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UNIVERSITY OF KANSAS PUBLICATIONS

MUSEUM OF NATURAL HISTORY

Vol. 10, No. 2, pp. 45-75, 6 pls., 1 fig.

December 20, 1956


Comparative Breeding Behavior of Ammospiza caudacuta and A. maritima

BY

GLEN E. WOOLFENDEN

UNIVERSITY OF KANSAS
LAWRENCE
1956


UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY

Editors: E. Raymond Hall, Chairman, Henry S. Fitch,
Harrison B. Tordoff

Volume 10, No. 2, pp. 45-75, 6 pls., 1 fig.
Published December 20, 1956

UNIVERSITY OF KANSAS
Lawrence, Kansas

PRINTED BY
FERD VOILAND. JR., STATE PRINTER
TOPEKA, KANSAS
1956


Comparative Breeding Behavior of Ammospiza caudacuta and A. maritima

BY

GLEN E. WOOLFENDEN




Transcriber's Note: There are three symbols used in the caption for
Figure b in Plate 1 which are not available in the Latin-1 character
set. They have been noted as follows:

For the black triangle --> [triangle].
For the black dot --> [dot].
For the five pointed star --> [star].




CONTENTS


                                               PAGE

INTRODUCTION                                     48

MATERIALS AND METHODS                            48

DESCRIPTION OF THE AREA                          49

FLORA                                            50

REPTILES                                         50

MAMMALS                                          50

PREDATORS                                        50

PASSERINE ASSOCIATES                             51

WINTER STATUS AND SPRING MIGRATION               51

TERRITORY                                        52

VOICE                                            58
  Song                                           58
  Calls                                          60

COPULATION                                       61

NESTS                                            62

EGGS AND INCUBATION                              65

YOUNG                                            65
  Growth                                         65
  Behavior                                       68

FOOD, FEEDING AND CARE OF THE YOUNG              71

ACKNOWLEDGMENTS                                  73

SUMMARY                                          73

LITERATURE CITED                                 74




INTRODUCTION


Taxonomically the Genus _Ammospiza_ has received the attention of
ornithologists for more than a century. Nevertheless, the behavior of
no species of the genus has been studied extensively. The papers of
Montagna and Tomkins are the only works that mention behavior and
natural history in any detail. There has been an increasing awareness
of the importance of ethological data and of their usefulness in
systematics. For these reasons, I made a comparative study of the
breeding behavior of the Sharp-tailed Sparrow (_Ammospiza caudacuta_)
and the Seaside Sparrow (_Ammospiza maritima_) in New Jersey in the
spring and summer of 1955.

The Seaside Sparrow is restricted to the Gulf- and Atlantic-coasts
of North America, breeding north to Massachusetts. The Sharp-tailed
Sparrow breeds south to North Carolina. The overlap of the
breeding ranges of the two species is therefore small. Furthermore
the forms breeding in the coastal states are restricted to tidal
marshes, and the geographically peripheral colonies of each species
are small. Irregular nesting is the case for the northernmost
colonies of the Seaside Sparrow, on Cape Cod (Griscom, 1944:317),
and the same is probably true for the colonies of the Sharp-tailed
Sparrow on Pea Island, North Carolina, as indicated by Montagna's
failure to locate any breeding birds in July, 1941 (Montagna, 1942b:
256). The center of overlap of the ranges of the two species is in
New Jersey where both forms are abundant and can best be studied
comparatively.


    MATERIALS AND METHODS

    The adult sparrows were captured and banded, and sometimes the
    nestlings were banded. The standard funnel trap, baited with seeds,
    proved useless for capturing birds of the Genus _Ammospiza_,
    although migrant Savannah Sparrows (_Passerculus sandwichensis_)
    readily entered. A Japanese bird net, twenty-five feet long, was
    used successfully. Eighty-five Sharp-tailed Sparrows and forty
    Seaside Sparrows were banded at two localities.

    All of the adult sparrows were banded with United States Fish and
    Wildlife Service numbered bands and colored celluloid bands. The
    colored bands I used were obtained from the Hinton Supply Company
    of New York City, which manufactures them for cage birds. The firm
    makes them in seven colors, sold at reasonable prices. With seven
    colors, the number of combinations, using only one colored band and
    one aluminum band per bird, is forty-two.

    In addition, I dyed many adults and all nestlings. Alcoholic
    solutions of Victoria Blue B S concentrate and Alizarine Red S
    concentrate were used. The males were dyed red, the females blue;
    various areas of the body were colored in order further to
    individualize the birds. Although the dyes disappeared in less than
    a month, the markings were helpful on many occasions.

