things 
 (see p. 26), a few bees discovered a honey-comb in my study, flying through the open 
 window. More and more nibblers came, some of them getting caught at the second closed 
 window. To prevent that, I placed the honey-comb in the opened window itself. When, 
 half an hour later, the bees were flying back and forth I drove them away from the honey 
 and closed the window. After perhaps twenty minutes I went into the bed-chamber above 
 the study, the windows of which stood wide open, and found it full of bees. By this time 
 I was observant, and after driving out the searchers and closing the window I went into 
 the garden and noted their behavior accurately. They attempted in vain to press in at the 
 window where I had fed them, then from time to time some flew to the next window and 
 tried their luck there, then further to the next window above, always keeping about a 
 handbreadth above the window-sill, the height at which the honey had been in the window 
 where they were fed. Thus I noticed bees seeking at all the windows of the house. 
 
 If the bees had actually associated impressions and connected the attainment of honey 
 with the form of a window, then it was to be supposed that they would extend their 
 search to the windows of the neighboring house about ten steps to one side, which, in fact, 
 happened. 83 
 
 MEMORY FOR THE FEEDING PLACE IN THE HIVE. 
 
 If an inner glass or wire-gauze door is kept closed, the back door of a hive may bo 
 opened without the bees getting out. Between these outer and inner doors there is usually 
 enough room to place a feeder. If this is filled with honey or sugar syrup, the bees can 
 reach the food when the slide in the glass door is opened, the outer door being closed. 
 If one is feeding for the first time, the bees may be guided to the honey by placing a few 
 upon it, or in some other way, for otherwise they do not transfer the food as quickly as 
 may be desired, because it takes some time for them to find it. To prevent the excitement 
 caused by feeding, this was done in the evening, and the empty dish taken out the next 
 morning, the slide of the glass door being again closed. But on the next evening, or the 
 one after, I noticed many times that, if I opened the glass door, the inmates .streamed 
 out so quickly that I had to hasten to close the outer x door in order not to crush any. Also, 
 if the feeding were done in the open air the bees often came hours and days afterward to 
 the place where they had once obtained honey. 
 
 83 Under the designation, "Instinct or Intelligence," I find in the American Bee Journal, Chicago, 
 1892, the following note: "If a dish of food is placed on the trunk of a tree, bees will soon search for 
 honey in the same place on all the trees in the neighbor-hood. A similar, thing., happens if bees art fed 
 from a window facing south. Soon bees are discovered seeking at the south side of all the neighboring 
 buildings." 
 
Natural History of the Honeybee. 29 
 
 It would be difficult to doubt) that here we have to do with memory processes; for 
 if bees are reflex machines, following ' ' chemotropically ' ' only adequate mechanical stimuli, 
 then we can not understand why, when the stimulus has not been present for a long time, 
 they react again and again as if the stimulus still existed. A plant never reacts helio- 
 tropically or chemotropically to a past stimulus, etc., if the respective stimuli are no longer 
 acting upon it. But here we see movements set loose, although the cause — the honey on 
 the feeding-spot in this case — no longer exists. We see bees once fed, often, after two days, 
 seek that place in vain; but then they modify their procedure; they learn that there is no 
 longer anything to be obtained there, and cease from further flight, often after a few 
 hours, as before mentioned. 
 
 CONDUCT OF BEES IN THE BUCKWHEAT SEASON. 
 
 If colonies stand in buckwheat, the flight is lively in the mornings until about ten 
 o'clock; then it lessens, and is entirely quiet for the greater part of the day, beginning 
 vigorously again the next morning. The buckwheat honey flows only in early morning; 
 so, as the nectaries dry up, the bees fly out a couple of times and then discontinue their 
 vain flight. In spite of the shimmering sea of) flowers, in spite of the strong fragrance, 
 only a few bees may usually be found after ten o 'clock in the buckwheat -field. 
 
 Here the stimuli of color and scent are constantly present, and there is also the habit 
 of daily flight to the same fields; and, in spite of that, we see that flying is discontinued. 
 Here, undoubtedly, as well as in the preceding feeding experiments described, the processes 
 of learning and remembering may play their important part. 
 
 ARE BEES ATTRACTED BY THE COLOR OF FLOWERS OR BY THE NECTAR? 
 
 This might be a good place to consider briefly this interesting question. While Plateau m 
 substantially advocated the view that bees are attracted chiefly by nectar and not by color, 
 it was above all Aug. Forel M who stood almost alone in opposition to this view, based 
 on many years of admirable experimentation. Recently some younger investigators have 
 been associated with him, who can experimentally verify his conviction that it is chiefly 
 the color which serves to attract. Upon the foundation of the work of Forel, Andreae, 87 
 Giltay, 88 Detto, 89 Kienitz-Gerloff, 90 we can establish as proven the statement that the honey- 
 bee, Apis mellifica L., is attracted substantially by the color of flowers, and not mainly 
 by the nectar. The color of the flower is indeed a gay flag which proclaims at a distance, 
 "Here there is something to sup. " 
 
 That bees observe flowers keenly, follows from the fact that single bees in foraging 
 practically never visit two kinds of flowers, but always hold to one kind. This may easily 
 be seen by examining the pollen-sacs on the return of the bees to the hive. One color of 
 pollen is always seen; a mixture of colors, I have observed but once. 
 
 PLACE PERCEPTION IN THE QUEEN. 
 
 In the literature of apiculture the observations concerning the memory of locality in 
 the queen, so far as the duration of this memory comes into question, diverge very far 
 from each other. Some say that she has capacity for remembering her hive, the outside 
 and surroundings of which she learned to know in her single virgin flight, for more than 
 three years; others say, for some days or weeks. The source of failure in these observations 
 is connected with the frequent unnoticed changes in queens going on unaware of the bee- 
 keeper. He often thinks that he still has the old queen in the hive when she has been 
 replaced for some time by a new one. 
 
 85 Plateau, Felix, Comment les Fleurs attirent les Insectes. Bull, Acad. roy. d. Belgique, 3 serie, 
 T. 30 ,1895; T. 32, 1896; T. 33, 1897; T. 34, 1897, Recherches experimentales sur la vision chez les 
 arthropodes, ibid., 1888, etc. 
 
 SG Forel, Aug., Recueil zoologiques Suisse, 1 serie, T. 4, 1886-88 (also as separate); Experiences et 
 remarques critiques sur les sensations les Insectes, partie I.-V. ; Munchen (Reinhardt) or Paris (Kling- 
 sieck), 1900, 1901; Die psyschilschen Fahigkeiten der Ameisen und einiger anderer Insekten, Munchen, 1901. 
 
 87 Andreae, Eugen, Inwiefern werden Insekten durch Farbe und Duft der Blumen angezogen. Beihefte 
 z. Bot. CentralbL, Bd. XV., Heft 3, 1903. 
 
 88 Griltay, E., Ueber die Bedeutung der Krone bei den Bluten and uber das Farbenunterscheidungs 
 vermogen der Insekten, I. Pringh., Jahrb. f. wiss. Bot., 40. 1901. 
 
 89 Detto, Carl, Blutenbiologische Untersuchungen, Tbeil I. u. II. Flora oder Allg. bot. Zeitung., 94, 
 Bd., Heft 2 u. 3, 1905. 
 
 **> Kienitz-Gerloff, Professor Plateau und die Blumentheorie I. u. II. Biol. Centralbl., 18, 1898, u. 
 23. 1903. 
 
30 Natural History of the Honeybee. 
 
 It is true that queens which fly away from the comb just taken out of the hive usually 
 get lost because they can not find their way home. 
 
 I made the following observation: When a queen flew away as just described, I stood 
 perfectly quiet, exactly as 'I was when she flew, in the hope that, if she had no memory 
 of her hive, she would surely return to the place from which she flew, since bees which 
 have not before oriented themselves always go back to the place from which they flew. 01 
 After barely a quarter of a minute, in fact, the queen sank down again, not on the comb, 
 to be sure, but upon a little piece of board which lay in the grass a step distant. There I 
 caught her easily and put her back in the colony. In keeping quiet and not changing the 
 surroundings I followed an old bee-keepers' rule, a practice developed from much obser- 
 vation like the above. 
 
 But it has, undoubtedly, been established many times by reliable observers that a 
 queen finds her way surely to the entrance of her hive a month after her marriage-fight. 
 
 A young queen often errs when the hives stand very close together, if she returns 
 hastily from the marriage-flight. If hives from which flight will take place are marked, 
 therefore, for example with a twig covered with leaves, 92 her return is very much more 
 assured. This is proof that the queen impresses accurately upon herself the appearance 
 of her dwelling. 
 
 MEMORY FOR LOCALITY IN SCOUTING BEES. 
 
 In the behavior of the so-called scouting bees we find one of the most conclusive 
 proofs against the vague hypothesis of an "unknown force." They prove most em- 
 phatically, in my opinion, that an orientation truly takes place through the sense of sight, 
 through memory pictures. 
 
