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Nineteen subjects were tested using the Innovation Arena: 11 long-term and 8 short-term captive cockatoos (Figure 4).

Figure 4: An overview of the number of tasks solved per session for each individual. a) Field group, b) Lab group. Red lines = female; blue lines = male. Subjects receiving the motivational protocol due to their reluctancy to interact with the apparatus were classified as not motivated and depicted with a gray background. Previously published in Supplementary Information of32. Please click here to view a larger version of this figure.
The Principal Component Analysis resulted in two components having Eigenvalues above Kaiser´s criterion38 (see Table 2 for PCA output). PC1 loaded on frequency of contacts with tasks, time spent in proximity (i.e., within the grid) of the tasks, and the number of tasks touched. PC2 was positively affected by the number of contacts with already solved tasks and negatively with the number of tasks touched, not solved. Such task-directed behaviors are frequently used for measuring motivation (see12 for a review). Therefore, we used PC1 and PC2 as quantitative measures for motivation to interact with the apparatus in our model. Together they explained 76.7% of the variance in apparatus-directed behaviors and both, as well as session, significantly influenced the probability to solve tasks (PC1: estimate = 2.713, SE ± 0.588, χ2 = 28.64, p < 0.001; PC2: estimate = 0.906, SE ± 0.315, χ2 = 9.106, p = 0.003; session: estimate = 1.719, SE ± 0.526, χ2 = 6.303, p = 0.001; see Figure 5; see Table 4).

Figure 5: Influence of control predictors on the probability to solve: (a) PC1, (b) PC2, (c) Session. Points show observed data, area of points indicates the number of observations for each data point, dashed lines show fitted values of model and areas symbolize confidence intervals of model. Previously published in32. Please click here to view a larger version of this figure.
Six out of the 19 subjects received the motivational protocol during the experiment (Lab: 1 out of 11; Field: 5 out of 8). PC1 of these birds, which we categorized as not motivated, ranged between -2.934 to -2.2, while positive values were found for all other motivated individuals (Table 3).
With the presented method we found no difference of group on the probability to solve the 20 technical problem-solving tasks of the Innovation Arena (estimate = −0.089, SE ± 1.012, χ2 = 0.005, p = 0.945; Figure 5; see Table 4 for fixed effects estimates; all birds included).
A post-hoc comparison of the model with one including an interaction term of group with session (estimate = 2.924, SE ± 0.854, χ2 = 14.461, p < 0.001) suggests lower probability to solve in the field group in earlier sessions but not in the later. This difference in earlier sessions might be due to the high number of less/not motivated birds in group field (individuals for which testing stopped due to the rule of not solving any task in 10 consecutive sessions received between 10 and 13 sessions).
Further, we found no difference between the groups regarding the overall difficulty of tasks (comparison of full model with all birds included, with a reduced model lacking random slope of Group within Task: χ2 = 7.589, df = 5, p = 0.18). However, visual comparisons of birds that never required a motivational trial, hint to some differences in ability for single individual tasks (see, e.g., the Button task in Figure 6).

Figure 6: Observed data of motivated subjects and fitted values of model per task and group: Boxplots show the proportion of successes per task for both groups (green = Field; orange = Lab). Bold horizontal lines indicate median values, boxes span from the first to third quartiles for birds. Boxplots illustrate data from motivated birds only (to improve visual clarity). Individual observations are depicted by points (larger area indicates more observations per data point). Red horizontal lines show fitted values. Fitted values originate from the whole data set. Included are illustrations of Bite (bottom left), Button (top middle) and Seesaw (top right) tasks. Previously published in32. Please click here to view a larger version of this figure.
These results demonstrate the feasibility of the methodology for comparative research even if the animals have different experiences and ecological circumstances. A comparison of innovative problem-solving abilities using only a single task, such as the Button task, might have yielded a false conclusion that long-term captive birds are better problem-solvers. This difference could be explained by the lab population’s experience with stick insertion experiments while the motor action might not be as ecologically relevant for wild populations. Such differences could potentially be more pronounced when different species are compared (see19). We were further able to test how motivation affects problem-solving ability, while at the same time comparing the results of the two groups while controlling for motivation.
The 20 technical problems of the Innovation Arena can therefore be used to detect group differences on particular tasks, but also to estimate the overall innovative ability of groups. In the case of the Goffin`s cockatoo, both groups can, i.e., have the ability to, retrieve many rewards, if they want to, i.e., are motivated to interact with the apparatus.
Table 1: Protocol for coding behaviors: Detailed description of coded behavioral variables. Previously published in32. Please click here to download this Table.
Table 2: Principal component output: Factor loadings above 0.40 are printed in bold. Previously published in32. Please click here to download this Table.
Table 3: Details on subjects and values of task-directed behaviors and principal components: Superscripts if measure loads go above 0.40 per PC. Previously published in32. Please click here to download this Table.
Table 4: Fixed effects results of the model for probability to solve. Previously published in32. Please click here to download this Table.
Supplementary File: Technical drawing of the Innovation Arena (InnovationArena.3dm). Dimensions might deviate slightly. Can be loaded, e.g., in 3dviewer.net, which is a free and open source 3D model viewer39. Please click here to download this File.