W. M. Wheeler.
The Phoresy of Antherophagus.
Psyche 26:145-152, 1919.

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PSYCHE
VOL. XXVI DECEMBER, 191 9 No. 6
THE PHORESY OF ANTHEROPHAGUS.1
August 16, 1919, while collecting Hymenoptera near Colebrook, . in northwestern Connecticut, I observed a worker humble-bee (Bmnbus vagans) behaving in an erratic manner on the flowers of a golden-rod. The insect was standing with straightened legs on the tips of its tarsi and repeatedly attempting to insert its pro- boscis into the flowers, but did not succeed because a small red beetle was firmly attached by its mandibles to the tip of the right maxilla and the tongue. The beetle, which proved to be a female of the Cryptophagid Antherophagus ochraceus Mels., did not release its hold in the cyanide jar, so that I am able to show it in its original position in the accompanying figure (Fig. 1). I failed to find any record of such behavior in our American Antherophagi (ochra- ceus, convexulus and suturalis), but a perusal of the accounts of the closely allied European species (nigricornis, silaceus and pollens) yields a satisfactory explanation of the peculiar activities described above.
In 1896 Lesne called attention to a number of small insects that habitually ride on larger insects. To this phenomenon he applied the term "phoresy" and showed that it is distinguished from ectoparasitism by the fact that the portee does not feed on the porter and eventually dismounts and has no further relations with the latter. The following year (1897) Charles Janet studied the known cases of phoresy somewhat more comprehensively, ex- panded the concept and distinguished no less than six different categories :
(1) Cases like that of the small flies of the genus Limosina which ride on the dung-beetle, Atenchus, and represent phoresy in its typical form as conceived by Lesne.
1 Contributions from the Entomological Laboratory of the Bueeey Institution, Harvard University, No. 162.




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(2) Cases in which the pork is conveyed to the neat of the porter, like the triungulin lam of certain beetles (Sita.ris, Melw, etc.) and the triungulinids of the Strepxiptera. (3) Cases like a few myrmecopbilous beetles (Thorictw) which attach themselves to the anteimie of ants for the purpose of accom- panying them on their peregrinations.
(4) Cases like the mitea of the genus Antennopharua which are not only carried but fed by the ants. These and the cases under (8) might be referred to ectoparasitisro. (6) Indirect phoresy, as exhibited by certain mites that ding to the surfaces of ant larvae and pup= which are in turn trans- ported by the ants.
(6) The cases of ants that carry in their mandibles their own young, other members of the colony or guests. In 1911 Banks published a valuable list of mme 17oases of pfaomty collected from the literature, and several others have been recorded by Warner (1903), Brues (1917a, 191%) and Kabattd (1917). Among the cases cited by these authors are those of certain small parasitic Hyraenoptera which attach themselves to the abdomens of Orthoptera or to the wings of Mantoidea in order to be on hand to oviposit in the eggs of their porters. Such cases really represent a seventh category of phoresy. Among the cases cited by Lesne and Janet and apparently over- looked by Banks, is Antheroph(wus, which attaches itself to the '
legs, mouthparts or antem of humble-bees for the purpose of being transported to their nests. The earliest observation of this habit seems to have been made by the British Coleopterist T. J.'~old. This author's two references to Antherophgus (1856 and 1871) were kindly sent me from London by ray friend, Mr. Hor- ace Doniatborpe. after I had vainly endeavored to find them in the Boston libraries. The first reference runs as follows: "Mr. Smith, in his admirable work on British bees, records the finding of Antkerophagus g1abe-r in the nest of Bmbus deshmellus. This season I met with an instance of the manner in which such iosecta may be transported thither. When hunting Bvmbi in September last, the peculiar motions of a neuter of B, sylvarwm attracted my attention: it was clinging to a. thistlehead, and wriggling and twisting its legs about in all directions. On getting hold of it I found that a large specimen of Adheropkagus nigncornb had



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seized the tarsus of a hind leg between its jaws, and was holding on like grim Death. I put both into my bottle, and the Artihero- phugus retained its hold until both were killed by the fumes of the laurel." The reference of 1871 is to this same find and occurs Fig. 1. Antherophugus ochracws Meis. attached to proboacis of Bmbus vagana sm.
on page 60 of Bold's " Catalogue of the Insects of Northumberland and Durham."
Redtenbacher (1858) records having taken three adult A. nigMc<ymIS in a humble-bee's nest, together with a number of larvae, which very probably belonged to the beetle. In 1863 Carus and Gerstaecker published the following note on the genus Antheraphagus: "The species live on flowers, attach themselves to humble-bees and permit the latter to transport them to their nests, probably for the purpose of oviposition; at any rate, small