 I can not here deny myself the pleasure of inserting the interesting information about 
 scouting bees which Baron v. Berlepsch, the so-called Bee Baron, 93 sent to the Bienen- 
 zeitung, VIII., No. 7, 1852. 
 
 "Annually, about swarming time one often sees bees in considerable number at holes 
 in the walls and crevices of old buildings, walls, and trees. They creep in and out, appar- 
 ently seeking something, run anxiously outside and back, flying to and fro. They buzz 
 about as if in front of the hive, and one must have considerable knowledge of bees to 
 be able to distinguish these so-called scouting bees from the true colony. Even if there is 
 no room behind the hole or crevice, I have often seen them forming a cluster before one 
 from six to eight inches long, two or three inches broad, but not at all deep. In so doing 
 they were always uneasy, which does not happen, as is well known, with bees in front 
 of a real hive. These bees are usually regarded as belonging to colonies about to swarm, 
 sent out to find a good place for quarters for the next swarm, hence the name scouting 
 bees or quarter-makers. I see these beeg every year at the crevices of the old knight's 
 castle, and in the garden wall ancfi the barn gable. These crevices are often hardly one 
 inch deep, and one-half inch wide, so that there is not room for even the smallest after- 
 swarm. My observation that, although indeed no year passed in which not one but most 
 of my after-swarms escaped, a swarm never alighted here, led) me to doubt the common 
 belief; and 1844 I determined to make very exact investigations andi experiments, and 
 to take careful note of everything. When, therefore, on May 12th of that year I first 
 saw bees on the edge of the wall, I had them sprinkled thoroughly with chalk toward 
 evening by my usual helpers, and the gardener and I stood in front of the hives to see 
 to which the bees belonged, and whether they were all from one or from more than one 
 hive. Soon coming in, they all entered No. 77 ('Solomon the Magnificent'). Early the 
 next day they were on the wall again^ and so it went on for four days. Each evening 
 they were powdered, and their return carefully noted (scouting bees had in the meantime 
 appeared at many other places). They belonged, undoubtedly, to the magnificent Solomon. 
 Finally on the 17th, about ten o'clock, the powerful ' Padishah' swarmed out with a for- 
 midable host, went in the direction of the scouting bees, but hung on a dwarf tree hardly 
 
 91 Orientation begins, therefore, at the moment of the flying out (see Box Experiment, p. 23 ) . 
 
 92 Dathe, Lehrbuch der Bienenzucht, 1892; Bensheim, p. 279. 
 
 93 .As is well known, v. Fiebold demonstrated Parthenogenesis in bees for the first time upon the 
 estate of v. Berlepsch in 1855 (see Wahre Parthenogenesis bei Schmetterligen und Bienen:" Leipzig, 
 1856, d. 110 and following). 
 
Natural History of the Honeybee. 31 
 
 twenty steps from the hives, fully exposed to the burning sun. I let them hang, and 
 remained standing near. About eleven o 'clock they again broke loose, soon alighting in 
 a somewhat shadier place at a short distance, remained there until three o'clock, broke 
 up very quickly for the third time, and went over the garden pavilion toward the open 
 fields. My assistants ran after them. I mounted a horse as quickly as possible, and 
 galloped after them. Before I came up to the assistants the swarm was lost to sight, and 
 we have it yet to see today. The scouting bees were still on the wall, went back again 
 in the evening to No. 77, and appeared early on the 18th. This reappearance was indeed 
 a disappointment, for, powdered on the evening of the 18th, they all entered No. 7 ('Dr. 
 Franzia'), and not one to 'Solomon.' On the 19th, quite early, I had the crevices in all 
 places carefully closed with lime, and put up a straw hive with an alighting-board. The 
 scouting bees came, at first crept behind the hive around the closed crevices, then took 
 possession of the hive. I tilted it frequently to see what the bees were doing inside. 
 1 saw little; they ran around anxiously as if they might} be cleaning out the* hive.* On 
 the same day at noon a swarm emerged from ' Franzia ' and, after changing its place twice, 
 finally alighted about four o 'clock under a shady linden-tree. Here it could be observed 
 comfortably. Single bees flew in all directions, each time mapping out the place with the 
 usual circling flight, then returned; and here the swarm hung over night. With the first 
 shimmer of morning red, there I sat again with my assistants, the garden doors open on 
 all sides as on the previous day, the groom (all the implements for capturing swarms on 
 his back, a true Cupid with arrows) at a short distance with two saddled horses. At 
 5:30 I saw more bees fly quickly toward the south without circling, and none returned. 
 At 7:30 the swarm broke away again, flying low and very slowly in a southerly direction, 
 and the leaders could be distinguished rather clearly. The groom hastily mounted a 
 horse, and I went running ipsissimis pedibus beside the head of the swarm nearly to the 
 end of the garden, ever more and more convinced thai} the swarm knew where it wished 
 to go. The assistant brought out the other horse. I threw myself upon it, hastened out 
 of the garden, and followed the- groom through thick and thin. We followed well at a 
 moderate trot for a quarter of an hour, but finally the swarm went so quickly, always 
 at a height of between four and nine feet, and in a southerly direction, that we hac!j to 
 ride indeed en carriere. We came to the next village, not three quarters of an hour's ride 
 distant, and the swarm went into a farmer's garden. I went over the hedge as if in hunt, 
 was in the middle of the swarm with the horse, and then saw it enter a hollow pear-tree. 
 This move took place with such celerity and surety that it seemed to me that the swarm 
 without a doubt had chosen this place in Seebach (through scouting bees). A capture 
 without smoke was not to be thought of, so I begged the owner to let me stay in his 
 Eden for a while (for I would smoke out the swarm for him the very day into a hive, 
 and leave it to him as his property.) After scarcely twenty minutes the bees began to 
 carry out little chips from inside the tree, went foraging, etc." 
 
 Can we explain this interesting procedure only upon the ground of an "unknown 
 force leading back to the hive"? Can we explain it without accepting the existence of 
 a memory, a capacity for orientation through the eyes, through memory pictures? 'I think 
 not in the least. 
 
 Must we not here acknowledge a means of communication, a method of understanding 
 which does not rest upon odor reflexes? I can not think of this going on without some 
 means of understanding, and I presume that the scouting bees lead the ' swarm by an 
 alluring sound which naturally can not be observed. 
 
 It is noteworthy that, of 60,000 to 80,000 bees, about fifty ta 1 a hundred serve as scouting 
 bees; but, peculiarly enough, only before the swarming of the unwieldy, usually low- 
 placed first swarm' (often with an old queen heavy with eggs), while the colony never or 
 very seldom sends out scouting bees before the issue of the light-footed after-swarm which 
 with the unfertilized easily flying queen generally alights high up. 
 
 Now and then the scouting bees make it very convenient for the bee-keeper by selecting 
 an empty hive and leading the swarm there.* 4 Dishonest people who wish to capture 
 
 91 Deutsche Bienenz. in Theorie u. Praxis, Nr. 9, p. 144, 1899. 
 
32 Natural History of the Honeybee. 
 
 swarms not belonging to them put out so-called bait skeps, or hives; and if the scouting 
 bees discover these they often lead the swarm there. 
 
 One of the highest colonies. ever found (300 feet high) was in the statue of Liberty 
 on the dome of the capitol in Austin, Texas. How high the scouting bees must have flown 
 to discover this strange place, 85 and what power of allurement the explorers must have 
 used to be able to lead the' colony to such an unaccustomed height! 
 
 THE EYES OF BEES. 
 Before we turn further to other interesting observations concerning the memory of 
 locality, let us inquire why there are such large well-developed eyes in the three kinds of 
 bees, if orientation through the sense of sight be denied. This is no idle question, for 
 wherever eyes are but little or not at all used, a gradual stunting takes place, etc.; thus, 
 for example, to restrict ourselves to social insects, in the ant, Solenopsis fiif/ax, the workers 
 possess eyes with six to nine facets, while the workers of Apis mellifica have about 4000 
 to 5000, the drones about 5500, and the queen about 5000 facets on each eye, besides the 
 three ocelli which are apparently for distinguishing near objects. 96 Bethe has since rejected 
 the statement that the ocelli are probably used for seeing near objects, as incompatible 
 with the 1 1 first principles of physiology. ' ' I thereupon replied that the renowned physiolo- 
 gist Johann Miiller was the first to advance this view, and that observations' in natural 
 history confirm and even substantially support it. I have communicated the details in my 
 " Stammesgeschichtlichen Entstehung des Bienenstaates ' ' (pp. 90-95), Leipzig, 1903. 
 
 Bethe credits bees with only "a slight capacity for receiving light stimuli" (Bethe, 
 p. 82), basing his view upon an experiment which I on the other hand can not consider 
 convincing. If a large screen be placed in their way, the routine-loving bees will follow* 
 the usual path of flight until within 1 to 1^ meters of the obstacle, then suddenly rise 
 and fly over it. But this does not prove conclusively that the bees had not long before 
 seen the immense screen (two and a half meters high and. three meters broad). They follow 
 freely the accustomed path as long as possible. Bethe himself says < * that a once used 
 correlation of movements may be retained for a long time" (Bethe, p. 92). 
 
