Olson • THE EXTINCT HAWAIIAN GENUS CIRIDOPS 
667 
neither a particularly wide gape nor any adaptations 
whatever for crushing or manipulating large objects 
with the bill. Munro (1892) records the last 
specimen being shot “while feeding on the lo|u|lu 
berries which abound in that place” he also noted 
that the fruits in that area were some 20 mm in 
diameter, which is much too large for a Ciridops to 
cat except for small pieces of exocarp and no such 
food was found in the stomach of that same bird. 
“No justification exists, however, for Ama- 
don's statement (1950: 223) that some 
drepaniids that rarely or never take nectar, such 
as... and Ciridops anna , still have a tubular 
tongue' [italics Bock's|. So little is known about 
the feeding habits of Ciridops , that no one knows 
whether or not this bird fed on nectar. In the 
absence of any contrary factual information. I 
would conclude from its tongue morphology that 
Ciridops fed on nectar at least during pan of the 
year" (Bock 1972: 75). 1 concur that it would 
have been unlikely that Ciridops did not take 
nectar when it was available, considering that 
many birds with no special adaptations for 
nectarivory are known to feed opportunistically 
on nectar (Fisk and Steen 1976. Franklin 1999). 
Speculation that Ciridops actually fed on fruit or 
nectar of Pritcluirdia palms based solely on the 
Hawaiian name is not supported by either the gut 
contents of the single fluid specimen nor by the 
morphological adaptations of the genus. 
Gizzard Contents and Tlteir Interpretation .— 
The gizzard in the alcoholic trunk specimen 
(BMNH 1939.12.9.58) had been slashed open by 
some previous examiner and could be seen lo be 
crammed with food. The contents were carefully 
removed and later identified by workers skilled in 
identifying food items from droppings obtained in 
field studies of Drepanidini. The overall insectiv- 
ory indicated for Ciridops was mentioned briefly 
by Scott el al. (1986: 156. and cited by Pratt 2005: 
275) based on communication from me concern¬ 
ing these findings: I adult lepidopieran (wing 
scales only), 13 adult psyllids (Hemipiera, jump¬ 
ing plant lice). 38 psyllid eggs, three adult psocids 
(Psocoptera. bark lice). 26 adult ‘Drosophila- like’ 
Diptera, 2 mites (Acari), I spider. 6 seeds of 
unknown fruit. I anther? (C. P. Ralph in litt. to 
Olson 30 Jun 1981). 
This collection of arthropods is similar lo the 
food items taken by other drepanidines in forest 
ecosystems on the island of Hawaii, including 
Apapane (Fancy and Ralph 1997), liwi (Fancy 
and Ralph 1998), Amakihi ( Loxops virens ; 
Lindsey el al. 1998). and Akepa (Lepson and 
Freed 1997). The first three are specialized nectar 
feeders and Akepa feed occasionally on nectar as 
well. Nectar, however, provides no protein so that 
even the most nectarivorous of birds, such as 
hummingbirds (Trochilidae), must feed on insects 
and spiders as well. 
The arthropods fed on by Ciridops and many 
other drepanidines are probably dispersed nearly 
throughout the forest ecosystems on Hawaii. 
Thus, it may not be so much a matter of 
specializing on a particular kind of prey as 
becoming adapted to extract widely available 
generalized prey from particular niches w ithin the 
ecosystem. Ciridops differed from all of its 
relatives in having strong feet and leg muscles 
that I hypothesize evolved for moving vegetable 
matter that accumulated in the axils of palm 
fronds. It may have fed on the same kinds of 
insects and spiders taken by Apapane and liwi. but 
it could gain access to them in places that could 
not be exploited by species whose hindlimbs were 
adapted solely for perching. 
DISTRIBUTION AND ECOLOGY OF THE 
GENUS CIRIDOPS 
Stejneger (1900: 72) wondered why Ciridops , 
along with Viridoitia. Loxioides . Rhodacanthis. 
and Cliloridops. should be confined to Hawaii and 
not have representatives on the other islands. 
Carlquist (1974: 129) considered that “the rarity 
and early extinction of some of the Hawaiian 
honeycreepers may have left gaps in our geo¬ 
graphical distribution; Ciridops might have oc¬ 
curred on islands other than the island of Hawaii, 
for example.” He could hardly have for,seen how 
the fossil record would completely upend every¬ 
thing previously thought to be known about the 
distribution of Hawaiian birds. The only direct 
evidence of the distribution of Ciridops comes 
from the scant historical record and from fossils 
(Fig. 8). 
Because of repeated hearsay reports of the Ula- 
ai-hawane, Wilson ventured that "I have little 
doubt that it will be found, perhaps in some 
numbers, in the upland region of the interior, 
which I was unable to explore....My friend Mr 
Francis Spencer, writing to me quite recently 
[presumably about 1892], says that his natives had 
seen the bird in the sw'ampy forest-region above 
Ookala [Keanakolu District according to Banko 
(1987: 248)] on Hawaii, and his description leaves 
