Notes on the Food Habits and Food Defense of the Acorn Woodpecker
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NOTES ON THE FOOD HABITS AND FOOD DEFENSE OF THE ACORN WOODPECKER MICHAEL H. MACROBERTS Hastings Reservation University of California Jamesburg Route Carmel Valley, California 93924 The Acorn Woodpecker (Melanerpes formi- specifics and heterospecifics; and 3) it com- C~VOW), the most social of North American pares some aspects of the food habits of the woodpeckers, is a common resident of mixed Lewis ’ ( Asyndesmus lewis), Acorn, and Red- oak woodlands from Oregon south through headed (Melanerpes eythrocephulus) Wood- California, east through Arizona, New Mexico peckers. Since the food habits of the Acorn and Texas, and south to Panama. These wood- Woodpecker have been the subject of several peckers live in year-round groups. Each group previous papers (see literature cited by Beal consists of 2-10 birds of both sexes and ap- 1911, Ritter 1938, and Bent 1939), a repetition parently all ages (Ritter 1938). In most of these earlier works is avoided. groups males are reported to outnumber females, but this apparent discrepancy in sex STUDY AREAS ratio may be explained by the fact that ju- The study entailed 40 hr of observation be- veniles possessthe adult male plumage during tween 20 October and 15 December 1966 of at least part of their first year (Ritter 1938). two Acorn Woodpecker groups 2.5 mi. NW of Territory and food defense, nest hole con- Orinda, California, near the southern end of struction, and the care and feeding of nestlings San Pablo Reservoir, and about 500 hr of appear to be accomplished jointly by all mem- observation from 11 April to 15 September bers of the group. Aside from these rather 1968 of 11 groups at the Frances Simes sketchy points, little else is known about the Hastings Natural History Reservation, Mon- social organization of these woodpeckers. terey County, California. The groups studied In its food habits, the Acorn Woodpecker is at Orinda lived in mixed oak-pine woodland, one of the most atypical of all woodpeckers. while those studied at Hastings lived in foot- During the fall and winter the birds feed hill and canyon-bottom mixed oak woodland. extensively on acorns and other mast which they store in prepared holes in the surfaces FOODS OF THE ACORN WOODPECKER of oaks and pines. This aspect of their ecology has been repeatedly described (see literature STORED ACORNS cited in Ritter 1938 and in Bent 1939), but The present study adds little to previous because of a predominance of interest in this descriptions of acorn storage, except to de- unusual habit, other aspects of the wood- valuate the relative importance placed by peckers ’ food habits have been largely ne- other investigators on it as a food source for glected by most naturalists. This is unfortu- the Acorn Woodpecker. nate since it has led to the assumption that About half of the 11 groups studied at the food habits of these birds are far less Hastings had exhausted their stored acorns diversified than is actually the case. In addi- before observations began there in April. This tion to storing and consuming dried acorns, was due largely to acorn failure in all but the the woodpeckers feed extensively on mature black oak (Quercus kelloggii) the previous and immature green acorns before storage year. Nevertheless, each group, irrespective begins during the fall, and are proficient fly- of the state of its acorn supply, maintained catchers and sapsuckers. its own territory and subsisted on other foods The purpose of this paper is threefold: 1) throughout the summer. Those groups which it describes some previously unreported or in- had acorn stores did not use these to any large completely described aspects of the food extent but, like those groups without stores, habits of the Acorn Woodpecker; 2) it de- subsisted on other foods. scribes the defense of these foods from con- It is apparent therefore that during the summer the woodpeckers can subsist on and 1Present address: Department of Zoology, Parks Road, Oxford, England. prefer foods other than stored dried acorns, The Condor, 72,:1Q6-204, 1970 11961 NOTES ON THE FOOD HABITS OF THE ACORN WOODPECKER 197 and that, even if stores remain from the previous winter, these may not be extensively used. IMMATURE AND MATURE GREEN ACORNS Ritter ( 1938:29,84) mentioned that the Acorn Woodpecker uses immature acorns as food, but the present study suggests that their im- portance in the birds ’ diet is greater than has previously been supposed. In the groups observed at Hastings, immature acorns were eaten (especially those of the live oak, Quercus agrifolia, the black oak, Q. kelloggii, FIGURE 1. Acorn storage branch (center, with the valley oak, Q. lobata, and the blue oak, centimeter