 In order to experiment to see that bees, instead of being 1 1 near-sighted, ' ' are very far- 
 sighted, stand at a distance of about ten steps from the hive at the time of the buckwheat 
 bloom, the nectar of which, as already mentioned, excites bees extraordinarily. The bees 
 coming out from the hive-entrance quickly, wholly indifferent as to the direction of the 
 wind, leave painful proof of their ability to see; and a man presents a considerably smaller 
 surface than the screen before mentioned. 
 
 "According to our knowledge, everything in living nature has a purpose" (Bethe, p. 
 19); therefore the very large eyes of bees must be for the purpose of guiding the insect 
 safely and well.* 7 
 
 The brain of the bee, very strongly developed in comparison with those of other 
 insects, is certainly not without purpose. The powerful optic lobes (Lohi optici) prove un- 
 questionably the great share that the eyes have in the nervous processes. 
 
 THE FLIGHT OF ORIENTATION. 
 How much bees need eyes in flying out from the hive, shows in a very clear way 
 from their striking behavior during the first flight. As the bees fly out they turn their 
 heads toward the hive, and in the continuous hovering up and down (resembling the 
 gnat's dance), the hive itself, the neighboring hives, and the bee-house are surveyed; and, 
 indeed, I repeat it, the head is constantly turned toward the hive, so that even a slight flying 
 backward is shown. This is the so-called first-play (Vorspiel) 98 which is not taken into 
 consideration at all by Bethe, and has not been considered by him as a specially charac- 
 teristic proof of the orientation by the sense of sight. After this short preliminary flight, 
 small and then larger orientation circles are taken, and thereby the near and distinct 
 surroundings are impressed on the memory. 
 
 95 American Bee Journal, 1892, Chicago. 
 
 ^Thos. Wm. Cowan, The Honey-bee, its Natural History, Anatomy, and Physiology ; London, 1890- 
 German by Gravenhorst; Braunschweig, 1891. a. c. ' 
 
 OT Up to the present time no one has doubted that the house-fly (about 5000 facets), or the dragon- 
 fly (about 12,000 facete), orients itself during flight solely through the excellently developed eves ° 
 
 « Dathe, 1. c, p. 146.. * ' 
 
Natural History of the Honeybee. 33 
 
 An old bee flies out from the entrance, if the forage is rich, directly and swift as a:i 
 arrow. It darts from the entrance through which it has often flown, and knows its path 
 of flight. A young bee flying for the first time must first make a flight of orientation. 
 This is a conclusive proof, that bees learn." 
 
 THE FINDING OF THE HIVE THROUGH THE SENSES OF SIGHT AND SMELL. 
 While the sense of sight is sufficient under ordinary circumstances for bees to find 
 their way home, still they use the sense of smell also under the following conditions: 
 
 If a colony has been brought from another district, and the slide is opened after the 
 hive is in place, then the bees flying out hasten away without orientation, for naturally 
 they do not know of- their change of place, and believe themselves in a well-known neigh- 
 borhood. In such cases they are seen striking out either in graded fight or mounting in 
 the well-known corkscrew lines without turning the eyes to the hive, as is always done 
 in the described flight of orientation. According to my observation, such bees, flying 
 without first orienting themselves, often come back in a surprisingly short time, because, 
 failing to see the customary landmarks, they immediately begin to seek orientation. If 
 the newly procured colony stand between others which look like it, then in finding their 
 way back there is what Herr Eoth, the leader of the bee-keepers' school at Baden, wrote 
 to me " a tasting with the sense of smell, which is extended to the neighboring hives. ' ' 
 This is a striking phenomenon easily observed. An orientation with the eyes goes on at the 
 same time, so that the real flight of orientation does not have to be carried on later. 
 
 As I have said, such oriented bees that fly out return quickly, often after^five or ten 
 minutes, with full burdens if the weather is mild and there is no wind. In cooler weather, 
 and with sharp wind, many go too far into unknown regions and do not find their way 
 back. 100 
 
 Bees remain out on flight seldom longer than one-half hour if the forage is near — 
 almost never over an hour if the forage is distant and in unknown surroundings. 101 They 
 then go home apparently willingly, with a half or a quarter of a full load, as one can easily 
 see in the tiny pollen-sacs and in the slight body. 
 
 Let us now hear from Dr. Bethe's description concerning a transported hive (1. c, 
 p f 92). "Lubbock placed some honey out in a room to which some wasps came: he then 
 closed the window which faced the nest. The wasps flew mainly against the window, 
 but finally flew through the other open window. After a few times the greater number 
 flew immediately to the open one. Since, however, a kind of ' accustoming, ' based upon 
 an unknown force, seems to enter in finding the way home in this experiment, I hoped 
 to succeed in getting results by observing accurately a new colony of bees. It was bought 
 in a village seven kilometers away, and was placed in the Institute garden about seven 
 o'clock in the morning. At first only a few bees hummed around in the air. 102 A great 
 number sat upon the flight-board and the front wall of the hive, as is generally the case 
 in excited colonies. Between nine and ten o'clock in the morning the first ones flew away. 103 
 
 99 The following observation proves how little bees acdustomed to the hive's place and surroundings 
 regard the place from which they fly in their swift exit. If the second hive-entrance, which is in the 
 upper honey-chamber (super) is opened for days bees will be seen leaving from this entrance but never 
 entering it." The entering takes place constantly in the usual path through the entrance below. Only by 
 and by bees begin to fly into the tipper one (cf. p. 40). 
 
 100 I quote the following from a private letter from Dr. Dzierzon, a distinguished bee-keeper to 
 whom we are indebted for the discovery of parthenogenesis in Apis mellifica. Dzierzon died last year 96 
 years old. "There is not the slightest doubt that bees find their way back to the hive in the way you have 
 described, guided by pictures of their dwelling and the near surroundings gathered as they flew away. 
 Instinct is the explanation in so far as they are led to observe accurately the position of their dwelling 
 and the nearest surroundings in the first flight out. On account o>£ this, as you know, in the first flight, 
 (Vorspiel), they turn around; and, forming small and then larger circles? gain an exact impression of 
 the neighborhood and the hive. I have often wondered at the rapidity with which they are able to do 
 this. I brought a hive from a distant apiary, and on opening the entrance a few bees flew out, probably 
 for water, not suspecting the removal. In the mild weather they were not lost, but came back to the 
 same place, and soon the flying after water was going on as if ijfothing had happened. Thus qiuickly a 
 part of the bees had become accustomed to flying in the new apiary," etc. Even thou,gh thiq letter con- 
 tains no new knowledge of the biology of the bee, yet it seems to me that its publication is appropriate, 
 in view of Bethe's opinions following. 
 
 101 As was mentioned before (see p. 19), the leader of the bee-keepers' school at Baden observed 
 that his bee's returned from a buckwheat-field six kilometers away, with full burdens, on the average in 
 thirty minutes. 
 
 102 It is a pity that we can not learn how these bees acted, whether they oriented themselves, etc. 
 
 108 Accordingly this appears to have been a*, cool autumn or spring day, else it is inexplicable why 
 the bees hesitated so long with their flying-out. Or was it windy or rainy weather ? Or were the bees 
 wearied with the transportation ? 
 
34 Natural History of the Honeybee. 
 
 They corkscrewed in widening circles into the air, as bees do when they have just been 
 set out. After three or four circles (in which note well they had never turned their eyes 
 toward the hive), 10 * they took their direction and flew quickly to the meadows. A gTeat 
 number of others followed, all corkscrewing into the air. 105 About three o'clock the first 
 ones returned laden with honey and pollen. 106 They came from the east (not from the 
 south whither most had gone), and flew from the place where I first caught sight of them 
 (5-6 meter distant) in a straight line to the entrance. If here a noticeable accustoming had 
 played a part, perhaps as a memory process, it would be impossible to think that the insects 
 would fly in such a straight line for the hive. A force then must be present, which draws 
 them like a magnet to this place in space, and the described ' accustoming ' is something 
 secondary, unnecessary for the occurrence of the reaction.' ' 
 
 Since we have not the slightest scientific proof whether these bees which returned at 
 three o 'clock, which ' ' flew in a straight line to the hive, ' ' did not belong to those which had 
 hummed around the hive in the early morning, and therefore had been able to notice the 
 position of the dwelling accurately, this experiment proves nothing, because we are not 
 informed concerning the method of the flying-out of the later bees. 
 
 Bethe speaks expressly of "accurate observations," so we must accept as a matter of 
 course that there were no errors in his observations. If, therefore, the foregoing really 
 happened as pictured, this experiment does not, I believe, present a conclusive proof for 
 but against the "unknown force." According to my long years of experience, a five or 
 six hours' roving and remaining away from the hive under the conditions described is 
 entirely impossible. We may here, from what is known of the natural history of the bee 
 in this direction, count upon the possibilities expressed before. Later on it turned out that 
 the observation was not an accurate, one. Cf. p. 90 of my Stammesgeschichtliche Entstehung 
 des Bienenstaates. Leipzig, 1903. 
 
