BREEDING AND MOLT SCHEDULES OF THE

RUFOUS-COLLARED SPARROW IN COASTAL PER6

JOHN DAVIS
Hastings Reservation
University of California
Carmel Valley, California 93924

The Rufous-collared, or Andean, Sparrow precipitation, and daily hours of sunshine as recorded
on a Campbell-Stokes type recorder were obtained for
(Zonotrichia caper&) occurs from the high-
Lima, 35 mi. NNW, and for Cafiete, 45 mi. SSE of
lands of the state of Chiapas, Mexico, to Cape the study area. Daylength was computed from The
Horn, Chile, its range spanning approximately Nautical Almanac.
73 degrees of latitude. Further, it is common Sparrows were caught in mist nets and each bird
over most of its range and is often the com- was marked with a distinctive combination of colored
leg bands so that individuals could be identified on
monest small passerine where it occurs. A
recapture. Of 523 birds caught, 133 were juveniles or
species with such characteristics is an ideal young first-year individuals, Of the remaining 390,
subject for a series of studies of breeding and 335 were either adults or mature first-year birds; 55
molt schedules at various latitudes where the were not aged. The age of juveniles or birds in part
populations are subjected to different photo- juvenal plumage was determined by feather characters.
The ages of birds that had comolet,ed the nostiuvenal
periodic regimes. Such comparisons make
or subsequent molts were determined by the ambunt of
possible an assessment of the relative and opacity (double-walling) of the skull after the method
changing importance of photoperiod and other of Miller ( 1946). Individuals with extensive clear
environmental features as proximate (timing) areas in the skull were classed as young first-year
factors in the scheduling of the important birds, those with small clear areas as mature first,-year
birds, and those with entirely opaque skulls as adults.
events in the annual cycles of a single species
The molt status of each individual was noted, and,
in different parts of its range. By so doing, a as an aid in determining sex, wing length was mea-
better understanding of the ultimate (adaptive) sured to the nearest 0.1 mm with a dial calipers.
factors underlying these various annual cycles (Males have significantly longer wings than do females
and the sexes arc about 90 per cent separable on the
in this and other species can be achieved.
basis of this character.)
The first study of the annual cycle of Zono- In addition to birds which were netted and released
trichia caper&s was made in 1958-59 by the after marking, 79 adult and mature first-year males
late Alden H. Miller (1959a, 1961, 1962) on a and 23 females of similar ages were collected for direct
population near Cali, Colombia, at latitude examination of gonads and stomach contents and
preservation of testes for subsequent microscopic
3” 30’ N. There the longest and shortest days
examination. Ten of the males and two of the females
of the year differ by only 12 min and photo- were collected on the study area and the others within
period may be disregarded as a proximate ?&-YA miles. Testes were fixed in Bouin’s fluid, stored
factor. More recently Wolf (1969) has reported in 70 per cent alcohol, prepared by the paraffin
on the annual cycle of this species near Las method, sectioned at 7 P, and stained with Mayer’s
hematoxylin and eosin. Each testis was assigned to
Varas, Costa Rica, at 10” 10' N, where longest
one of the six spermatogenetic stages described by
and shortest days of the year differ by 76 min. Bartholomew (1949), as follows: stage 1, resting
The present paper reports on the annual cycle spermatogonia only; stage 2, spermatogonia dividing,
near Chilca, Department of Lima, Peru, in the but only a few spermatocytes present; stage 3, many
coastal desert at 12” 30’S, where longest and spermatocytes; stage 4, spermatocytes with spermatids;
stage 5, spermatids with a few sperm; and stage 6,
shortest days differ by 91 min.
full spermatogenetic activity with many sperm.
The sex of banded birds was determined primarily
MATERIALS AND METHODS
on the basis of the presence or absence of a cloaca1
Field work was carried out between 9 October 1968 protuberance (Wolfson 1952). Breeding females were
and 9 August 1969’ at Hacienda San Javier Alto, 3 km easily distinguished by the presence of an incubation
NE Chilca, 9 km due E of the coast at an elevatioa (or “brood”) patch and the breeding status of a
of 25 m. This hacienda comprises about 150 ha female could be determined by the nature of the
planted mainly to citrus groves and cornfields. My patch: defeathered, vascular, edematous, or recovery
work was confined to a 20-ha area of citrus orchards Cstages l-4 of Bailev 1952). As regards the breeding
and cornfields. Although there was no natural source stat; of males, some workers have-used testis length
of fresh water anywhere on the hacienda, extensive as determined by laparotomy as an indicator of level
deep well irrigation provided an abundance of surface of gonadal activity since in this species, as in pas-
water for the sparrows during the study period. serines generally, there is a significant correlation
A Taylor max-min registering thermometer and a between testis length and the six stages of spermato-
Tempscribe thermograph were maintained in a stan- genetic activity previously described. In this study
dard instrument shelter near the study area and a the breeding status of individual males was assessed
Taylor “Clear-Vu” rain gauge was placed nearby. In on the basis of the size of the cloaca1 protuberance
addition, official government data on temperature, as measured aloag the anterior wall, and its shape

t1271 The Condor, 73:127-146, 1971

128 JOHN DAVIS

IO-

2
Yc = 2.227+.95046 X 0 N D J FMAMJJA
21, , , , , , , , , ( FIGURE 2. Male breeding cycle of Rufous-collared
2 4 6 8 IO Sparrows near Chilca, based on mean length of the
Cloocal protuberance ,mm
cloaca1 protuberance of monthly samples of banded
adults only. Horizontal line indicates mean, vertical
FIGURE 1. Regression of testis length on length of line indicates range, and rectangle indicates two stan-
cloaca1 protuberance of 62 adult male Rufous-collared dard errors on either side of the mean. Sample size
Sparrows collected near Chilca. is given at top of each vertical line. When an individ-
ual was captured more than once in a single month,
only the first measurement was used.

and consistency (changing from tubular to ovoid to

globular, and from flaccid to turgid, as it develops).
Cloaca1 protuberance is felt to be a more useful from October through December parallels the
indicator of breeding status than testis length for the steady increase in this period in the percent-
following reasons. First, there is a very high correla-
tion between the length of the protuberance and testis
ages of males in effective bree’ding condition
length in a sample of 62 fully adult males collected (with a protuberance length of 7 mm or more).
on o’r near the study area (r = 0.85; P < .OOl; 99 per Co’ntinued growth of the protuberance even
cent confidence limits, 0.73-0.92,; see also fig. 1). after individuals have reached breeding condi-
Second, there is a very high correlation in the same
tion is indicated by the steady increase in
sample between protuberance length and the six stages
of spermatogenetic activity (r = 0.75; P < .OOl; 99
mean length through March. The noticeable
per cent confidence limits, 0.X-0.86). In other words, decreases in size of protuberance and in per-
the length of the protuberance conveys much the centage of breeding males in April coincided
same type of information as does testis length. More with the start of the complete postnuptial molt
importantly, Wolfson (19854) pointed out that the
in a number of individuals. May and June
testis may be fully active spermatogenetically when
the protuberance is at a low level of development.
were months in which nearly all birds were
Thus the possibility is always present that one is deal- either molting or had recently completed the
ing with a male with fully active testes but without postnuptial molt; only small percentages of
motile sperm in the glomera and thus not in a state males were in breeding condition and mean
of breeding readiness. For this reason the nature of
protuberance length was small.
the cloaca1 protuberance is a more useful indicator of
actual breeding condition than is testis length or
histology. In the population under study, any pro-
TABLE 1. Percentages of male Rufous-collared Spar-
tuberance measuring 7 mm or more was fully devel-
rows with cloaca1 protuberance 7-10 mm long.
oped, with rare exceptions noted later. Miller (1958)
stated that in his population at Cali, males that were
No. of
breeding or had recently bred had a protuberance Cloaca
5-7 mm long. Month adults first-rear 7-10 mm %
Although it is possible to obtain motile sperm from
the glomera before the protuberance is fully developed
Oct. 40 6 21 45.7
(Albert Wolfson, pers. comm. ), any male with a fully Nov. 46 13 45 76.3
developed protuberance is definitely in breeding con- Dec. 28 10 35 92.1
dition; thus, this criterion is conservative and will Jan. 26 2 26 92.9
prevent ascribing breeding readiness to males before Feb. 65 16 78 96.3
they have actually achieved it. Mar. 15 6 20 95.2
Apr. 19 0 14 73.7
MALE CYCLE May 19 0 3 15.8
The main features of the male cycle are indi- June 21 2 3 13.0
cated in figure 2 and table 1. The increase July 17 7 10 41.7
in mean length of the cloaca1 protuberance Aug. 7 2 1 11.1
 ANNUAL CYCLE OF RUFOUS-COLLARED SPARROW 129

