Limited Spread of Innovation in Kea - Kea Conservation Trust
Limited Spread of Innovation in Kea - Kea Conservation Trust
Limited Spread of Innovation in Kea - Kea Conservation Trust
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more than half way was significantly larger <strong>in</strong> opportunists<br />
than scroungers (median (quartiles) <strong>of</strong> opportunists: 0.33<br />
(0.13–0.53); scroungers: 0 (0–0.05), Mann–Whitney U test:<br />
U = 2.0, N1 = 4, N2 = 8, P = 0.022).<br />
Birds watch<strong>in</strong>g b<strong>in</strong> open<strong>in</strong>g<br />
With one exception there were always scroungers present<br />
at the kitchen area at the time <strong>of</strong> the 41 successful b<strong>in</strong> open<strong>in</strong>gs<br />
we observed (median number <strong>of</strong> scroungers present:<br />
4, quartiles 2–8). In 17 <strong>of</strong> these open<strong>in</strong>gs, we saw clear<br />
signs that other keas watched the successful performance:<br />
<strong>in</strong> six events one other opportunist was watch<strong>in</strong>g, <strong>in</strong> 11<br />
events one scrounger, and <strong>in</strong> two events, each, two and three<br />
scroungers. With 186 bird counts for scroungers, calculated<br />
from the data earlier, and 49 bird counts for additional opportunists<br />
present at the kitchen area at time <strong>of</strong> successful<br />
b<strong>in</strong> open<strong>in</strong>g, this gives a very similar proportion <strong>of</strong> counts<br />
for opportunists (0.122) and scroungers (0.113) watch<strong>in</strong>g<br />
successful b<strong>in</strong> open<strong>in</strong>g.<br />
It was not systematically recorded which birds watched<br />
unsuccessful open<strong>in</strong>g attempts, but it happened frequently.<br />
Beside the three events reported earlier when two birds<br />
attempted to lift the same lid at the same time, there were<br />
another 12 events when a watch<strong>in</strong>g bird approached and<br />
attempted to lift the same lid after the first bird had released<br />
it. On four other occasions, watch<strong>in</strong>g birds approached and<br />
attempted to lift the lid <strong>of</strong> an adjacent b<strong>in</strong> rather than the<br />
same lid.<br />
Discussion<br />
We found that only 14% (5/36) <strong>of</strong> the observed <strong>in</strong>dividuals<br />
<strong>in</strong> Mount Cook Village successfully opened the lids<br />
<strong>of</strong> rubbish b<strong>in</strong>s. Another 47% (17/36) <strong>of</strong> the observed <strong>in</strong>dividuals<br />
were seen to scrounge from b<strong>in</strong>s opened by the<br />
opportunists or from food scraps taken out <strong>of</strong> the b<strong>in</strong>s.<br />
Thus, <strong>in</strong> total 61% <strong>of</strong> the <strong>in</strong>dividuals had some pr<strong>of</strong>it from<br />
b<strong>in</strong> open<strong>in</strong>g. In the follow<strong>in</strong>g, we discuss possible reasons<br />
for the low frequency <strong>of</strong> opportunists.<br />
Implications <strong>of</strong> scroung<strong>in</strong>g opportunities<br />
The question <strong>of</strong> whether scroung<strong>in</strong>g opportunity <strong>in</strong>hibits<br />
or facilitates other birds to learn new behaviour is <strong>of</strong> crucial<br />
relevance for the spread <strong>of</strong> <strong>in</strong>novations (Giraldeau and<br />
Lefebvre 1987; Caldwell and Whiten 2003; Laland 2004).<br />
In contrast to other forag<strong>in</strong>g tasks <strong>in</strong> which producer and<br />
scrounger tactics are not fully compatible (Barnard and<br />
Sibly 1981), such a constra<strong>in</strong>t is unlikely for b<strong>in</strong> open<strong>in</strong>g<br />
as no split attention is required to detect opportunity for<br />
either <strong>of</strong> the two tactics. Thus <strong>in</strong> terms <strong>of</strong> the model <strong>of</strong><br />
Vickery et al. (1991), the sum <strong>of</strong> relative abilities for the<br />
two tactics fully overcompensate each s<strong>in</strong>gle relative ability;<br />
a condition <strong>in</strong> which a population <strong>of</strong> pure opportunists<br />
is favoured by the model. Yet this was not the case <strong>in</strong> b<strong>in</strong><br />
open<strong>in</strong>g performed by the kea.<br />
Also, the small frequency <strong>of</strong> successful b<strong>in</strong> openers <strong>in</strong><br />
the local population was unlikely to be simply the result <strong>of</strong><br />
scroung<strong>in</strong>g opportunity. Opportunists had a higher pay<strong>of</strong>f<br />