 If, indeed, a magnetic force existed to draw bees homeward, then without doubt the 
 bees which flew out would have succeeded in reaching home inside of ten minutes or at 
 most an hour after leaving. 
 
 bethe's tree experiment. 
 
 This is the place, perhaps, in which to examine more closely Bethe 's tree experiment. 
 "In the fall of 1896 I put my hives in a small house which lay toward the east. In front 
 was a plane-tree, seven meters high, whose branches approached the house within about 
 one and a half meters. The top of the tree was eight meters wide and began three meters 
 above the ground, so that the little bee-house was completely shaded. Now, because the 
 bees flew very badly in the spring of 1897 on account of a lack of sun, I decided to have 
 the tree cut down, and this was done in the morning about 10:30 (June 11, 1897), just 
 when most of the bees were out. Thereby v were the surroundings of the little house com- 
 pletely changed: instead of the high tree there was a large open quadrangle fifteen meters 
 square. The bees had been coming perpendicularly between the tree and the house to their 
 hives, but those returning (which had flown out long before the change, since they all 
 had pollen on their legs) immediately after the fall of the tree flew directly in a slanting 
 line to the entrance. They flew, therefore, through the space in which the tree had stood ' 
 a second before, and showed not the slightest uneasiness; they were not 'surprised,' as, 
 without doubt, many authors would report with astonishment" (Bethe, p. 85). 
 
 Bethe decides from this that finding the way home can not depend upon memory 
 pictures. 
 
 From this statement of Bethe 's we get the idea that there were no obstacles present 
 but the plane-tree. Up to where the branches began, therefore, there was a free path 
 three meters high. No one who is familiar with the practices of bees will doubt that the 
 
 504 Bethe appears to consider this the normal flight out; above all, nowhere does he mention the 
 entirely different flight of orientation. 
 
 105 As none of these tees flew off in a straight line, it is natural to suppose that these were greatly 
 irritated by the transportation or suffered by the lack of air during the transportation. The thick* gathering 
 on the outside points to the latter supposition, ss dees ako the late flying-out. 
 
 loe Therefore after five or six hours! Doubtless almost all the tees which flew in the forenoon 
 were lost, or returned again to the known neighborhood, because the village was only seven kilometers 
 distant, for under the conditions cited an absence of five or six hours can not te accepted as true. 
 
Natural History of the Honeybee. 35 
 
 bees used this large space for flying, therefore flew under the branches. A bee always 
 flies low to the forage unless obstacles are present or the distances are extraordinary; 
 and there is not the slightest discernible reason why the bees should not have used this 
 comfortable path if they flew mainly in the direction of the tree. It would have been 
 abnormal if they had not done so. That Bethe noticed the bees flying in under the branches 
 or through the top of the tree only after the tree was down is easily explained, since their 
 flight was much more visible, and the tree had not been cut down for the sake of obser- 
 vations but merely to remove the shade. The opinion might, therefore, be allowed that 
 the special kind of flying in had not before been accurately noticed. That bees fly through 
 bushes standing not very close together, or through trees which put out their leaves late 
 (as does the plane-tree), I have often observed. They maintain the accustomed flight as 
 long as possible. But there is no contradiction in saying that bees, and perhaps very many 
 of them, came down * ' perpendicularly between the tree and the house to the hive. " But 
 they surely did not fly out over the tree, but to the other side over the roof of the "little 
 house, ' ' or sidewise over the roof to the ' ' meadows ' ' lying in the ' ' south. ' ' I am strength- 
 ened in this opinion, by the following assertions of Bethe: "I have said that the bees 
 returning after I had cut down the plane-tree in front of the apiary, flew immediately 
 through the place where it had stood instead of, as before, coming down perpendicularly 
 in corkscrew lines. The outgoing bees conducted themselves in an entirely different way. 
 After the fall of the tree they corkscrewed upward penpendicularly just as if the tree still 
 stood. When I saw the bees fly for the last time in this year (it was on September: 14, 
 therefore about three months after the tree had fallen, June 14), all the outgoing bees 
 still flew upward perpendicularly in front of the house, as if the tree still stood. I am 
 wondering what they will do next year. There can be here no question of any perceptions 
 of the senses, of any deliberations ' ' (Bethe p. 92). 
 
 Bethe here overlooks the fact that most likely all the bees which flew September 14th 
 had never seen the tree, since in summer bees live at most six or seven weeks. 107 The 
 cause of this singular flight was, therefore, certainly not the tree which fell fourteen 
 weeks before. There can, therefore, be no question in this case of "perceptions of the 
 senses, " as Bethe says with another significance. And then "the next year"! - 
 
 Even if the foregoing observations are accurate, it is clear that the peculiar flight of 
 the bees has not been necessarily because of the tree, and therefore the demonstrative power 
 of these observations is very doubtful. 
 
 Exact judgment is not possible in this case, since the other obstacles further away 
 perhaps, as there undoubtedly were, are not mentioned. 
 
 I made the following control experiment: Close to my hexagonal bee-pavilion garden- 
 paths strike out to the east and southwest. In order that passers-by would not be troubled 
 by the bees, I was forced to plant bushes and treea so thickly in front, one-half to one 
 and a half meters, that the bees were compelled to fly high immediately. The majority 
 did me this kindness; but those which developed early in the spring, before the leaves 
 came out, picked out larger or smaller openings between the twigs or between the bushes 
 as pathways, and held to these paths even when the space became almost entirely covered 
 with leaves. Because of a birth-tree standing on the southwest side, six meters high 
 and about two meters wide, the lowest branches beginning only about a meter from the 
 ground (therefore a rather bush-like growth), the flying bees from a colony standing a 
 meter and a half away, and over one meter high, were forced to divide. Some of them 
 flew to the left, others to the right, since the young trees standing next had not yet 
 effected a closing. The birch was cut down, and I noticed as follows: No hesitation of 
 
 107 "How great the death-rate is, especially at the time of forage, the following experiments prove: 
 I made several artificial swarms from pure German bees, with a pure Italian queen. In six weeks 
 there was not a single German bee left. July 17th I took a rather old German queen from a hive and 
 put in an Italian the third day after. Notwithstanding the fact that for three weeks the German brood 
 emerged, after six weeks, scarcely every twentieth bee was German." — Dzierzon in Bienenzeitung, IX., 
 No. 23. "The length of life of workers is usually very short. Those reared, in the spring and summer 
 live often hardly six to eight weeks." — Ludw. Huber, Die neue, nutzlichste Bitoenzucht, 13. Aufl., 
 Lahr, 1900, p. 16. In rich forage the bees live often only two or three weeks. Many personal obser- 
 vations sustain the foregoing. Queens usually reach the age of four or five years; but cases have been 
 known of queens which were six or seven years old (Bienenzeitung, 1882, p.* 78). 
 
36 Natural History of the Honeybee. 
 
 the bees in any way remarkable took place, as regularly happens if the surroundings of 
 the hive itself are changed. The explanation, I think, lies in the fact that the bees in 
 flying out undertook no special inspection of the surroundings — they know their own circle 
 of flight. 1 ' 8 The in-flying bees could adjust themselves by so many other landmarks, all of 
 which are still present in the immediate neighborhood of the hive, that they were not con- 
 fused by the absence of one landmark in the proximity of the hive, since by that time the 
 hive itself was recognized. 
 
 The flight itself remained forked for many days. Some few, however, a few minutes 
 after the fall of the tree, went out through the space where it had been. A(S the colony 
 was very weak, and usually only a few bees were seen under way, I can not say whether 
 these bees had before flown through the spaces between twigs, for an hour passed before 
 I saw another fly through the now open space. These few which flew through might easily 
 have been overlooked by me before. 
 
 But even if all the bees- had suddenly flown out in a straight line, I could see in that 
 no confirmation of an unknown force, with so many proofs against it, but only a special 
 capacity for orientation. 
 
 SPECIAL CAPACITY FOR ORIENTATION IN BEES. 
 
 That bees orient themselves indeed in a special way follows from many things. Lub- 
 bock lu9 also is of this opinion. Nevertheless in general one is inclined to apply human 
 standards to them. A simple experiment shows that bees have their special estimation of 
 position. 
 
 If the height of a hive is changed so that the entrance is suddenly made, let us say, 
 thirty centimeters higher or lower, then we see that the bees fly exactly. to the spot where 
 the entrance was before. Hours and often days pass before a smooth graded flight is to 
 be noticed to the new height. To my mind it follows from this that in fact, as Lubbock 
 also believes, the bees ' ' know how to find their way by the relative position of the well- 
 known entrance to the surrounding trees and other large obstacles, and to the hive itself. " 
 The principal thing is the relative position of the entrance to the ground, for trees and 
 large obstacles can be lacking and yet the entrance be reached, for the bee has an extremely 
 fine ability for the appreciation of height. The entrance itself is not regarded at greater 
 or less distances, as the experiment given above proves, for first an observation of it takes 
 place in the immediate proximity. This ability to judge distances goes so far that, under 
 normal conditions, bees are accustomed to fly, for example, always in and out of the same 
 corner of the hive-entrance. If a bee is powdered on entering at the outermost right 
 corner of the entrance, it will be seen that, in unmolested flight, it uses the same corner 
 constantly in a straight line, even if the entrance is only ten centimeters wide. If the 
 eu trance is then stopped up all but about the breadth of an inch, they will, in spite of that, 
 try to press in just at their points, and find the open part only after a shorter or longer 
 search. 
 