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148 Psyche [December
larvae resembling those of Cryptophagus are sometimes found among the beetles in the nests of humble-bees." Eichhoff (1866) after examining several Bombus nests states that A. nigricornis was nearly always present and that single specimens of silaceus and pallens occurred in the same situations. Gorham (1869) captured A. pdllens together with certain species of Cryptophagus in a nest of Bombzis pratorum. Perris (1869-'YO), while collecting in the Pyrenees, took an A. nigricornis attached to the antenna of a B. montanus, and though he did not know of the observations of Bold and Carus and Gerstaecker he nevertheless drew the same infer- ence from his observations as the two German authors. Bugnion (1869-'70) in a letter to Perris recorded the following observation: "While collecting at Angeiades (alt. 1900 m.) in the Alps of Vaud, in the month of August 1866, I took a Bombus which had an Antherophagus pallens Oliv. attached to its proboscis by the mandi- bles." Seidlitz (1869-'70), commenting on Perris' observations, records the occurrence in a museum collection of three Bombi each with an Antherophagus attached to an appendage. In 1875 Perris published a detailed description of the larva of A. silaceus taken from the nest of B. sylvarum. Hoffer (1883), Fowler (1889), Sharp (l899), Wagner (l907), Reitter (1911) , Sladen (1912) and Reuter (1913) all give brief notices and Wagner publishes a figure of A. nigricornis attached to the bee's proboscis. The accounts of the North American A. ochraceus though meager go back to 1864 when Packard recorded its capture by F. W. Put- nam in several Bombus nests in Massachusetts and Vermont. Packard figured the beetle in this paper and the beetle and larva in two of his well-known books (1872, 1873). J. B. Smith (1909) and Blatchley (1910) mention the occurrence of A. ochraceus on various flowers and the former notes its occurrence in Bombus nests. Casey (1900) in his taxonomic revision of the Crypto- phagidae says nothing about the habits of the beetle, though he makes the following significant remarks on the genus (p. 87): "This is one of most isolated genera of the family and contains by far the largest species, Haplolophus being the only other which approaches it in this respect. The emargination of the clypeus, very deep in the male but feeble in the female is apparently a unique character in the family," etc.
The observations recorded at the beginning of this paper to-



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19191 Wheeler-The Phoresy of Antherophagus 149 gether with Packard's show that our species are very similar in habits to their European cousins. Though possessed of well- developed wings and able to fly about and take up their position on flowers, Antherophagus does not seek out the Bombzis nests but compels the bee to carry it to the place in which its eggs and larvae are to develop. As Sharp says, "we must presume that its senses and instincts permit it to recognize the bee, but do not suffice to enable it to find the bee's nest." The structure of the mandibles and the peculiar notch in the clypeus are clearly adaptations to firmly grasping the more or less cylindrical joints of the bee's appendages, and the red color of the integument and investment of golden yellow hairs, so very suggestive of conditions in many myrmecophilous beetles, may account for the fact that the Anthe- rophagi live unmolested in the Bombus nests. The feeding habits of the adult and larval Antherophagus seem not to have been actually observed by any of the authors mentioned in the preceding paragraphs. Packard' (1864) believed it "prob- able from the fondness, which these insects manifest for the sweets of flowers, that they visit the nests of the bees for the purpose of consuming the honey stored up within them." In 1873, however, he inferred that the beetle "probably feeds upon the wax and pollen," a statement which seems to have been suggested by the generic name given by Latreille. The views of the various authors concerning the feeding habits of the larva are, with one excep- tion, practically unanimous. Perris (1875) says: "The larvae of Antherophagus probably play the same role in the humble-bee nests as do Cryptophagus pubescens and scanicus in the nests of wasps. I do not believe that they devour the honey stored up by the bees or that they attack the bee larvae, not one of which showed the slightest lesion; I am convinced that they live on the feces of the inhabitants and that they are, properly speaking, merely scavengers." In the same paper he calls attention to the larva of Cryptophagus dentatus Herbst which lives under chestnut bark in company with the larva of Dryocoetus villosus and feed on its excre- ment. Lesne (1896) states that the Antherophagus larvae "live as mutualists rather than as commensals in the Bombus nests." In contrast with this rather vague and colorless statement, Wagner (1907) paints a lurid picture of the activities of the beetle and its larvae. After describing the transportation of the beetle on the



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150 Psyche [December
bee's proboscis, he continues:
"The humble-bee evidently feels
decidedly uncomfortable if she does not actually suffer pain. She
crawls over the combs of the nest, extrudes her proboscis and makes a series of movements for the purpose of getting rid of her burden, but in vain. Other bees come up, 'affectionately' pal- pate her with their antennae and pass on without the slightest attempt to help their 'comrade,' without the feeblest movement towards assisting her to ward off the dangerous enemy. And dangerous the beetle certainly is for from the eggs it lays hatch larvae which, by destroying both the wax and the cocoons, will cause enormous devastation in the nest." It is difficult to esti- mate how much of this is observation and how much is imagina- tion. Subsequent writers return to the opinion of Perris and regard the Antherophagus larvae as harmless scavengers. Thus Reitter (1911) asserts that they "probably live on the excrement of the inhabitants of the nest" and Renter (1913) that they "live on all sorts of refuse." Sladen (1912), who has a very intimate 'knowledge of the humble-bees and their nest-mates, classifies the Antherophagi "among the smaller and less important inhabitants ,of humble-bees' nests." We may conclude, therefore, that the larvae of these beetles are in all probability merely scavengers in the Bombus nests and hence closely resemble the larvae of Crypto- phagus in habits as well as structure.
Banks, N. Cases of Phoresie. Ent. News 22, 1911. pp. 194-197.
Blatchley, W. S. An Illustrated Descriptive Catalogue of the Coleoptera or Beetles (Exclusive of the Rhyncho- phora) known to occur in Indiana.
Nature Publ. Co.,
Indianapolis, 1910.
Bold, T. J.
Note on Antherophagus nigricornis and Born- bus sgharum.
Zoologist 14, 1856, p. 5003.
Bold, T. J.
A Catalogue of the Insects of Northumberland and Durham [Revision of Coleoptera]. Nat. Hist. Trans. Northumberland and Durham 4, 1871, p. 60. 1917a. Brues, C. T.
Adult Hymenopterous Parasites Attached
to the Body of Their Host.
Proc. Nat. Acad. Sc. 3,
1917, pp. 136-140, 1 fig.