The towhee cycle is noticeably well-ordered

6 1 : : .._.-
:-:-:_.-.__ f
:_:-.-:._._. %!_._:.: ._. compared to that of Z. capensis; nearly all
birds regressed to stage 1, and when gonadal
5
. . ._.- :-. development started in mid-winter, it started
4
from a stage 1 baseline. Thus, the range of
variation was noticeably small at any given
point of time, and a number o’f to’whees had
testes in a “regression” stage, in transition from
._A.. -. :.
..-
full spermatogenetic activity to the resting
state. Such testes had the lumina of the
I
. tubules filled with degenerating cells and other
I
14 13 19 9 I/ 4 3
Apr
21 3 18 298
June July
9
plug
detritus. No male was collected near Chilca
Oct. NO”. Dec. Jan kb Mar. WV
with testes in this stage, another indication that
FIGURE 3. Stages of gonadal activity of 79 male very few males regressed completely after the
Rufous-collared Sparrows collected near Chilca.
breeding season ended there.
Whether any male Zonotrichia actually
reached full breeding condition in May is
Although few males reached actual breeding questionable, and apparently only a few males
condition in May and June, the testes of a progressed that far in June and July. Males
great majority of males examined histologically with cloaca1 protuberances 7 mm long were
showed considerable activity (fig. 3). This examined on 2, 22, and 26 May and on 5 and
probably resulted from two factors. First, the 9 June. The protuberance examined on 26
noticeable rarity of males with completely May is not described in my notes; the others
regressed (stage 1) testes suggests that the were described as either flaccid or ovoid, and
gonads of most males do not regress com- thus not typical o’f the fully developed struc-
pletely after the breeding season but return ture.
to either stage 2 or 3. Indee’d, of the three The only males in undoubted breeding con-
males that had stage 1 testes, the one collected dition after the postnuptial molt were exam-
on 2 June lacked the right testis and the left ine’d between 30 June and 4 July. Of 13 males
testis was hard and lumpy and obviously path- examined in this period, 11 had cloaca1 pro-
ologic. Thus, only two of 78 normal males had tuberances 7-10 mm long. Eight adults and
completely regressed testes. Incomplete re- one unaged bird had fully developed pro-
gression has been described in other species tuberances; they were “nearly globular” and
of tropical birds. Moreau et al. (1947) noted “ovoid” in two’ mature first-year males. The
that the annual range of testis size was far very restricted perio’d in which breeding birds
greater in temperate zone species than in a were examined is undoubtedly the result of
number of tropical species which they inves- sampling error, as a female with a stage 1
tigated. As Lofts and Murton (1968) pointed incubation patch and a large, yolky egg in the
out, this discrepancy may be accounted for in oviduct was collected on 18 June, and the last
part by the fact that tropical species with long females with active incubation patches were
breeding seasons often do not regress com- examined on 15 July. The period in which
pletely as do, northern birds and the gonads in breeding males occurred after the postnuptial
such species remain relatively large. Second, molt probably extended from about mid-June
experimental work done by Miller with birds to abo’ut mid-July. Relatively few males were
from his study area in Colombia (Miller 1959b, involved and this was a minor effort compared
1965) and by Miller and 0. P. Pearson (MS) to the main breeding period from December
on further samples from the same locality through March. It was apparently over by
indicated that the refractory period in those mid-July, as only two of 17 males collected
birds was either negligible or lacking. The between 14 July and 9 August had testes in
same was apparently true of the Chilca pop- stage 6, one collected on each of those dates.
ulation. Most males sho’wed a strong gonadal The latter bird had the only cloaca1 protuber-
recrudescence, even to stage 68,as soon as the ance measuring as much as 7 mm; in the other
postnuptial molt had been completed, or in
16, measurements ranged from 2 to 5 mm,
some cases when the terminal stages of molt
averaging 3.3 mm.
had been reached.
Miller ( 1959a ) , working at 3” 30’ N in Co-
A marked difference may be noted between
the present cycle and that of a small, North lombia, followed the annual cycles of a num-
Temperate Zone passerine such as the Rufous- b’er of colo8r-banded males over long periods
sided Towhee (Pipilo eythrophthalmus) in and showed beyond doubt that many males,
central coastal California (Davis 1958, fig. 1). perhaps most, came into breeding condition
130 JOHN DAVIS

TABLE 2. Cloaca1 protuberance length” and type, and testis stage in 16 male Rufous-collared Sparrows before
start and after completion of postnuptial molt.

Cloaca before molt Cloaca after molt

Bird no. date (mm) date (mm) Cloaca1 type Testis stage

18 18 Oct. 9 12 July 5 flaccid 4

26 21 Oct. 9 23 June 5 tubular
41 23 Oct. 7 22 May 5 flaccid
63 18 Nov. 9 22 May 5 6
88 15 Jan. 9 9 Aug. 5 5
105 8 Nov. 9 1 July 10 globular
107 14 Feb. 9 22 May 7 ovoid
17 June 6 tubular
125 12 Nov. 7 19 June 5 tubular
22 July 3’ flaccid
224 30 Dec. 9 9 July 5 ovoid
254 15 Jan. 8 23 June 3 flaccid
267 31 Jan. 10 4 July 9 globular
276 4 Feb. 10 4 July 9 globular
297 6 Feb. 9 20 May 3 2,
305 10 Feb. 10 14 July 5 flaccid 6
338 25 Feb. 9 30 May 5 tubular
344 2.5 Feb. 10 2 July 8 globular
a When a bird was captured more than once before or after the postnuptial molt, maximum protuberance length is given.

twice in a single year. My records of individ- carry the stage 4 patch until it is refeathered
ual males are much more limited, and are in the postnuptial molt, or they may breed
summarized for 16 males examined before and again, the patch again becoming vascular and
after the postnuptial molt (table 2). Since then edematous ( Bailey 1952). Three females
males regress from breeding condition after the definitely bred again after having a stage 4
onset of this molt, it is obvious that males 105, patch. Number 256 had a stage 4 patch on
267,276, and 344 achieved full breeding condi- 16 January and a stage 3 patch on 18 February.
tion twice within the period of study. Number Number 270 showed the same sequence on 31
107 may have come into breeding condition a January and 19 February, and number 299 the
second time by 22 May, although his protuber- same on 68February and 13 March. It is evi-
ance was not fully developed at that time. By dent that December was the month of most
17 June he had regressed somewhat. The intense breeding, as virtually all females exam-
remaining 11 males ha’d failed to reach breed- ined on any day were in breeding condition
ing condition after the complete molt. Num- whereas an appreciable proportion of females
ber 125 clearly regressed in late June or in on any day in January, February, or March
July from the level of gonadal activity that he were either between nestings or were through
had reached on 19 June. Apparently most, if breeding for the season.
not all, males show a pronounced recrudes- Three females had breeding seasons lasting
cence of gonadal activity after the postnuptial at least four months. Number 1 had a stage 1
molt, but few reach effective breeding condi- incubation patch on 9 October, and on 19
tion. February she had a stage 3 patch regressing
to stage 4, indicating that she had recently
FEMALE CYCLE finished breeding. Number 8 had a stage 1
The female cycle is indicated in figure 4. In patch on 10 October and a stage 2 patch on
October 57.5 per cent of the females examined 10 February. Number 38 had a stage 3 patch
were breeding. The percentage rose to 68 in on 23 October, a stage 3 patch on 5 February,
Novemb’er and to 96 in December, when only and a stage 3 patch regressing to stage 4 on
one female examined lacked an incubation 20 February. Miller and Miller (1968) estab-
patch. In these first three months the female lished an incubation period of 12 days and a
cycle was quite similar to that of the males, mean fledging period of 11 days in the popula-
From January through March the cycle is dif- tion near Cali, Colombia; they also found that
ficult to interpret because of the increasing females may spend up to 33 days with depen-
proportion of females with stage 4 patches. dent young after fledging. Assuming a similar
Such females may either stop breeding and schedule in the Chilca population, these fe-
 ANNUAL CYCLE OF RUFOUS-COLLARED SPARROW 131

0
o-x-8 ; .-. : (‘ ._t,

0
y-x
9
X.0..
XXO...
x
:
:,
0 y----_-r
6 4
.-.-O

I 5
I
5 3
I
8
l
.
.
.-.