 This accurate adherence to a settled path of flight has something to do with the rela- 
 tive position of the hive or the entrance, but not with the hive itself nor the entrance itself 
 at all; for if the hive is taken away entirely the bees will continue to fly to the accustomed 
 place at the accustomed height. In such steadfast holding to the path, all disguises such 
 as Bethe 's have little proving value (Bethe, p. 85), in so far as (and Bethe leads on in 
 part to this end) it should prove that bees are guided on their way, not by memory 
 pictures. The simple consideration that bees fly back exactly to the place where the 
 entrance has been, even though the hive has been removed many days before, makes it 
 clear that their flight will not necessarily be led astray if the hive or background be com- 
 pletely disguised, as was done by Bethe; therefore the spreading of colored papers on 
 
 108 The careless flying-out of the bees, which is often fatal, is well known to the bee-keeper. On 
 this is based the old practice that colonies which are to be sent to buckwheat-fields are always brought 
 out to these fields before the buckwheat blooms, so that the bees can learn thoroughly the new locality. 
 If the bee-keeper does not stick to this rule the proximity of the strongly fragrant fields is so stimulating 
 that the bees pour out without any orientation, or only a hasty one, and then they get lost, etc. The 
 colonies fly themselves "bald" (cf. p. 32*). 
 
 100 Lubbock, The Senses, Instincts, and Intelligence of Animals; London, 1889; German translation, 
 Leipzig, 1889. . 
 
Natural History of the Honeybee. 
 
 37 
 
 the ground in front of the hive can not, of course, influence the valuation of the height 
 particularly. If it is wished to draw the conclusion from this that bees are not led on 
 their paths through memory pictures, that seems to me to be anthropomorphism of the 
 
 best kind. 110 
 
 It is no contradiction to this that bees notice very closely the appearance and form 
 of the hive, as I have said before. One needs only to fasten a piece of colored paper above 
 the entrance or to change the form of the hive to notice immediately a hesitation in the 
 flight toward it; a closer survey is taken while flying in. 
 
 A bee, therefore, flies to the point of the entrance without being able to see it or 
 the nearest surroundings of the entrance, on the basis of its orientation in their neighborhood; 
 on the other hand, when it comes closer it generally notices the surroundings of the entrance, 
 for it becomes aware of any changes in the outside of the hive. The quicker it hurries back 
 to the hive (for example in a heavy honey-flow), the less the appearance of the hive is 
 noticed; and if attention is gained by a very striking change, the hesitation or loitering to 
 be seen is much less. 
 
 THE INFLUENCE OF WEATHER CONDITIONS ON THE SENSE OF SIGHT^, THEREFORE ON 
 
 THE ABILITY TO ORIENT. 
 
 If bees are thrown up into the air at dusk, not far from their dwelling, they do not 
 find their way back. They make several small circles and then fall to the ground, i ' fright- 
 ened and lost. ' ' Bethe has made this experiment, but we are not told why the ' ' unknown 
 force" does not act under such conditions. That simply the sense of sight is here deter- 
 minative in regard to orientation may be clear from the following observations: 
 
 "A fruit-grower in England was astonished to find the trees much more heavily 
 laden with fruit in the part of his large garden where there were some colonies of bees 
 than in the other parts. He then remembered that it was extremely foggy during the 
 blooming of the fruit-trees, and the bees, in consequence, flew only short distances. ' ' U1 Any 
 one who has observed bees for a long time knows how strongly sudden dark clouds influence 
 flight. In such cases we often see an amusing sight during the honey-flow. At the 
 approach of a storm the hundreds of thousands of bees that have hastened to the fields, 
 rush in wild haste to their sheltering homes. With pollen-sacs hardly begun to be filled they 
 stream on in blind haste in a ceaseless train and l ' dash ' ' into the hives. If it happens that 
 the clouds are in the background, and the sun is shining on the thousands of whirring wings, 
 the appearance is most peculiar. 
 
 That we have to do with orientation through the eyes in this journey home at the 
 approach of a storm, and not with an ' ' unknown Yorce, ' ' follows from the fact that, in the 
 
 i ' mad haste ' ' mentioned, many blunders are made, and many bees enter strange hives. 
 1 l unknown force ' ' which ' ' draws like a magnet ' ' would not permit such errors. 
 
 An 
 
 110 American bee-hunters base their methods of capture on the above-mentioned fact, that, if bees 
 have once flown in, they regulate their paths of flight with extraordinary accuracy, and constantly fly in 
 almost a straight line. "In a clearing of the woods a little fire is made and an old comb held over it. 
 Attracted by the odor, bees come and taste the honey inside. After sufficient orientation they begin by 
 degrees to take up a straight line of flight. The hunter follows this line and marks it with any land- 
 marks at all. Now the. bait is placed at the other end of the clearing. After a short time the bees 
 find it again, and the line of flight is again marked. Then, where the! two lines cross, the colony is 
 found. Autumn is most favorable for this, for since then no other forage attracts the bees and leads the 
 hunter into error." — American Bee Journal, Chicago, 1893. 
 
 The bee-keeper also uses this characteristic of bees to discover a robbing colony. "The bees always 
 fly in a straight line from the place where they find food, back to the hive. The bee-keeper, therefore, 
 goes in this direction and soon stands before the hive where the robbers belong." — Ludw. Huber, 1. c.„ 
 p. 35, 1900. 
 
 i- 1 Bienenwirtschaftl. Centralblatt, 1892, Hannover, p. 205. 
 
38 Natural History of the Honeybee. 
 
 A very important bee-keeper, Herr Dathe, in Eystrup, submits the following: In his 
 extensive apiary of about five hundred colonies there were various stands of box hives. 
 The line of flight went on as indicated in the figure above in various directions. The 
 arrows gif e the point of the entrances — that is, the direction of the flight in and out. 
 
 Just before thunderstorms, etc., Dathe made numerous observations, that in their hasty 
 return many bees threw themselves on the wrong stand; and such as were accustomed to 
 enter at the side (c) made great exertions in vain to get into the hive at the stands 1 and 3 
 at the corresponding place (b), although there was no entrance there. In the same way 
 the li a ' ' bees sought their hive-entrance on the stands 2 and 4 at " d. ' ' 
 
 A similar thing is seen in a violent wind, which throws the bees out of their accustomed 
 paths of flight. 
 
 It follows from this that a memory picture of the hive is present, as well as a valuation 
 of the right place and height, even if on the wrong stand. An explanation of this error 
 is, I might say, natural aa soon as one ascribes the orientation to the sense of sight. It is 
 difficult to think of it on the ground of the unknown force. 
 
 I owe further interesting observations to the already mentioned leader of the Baden 
 Bee-keepers' School, Herr Both. This leads us at the same time to the special reaction 
 of bees toward colors. 
 
 INFLUENCE OF COLORS ON BEES. 
 
 I shall here mention only two observations, and for the rest, in regard to the influence 
 of color on bees, I can only allude to the well-known experiments of Lubbock 1 ^ and 
 Hermann Miiller, 133 as well as to those of Bethe (1. c), etc. 
 
 We have seen above that flight becomes very unsafe in the dusk; therefore it is 
 evident that gloomy weather influences considerably the ability to orient. 114 
 
 "One of my former neighbors, " says Roth's contribution, "painted the gable of his 
 house over the apiary with sky-blue (luftblau) color. The very bees which always flew over 
 the gable, on the next dark day bumped against it with their heads, trying to ♦fly through it. ' ' 
 
 A teacher, Stahelin, made the following observations: A weak after-swarm, mostly of 
 young bees from a hive painted blue, dispersed 115 among the masses of humming bees which 
 were just taking their flight of orientation out of the other hives (which, as is usually 
 the case in Germany, Switzerland, and Austria, were standing close together), and settled 
 here and there in little clumps. After a short time they flew back to the bee-house; but 
 only a. few found the right hive; the rest flew to other colonies, and to which? Only where 
 a blue door invited them did they attempt an entrance, but nowhere else. Unfortunately 
 they were so hostilely received that the ground in front of all hives marked blue was 
 covered with dead bees. 116 
 
 These cases also show nothing of an "unknown force, M but, instead, exhibit the great 
 significance to bees of the sense of sight, and the errors suffered through deceptions of 
 the eyes. 
 