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19191 Wheeler-The Phoresy of Antherophagus 151 1917b. Brues, C. T. Note on the Adult Habits of Some Hymen- opterous Egg-parasites of Orthoptera and Mantoidea. Psyche 24, 1917, pp. 195-196, 1 fig.
1869-70. Bugnion, E. (Letter to Perris on Antherophagus) L' Abeille 7, 1869-70, pp . XXVI-XXVII.
1863. Carus, J. V. and Gerstaecker, A. Handbuch der Zoologie, Vol. 2. Leipzig, W. Engelmann. 1863.
1900. Casey, T. L. Review of the American Corylophidse, Cryptophagidse, Tritomidse and Dermestidse, with other Studies. Journ. N. Y. Ent. Soc. 8, 1900, pp. 51-172, 4 figs.
1866. Eichhoff. Sammelberichte, Berlin, Ent. Zeitschr. 1866, p. 293-295.
1889. Fowler, Canon. The Coleoptera of the British Islands. London, Reeve & Co., 1889.
1869. Gorham, H. S. Leptinus in bees' nests. Ent. Month. Mag. 6, 1869, p. 89.
191 1. Grouvelle, A. Col6opteres Cryptophagides recueill& au Mexique par M. L. Diguet dans les nids du Bombus ephippiatus Say. Bull. Mus. Hist. Nat. Paris, 1911, pp. 97-100.
1883. Hoffer, Ed. Die Hummeln Steiermarks, 1, Halfte, 1863, p. 46.
1897. Janet, C. Etudes sur les Fourmis, les Guepes et les Abeilles. Note 14, Rapports des Animaux Myrmko- philes avec les Fourmis. Limoges, V. H. Ducourtieux, 1897.
1896. Lesne, P. Moeurs du Limosina sacra. Phenomenes de transport mutuel chez les animaux articules, Origine du parasitisme chez les insectes Dipteres. Bull. Soc. Ent. France 45, 1896, pp. 162-165.
1864. Packard, A. 8. The Humble-Bees of New England and Their Parasites, with notes of a new species of Anthero- phorabia, and a new genus of Proctotrupidse. Proc. Essex Inst., 4, 1864, pp. 107-140, PI. 3. 1872. Packard, A. S.
Guide to the Study of Insects, 3rd Ed.
Salem, Naturalists' Agency, 1872.
1873. Packard, A. S. Our Common Insects. Salem, Natural- ists' Agency, 1873.




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152 Psyche [December
1869-70. Perris, Edm. (Note on Antherophagus nigricornis) L'Abeille 7, 1869-70, pp. IX-X, XXV-XXVII. 1875. Perris, Edm. Larves des Col&opt&res. Ann. Soc. Linn. Lyon, 22, 1875, pp. 259-418.
1917. Rabaud, Et. Sur les Hymhopt6res parasites des oothh- ques d'orthoptkres. Bull. Soc. Ent. France, 1917, p. 178. 1858. Redtenbacher, L. Fauna Austriaca. Die Kafer. Wien, Carl Gerold's Sohn, 1858, p. 360 nota.
1911. Reitter, Edm. Fauna Germanica. Die Kafer des deut- schen Reichs. Stuttgart, K. G. Lutz. 1911. 1913. Renter, 0. M. Lebensgewohnheiten und Instinkte der Insekten bis zum Erwachen der sozialen Instinkte. Berlin, Friedlander and Sohn. 1913.
1869-70. Seidlitz, G. (Note on Antherophaqus niqricornis) L'Abeille 7, 1869-70, p. LXII-LXIII.
1912. Sladen, F. W. L. The Humble-Bee. Macmillan & Co. 1912.
1909. Smith, J. B. Insects of New Jersey. 1909. 1899. Sharp, D. Insects, in Cambridge Natural History 6, 1899. 1907. Wagner, W. Psychobiologische Untersuchungen an Hum- meln mit Bezugnahme auf die Frage der Geselligkeit im Tierreiche.
Zoologica 19, 1907, p. 145 (Fig. 117).
1903. Warner, W. V. (Note on Dichrornorpha viridis). Proc. Ent. Soc. Washington 5, 1903, pp. 308, 309. 1877. Xambeu, P. (Note on Podaqrion pachymerus). Bull. Soc. Ent. France 1877, p. LXIX.




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