5
..o.
. .
:

3
. .
1
._;

7
Oct. Nov. Dec. Jon. Feb. Mor. Apr. MoY June July Aug.

FIGURE 4. Female breeding cycle of Rufous-collared Sparrows near Chilca. Stage 0 indicates absence of an
incubation patch. Stages 14 indicate the four stages of incubation patch described by Bailey ( 1952). Stage 5
indicates a refeathering patch. When an individual was captured more than once, at least 14 days elapsed be-
tween successive records.

males had time for two successive nestings AGE OF BREEDING BIRDS
separated by a lengthy period of attendance Only one recapture record provides any infor-
on dependent young. A fourth female, number mation on this point. A male, number 52, was
120, had a stage 3 patch on 11 November and in juvenal plumage on 24 October. On 8 Jan-
was seen carrying nest material on 7 January. uary he had completed the postjuvenal molt.
She undoubtedly attempted two broods. Miller (1961) found that Colombian birds
The latest breeding, female examined had a complete’d this molt at four months of age and
stage 3 incubation patch on 13 March. On 13 that the molt occupied a mean of 70 days. If
May a pair of adults accompanied by a single the Chilca population was similar in this re-
juvenile was seen. One adult held a green gard, number 52 would have been about four
caterpillar in its bill, obviously for delivery to months old on 8 January. At that time there
the young bird. This juvenile probably came was no sign of gonadal activity bat on 7 Feb-
from an egg laid sometime in April. The next ruary his cloaca1 protuberance measured 5 mm
breeding female was collected on 18 June; she and on 14 February, 7 mm. He apparently
had a stage 1 patch and a large, yo’lky egg in came into breeding condition at about five
her oviduct. The bree’ding season for most months of age. This was the earliest age for
females ended in March, about a month before which Miller (1959ia) found evidence of breed-
most males regressed from breeding condition, ing condition in a Colombian sparrow. On 12
although there was some breeding in April, as March, number 52 was starting a complete
just noted. Between I8 June and 6 August, molt and his cloaca1 protuberance was only
ten of 23 females examined had active incuba- 5 mm long an’d tubular. He had reached
tion patches, the latest on 15 July. Thus, there breeding condition so late in the season that
was a second perio’d of breeding separated
it is doubtful that he had actually mated.
from the first, main peak by three months in
When he was collected on 20 May, he had
most birds and by at least two months in some.
finished a complete molt and his testes were
Apparently most females did not breed at that
in stage 6.
time, although the sample is too small to give
a good idea of what proportion of females Of 82 breeding males of known age exam-
actually bred in winter. As in the males, the ined in October, November, and December,
second breeding period was much shorter than 62 (75.6’ per cent) had fully ossified skulls. If
the first. the complete double-walling of the skull takes
132 JOHN DAVIS

a maximum of 11 months, as established by a rather limited area. Number 26, which had
Miller and Miller (1968) for Colombian birds, a song peculiarity permitting easy identifica-
about one-fourth of the breeding males in this tion, remained in the same area throughout the
period were less than one year old. Of 56 study period. Further, the fact that adults and
breeding females examined in the same period, mature first-year males have active gonads
35 (62.5 per cent) had fully ossified skulls. outside the molt period makes it unlikely that
The difference between first-year to adult they would enter flocks readily.
ratios for males and females is not significant
in a chi-square test. MOLT
Two molts were evident in adults and
SINGING AND FLOCKING BEHAVIOR mature first-year birds in the Chilca popula-
Singing behavior showed a high degree of tion The major breeding period was preceded
correlation with the testis cycle. Miller and by a partial molt in some birds and followed
Miller (1968) found that singing was depressed by a complete molt in all. This schedule dif-
during the molt periods and elevated during fers from that described by Miller (1961) for
the two peaks of breeding in their population. his population in which there was a complete
In the Chilca population, singing was at a high molt following each of the two breeding
level from early Octolber to mid-March. I was periods. Thus, and this was the striking fea-
absent from the area 19-31 March. Singing ture, birds of breeding age replaced the entire
fell off noticeably in the first half of April, a plumage twice a year. The Chilca schedule
time when many birds were starting the post- was similar to that described by Wolf (1969)
nuptial molt. By the end of the month only a for the Costa Rican population, but there were
few birds were singing short bouts. In early important differences between the prenuptial
May a few birds resumed singing and on 20 molt described by Wolf and the corresponding
and 21 May several more birds had started to molt found in my population.
sing. On 11 June there was a very noticeable
“PRENUPTIAL” MOLT
resurgence of singing and by mid-June I esti-
mated that the song o#utput was at the level In October, November, and December 2,18
noted in the breeding season. By mid-June, adult and mature first-year birds, both breed-
many males had either finished the molt or ing and non-breeding, were examined. Of
were in its terminal stages and, as indicated these, 168 were aged on the basis of skull
in birds collected or examined, a strong go- characteristics; 50 were not aged but were
nadal recrudescence was under way in the assumed to be adults or mature first-year birds
males. On 12 and 14 July no singing was heard because they showed signs of gonadal activity.
and there was a virtual absence of song until Of the 218, 108 (49.5 per cent) showed either
7 August. The mean stage of the testes of nine some active molt or very fresh feathers indica-
birds collected 12-30 July was 2.6. During my tive of recent molt so’mewhere in the plumage.
last three days in the area, 7-9 August, several This molt must have started in some birds in
males began to sing much of the time, although September, as male number 9, caught on 10
song bouts were rather short. The testes of October, was replacing primaries 1-5, and
nine males collected in this period were at a males 11 and 13, caught on 11 October, were
mean of stage 4. replacing primaries 14 and 1-6, respectively.
Flocking, which occurs in many passerines Birds showing active molt were very rare in
outside the breeding season, was noted only December and feather replacement had ended
once. On 15 May a flock of at least 30 spar- in nearly all birds by the end of November.
rows was seen working through a tangerine There was a highly significant difference in
grove. Three were collected; all were young the proportions of males and females taking
first-year birds and the gonads of the two birds part in this molt. Of the 130 males, 78 showed
autopsied were very small and completely in- active o’r very recent molt, but only 30 of the
active. It seems likely that the flock was 88 females (x2 = 14.09; df, 1; P < 601). Con-
composed mostly or entirely of young birds. sidering only the 168 birds of known age, there
This group was in the area for several weeks was a significant difference in the proportion
although it became smaller as time passed. of adults to first-year birds among the molting
Miller and Miller (1968) reported that flock- males; 20 of 27 first-year males showed signs
ing was rare in their population and that males of molt, but only 34 of 72 adults (x2 = 5.71;
defended territory throughout the year. My df, 1; P < .02). Among the females, 11 of 22
data on this point are scanty, although my first-year birds and 16 of 47 adults showed
recapture records indicate that adults and signs of molt; the difference is not significant,
mature first-year birds tended to stay within possibly because of the relatively small sample
 ANNUAL CYCLE OF RUFOUS-COLLARED SPARROW 133

TABLE 3. Number of primaries replaced by Rufous- 5-5 half-grown. On 6 February l-5 were full-
collared Sparrows showing discontinuities. grown and easily distinguishable from the
No. of primaries
older primaries 6-9. Number 38, a mature
revlaced M&S Females Total first-year female, was replacing primaries l-3
1 3 0 3 on 23 October; 2 and 3 had barely erupted.
2 10 1 11 On 7 November, 2 and 3 were full-grown. On
3 15 4 19 5 and 20 February no further molt had oc-
4 11 5 16 curred. On 15 July she had undergone a
5 16 6 22 complete postnuptial molt. In her case there
6 11 4 15 was clear overlap of molt and breeding. When
7 1 1 2 caught on 23 October she had a stage 3 incuba-
8 2 1 3 tion patch; in addition to primary molt she
also showed light body molt and some under
tail covert molt. Number 63, an adult male,
size. Thus, the overlap between molt and the was replacing primaries 14 on 29 October;
onset of breeding condition was less for the 44 were half-grown. On 18 November 44
adults, which made up the bulk of the breed- were full-grown; on 10 January and 4 February
ing population, and for the females, which no further molt had occurred. On 22 May he
have much greater demands on their resources was in completely fresh plumage.
in the breeding season than do the males. Birds 38 and 63 not only illustrate arrested
This molt invariably involved the primaries molt but also indicate that in the ensuing post-
and the feathers of the old and new genera- nuptial molt, replacement started all over again
tions were easily distinguishable and formed with primary 1 and not at the point at which
what Ashmo81e( 1963) termed “discontinuities” molt had been arrested. It is clear that we
in the primary series. Replacement always are dealing with two separate molts and not
started with primary 1 and proceeded distally; with a single, suspended molt. In addition to
most birds replaced 2-6 of these feathers these birds, which were actually molting when
(table 3). Primary molt was usually accom- first caught, there were many which showed
panied by light to heavy molt of the crown no active molt but had obvious bilaterally
feathers and/or upper and under tail coverts. symmetrical discontinuities in their primary
Body molt was uncommon and was always series indicative of arrested molt (table 3).
light. Rectrix molt was found in only 13 per Such birds could be distinguished at least until
cent o’f the birds and ranged from replacement mid-March. Three males with discontinuities,
of the central rectrices (l-l) to replacement caught on 2 December, 15 January, and 6
of the entire tail. February, replaced all primaries in the com-
This molt differed markedly from the pre- plete postnuptial molt, as did several males
nuptial molt described by Wolf (1969), in and females which were still actively molting
which replacement of remiges was rare and when first caught and which had replaced
never involved the primaries. Since primary varying numbers of primaries. All these birds
replacement came early in the complete molts indicate that the molts preceding and follow-
describ’ed by Miller (1961) and also in the ing the major breeding period were completely
postnuptial molt of the Chilca population, this separate molts.
limited molt should not be considered a pre- Toward the end of my studies at Chilca,
nuptial molt but an incipient second complete between 22 July and 9 August, eight of 12
molt which was arrested by the onset of breed- mature first-year and adult males and two of
ing. four females of similar ages showed active
Miller (op. cit.) noted four cases in 2. molt. Of the 10 molting birds, three were
capensis in which molt was arrested by breed- taken in July and the others between 5 and 9
ing activity, and similar arrest has also been August. There was crown molt in five birds,
described in some seabirds (e.g., Sterna fus- chin molt in five, throat molt in eight, body
cata, Ashmole 1963)) in a tropical ground-dove molt in two, and tail covert molt in one. This
(Columbgallina talpacoti, Snow and Snow
was apparently the beginning of the incipient
1964), and in several species of cuckoos of
complete molt between the end of the second
the genus Clamator (Friedmann 1948; Payne
breeding period of 1968-69 and the beginning
1969).
Three birds in the Chilca population clearly of the major breeding period of 1969-70. If so,
showed arrested molt. On 9 October number it was about a month earlier than in the pre-
2, an adult male, had replaced left primary 1 vious year; as noted previously, some birds
and right primaries l-2. On 29 October pri- must have started this molt in 1968 as early as
maries 1-5 had been replaced bilaterally with September. There is evidence that breeding
134 JOHN DAVIS

. ::H: io
C

II
8 .O .