 The disproof of Bethe'* theory by Forel appeared independently shortly after the 
 publication of my paper (Forel, Aug., Sensations des Insectes, IV., Paris, 1901). Forel 
 reaches the same conclusions that I do (cf., also, Forel, Aug., Nochmals Herr Dr. Bethe 
 und die Insekten-Psychologie, Biol. Centralbl., Bd. XXIII., 1903). The confirmation of 
 my work by this important investigator is of special value to me. The quintessence of 
 Forel 's views about all these matters will be found in his paper: Die psychischen Fahigkeiten 
 der Ameisen und einiger anderer Insekten, Miinchen, 1901, which, has been translated by Wm. , 
 M. Wheeler: Ants and some other insects. An inquiry into the psychic powers of these 
 Animals with an appendix on the peculiarities of their olfactory sense. Chicago, 1904. 
 
 112 Lubbock, Ants, Bees, and Wasps. International Scientific Series: New York, 1883; German trans- 
 lation, Leipzig, 1883. 
 
 1]3 Herm. Muller, 1. c. 
 
 114 The similar behavior of carrier pigeons may be compared with this (H. E. Ziegler, Ueber die 
 Geschwindigkeit der Brief tauben, 1. c). 
 
 115 Because the swarm tone which holds the swarming bees together (see p. 16) had been swallowed 
 up by the loud buzzing of the other colonies which were giving out many bees in their '"first play." This 
 shows the possibility of communication through hearing? 
 
 118 Schweiz. Bien. Ztg., Aarau. Jahrg. 1893. 
 
Natural History of the Honeybee. 39 
 
 In my ' ' Stammesgesehichtlichen Entstehung der Bienenstaates ' \ (Leipsig, 1903), I have, 
 in Appendix 4 pp. 90-95), a "concluding word concerning Bethe's investigations on bees. 
 The significance of ocelli in bees, ' * in which I confirm my views on their ability to see, and 
 verify it in part through experiment. Many other pertinent questions are there touched 
 upon. 
 
 Full of importance in consideration of the ability of insects to orient by ^heir eyes is the 
 following work of the Peckhams. The observations of the Peckhams^on the sense of orien- 
 tation in wasps supports my work on bees. Unfortunately this entire excellent investigation 
 (On the Instincts and Habits of the Solitary Wasps, by George W. Peckham and Elizabeth 
 O. Peckham, Madison, Wis., 1898), became known to me only after a long time after the 
 German edition of this paper had been published. I value these observations more highly 
 than the very admirable but frequently visionary work of T. H. Fabre (Souvenirs ento- 
 mologiques, Serie 1-9. Paris, 1879-1905). 
 
 The works of Bouvier on solitary wasps (Les Habitudes des Bembex, Monographic 
 Biologique, Paris, 1901); of K. Escherich (Die Ameise, Braunschweig, 1906), and the obser- 
 vations of Charles Ferton (Notes sur 1 'instincts des Hymenop teres, Melliferes, et Ravis- 
 seurs, 3 serie, Paris, 1905, fall in this province. All these works give many beautiful 
 explanations, and confirm my statements more or less. 
 
SOME FURTHEB CONTRIBUTIONS TO THE NATURAL HISTORY. 
 
 THE FLIGHT OF BEES INTO A ROOM. 
 
 If honey is placed in the part of a room opposite the windows, and the windows 
 opened, bees will soon appear at the openings. The flying-in is accomplished in a very 
 characteristic way. Very slowly, as if cautiously exploring, they risk going into the inner 
 part of the room where the light is subdued. Timidly the advance is made, back and 
 forth; here and there goes their flight until the honey is reached. 
 
 When I was occupied with investigations on bees' eggs one summer, some honey-comb 
 was standing in the laboratory of the Institute. One day, on opening the window, some 
 bees flew in with remarkably certain flight to the wall opposite the windows. That puzzled 
 me, and I called the matter to the attention of the assistant, Dr. L. Schultze, who was 
 present, with the remark that these bees must have been nibbling sweets in another room, 
 because their flight' was so sure. The only honey in the Institute was in the room of the 
 assistant, and that window proved to have been closed. I knew from their coloring that 
 these Were strange bees, not those from the apiary in the Institute garden. The enigma 
 was soon solved. In a villa, scarcely twenty steps away from the open window, a bee- 
 keeper had a couple of colonies on the loft under the roof. The bees flew through the 
 roof window, and were, therefore, accustomed to flying into a dimly lighted room. 
 
 This modified conduct proved to me that bees undoubtedly learn. The bees from an 
 outdoor apiary would certainly have acted differently. 
 
 THE BEHAVIOR OF ROBBING BEES. 
 
 Robbing bees look like robbers very soon, for they soon lose the hairy covering, because, 
 with their foreign hive-odors, they are pulled and hauled about when they force themselves 
 into strange colonies, and also because energetic licking must be done, as they get soiled 
 with honey. The colored hairs disappear, and the dark hard chitin shows, as can be seen 
 in the spring on old bees too, which have lived over the winter. 117 In summer, when most 
 of the bees are not old (see p. 35), robbers can be recognized in most cases by this 
 appearance. 
 
 Since the loss of hairs on bees gives them the appearance of being smaller, these "black 
 bees" were formerly taken for a special kind of bee. 118 
 
 It is of special interest to watch the conduct of these robbing bees, for they depart 
 completely from the usual behavior of workers. 
 
 Their flight to a strange hive, when they are just starting in their profession, is very 
 uncertain, * ' timid, ' ' and ' i anxious. ' ' n9 Even the boldest act as if they did not dare to 
 follow their desire to rob, as if they knew the danger in which they were placing them- 
 selves. But if they once succeed in entering a hive they soon become bolder, and "those 
 which have carried on robbing for a long time may be recognized by their hasty practiced 
 manner of entering. ' n20 Here, again, we have a beautiful example of capacity for learning. 121 
 
 TH E ORIGIN OF HOSTILE C ONDUCT. 
 
 While solitary bees, such as Chalicodoma muraria or Andrena ovina or Anthrophora 
 personata have apparently no knowledge of their enemies, we see in Apis melliftca that this 
 
 *« Dathe, 1. c, p. 180.\ - ^ 
 
 118 Bienenzeitung, No. 5 and No. 11, I.: Nos. 13, 14, 15, V.: further, Dr. Magerstedt, Der prak- 
 tische Bienenvater, 2. Aufl., p. 154: Sondertehausen, 1845, etc. 
 
 119 Lud. Huber, Neue nutzl. Bienenzucht, 13. Aufl., Lahr, 1900. 
 
 120 Dathe, Lehrbuch der Bienenzucht, 1. c, p. 180. 
 
 m "The nibblers are often so told that they stop the tees roming from the field at the entrance, 
 and the latter, therefore, getting the idea that they have flown into the wrong hive, reach them their 
 probosces and thus let them steal the honey from the mouths. Such robbers are like swindlers in a 
 great city, who cloak themselves as police officers, and under this mask carry out their fraud." Dathe, 
 1. c, p. 183. 
 
Natural History of the Honeybee. 41 
 
 species understands how to make its enemies for the most part harmless in the hive. 
 Carriere m thinks that this knowledge and ability to exterminate them is formed in Apis 
 mellifica beeause it represents "a permanent community, and therefore a ' tradition ' could 
 have been developed.' ' I can not share his opinion, which supposes entirely too much 
 human intelligence. I believe_ that_here the principle of selection has acted also. For^ 
 colonies^ which collect a supply of honey it is naturally of great significance that there 
 should be means of defense against the entrance of robbers, etc. Therefore the instinct 
 developed, by and by, to react hostilely toward those ' entering with a foreign hive odor 
 or with unusual behavior. 
 
 THE VANISHING OF INSTINCTS WITH THE DECREASE IN THE STRENGTH OF A COLONY. 
 
 Only a strong colony resists its enemies, such as robbers, wax-moths, etc. — not that 
 the weaker colony is not able to do so very well, for there are still plenty of fighters, but 
 it is a fact that its " spirit,' ' its il energy,' ' disappears as a colony weakens. The narrow 
 entrance could be defended with ease; but such colonies often do not place bees at the 
 entrance at all. They let single robbers and single wax-moths, etc., quietly alone, although 
 they could kill them with no trouble. 
 
 The more reduced a colony becomes, the more do all instincts fail. The hive is no 
 longer cleaned, flights are more and more discontinued. Languidly and ' ' apathetically ' ' the 
 bees sit around on the comb. If, finally, almost all are taken away, so that only the queen 
 remains with a handful of bees, even honey which has been placed in the hive remains 
 untouched. 128 
 
 In experimental observations, particular attention should be given to this characteristic 
 failure or weakening of instincts with the decreasing strength of a colony, for otherwise 
 the results obtained are never exact. A_weak colony acts entirely differently from a strong 
 one, and the reactions are often in the opposite direction; also the varieties of Apis mel- 
 lifica must be taken Jnto consideration, as a small colony of heather bees of North Germany, 
 for_ example, acts differently from a weak colony of bees from middle or southern Ger- 
 many, and these, again, differently from Carniolan or Italian' bees under the same circum- 
 stances. In short, one must be entirely familiar with all the factors pertaining thereto in 
 order to obtain results free from error. 
 
 REACTIONS TOWARD FLIGHT. 
 
