TL;DR: Clark's Nutcracker (Nucifraga columbiana) is one of four species of corvids that commences harvesting, transporting, and caching the seeds of pinon pine (Pinus edulis) in mid—August.
Abstract: Clark's Nutcracker (Nucifraga columbiana) is one of four species of corvids that commences harvesting, transporting, and caching the seeds of pinon pine (Pinus edulis) in mid—August. Caching continues until inclement weather intervenes or the seed crop is depleted. Nutcrackers are efficient at collecting seeds from the trees because the birds discriminate edible and aborted seeds, select cones with higher than average number of good seeds and appear to concentrate on those trees that produce cones with large numbers of good seeds. The use of a sublingual pouch allows the nutcracker to carry up to 95 pinon pine seeds per trip (x = 55). A pouch full of seeds may be carried as far as 22 km from the collecting area (the pinon—juniper woodland, elevation ~ 1800 m) to communal caching areas where the seeds are buried in clumps in subterranean caches at a depth of 2 to 3 cm. These caching areas are on steep, south—facing slopes that are usually free of snow by late winter. Although solitary nesters, the birds, during the nonbreeding season, often form flocks when harvesting seeds. A flock of 150 nutcrackers cached between 3.3—5.0 X 106 pinon pine seeds in the autumn of a good cone crop. Each bird caches between 2.2 and 3.3 X its needed energetic requirements for the insect—free portion of the year. Some of the unrecovered seeds germinated and grew. The pinon pine reaches its highest density in relatively dry areas below the elevational range of other members of the genus Pinus, but ranges up to 3,100 m into the mixed coniferous forest. This pine has developed a number of traits that enhance its efficiency to be located and dispersed by the birds. The seeds are unusually large, of high energy value, and have relatively thin seed coats. The large size improves the germination potential of the seed but also attracts predators and dispersal agents. The seed coat color is markedly different for edible and aborted seeds, thus labeling them for visually oriented seed harvesters. Most pinon pine cones are oriented outward and upward which increases illumination of the seeds and tends to hold them in the cone. The pinon pine displays the seeds in its cones more readily than wind—dispersed pines by opening the seed—bearing cone scales to a uniform angle. Pinon pine seeds are retained for a long period of time in the cone because they are held in deep depressions on the cone scales by small flanges. This constellation of features indicated an evolutionary strategy on the part of the pinon pine to exploit and encourage members of the seed—caching guild to disperse the seeds. Deposition sites are presumably more favorable for germination than those obtained by simply broadcasting the seeds over a wide area. Individual seedlings potentially experience immediate intraspecific competition because the seeds are cached in clumps. Only one seedling, presumably the strongest, fastest—growing individual, survives. The benefits of this system to the nutcracker are that the bird has a supply of energy for the most unproductive months of the year. The bird also breeds in late winter or early spring, using the stored seeds for reproductive energy.
TL;DR: Experimental results suggest that seeds retrieved from nutcracker caches are as viable as seeds extracted by hand from cones and that seedlings originating in nutcracking caches have a good survival rate.
Abstract: (1) Clark's nutcrackers (Nucifraga columbiana) store a mean of only 3.7 whitebark pine (Pinus albicaulis) seeds per cache, which reduces competition for moisture and space. The mean depth at which seeds are stored, 2.0 cm, is compatible with germination requirements, and many sites selected appear suitable for seed germination. (2) One nutcracker stores about 32 000 whitebark pine seeds at subalpine elevations each year, which represent 3-5 times its energetic requirements. Although parent nutcrackers feed stored seeds to juveniles, in some years there is probably an excess of seeds stored by the population as a whole. (3) Experimental results suggest that seeds retrieved from nutcracker caches are as viable as seeds extracted by hand from cones and that seedlings originating in nutcracker caches have a good survival rate: 56% over the first year and 25% by the fourth year. (4) In comparison with Clark's nutcracker, alternative disperal agents, i.e. rodents and cone disintegration, disperse fewer seeds, disperse seeds shorter distances from parent trees, place seeds in sites less suitable for germination, and/or make large seed caches which lower reproductive potential. Nutcrackers disseminate seeds throughout the subalpine habitat and are, in part, responsible for the 'pioneering' status of whitebark pine. (5) The evolution of wingless seeds and indehiscent cones in the Cembra pine group was probably a consequence of seed dispersal by an ancestral nutcracker form. It appears that the Clark's nutcracker-whitebark pine interaction is both coevolved and mutualistic.
TL;DR: Five hypotheses of cache recovery behaviour in Clark's nutcracker (Nucifraga columbiana) were examined experimentally and it was found that seed-caching nutcrackers relocated caches using large objects as remembered visual cues.
TL;DR: Pinyon jays displayed significantly lower error rates than did nutcrackers or scrub jays after reversal of reward contingencies for both spatial and color stimuli and showed significant transfer between color and spatial tasks, suggesting a generalized learning strategy.
Abstract: In serial reversal learning, subjects learn to respond differentially to 2 stimuli. When the task is fully acquired, reward contingencies are reversed, requiring the subject to relearn the altered associations. This alternation of acquisition and reversal can be repeated many times, and the ability of a species to adapt to this regimen has been considered as an indication of behavioral flexibility. Serial reversal learning of 2-choice discriminations was contrasted in 3 related species of North American corvids: pinyon jays (Gymnorhinus cyanocephalus), which are highly social; Clark's nutcrackers (Nucifraga columbiana), which are relatively solitary but specialized for spatial memory; and western scrub jays (Aphelocoma californica), which are ecological generalists. Pinyon jays displayed significantly lower error rates than did nutcrackers or scrub jays after reversal of reward contingencies for both spatial and color stimuli. The effect was most apparent in the 1st session following each reversal and did not reflect species differences in the rate of initial discrimination learning. All 3 species improved their performance over successive reversals and showed significant transfer between color and spatial tasks, suggesting a generalized learning strategy. The results are consistent with an evolutionary association between behavioral flexibility and social complexity.
TL;DR: This work demonstrates that birds other than the nutcracker, rodents, and other mammals do not have the requisite behaviors to systematically disperse or establish whitebark pine, and that the pine is therefore dependent on the Nutcracker for its regeneration.
Abstract: Whitebark pine (Pinus albicaulis) is known to have its seeds harvested and cached in the soil by Clark's Nutcracker (Nucifraga columbiana), and unretrieved seeds are known to be capable of germinating and establishing new pines. Many other vertebrates also harvest and feed on these seeds, however, and the roles of these animals as dispersers and establishers of whitebark pine has been uncertain. This work demonstrates that birds other than the nutcracker, rodents, and other mammals do not have the requisite behaviors to systematically disperse or establish whitebark pine, and that the pine is therefore dependent on the nutcracker for its regeneration. These findings support previous suggestions that Clark's Nutcracker is a specialized frugivore that has profoundly influenced the ecology and the evolution of whitebark pine.