IO
. 0 ! 0

9
0 . .O 0 .

8
0

7
. : . .

6
. . 0 .

5
. . . 0 0

4
. . .O

March April May June

FIGURE 5. Period of the postnuptial molt in the Chilca population of Rufous-collared Sparrows, based on 84
adult and mature first-year males (dots) and 51 females of similar ages (circles). N indicates molt not started
and C indicates molt completed. Stages l-11 are those described by Miller ( 1961); stages Z-10 refer to replace-
ment of primaries 1-9.

in the second period of 1968 persisted at least co’mplete and the molt is similar to the com-
until August, and perhaps into early Septem- plete molts described by Miller (1961) and
ber, as a full-grown juvenile was caught on by Wolf (1969).
9 October and another was seen on 10 October.
POSTJUVENAL MOLT
It is impossible to ascertain the limits of the
second breeding period of 1968 with any The juvenal plumage, figured by Miller ( 1961:
degree of precision. 145, fig. l), is replaced by adult type plumage
at the incomplete postjuvenal molt, in which
POSTNUPTIAL MOLT
the remiges are either not replaced at all or
The chronology of the postnuptial molt is are only partly replaced. Of seven juveniles
shown in figure 5, using the 11 stages of molt with adequate recapture records, six showed
described by Miller (1961). Feather replace- no primary molt and the seventh showed
ment in adults and mature first-year birds was partial primary replacement. This bird was
 ANNUAL CYCLE OF RUFOUS-COLLARED SPARROW 135

TABLE 4. Stomach contents of 97 Rufous-collared Sparrows.

Green material SOlanUm Amaranth Other seeds Insects

Month N n % n % n % n % 11 %

Oct. 5 4 80 3 60 0 0 1 20 2 40
Nov. 5 4 80 3 60 2 40 0 0 0 0
Dec. 3 1 33 2 67 2 67 0 0 0 0
Jan. 3 0 0 2 67 3 100 2 67 1 33
Feb. 2, 2 100 1 50 1 50 1 50 0 0
Mar. 3 1 33 3 100 2 67 2 67 1 33
Apr. 13’ 1 8 3 23 8 62 5 38 3 23
May 15 3 20 12 80 4 27 3 20
June 16 37 : 3: 11 69 4 25 3 19
July 21 : 19 7 33 19 90 0 0 2 10
Aug. 11 5 45 1 9 11 100 1 9 1 9

Totals 97 31 32 30 31 71 73 20 21 16 16

first caught on 24 October in juvenal plumage of the five birds of known sex were males.
with all rectrices but left 5-6 full-grown. On The female may simply have been starting her
8 January the plumage was entirely of the postnuptial molt early. The most significant
adult type; primaries I4 were new and full- birds were 62 and 113, as they may have rep-
grown, 5-9 were old. On 12 March his first resented a small group in which the incipient
complete molt had started. Primaries 1-5 were complete molt was not arrested but reached
new (the second replacement for l-4) and completion. This suggestion is strengthened
the three inner pairs of rectrices were being by the fact that the cloaca1 protuberance of
replaced. Body molt had just started. The 62 was only 3 mm long and that of 113 only
skull still had large “windows.” This bird was 5 mm long. In these individuals molt may
collected on 20 May. It was in completely have delayed the attainment of full breeding
fresh plumage and all primaries and rectriccs condition. This seems most likely for 62, which
were full-grown. The skull was about 40 per had the smallest cloaca1 protuberance of any
cent opaque. In general, the postjuvenal molt adult examined in October and November.
appears to be similar to’ that described by These two males are also’ significant in that
Miller (op. cit. ) as “incomplete, most notably they were the only birds examined which
failing to involve the remiges.” might have had two complete molts within one
year. If so, such a schedule must be very rare
ANOMALOUS MOLT in the Chilca population. Including these two,
A few adults showed heavy general molt at 132 adult and mature first-year males were
times when such molt was not evident in the examined in October, November, and Decem-
rest of the population. A male, 62, had re- ber, and the incipient complete molt proceeded
placed all primaries and was molting heavily to’ completion in only two ( 1.5 per cent).
on 29 October; primary 9 was about one-fourth
grown. Male 113, caught on 8 November, had FOOD HABITS
replaced all primaries; primary 9 was half- The stomach contents of 71 adults or mature
grown. All rectrices were new and all but the first-year males, 22 females of similar ages,
outermost were full-grown. The remainder of and four young first-year males were examined
the plumage was fresh. Number 250, a male macroscopically. Since no pronounced varia-
caught on 13 January, had replaced primaries tion in diet among these age and sex groups
l-7 left and 1-8 right, primary 8 being half- was noted, the data presented in table 4 are
grown. All remaining plumage was fresh and pooled. The diet was predominantly vegetable
this male was apparently in the late stages of throughout the study period. Seeds of ama-
a complete molt. Numbers 301 (a male) and ranth (Amaranthus sp. ) were by far the most
333 (sex undetermined) were also apparently important single food item and were found in
near the end of a complete molt on 7 and 24 73 per cent of all stomachs examined. In 29
February, respectively. Female 373, with a stomachs (30 per cent) they were the sole
refeathering incubation patch, was about half- item present. Insects were found in only 16
way through a complete molt on 28 February. per cent of all stomachs and were usually
Without knowing the previous histories of represented by small fragments. In only two
these birds it is impossible to account for the of 97 stomachs were they a major component,
unseasonable nature of their complete molts. one examined on 18 April, one on 17 June.
Significantly, all were adult and all but one Although the greatest percentages of stomachs
136 JOHN DAVIS

insects] were not measured at our field station,

the subjective impression gained was that they
were always present in abundance.” In this
study, an abundance of see’ds and green mate-
rial was evident at all times, but it is apparent
that there were major fluctuations in the insect
population.