scale) and sapsucking branches (right and Q. d0ugZa.G) beginning in late July and con- left). Note differences in diameter of acorn storage tinuing through early September, without any and sap holes. The sapsucking branches used for this storing being involved. The immature nuts photograph were dead and desiccated and con- sequently nonfunctional at the time they were removed were picked, carried to “anvils,” split open, from the tree. and the meat immediately consumed. In the second week in September, the wood- peckers at Hastings began storing mature The sap holes were more or less uniformly green acorns. During this time they con- spaced. The distance between holes was ap- tinued to feed extensively on acorns, but in- parently independent of branch diameter, with stead of immediately eating all they picked, a mean nearest-neighbor measurement (mm) they stored some. These immature acorns of 24.7 (SD = 5.2, N = 131) and 21.6 (SD = 5.4, (and later, mature green acorns) were the N = 107) in two independent samples. This major food source of all the groups studied is approximately the same distance as between at Hastings throughout August and early acorn storage holes drilled in dead smooth sur- September, regardless of whether or not a faced trees (e.g., sycamores or valley oaks). particular groups’ acorn stores from the pre- The mean nearest-neighbor measurement vious year had been exhausted. (mm) between storage holes was 24.2 (SD = 4.3, N=73) and 20.5 (~~=3.6, N=llO) SAPSUCKING in two independent samples. The mean of the A third major food source is sap. The six entire set was 22.7 (SD = 5.0, N = 421). Vari- most studied Hastings groups had one or two ance analysis of the four samples shows that sap trees which they defended from other neither sap nor storage holes can be dis- birds. (It is not known whether the other tinguished by separation distance (F = 2.8, five groups maintained sap trees. ) In each P < 0.05). This suggests that a unitary be- case, these were live oaks (Q. agrifolia) lo- havioral mechanism may be involved in de- cated within 100 m of the groups’ main storage ciding the spacing pattern of storage holes tree. Live oaks were never used for acorn as well as sap holes. Figure 1 shows a sap- storage. sucking and an acorn storage area for com- The woodpeckers drilled small holes 5-15 parison. mm in diameter and 3-19 mm deep on the Since there are two possible ways that sap upper surfaces of middle and upper canopy and storage holes can be excavated to produce branches of live oaks. The branches in one a uniform spacing pattern, I shall briefly de- tree into which sap holes were drilled were scribe the process used by the woodpeckers. 2.5-15 cm in diameter. The diameter and On a certain section of limb, a bird will begin depth of the sap holes are related to the di- excavating a hole. After this hole is begun, ameter of the limb in which they are ex- but before it is finished, this bird or another cavated. Large limbs support wider and one will begin excavating a second hole at deeper holes than do small limbs. This is due the “correct” distance from the first. Other to the depth of the bark since the sap holes holes are excavated in a similar manner, each always penetrate through the bark but not into adjacent to, but at the “correct” distance from, the wood. In one particular live oak, about its nearest neighbor. Since hole construction 1800 sap holes were counted, although only a is a relatively slow process, many birds will part of the tree was used by the birds for this have participated in the excavation of a purpose. particular hole before its completion. Con- 198 MICHAEL H. MACROBERTS sequently, since the birds are continually Woodpeckers engage almost exclusively in making new sap and storage holes, there are “specific flights” or catch only one insect per numerous holes in various states of prepared- flight. In addition, the Acorn Woodpecker ness. never forages for insects directly on the The woodpeckers repeatedly visited and ground or in low brush as is common in the probed into these sap-filled holes. I would Lewis ’ Woodpecker. estimate that during the height of the sap- I feel, however, that previous researchers sucking period (June and July) each hole have underestimated the importance of fly- was visited, on an average, 4-10 times per catching as a foraging method of the Acorn hr, but since different holes were differentially Woodpecker. At Hastings flycatching was a used, it is difficult to make a more precise common activity throughout the study, espe- estimate. In the process of excavating these cially during the nestling period when the sap holes the woodpeckers may eat some of newly hatched birds were fed on insects the excavated bark, but whether or not this caught in this manner.