 We must certainly notice further a reaction of bees which thus far has been thought 
 to be slight, but the disregard of which can lead to false conclusions; that is, the sharp 
 reaction toward the manner and way of flying to the entrance of the hive. In many cases, 
 therefore, it may not be the special odor which betrays the robbers, but as said, their 
 il timid" loitering flight. The bees which defend the entrance and flight-board may often 
 be seen to baffle the attempts of robbing companions by flying at them and seizing them 
 in the air before there can be, in my opinion, the least perception of the odor of these 
 single bees, because of the strong evaporation from the hive entrance. 124 It is a reaction 
 toward the special kind of flight. In connection with this is this fact: that if the robbers 
 are more cunning, and go flying smoothly into the entrance, a defense is hardly noticeable, 
 and often does not take place at all. 
 
 If the experiment is made o£ bathing bees in water or weak alcohol to remove the 
 adherent hive-odor, and, after drying, the bees are allowed to run into their hive, they 
 can enter the accustomed opening safely on account of their quiet and sure manner of 
 entering, and no definite reaction of the inmates' is noticeable. If the bath is thorough, 
 there will be no definite reaction set free when entering a strange hive either, not only 
 because the foreign odor is taken away, but also because the bath destroys former mem- 
 
 122 Carriere and Burger, Entwicklungsgeschichte der Mauerblene. Abhd. der Kaiserl. Leop. Carol, 
 deutsch. Akad. der Naturf. Bd. LXIX., No. 2. 
 
 123 And yet, as is well known, the attraction of honey for bees surpasses everything else. If the 
 abdomen is quickly severed from the thorax while a bee is sucking on a drop of honey, it sucks on quietly 
 for a long time, while the sucked-up portion flows out behind. 
 
 124 I spoke before (see p. 14) of how little bees are able in open air to perceive the strong odor of 
 the queen at approximately short distances. 
 
42 Natural History of the Honeybee. 
 
 ories (see p. 27 ), 125 and the respective bees no longer know their own hives; so without 
 hesitation they run in, and their quiet entrance is often little or not at all noticed. 
 
 If a queenless colony is allowed to run into a queen-right one, or if it wanders over 
 of itself, a broad stream of bees joyfully "humming" enters the strange hive. These 
 usurpers of the strange dwelling, with such a sure (and I might say matter-of-course) 
 manner must in most cases have a somewhat " disconcerting ' ' and "embarrassing" effect 
 on the inmates, for seldom is a defense attempted. It appears, therefore, that the hive- 
 odor is not the only absolutely certain sign of recognition, but that the kind of flight, the 
 particular conduct on flying in, plays an important part in the recognition of friend or 
 foe, or at all events in the setting free of hostile reactions. 
 
 If daubed bees, which have, in addition to the foreign hive-odor, a still stronger odor 
 of the substance used in daubing, are shaken on to a colony, hostile reactions are never 
 set free, for the bees shaken on lie entirely quiet or move only very slowly. 
 
 THE FORMATION OF THE CELLS IN THE COMB. 
 
 The honey-comb, apparently so "ingenious," can hardly be brought in as proof of the 
 higher faculties of bees, since it is only the result of mechanical force, aa Mullenhoff m 
 demonstrated, and as is well known. I can not reconcile myself to many statements in 
 Mullenhoff's explanation, for they do not entirely coincide with my observations. But a 
 doubt can scarcely be raised against the foundation of^Mullenhoff 's theory of the mechan- 
 ical acting principle. According to his theory, a so-called one-sided comb can not produce 
 the pyramidal cell-floors formed of three rhombs, yet we see them frequently in such 
 comb. Also the case can not be so represented as tho bees worked with semi-liquid material 
 in order to explain thereby the tension of Plateau's "liquid membrane," or the tension of 
 the floors and sides of the cells. In a cluster of bees the temperature is, usually from 
 28 to 34° C, while the melting-point of wax is 62° C. Of course, the wax at 30° is weak 
 and easily molded, yet there remains for the instinctive activities of bees the act of thin- 
 ning and smoothing of the floor and walls, as independent productions, as well as the 
 special sizes of cells. Mechanics does not explain why drone-cells are so much larger 
 than worker cells, but the form and the width of the angles have certainly nothing to do 
 with the "dexterity" of the bees. I can here only refer to the interesting work carried on. 
 
 Miillenhoff's statement that, in the cells thus formed, a "drop of the secretion from 
 the poison-gland" is added for the preservation of honey "gathered not for immediate 
 use, ' f is anthropomorphically quite clear, but does not correspond with the facts in the case. 
 
 This statement has been proven untenable by Schonf eld 127 and by Planta 128 through 
 chemical analysis. I must also call into question Mullenhoff's view that "the form of the 
 body of the animal depends chiefly upon the form of the cell in which/ it developed ' ' (1. c, 
 p. 616). For the purpose of other observations I placed a colony only upon drone comb, 
 so that only the large drone-cells were available for the queen. After long hesitation she 
 began to lay eggs. From the drone-cells, besides a number of drones, mostly workers of 
 normal ■ size crept out. The nourishment does not here come into consideration, for in 
 all cases that was as usual. Drones also which have hatched in queen-cells are very fre- 
 quently of normal size. The view is not correct, that the cell 1 1 surrounds the entire body 
 of the insect like an enclosing caoutchouc sheath, hermetically sealed, and, accordingly, it 
 must be the cell which determines the form which the body of the young developing insect 
 shall assume." The larva has room for motion, else it would not be in condition to spin 
 its cocoon in there. 
 
 That drones are smaller if reared in worker-cells is not positive proof of Mullenhoff's 
 theory. These are, I might say, necessary relations, and do not permit the conclusion that 
 we "get a suggestion concerning the causa eflicienSj which may change the body-form of the 
 insect from the action of the molecules of the cell constituents." If bees go over to the 
 building of drone comb, which condition is noticed regretfully by bee-keepers, because then 
 
 125 1 must mention the fact that memory pictures are not always effaced by bathing. It seems that 
 the length of time and the temperature of the wafer are particularly determinative. My experiments in 
 this direction are not yet concluded. 
 
 126 K. Mullenhoff, Ueber die Entstehung der Bienenzellen, Pfluger's Archiv f. d. ges. Physiologic, Bd. 
 XXXII., Bonn, 1883. - ' . 
 
 127 Gravenhorst's illust. Bienenzeiting, Nr. 5 u. 6. Aaran, 1893 
 
 128 Schweizerische Bienenzeitaing, Nr. 5 u. 6, Aarau, 1892. 
 
Natural History of the Honeybee. 43 
 
 only consumers and not producers are reared, artificial foundation with the floors of worker- 
 cells pressed in may be hung in the hive. The instinct is thereby diverted, and the bees 
 erect worker-cella according to the wishes of the bee-keeper. In the hope of increasing 
 the size, and at the same time the capacity for production in workers, colonies have been 
 placed on foundation on which had been impressed cell-bottoms ii> size between those of 
 workers and drones. The bees built cells corresponding in size on this; but in spite of the 
 extra room for the developing brood the size of the workers remained exactly the same. 
 
 THE PLAY INSTINCT IN BEES. 
 
 Although I am decidedly of the opinion that there is to be noticed in the gay circling 
 of swarming bees at the same time the activity of certain play instincts, still proof is 
 certainly difficult, inasmuch as all the proceedings which come into consideration may be 
 led back exclusively to the swarming instinct, and can be explained only as an expression 
 of that instinct. After all, this strong pressing-forth from the entrance, this sounding of 
 the "joyful"- dear swarm-tone, which (as already mentioned) drags into the confusion 
 colonies nearly ready to swarm, these waves up and down in the sunshine in ' l drunken 
 lust" (swarm-dizziness\, all exhibit the playing activity of a certain surplus energy, and 
 therewith one of the fundamental elements of play. Schiller, as well as Herbert Spencer, 
 sees in play the expression solely of overflowing energy. I refer, therefore, to the ingenious 
 and interesting work of Karl Groos, 328 who, correctly, does not find complete satisfaction in 
 this interpretation, and who alludes to the great biological significance of the instincts 
 which are early manifested in pHy, and which perfect themselves in play. Groos does not 
 mention bees in his very comprehensive work, and yet we may mention, besides the swarm- 
 dance, where the play character may be doubted, a singular activity of bees, which, in my 
 opinion, shows undoubtedly true playing. Especially on warm summer evenings, but at 
 other times also, now and then on the outer walls of hives which are in good condition, near 
 the entrance or on the flight-board rows of bees may be seen moving to and fro in a 
 peculiar rhythmic way. As the head is held down it looks as if the ground were being 
 gnawed or licked in rhythm; but nothing of this kind takes place. A very characteristic 
 "contented" humming is audible. We doubtless have here in play the expression of a 
 feeling of comfort. 
 
 Single bees are often seen not fitting into the common rhythm of the line, but I have 
 frequently noticed a gradual adaptation to the prevailing manner. It seems that there 
 must be a special pleasurable sensation in this common rhythmic movement or else it 
 itself and the apparent mutual adaptation would not take place. 
 