BIRDS OF THE LOMAS

Lamring (1967) wrote of the barren hills of
0
29 I/ 25 19
,
13
I
27 15 15 IS 222859
I the coastal desert, “In areas of dense winter
act NO” Dee Jml Feb Apr May July piug
JYne
fogs, grasses, bushes, and other plants manage
FIGURE 6. Numbers of insects per 10 sweeps of an to thrive on trapped and condensed moisture.
insect net on a standard sampling area near Chilca, on Such areas of fog vegetation are known as
dates indicated. Zomas.” Koepcke (1954, 1963) gives excellent
descriptions of the lomas and their vegetation
types. She notes that in the summer sunny
containing insects were recorded in October, season, when the vegetation of the lomas is
January, and March, samples in tho8semonths dry and sparse, Rufous-co811aredSparrows are
were very small. With larger samples from rare or absent, but that when the lomas are
April through early August, the relatively high covered with green vegetation in winter, the
percentages recorded in April, May, and June sparrows come to such areas in large numbers
may represent increased insect consumption and breed there commonly. I made the follow-
in the molt period. Further, the two stomachs ing observations in the Lomas de Atocongo,
containing large amounts of insect material 15 mi. SE Lima, an area of fog vegetation on
were taken in that period. a steep, west-facing slope.
Although adults and independent young On 21 November, I found scattered shrubs
birds rely primarily on seeds and green mate- still in leaf, a few small trees, some forbs, and
rial, nestlings and dependent young are appar- scattered small patches of grass. Seven singing
ently fed an insect diet. On several occasions male Zonotrichia were counted from the base
adults were noted flying with large insects or of the slope to the crest, a distance of perhaps
insect larvae in their bills, undoubtedly for % mi. By 10 January the shrubs had been
delivery to nestlings or fledglings. Miller and browsed back severely by goats, no grass was
Miller (1968’) and Wolf (1969) stated that present, and the few sparse annuals remaining
nestlings and fledglings were fed insects; this were dry and covered with dust. No sparrow
is usual in emberizine sparrows. was seen or heard along the route previously
Food for adults and independent young traversed. On 12 June, the lower third of the
birds was always readily available on the slope was still very dry, with little green vege-
study area. Both Solunum nigrum and Ama- tation present, and little regrowth of the
ranthus sp. grew abundantly along the irriga- browsed shrubs. Hoswever, on the steep upper
tio’n channels paralleling the ro8wsof fruit trees two-thirds of the slope, there were extensive
and they formed luxuriant growths beneath patches of grass and herbaceous vegetation
the trees themselves. The same was true of and the shrubs had leafed out. On the upper
purslane ( Portulaca obracea), which may third there was nearly continuous ground
have been an important source of green mate- cover and the vegetation was wet with droplets
rial. In addition, forbs of other species and of condensed fog. Many of the shrubs in this
grasses were present along roadsides and on area were nomwover 5 ft high. One sparrow
the shoulders of main irrigation canals. was heard singing steadily on the census route.
To get some idea of seasonal fluctuations in Despite the noticeable regrowth of vegetation,
insect abundance, 10 sweeps with an insect net no influx of sparrows had yet occurred. On
were made at intervals of about one month in 11 July there was yet a further advance in
a patch of Solarium, Amaranthus, and tall grass the vegetation. The shrubs had leafed out
on the study area. The results of this sampling more and there was a thick carpet of grass,
are shown in figure 6. Insects were relatively moss, and low annuals on the upper two-thirds
numerous in October, November, and Decem- of the slope. The lower third was densely
ber, in the first part of the major breeding covered with a blanket of blooming amaryllis
period, and again in April and May, the period (Hymenocallis sp.) and was thus unsuitable
of heaviest molt. Miller (1962) stated, “Al- for the sparrows. The soil all the way up the
though these food sources [grass seeds and slope was so muddy that footing was difficult
 ANNUAL CYCLE OF RUFOUS-COLLARED SPARROW 137

and near the crest the fog or low clouds were predators, and favorable weather conditions,
swirling down to the ground. At least seven would be of critical importance. As Lack
steadily singing males were recorded, and (1968) notes, at times a compromise must be
perhaps as many as nine. Two pairs were seen made. In temperate regions, for example, nest
and one chase was observed. On 11 August, a predators, especially snakes, would be most
conservative count of 11 singing males was abundant when weather conditions are best
made with every attempt to avoid duplicate for nesting.
counts. One juvenile with a full-grown tail However, Pitelka (1957, 1958) was the first
was noted. Many sparrows were seen; I esti- to point out that molt may b’e of considerable
mated a total of at least 25 pairs on the slope. importance in the evolution of avian breeding
The vegetation was lush and damp and again, schedules because it demands so much energy
at the crest, there was cloud cover down to that molt and breeding cannot be carried on
ground level. simultaneonsly but must be mutually exclusive,
Two visits were made to the Lomas de or virtually so. There are therefore two alter-
Lachay, 66 mi. NW of Lima. On 1 March natives involved. If the ultimate factors in the
I noted three single sparrows and a flock of evolution of an annual cycle pertain to breed-
lo-12 in a grove omfintroduced pines and ing schedule, then breeding coerces the timing
eucalyptus. There was some leaf litter and dry of molt by preempting a certain part of the
grass beneath the trees. One bird sang a single year in which molt cannot occur. If the ulti-
song and another, or po8ssibly the same one, mate factors pertain to molt, then the reverse
sang steadily for 5 min. The presence of a is true.
flock suggested that the birds were not breed- Most students of avian breeding schedules
ing at that time. On 13 July the ground in agree that an adequate food supply for nest-
the same area was covered with a dense green lings and fledglings is probably the single
growth of grasses, forbs, and leafy shrubs. most important prerequisite for breeding suc-
Sparrows were abundant and an estimated 200 cess. Since growing young need a high protein
were seen. Several chases and numerous dis- diet, and since insects are nearly always the
placements were noted. The impression was most abundant source of protein readily avail-
that of a large population at the beginning of able to most passerines, it is not surprising that
the nesting season. these birds usually feed their young a diet
The derivation of the populations of the mainly, if not exclusively, of insects. Molt also
lomas is unknown, as is the gonadal status demands a rich diet to support the synthesis of
of individuals in that part of the year spent a new feather coat, and again, availability of
outside the lomas. Such questions will be insects is an important adjunct to survival in
answered only by extensive color-banding of the molt period for many birds. However,
individuals breeding in the lomas and re- molt makes varied demands, and some depend
covered elsewhere. However, in view of the on factors other than diet. There is no doubt
extremely low recovery rates of North Amer- that a heavily molting bird suffers from reduc-
ican passerines (see, for example, Cortopassi tion of insulation against heat, cold, and mois-
and Mewaldt 1965), and the virtual absence ture. In a temperate or a cool tropical area
of ornithologists in Peru, it seems doubtful that this could be critical at night, when tempera-
any relevant information will be forthcoming tures are lowest and foraging is impossible.
in the foreseeable future. The important point Again, heavy molt undoubtedly impairs loco-
relative to the present study is that sparrows motion to some degree just at a time of stress
are absent from the lomas when breeding is at when efficient foraging and effective evasion
its height in the coastal river valleys and breed of predators would be at a premium. Although
commonly in the lomas when breeding is in- dietary considerations are undoubtedly the
frequent elsewhere. mo’st important in both breeding and molt,
weather and shelter factors are also important
PROXIMATE AND ULTIMATE and should not be overlooked.
FACTORS IN THE ANNUAL CYCLE Previous studies of the annual cycle of Zono-
Avian breeding has traditionally been thought trichiu cape&s have evoked opposing view-
of as being timed to occur in that part of the points. Wolf (1969) assumed that the ultimate
year in which the prerequisites fo’r successful facto’rs in the timing of the annual cycle were
nesting would be optimal and nesting success associated with breeding and that molt “is
would be highest. Such prerequisites as ade- relegated, evolutionarily, to periods that are
quate nest sites and supplies of nest materials, unfavorable for breeding.” Miller ( 1962), on
an adequate food supply for nestlings and the other hand, believed that molt was the
fledglings, a sufficiently low incidence of nest important factor in the annual cycle of his
138 JOHN DAVIS

Hat. San Javier Alto

, ’ 3 km NE Chilca

A Ca?iete

1‘1 i2 20 24 i8 i8 i5 i3 i0 i
Oct. Nov. Dec. Jan. Feb., Mar. Apr. May June Aug.
FIGURE 7. Mean weekly temperatures (“C) for Lima, Cafiete, and Hacienda San Javier Alto.

population and that it coerced breeding. In fore critical in regulating food supply, both as
the Chilca population there were two distinct reg,ards green and seed-bearing vegetation and
breeding periosds, a major period extending the insects which lay their eggs and develop
from October to mid-March in the females and on such vegetation or in the soil. Fresh water
into April in the males, and a second, lesser for direct consumption was available in abun-
period, restricted both in extent and in the dance throughout the study period and this
numbers of individuals involved, from about supply was independent of precipitation.
mid-June to mid-July. The major period was Thus, any possible effect of the extremely low
preceded by an incipient cosmpletemolt which precipitation was overwhelmed by agricultural
was arrested by the onset of breeding condi- practice.
tion, and followed by a complete molt which Photoperiod also seems to be of no impor-
occurred from April through June. We may tance. Males came into breeding condition in
now consider possible proximate and ultimate October and Novemb’er while daylength was
factors involved in this cycle. increasing. From 1 October to 30 November
In previous studies of avian breeding, and daylength increased from 12 hr 13 min to’ 12
molt cycles, the environmental variables most hr 46 min. Whether such a minor increase
frequently considered have been the photo- over such a long period would have any effect
periodic cycle, the annual cycles of tempera- is questionable. Most of the major breeding
ture and precipitation, and seasonal fluctua- period, from late December to April, was in
tions in food supply. Sunshine has been a period of decreasing daylength. The rapid
consi’dered in rather few studies but will be gonadal recrudescence which followed the
considered here. postnuptial molt also occurred while the days
Precipitation seems to be of no importance were getting shorter. The second, lesser breed-
as a proximate factor in the Chilca cycle and ing period started while daylength was still
may be dismissed at the outset. Between 10 decreasing and when days were shortest, and
October 1968 and 9 August 1969, a total of it extended into the first two or three weeks
15.2, mm of precipitation was recorded. The of the perio’d of increasing, daylength. Be-
greatest amount in any 24-hr period was 1.8 tween 158 June and 158 July, daylength in-
mm. The regular irrigation which was carried creased from 11 hr 25’ min to 11 hr 30 min; this
out was far more important in regulating soil could hardly have been a factor in causing the
moisture than was precipitation and was there- onset of breeding in some birds in that period.
 ANNUAL CYCLE OF RUFOUS-COLLARED SPARROW 139

II 29 20 17 21 28 25 23 27 7
Oct. Nov. Dec. Jan. Feb. Mar. Apr: May June Aug.
FIGURE 8. Hours of sunshine per week at Lima and Cafiete.