than scroungers <strong>in</strong> terms <strong>of</strong> both the number <strong>of</strong> observed<br />
feed<strong>in</strong>g events at the kitchen area, and the proportion <strong>of</strong><br />
feed<strong>in</strong>g food from the b<strong>in</strong>s rather than <strong>of</strong> the food scattered<br />
around the b<strong>in</strong>s. Because food was left outside the b<strong>in</strong>s after<br />
tak<strong>in</strong>g the preferred energy-rich parts like fat and meat, it is<br />
reasonable to assume that food left outside the b<strong>in</strong>s was less<br />
nutritious for keas than food chosen and taken out <strong>of</strong> a b<strong>in</strong>.<br />
Thus, it would have been advantageous for scroungers to<br />
open the b<strong>in</strong>s themselves. This is also demonstrated by the<br />
fact that 59% (10/17) <strong>of</strong> the scroung<strong>in</strong>g <strong>in</strong>dividuals were<br />
seen to attempt to open the b<strong>in</strong> lids.<br />
It may be that a scrounger’s pay<strong>of</strong>f be<strong>in</strong>g considerably<br />
smaller than the opportunists’ pay<strong>of</strong>f is still pr<strong>of</strong>itable because<br />
by scroung<strong>in</strong>g only, the bird does not have to pay the<br />
energetic costs <strong>of</strong> acquir<strong>in</strong>g the technique how to open b<strong>in</strong>s.<br />
Accord<strong>in</strong>g to this argument, less successful scroungers<br />
would be expected to do more open<strong>in</strong>g attempts than more<br />
successful scroungers because, <strong>in</strong> comparison to more successful<br />
scroungers, their ga<strong>in</strong> is more likely to outweigh<br />
their costs. Our f<strong>in</strong>d<strong>in</strong>g is contrary to this: unsuccessful<br />
<strong>in</strong>dividuals who were regularly seen at the kitchen area<br />
(dur<strong>in</strong>g 10 sessions at least), who had a low dom<strong>in</strong>ance <strong>in</strong>dex<br />
and were seen less <strong>of</strong>ten to scrounge directly from open<br />
b<strong>in</strong>s were seen less <strong>of</strong>ten to perform open<strong>in</strong>g attempts than<br />
unsuccessful <strong>in</strong>dividuals with a high dom<strong>in</strong>ance <strong>in</strong>dex and<br />
high forag<strong>in</strong>g <strong>in</strong> b<strong>in</strong> ratio. Given the fact that dom<strong>in</strong>ance<br />
<strong>in</strong>dexes do not differ between scroungers and opportunists,<br />
it is unlikely that the low frequency <strong>of</strong> opportunists is the<br />
result <strong>of</strong> a simple asymmetry <strong>in</strong> competition for access to<br />
closed b<strong>in</strong>s.<br />
In summary, <strong>in</strong>creased scroung<strong>in</strong>g opportunity can<br />
facilitate certa<strong>in</strong> aspects <strong>of</strong> f<strong>in</strong>d<strong>in</strong>g solutions as it was<br />
found <strong>in</strong> ravens (Fritz and Kotrschal 1999) and marmosets<br />
(Caldwell and Whiten 2003), for example to manipulate<br />
b<strong>in</strong> lids rather than to prevent such attempts (Giraldeau<br />
and Lefebvre 1987).<br />
Lid open<strong>in</strong>g as mechanical problem<br />
Pfeffer et al. (2002) found a similar <strong>in</strong>dependency <strong>of</strong> social<br />
status from <strong>in</strong>novator status <strong>in</strong> an operant task for freerang<strong>in</strong>g<br />
graylag geese as we did <strong>in</strong> lid open<strong>in</strong>g. In contrast,<br />
the study <strong>of</strong> Kothbauer-Hellman (1990) showed an impact<br />
<strong>of</strong> the social status <strong>of</strong> titmice that opened milk bottles <strong>in</strong> a<br />
flock. This difference may be expla<strong>in</strong>ed by the fact that b<strong>in</strong><br />
open<strong>in</strong>g provides a more challeng<strong>in</strong>g mechanical problem<br />
than open<strong>in</strong>g <strong>of</strong> milk bottles with one strike <strong>of</strong> the beak.<br />
The fact that only 9% <strong>of</strong> all open<strong>in</strong>g attempts resulted <strong>in</strong><br />
complete lid open<strong>in</strong>g demonstrates that the <strong>in</strong>vention <strong>of</strong> b<strong>in</strong><br />
open<strong>in</strong>g is new for most <strong>in</strong>dividuals and that it is not easily<br />
solved by apply<strong>in</strong>g the bird’s usual behavioural repertoire<br />
<strong>of</strong> forag<strong>in</strong>g techniques. <strong>Kea</strong>s performed <strong>in</strong>appropriate<br />
open<strong>in</strong>g attempts at b<strong>in</strong> lids that were blocked <strong>in</strong> some