 The whole interesting phenomenon seems to me clear, if we accept the idea of the 
 expression of a play instinct. It is known to every experienced bee-keeper, generally under 
 the name of swinging or planing (Hobeln). 180 
 
 129 Karl Groos, Die Spiele der Tiere, Jena, 1896. (2. Aufl. 1907.) 
 
 130 Schmid und Kleine, Leitfaden einer Ration. Bienenzucht, Nordlinger, 1865, etc. 
 
CONCLUSION. 
 
 From the very rich material which the natural*history of the honey-bee affords I have 
 been able to take into consideration in the foregoing pages only a few essentials. But 
 these may have shown us, I hope, that it is superfluous to suppose in bees the presence 
 of an < ' unknown force, ' ' since the biological proceedings are made clear easily and naturally 
 through the known forces. 
 
 It is unquestionable to me that the senses of bees are similar to those of men, and 
 that especially the senses of sight, hearing, and smell play an important part. As Wundt 
 has already shown, we certainly are "referred essentially to outer observations in animals; 
 what they teach us is not a total dissimilarity in the capacity of mind, but the most essen- 
 tial conformity with the psychic processes which we observe in man, and which we know 
 chiefly from observation of ourselves/* ' 31 There is very great danger, therefore, of ascrib- 
 ing anthropomorphic characters to lower animals, and or* applying human motives to them, 
 or of comparing their motives with the higher human standards. Thus I can not agree 
 with Lubbock's experiments on bees (1. c.) in so far as the proof of psychic faculties comes 
 into consideration. The anthropomorphic method of treatment must lead the results astray. 
 It is very characteristic of this method of treatment that Lubbock entertains great doubt 
 as to the existence of a sense of hearing in bees, because he has played a violin, whistled, 
 played a flute, and shouted before- them, and never observed that they took the slightest 
 notice. If, however, bees hear, they react only to such acoustic stimuli as have biological 
 significance for them; for example, to the various sounds which they produce themselves. 132 
 
 *». * lu W i! indt ' Vorles .- uber d . ie Menschen und Tierseele, Leipzig, 1863, p. 318. In passing I wish to say 
 that the bee community as pictured by Wundt (second/ edition, p. 453) contains many errors. I shall 
 mention briefly here that the queen does not lay either fertilized or unfertilized eggs "from the berinnine " 
 and that the workers do not "carry a larva from the usual cell into an unocfcupied queen-celL where 
 she is made a queen ty good food." Bees are incapable of tbjat. Just think of carrying a soft larva 
 rl , i lto » c - han £ in £ down perpendicularly, and open beneath! The bea does it more comfortably 
 g r s TJ widening a worker-cell into queen-cell. These so-called "made-over" cells are always recognizable, 
 lnf*n2? ?? r? sh °w, the prismatic joining of the worker cells, while the true qjueer^cell shows I round 
 iW with ™ w < ho i mcorr 1 ect ". ,f hat T the fi rst queen to emerge remains in the hive while "the others 
 n™ 1 w of t^, workers - Just the opposite is the case. The facts about drones are also 
 incorrect in part. Further, one can not disaccustom "the tame bees from swarming" "by enlarging the 
 thl? w° rd " lg ir? / h81r n «* essi ty-" Swarming is sometimes prevented by this enlacing- but T colony 
 "cu & tom" or ™iL™ SWarm ' ™ armS ° Ut + .° f the largest hive - Jt is ' therefore, not ablate to speak of 
 d^noMnllr JtF^T' Moreover there are no "tame" bees. The views concerning robbing, also, 
 do not tally with my observations. Wundt says that, as> there is an instinct in bees to imitate as in 
 
 acouirpd er fn 1S ; a rW Can C °r ClUde th T fr ° m that - " nat ever * colon y be * in « * n ™ ^ butThat 'custom^ 
 e l »k? e J, gen w? n8 ( ,? an * t ? communicated, not only through hereditary dispositions" or organi- 
 zation but also through the direct influence of the older bees upon the younger" I can not acre© 
 
 doubt diJ'Z- o? a^r'r^ T ft? Winter ] n exceptional caLs (see p 27) they nevertSeless^ To 
 doubt die out, on account of the short-hveness of the worker-bees (see p. 35), and the possibility of 
 heredi tary transmission is excluded too, because workers are sterile. The second powSSliS that indi 
 n«t hi y .^?5S e i h ? bltS are tra \ smitted trough the influence of older bees on younger ones can, as well; 
 veiv v.ln«l£ tZ *™ e> aS W 1 i e 1 Seen < from w >* f0lk)WS ' Tt is an » ld hee-keeper'l trick to introduce a 
 llli nihihrt « an abs ? lute y saf . e wa y b y establishing a colony of brood-combs with just-emerging 
 were ne7er with oldpfL^r/vf glV - Cn lmn " di £ el y with ™t further precautions. The young^ tecs w*ch 
 1 Imltati™ nr ItZh^t \ he fJ G ? CXaCtly . the same way as others at cnce ' the y feed * gather, build, 
 can /o? L wili f ng i by £ lder t ?, es w £ ot present at a11 ' * et the least difference from other colonies 
 has existed ?h Philf Kogevnikow Biol. Centralbl., 1896). As long as information about bees 
 our verv pvps ^ ^^™ //♦ £ ?en c ? m ^ te ""mutability It is, therefore, not true that "we see under 
 tno view that til ,w Q ,f S i t + akmg * plaC l f- m l the hab . lt ? of bees that there is no reason for nbt accepting 
 ablliti^pX t t^?£&£ 1 ^ l Zm? ° rigmated gradUally ' ParUy thT ° USh heredHary phy6iCa1 
 ^ All further conclusions of Wundt's about the genesis of the bee colonv (1 c n 455^ sinro thpv 
 wiv Wund?s rt th^ "continuity with the past," can^ardly be ^M ^J^nilii^^n^e £Z 
 Z^lJn^A ■ theor y u of origin of instincts with reference to bees can not be supported- yet as Wundt 
 
 th^^tt&^J^^f^S^^f /JSI??* *£ an be refuted sati « fa ^orily upoTthe abole b2.iT In 
 ™w^ w edition of this work (1906) the greater part of the incorrect biological statements are 
 
 omitted, but new errors are found upon which I can not here enter. uiuiogicai siaiemenis are 
 
Natural History of the Honeybee. 45 
 
 Compare herewith the interesting experiments Wm. M. Wheeler has made about the hearing 
 capacity of ants already spoken of on p. 17. 133 
 
 We obtain sure results in all these questions only by observing the conduct of the 
 insects among themselves, or at least in experiments with artificially produced tones which 
 approach those produced by the insects in question. ... 
 
 The anthropomorphic apprehension relative to the question of consciousness has brought 
 it about that there has been ascribed to bees a consciousness similar to the human con- 
 sciousness, and accordingly the most varied human feelings. In the settling of this kind 
 of question the utmost caution must be used. On the basis of my observations I am of 
 the opinion that bees possess either no consciousness at all, or one of only the lowest 
 degree of development. The question of consciousness is left to subjective estimates; but 
 the question whether an animal learns, and can acquire experience or not, may be deter- 
 mined objectively. The question is whether, in addition to the inherited (kleronome) 
 n«rve-paths, acquired (enbiontische) associations are formed (H. E. Ziegler). We see that 
 bees show signs of an admirable memory in their orientation, and also in other activities; 134 
 further, I believe I have shown that the bee possesses a perception for color and form, 
 and develops a rich capacity for communication by means of its well-developed ' ' language ; ' ' 
 that, further, it is able to gather experiences, to learn, and to form associations of impres- 
 sions, etc. I can not agree with Bethe, therefore, in his denial that the bee has capacity to 
 gather experience, and thereby to modify its actions (see p. 3). The bee is evidently much 
 more than a reflex machine. 
 
 Zoological Institute, Oldenburg, i. Gr., 1906. 
 
 133 A clergyman, J. A. O'Neill, has discovered a% interesting method for catching bee-ants (Mutilla) 
 without difficulty. " This hymenopterous insect belongs to the musical ones. The male and female 
 produce a clear sound which is caused by the rubbing of a sharp ledge on the third abdominal segment 
 against a delicately furrowed triangular spot on the surface of the fourth segment. If one now finds 
 a female mutilla and places it in his hand so that it can make this schind, the males in the vicinity 
 come immediately, and are so fascinated that they sit on the hands of the capturer and can easily be 
 seized (see L. Permguey in the Jahrbuchern des Sudafrikanischen Museums; Reference in Prometheus 
 No. 540, 1900). ' 
 
 134 ^m m Wheeler, Ethological Observations on an American Ant. Journal fur Psycholoeie and 
 Neurologic Bd. II. Heft 1 und 2. Berlin, 1903. 
 
 135 Concerning the wonderful capacity for memory in bumble-tees. Ed. Hoffer, of Graz submits 
 an interesting communication! in " Kosmos," Zeitschr. f. d. ges. Entwicklungslehre, XVIII. Bd., p. Ill 
 Stuttgart, 1886. From that we learn that' in the bumble-bees also, memories outlast the winter. ' ' 
 
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