The most convincing evidence against the enda. The males came into breeding condition
importance of photoperiod as a proximate in October and November in a period of rising
factor in this study is derived from a compari- temperatures, and relatively high temperatures
son of the breeding schedule of 2. capensis (22” C or above) prevailed over the major
near Chilca with that reporte’d for this species breeding period and over the first three weeks
at Acolla, Provincia de Jauja, Department of in May, i.e., over most of the molt period. A
Junin, Peru, on the east slope of the Andes at second, minor peak of elevated temperatures
3400 m, by Blancas Sanchez (1959). Acolla is occurred in the last two weeks of July, starting
at 11” 46/ S, or 44 mi. N of Chilca latitudinally, nearly at the end of the second breeding
and total daylength at Chilca exceeds that at period. Temperature has been shown to in-
Acolla by only 4 min a year. Blancas Sanchez fluence testis development experimentally in
reported that 2. capnsis, abundant at Acolla, north temperate birds if a stimulatory day-
nested there in March, April, and May, starting length is provided (see Farner and Wilson
precisely at the time that the sparrows near 1957:259-260 for discussion). In the study
Chilca were finishing. Further, the entire nest- area daylength ranged from 11 hr 22 min to
ing season at Acolla lay entirely within the 12 hr 53 min and such a regime would prob-
period of decreasing daylength. Thus, photo- ably permit temperature to act as a proximate
period at these latitudes must be so weakly factor. Although temperature may have been
coercive on the bree’ding of this species, if it of so’me influence in relation to the first breed-
is coercive at all, that it is overridden by other ing period it failed to correlate with the second
factors. and could not have been important as a proxi-
Temperature cycles for Hacienda San Javier mate factor at that time.
Alto, for Lima, 35 mi. NNW, and for Cafiete, Sunshine shows a high degree of correlation
45 mi. SSE, are shown in figure 7. The general with both breeding periods. Sunshine records
similarity in pattern among the three cycles is for Lima and Caiiete are shown in figure 8.
apparent, although the absolute values of the The general similarity of the two cycles is
Caiiete cycle are closer to those for the haci- evident. Since there is a rather close corre-
140 JOHN DAVIS

spondence between sunshine and temperature mate factor, it has been found to be of possible
on the Peruvian coastal desert, as may be seen importance in some studies (Baker 1938;
by comparing figures 7 and 8, and since the Threadgold 1956, 1960). Marshall (1951)
temperature cycles at Cafiete and the hacienda found that sunshine and temperature were of
are nearly similar, the sunshine cycles at these considerable importance in accounting for the
two localities must be much the same. As difference in timing of the gonad cycles of
with temperature, the males were coming into certain passerines from one year to the next
breeding condition in a period of increasing at Oxford, England, and he stated that for
incidence of sunshine, the incidence of sun- certain birds resident in areas in which the
shine was high throughout the first breeding winter sky is overcast (true of Chilca), sun-
period, and it remained high until mid-May. shine may be a principal long-range environ-
The sunny period thus included at least the mental influence.
first half of the molt period. Further, there In the permissive climate which prevailed
was a lesser, but quite noticeable, period of on the study area, and with an abundance of
increased sunshine between 20 June and 18 food and water present throughout the year,
July which correlated with the second breed- the energy deman’ds on the males were slight
ing period. outside the molt period. Kendeigh (1941) has
Finally, we may consider fluctuations in pointed out that, striking though the develop-
insect abundance as indicated in figure 6. ment of the testes from the resting to the fully
October, November, and December were active state may be, these organs at maximum
months in which insect numbers were rela- size do not exceed 2-3 per cent of the weight
tively high; this includes the period in which of the bird. It is not surprising that, with a
the Zmotrichia population was coming into refractory period apparently negligible or lack-
breeding condition, and the first month in ing, there was a pronomuncedresurgence of
which nearly all birds were in breeding con- gonadal activity as soon as the energy demands
dition. There was a noticeable decrease in of molt had terminate’d. Thus, the proximate
insect numbers in January and February, de- facto’r or factors which operated in October
spite the fact that actively breeding females and November acted on males which were
were found as late as 13 March. However, already at an advanced level of gonadal
females with stage 4 (recovery) incubation activity. This contrasts with the situation in
patches began to appear in early January and temperate zone males in which nearly all
the intensity of bree’ding began to wane from individuals are in a state of complete gonadal
that point on. It is possible that the decrease inactivity at the time the important proximate
in insect numbers coincided with less frequent factor, increasing photoperiod, starts to be
breeding in January and February. The in- effective. Such males must be brought from
crease in insect numbers in April and May the lowest level of gonadal activity to the
coincided with the first two-thirds of the molt highest and, as Wolfson (1942) noted in the
period and, as previously noted, examination Oregon Junco (Junco oreganus), the earliest
of stomach co’ntents suggests that adults and stages of gonadal development are the slowest.
mature first-year birds were actually consum- As regards the females, in most species
ing more insects in those months. The low which have been investigated, and nearly all
numbers of insects in June, July, and the first breed in temperate areas, they have a far less
nine days of August includes the latter part pronounced response to photoperiod than do
of the molt period and the entire second breed- males. Experimental studies sugg,estthat they
ing period. respond to the visual stimuli presented by the
As regards the males, increasing incidence presence of the necessary prerequisites for
of sunshine seems to be the proximate factor nesting and to the behavior of the male. It is
of major importance. Rising temperatures may difficult to think of an environment more
act in concert with it. The apparent failure stable and generally more favorable for nesting
of many males to reach effective breeding than that in the study area. Predators were
condition in the second breeding period may rare; the only potential nest predators actually
have resulted from the fact that although sun- noted on the area were a pair of Sparrow
shine increased in this period, it never reached Hawks (Falco sparoeriw ) that flew over occa-
the high levels attained in October and No- sionally. Norway rats (Rattus norvegicus)
vember. It may also be that in part the low were seen rarely around buildings but never
level of breeding was caused by the failure in the study area. Snakes were conspicuous
of temperatures to rise concomitantly, as they by their absence and no cats were present.
did in October and November. Although sun- The lowest temperature recorded in 10 months
shine has been largely neglected as a proxi- was 12.8” C and the highest was 32.8; wind
 ANNUAL CYCLE OF RUFOUS-COLLARED SPARROW 141

Northern Hemisphere

prenuptial molt

Costa Rica

dry season dry season

Colombia

Southern Hemisphere

I I I I I I
Oct. l Nov. ’ Dec. I Jan. I Feb. ’ Mar. A pr. May June July Aug. Sept.

FIGURE 9. Diagrammatic representation of the occurrence of breeding and molt periods of Zonotrichia ca-
pensis at Las Varas, Costa Rica (Wolf 1969)) near Cali, Colombia (Miller 1961, 1962), and near Chilca, and
nesting season at Acolla (Blancas Sinchez 1959). Diagrams marked “Molt” refer to complete molts. Cross-
hatched bar in Chilca diagram refers to incipient, arrested complete molt.

and precipitation were never prominent. Thus, nificantly in the 10 months of study was the
weather was permissive to nesting throughout level of insect abundance, which would pre-
the study period. Nest sites were abundant in sumably reflect variation in the availability
the thousands of citrus trees and nest materials of food for nestlings and fledglings. The proxi-
were common along irrigation ditches and on mate factors timing the onset of breeding
the shoulders of irrigation canals. Ample condition in the females thus appeared to be
supplies of foo’d and water for adults and sunshine, which might have been a potent
independent young birds were present at all psychological stimulus, and the increasing
times. The only prerequisite that varied sig- numbers of insects, presumably directly evi-
142 JOHN DAVIS

dent to the females and another potential

visual stimulus.
If the ultimate factors involved pertain to
nesting, the conclusion seems inescapable that
the breeding schedule was timed to take ad-
vantage of increased availability of insect food
for the young. Weather factors may also have
been involved secondarily, since the major
breeding period fell entirely within the warm-
est and sunniest part of the year. However,
such factors were undoubtedly of lesser impor-
tance since the climate was generally so mild.
The annual cycle near Las Varas, Costa
Rica (Wolf 1969) is shown in figure 9. Cor-
relations with rainfall and temperature were
not consistent and it is difficult to point to any
proximate factor of outstanding importance.
Photoperiod was apparently not coercive as
the onset of breeding was about one month
later at a station at 2200 m than at a nearby
station at 1800 m. However, in his habitat
descriptions he stated, “In the early morning
there is usually a cloud bank hanging over the
eastern slopes of PO& [the higher station]
while the western side of Barba [the lower
station] tends to be relatively free of such
clouds until later in the day.” Here is another
suggestion that sunshine and/or temperature
may be important proximate factors in cool
tropical areas near the equator. JFMAMJJASOND
The cycle near Cali, Colombia (Miller 1961, FIGURE 10. Mean monthly rainfall (mm) at Cali,
1962), is also shown in figure 9. Breeding Colombia, July 1934-December 1954.
periods d ‘ id not correlate with the weak sea-
sonal photoperiodism which amounted to only
12 min difference between the longest and the schedule at Chilca. Wallace noted that in
shortest days of the year. No correlation with the fall and winter of 1955-56 at PopayLn,
temperature was evident but there was a high unusually heavy and prolong,ed rains virtually
correlation with rainfall. Miller assumed con- eliminated the usual dry season and may have
stant adequacy of food for both maintenance interfered with fall breeding; they may also
and breeding at all times. This may have been have caused earlier breeding in the spring
true, but the possibility cannot be dismissed than was evident in Miller’s population. This
that there might have been periods of in- was apparently an unusual year. I have
creased insect abundance following the peaks summed up monthly rainfall for Cali, Colom-
of rainfall and that the peaks of nesting were bia, near Miller’s study area but at lower
timed to take advantage of them. elevation, from July 1934 through December
In this connection, George J. Wallace (MS) 1954. The data (fig. 10) clearly indicate a
kept notes on the nesting of 2. capensis at pronounced bimodality of rainfall and the
PopayLn, Colombia, 2” 3’N and at 2092 m, rainiest periods coincide almost exactly with
from September 1955 to July 1956. The only those described by Miller. Apparently, the
active nests were found from April to June, schedule Miller described is the usual one at
although the presence of a few streak-breasted his study area.
juveniles, some attended by adults, in the fall Miller (1961, 1962) postulated molt as the
and early winter, indicated limited nesting at major factor determining the annual cycle near
that time. This suggests that in the year in Cali. In the females the two complete molts
which Wallace observed there was a major followed the two peaks of nesting; in other
breeding period in spring and early summer words breeding and molt were mutually exclu-
and a minor breeding period in the fall and sive. In the males, the separation of these
early winter. This schedule would agree with two activities was not as clear-cut, presumably
that reported by Wolf in Costa Rica and with because the energv demands imnosed on the
 ANNUAL CYCLE OF RUFOUS-COLLARED SPARROW 143

males by breeding activities were less than schedule geared to rainfall as a proximate
those imposed on the females, and some over- factor might benefit both molt and breeding.
lap was possible. Nevertheless, it was appar- Investigations of food supply in relation to
ent (Miller 1961) that males in the middle rainfall in the area occupied by this population
stages of molt, when energy demands were would be of the greatest interest.
greatest, were rarely in breeding condition and The breeding schedule of Z. capcnsis at
that regressed testes occurred relatively rarely Acolla, Peru, is also of interest as regards molt.
in non-molting males. The onset of molt oc- Blancas Sanchez (1959) noted that the birds
curred three or four weeks after the cessation there nested in March, April, and May. Al-
of rainfall at the end of May and at the end though he gave no information on the time
of December. of molt, we may assume that it began shortly
Miller concluded that molt was triggered by after the end of the breeding season and
the psychological response of the birds “to the probably started sometime in June for most
changed condition of wetness in the grass” in birds. In his descriptions of the four seasons
which they foraged and to a greater incidence at Acolla, Blancas Sanchez notes that winter
of sunshine. The argument that the molt sched- (22 June-22 September) is the dry season,
ule coerced the breeding schedule is made with no rain. This would also be the season
more plausible by Miller’s assumption of a of most sunshine. Unfortunately, he presents
constant food supply throughout the year. few temperature data. The mean minimum
Presumably other prerequisites for successful for 7-16 May was 7.5” C and for 7-22 August,
nesting were also assumed to be in constant 6”. These few data suggest that the cold stress
supply. This would almost automatically on the population would not be appreciably
ascribe ultimate factors to molt. greater in winter than in fall, when breeding
Miller noted that the population was on a occurred. Again, molt appeared to be timed
schedule in which molt occurred mainly in the to coincide with the season of greatest dryness
dry seasons and provided fresh plumage for of vegetation and most sunshine.
the ensuing wet periods. He stated, “This may Snow and Snow (1964) hypothesized that
have some general advantages, although WC the breeding seasons of the numerous species
have not seen these sparrows badly water- which they studied in Trinidad (10” 40’ N)
soaked from periods of heavy downpour.” were “ultimately adapted to take advantage
However, there are obvio8us advantages to of the periods of greatest food supply and, in
such a molt schedule. Miller stated that fre- some species, to the period of greatest safety
quent foraging in coarse, perennial grasses by of the nest.” They concluded that none of the
the birds in his population wore the plumage proximate factors considered (daylength, rain-
heavily and that the individual, whether it has fall, temperature, and seasonal fluctuations in
bred or not, must molt twice a year or become food supply) was important in controlling the
incapacitated. The correlation of molt with whole annual cycle. Further, they pointed out
rainfall thus assured adequate spacing of the that despite the great variation in time of
two molts. If they were separated by too short onset of breeding, nearly all of the species
an interval, the lengthy period between the which they studied molted in the same months
second and third of a sequence of three molts of the year. They therefore suggested that the
would place far too much wear on the plu- molt schedule was the major factor in timing
mage. Further, foraging in wet grass, which the annual cycle, with the breeding season
might be disadvantageous for a heavily molt- timed in each species according to the specific
ing bird, would be largely avoided if the molt refractory period which would determine the
were timed mainly to those periods in which amount of time elapsing between the end of
the grass dries out each day. molt and the onset of breeding. The annual
Another possibility is that there was a bi- variation in breeding season for a given species
modality in the food supply for the young would result from modification by “environ-
associated with the bimodality of rainfall, and mental factors which sometimes may be ob-
that rainfall adjusts to best advantage the tim- vious . . . but more often are too subtle to be
ing of both molt and breeding. With the apparent to the observer” (Snow and Snow, op.
relatively great outlay of energy that the indi- cit. ). This is something of a compromise, in
vidual, especially the female, makes in the which the ultimate factors are still related to
course of two breeding efforts and two com- breeding but with molt itself acting as the
plete molts a year, the premium would be on major timing factor. This, of course, demands
the proper timing of all major energy-demand- a set of proximate factors for molt. There is
ing activities, or at least on the best com- some evidence that the beginning of the wet
promise possible. Thus, the evolution of a season is the major proximate factor timing
144 JOHN DAVIS

the molt of Manacus manacus in Trinidad, but to coincide with an increased abundance of
information is lacking for other species. insects. But this is no more compelling a
As regards the Chilca population, one may correlation than is the timing of the onset of
assume, as I have, that the ultimate factor in breeding condition and the month of most
the timing of the entire annual cycle is the intense breeding (December) also to occur at
coincidence of breeding with insect abundance. a time of increased insect abundance.
However, to consider the opposite possibility, In evaluating molt as a major factor in the
let us assume that my insect sampling, ad- timing of nesting (as sugg,ested for Trini’dad
mittedly minimal, was inaccurate, and let us by Snow and Snow 1964), in October and
assume, as Miller did, that a food supply ade- November at Chilca, when these two activities
quate for both breeding and maintenance was were actually competing for the energy re-
present at all times. The other important sources of the individual, it was molt that was
prerequisites for successful nesting, as noted arrested rather than breeding. The only pos-
previously, were indeed present at all times, sible role that molt might have had in the
so we are assuming, that effective bree’ding timing of breeding would have been to ter-
could occur at any time of year. Thus, ulti- minate it. Here we are on dangerous ground
mate factors wo8uldpertain to molt rather than for we are dealing with a population that is
to reproduction. apparently uninfluenced by the slight photo-
In this case, with precipitation virtually non- periodic changes in its environment, and one
existent, the foraging milieu would be dry that appears to have a negligible refractory
save for perhaps 30-35 daylight hours a year. period. Considering all of the evidence in
Temperature would then be the ultimate factor hand, it seems most likely that the ultimate
of importance, with molt occurring in a warm, factors involved in this cycle pertain to breed-
sunny season so that individuals in heavy mo’lt ing and not to molt, that the ultimate factor
would not have to contend with overcast skies of primary importance is the timing of nesting
and relatively low temperatures. The period to an adequate supply of insect food for the
of relatively high mean temperatures, high young, and that the sunshine and temperature
mean minima, and high incidence of sunshine cycles are the proximate factors of greatest
starting in October was amply long to include importance.
most of the nesting effort and most of the The birds breeding in the lomas must re-
period of molt (fig. 8, 91). This is in sharp main an intriguing mystery. There is consider-
contrast to the situation at high northern lati- able anthropological evidence indicating that
tudes (Morton et al. 1969) in which birds have the lomas were much more widespread in the
a short season in which to breed and molt. central coast, at least up to 6000 B. C. (Lan-
The lack of urgency of molt in the Chilca ning 1967). The numbers of birds presently
population was exemplified by the fact that involved are apparently only a fraction of
the females finished breeding at about mid- those involved in earlier times. Whence they
March but did not start the postnuptial molt are derived, and what they do away from the
until April (fig. 5). In contrast, at Point lomas, are unknown. One may hypothesize
Barrow, Alaska, 71” 20’ N, female Lapland only that for these birds, the breeding period,
Longspurs (Calcarius lapponicus) losing a which is a second, minor reproductive effort
nest or young after 20 June will, with rare for populations resident elsewhere along the
exceptions, not renest but start the postnuptial coast, is a major, and possibly the major,
molt ( Frank A. Pitelka, pers. comm. ) . breeding period.
If the ultimate factors in the Chilca cycle
did in’deed pertain to molt and involved high SUMMARY
temperatures and high incidence of sunshine, The annual cycle of a population of Zono-
then molt was not timed optimally, as the trichia cape&s was studied in the coastal
weather started to become cooler and more desert of Peru at 12” 30’S from early October
overcast in May, well before the molt period 1968 to early August 1969. Both sexes came
was over. December, January, February, and into breeding condition in October and No-
March were the months of most sunshine and vember, and by Decembmervirtually all birds
January, February, March, and April were were breeding. Females stopped breeding at
the months of highest mean minima. Thus, mid-March and males in April.
January, February, and March would have In October and November an incipient
been the months best suited to molt. The most complete molt involving the primaries as well
compelling reason for the occurrence of molt as other parts of the plumage was arrested by
in April, May, and June, on the basis of the the onset of bree’ding condition. The post-
evidence in hand, would be the timing of molt nuptial molt started in April in both sexes and
 ANNUAL CYCLE OF RUFOUS-COLLARED SPARROW 145

was completed in some birds by the end of CORTOPASSI,A. J., AND L. R. MEWALDT. 1965. The
May and in nearly all birds by the end of circumannual distribution of White-crowned Spar-
rows. Bird-Banding 36: 141-169.
June.
DAVIS, J. 1958. Singing behavior and the gonad
Testes regressed incompletely at the begin-
cycle of the Rufous-sided Towhee. Condor 60:
ning of the postnuptial molt but rapidly re- 308-336.
turned to high levels of spermatogenetic FARNER, D. S., ANIY A. C. WILSON. 1957. A quan-
activity when the molt had been completed. titative examination of t,esticular growth in the
A second breeding period involving relatively White-crowned Sparrow. Biol. Bull. 113:254-
267.
few birds occurred from about mid-June to
FRIEDMANN, H. 1948. The parasitic cuckoos of
about mid- July. Africa. Washington Acad. Sci. Monogr. no. 1.
Precipitation and photoperiod were unim- KENDEIGH, S. C. 1941. Length of day and energy
portant in timing the annual cycle. Increased requirements for gonad development and egg-
incidence of sunshine appeared to be of pri- laying in birds. Ecology 22:237-248.
mary importance in bringing both sexes into KOEPCKE, M. 1954. Corte ecol6gico transversal en
10s Andes de1 Per& central con especial con-
breeding condition, with temperature probably
sideraci6n de las aves. Parte I. Mem. Mus. Hist.
of secondary importance. Nat. “Javier Prado” (Lima), no. 3.
Greater insect abundance in October, No- KOEPCKE, M. 19’63. Anpassungen und geographische
vember, and December may also have been Isolation bei Vageln der peruanischen Kiisten-
important as a proximate factor for the females lomas. Proc. XIII Int. Ornithol. Congr., Ithaca,
P. 1195-1213.
and was probably the most important ultimate
LACK, D. 1968. Ecological adaptations for breeding
factor in this cycle.
in birds. Methuen & Co. Ltd.. London.
LANNING, E. P. 1967. Peru before the Incas. Pren-
ACKNOWLEDGMENTS tice-Hall, Inc., Englewood Cliffs, New Jersey.
This project would not have been possible without LOFTS, B., ANI) R. K. MURTON. 1968. Photoperiodic
the continued support and encouragement of Oliver P. and physiological adaptations regulating avian
Pearson. He also read the manuscript critically and he breeding cycles and their ecological significance.
generously made available unpublished experimental J. Zool., London 155:327-394.
data on the refractory period of Z. capensis. I wish to MARSHALL, A. J. 1951. The refractory period of
thank Frank A. Pitelka for critically reading the manu- test% rhythm in birds and its possible bearing on
script. I am most grateful to Don Manuel de1 Solar for breeding and migration. Wilson Bull. 63:238-
allowing me to work at Hacienda San Javier Alto and 261.
for extending every kindness and courtesy. Many MILLER, A. H. 1946. A method of determining the
people helped me in various ways, especially Maria age of live passerine birds. Bird-Banding 17:
Koepcke, Emanuel Plenge, Hernando de Macedo, Luis 33-35.
Gonzales M., and Luz Sarmiento. Plant specimens MILLER, A. H. 1958. Reproductive periods in birds
were identified by Dr. Osscar Tovar S. Weather data near the equator. Caldasia 8:295-300.
were obtained through the courtesy of Sr. Santiago MILLER, A. H. 1959a. Reproductive cycles in an
Vallejo Espinosa, Chief, Departamento de Climatol- equatorial sparrow. Proc. Natl. Acad. Sci. 45:
ogia, Direcci6n General de Meteorologia, Lima. Dr. 1095-1100.
George J. Wallace kindly put at my disposal his manu- MILLER, A. H. I959b. Response to experimental
script on Z. capensis at Popayiin, Colombia. I wish to light increments by Andean Sparrows from an
thank especially Manuel Plenge and J. Murfree Butler equatorial area. Condor 61: 344-347.
for their continued help in many matters throughout MILLER, A. H. 1961. Molt cycles in equatorial An-
my stay in Per&. Accompanying illustrations were dean Sparrows. Condor 63: 143-161.
made by Gene M. Christman, and tissues were pre- MILLER, A. H. 1962. Bimodal occurrence of breed-
pared by Rod Jackson. This study was supported in ing in an equatorial sparrow. Proc. Natl. Acad.
part by NSF Grant GB-7917 to Pearson. Sci. 48:396-400.
MILLER, A. H. 1965. Capacity for photoperiodic
LITERATURE CITED response and endogenous factors in the reproduc-
tive cycles of an equatorial sparrow. Proc. Natl.
ASH~~OLE, N. P. 1963. Molt and breeding in popu- Acad. Sci. 54:97-101.
lations of the Sooty Tern, Sterna fuscata. Postilla, MILLER, A. II., AND V. D. MILLER. 1968. The
no. 76. behavioral ecology and breeding biology of the
BAILEY, R. E. 1952. The incubation patch of pas- Andean Sparrow, Zonotrichiu cupensis. Caldasia
serine birds. Condor 54: 121-136. 10:83-154.
BAKER, J. R. 1938. The relation between latitude MOREAU, R. E., A. L. WILK, AND W. ROWAN. 1947.
and breeding seasons in birds. Proc. Zool. Sot. The moult and gonad cycles of three species of
London 108 (ser. A) :557-582. birds at five degrees south of the equator. Proc.
Zool. Sot. London 117:345-364.
BARTHOLOMEW, G. A., JR. 1949. The effect of light
MORTON, M. L., J. R. KING, AND D. S. FARNER. 1969.
intensity and day length on reproduction in the
Postnuptial and postjuvenal molt in White-
English Sparrow. Bull. Mus. Comp. Zool. 101:
crowned Sparrows in central Alaska. Condor 71:
433-476.
376-385.
BLANCAS S~~NCHU, F. 1959. Comunidades y cam- PAYNE, R. B. 1969. Overlap of breeding and molt-
pos de vida de Acolla y sus alrededores. Mem. ing schedules in a collection of African birds.
Mus. Hist. Nat. “Javier Prado” (Lima), no. 7. Condor 71: 140-145.
146 JOHN DAVIS

PITELKA, F. A. 1957. Population controls in birds WOLF, L. L. 1969. Breeding and molting periods in
and other animals. Ecology 38: 176-177. a Costa Rican population of the Andean Sparrow.
PITELKA, F. A. 1958. Timing of molt in Steller Jays Condor 71:212-219.
of the Queen Charlotte Islands, British Columbia. WOLFSON, A. 1942. Regulation of spring migration
Condor-60:38-49. in juncos. Condor 441237-263. _ _
SNOW. D. VV.. AND B. K. SNOW. 1964. Breedina WOLFSON. A. 1952. The cloaca1 urotuberance. A
seasons and annual cycles of Trinidad land-bird; means for determining breeding condition in live
Zoologica 498:l-39. male passerines. Bird-Banding 23’: 159-165.
THREADGOLD, L. T. 1956. The anrual gonad cycle WOLFSON, A. 1954. Notes on the cloaca1 protuber-
of the male Jackdaw Corvus monedulus. Quan- ance, seminal vesicles, and a po’ssible copulatory
titative aspects. Cellule 58:45-54. organ in male passerine birds. Bull. Chicago
THREADGOLD, L. T. 1960. A study of the annual Acad. Sci. lO:l-23.
cycles of the House Sparrow at various latitudes.
Condor 62: 190-201. Accepted for publication 5 October 1970.