    When an adult bird was captured I always sexed it and ordinarily
    weighed and measured it. The nestlings were weighed and measured
    daily at intervals of 24 hours. I built a corral of hardware cloth
    around one Sharp-tailed Sparrow nest in order to measure the young
    after they left the nest. The sex of any adult was ascertained by
    examining the cloacal area, as described by Salt (1954:61-75). Sex
    as determined by this method was corroborated by internal
    examination of the specimens collected.

    A pan balance accurate to one-tenth of a gram was used for
    weighing. The adults were weighed in a cloth sack, the sack being
    weighed each time to prevent error owing to variable moisture and
    other factors.

    Dragging the marsh with a rope was ineffectual in finding nests.
    The birds flushed long before the rope neared them. I found nests
    of the sparrows by using a blind. From a blind I would determine
    the approximate location of a nest by watching the movements of the
    adult birds. Then I would either make a direct search of the
    vegetation or move the blind closer to find the actual site.

    Many hours were spent in blinds. I had two in operation throughout
    the breeding season, and it was from these that most of the data on
    behavior were accumulated. Observations were made by means of a 7 x
    50 coated binocular and on occasions by means of a 19.6x spotting
    telescope.


    DESCRIPTION OF THE AREA

    The intensive work was carried out on the marshes west of the town
    of Lavallette in Ocean County, New Jersey. Further observations
    were made at other localities in the county, in particular at the
    Chadwick marshes (plate 6), one mile north of the Lavallette site,
    where many of the Ammospizas were banded. The breeding Ammospizas
    of the localities are the nominate races, _A. c. caudacuta_
    (Gmelin) and _A. m. maritima_ (Wilson).

    Characteristic of the sand beaches of the Atlantic coast of the
    United States are offshore bars which, when exposed, form long bays
    parallel to the coastline. These bays become surrounded by marshes
    that in turn are inhabited by the two species of _Ammospiza_. The
    birds prefer the marshes closest to the ocean (plate 6). I made
    trips to the marshes on the mainland side of upper Barnegat Bay and
    found only a few Sharp-tailed Sparrows and no Seaside Sparrows in
    residence.

    The island of the Lavallette marshes that I worked on was
    approximately 1400 feet long and 600 feet wide. One-third of the
    east central end of the island was covered with sand fill, pumped
    there several years before the study was begun (plate 1, fig. b).
    The island was also ditched. The four east-west ditches are spaced
    125 feet apart; the two ditches perpendicular to these are 340 feet
    apart and are situated in the western portion of the island. These
    ditches, originally dug as a means of decreasing the mosquito
    population, are one foot wide and almost three feet deep. The
    excavated earth is deposited in a row paralleling the ditch. The
    entire island, excluding the sand fill is not more than two feet
    above normal high tide. In August, 1955, abnormally high water, a
    result of hurricane "Connie", rose four to five feet and covered
    all but the tops of the bushes and a few mounds of sand. Low tides
    expose no mud flats for the edges of the marsh are nearly vertical
    banks and the water along the edges is more than one foot deep.

    The average temperature for July, compiled over a 34 year period at
    the Asbury Park weather station is 72.6°F. The average
    precipitation from May through August, acquired over the same
    length of time, is between 3.5 and 4.5 inches per month.

    In spring and summer the prevailing winds are from the south and
    southwest. Therefore, the south and west shores of the island are
    subject to greater inundations by water. The fact that the island
    is unprotected by neighboring islands from the open expanse of the
    bay on this side is also of importance in this respect. The north
    and east shores, on the lee side of the island, are guarded from
    the open bay by nearby land. The exposed southern shores, where
    there was open mud and sparse patches of cord-grass, were the
    preferred feeding areas of the Seaside Sparrows. Lack of exposed
    and open feeding areas may account for the absence of this species
    in areas that otherwise seem to fulfill the requirements of the
    species.

    Two major drift lines were present on the island: one within a few
    feet of the waterline consisted mostly of dead eel grass (Zostera
    marina), and the other, situated close to the cattail strip,
    contained a variety of flotsam (pl. 2, fig. a).


    FLORA

    The vegetation on the island consisted chiefly of smooth cord-grass
    (_Spartina alterniflora_), black grass (_Juncus gerardi_), cattail
    (_Typha_ sp.), and marsh-elder (_Iva frutescens_). Other plants
    identified on the area were: common reed grass (_Phragmites
    communis_) and slender grass wort (_Salicornia europea_). Black
    grass grows on the inner, dryer portions of the marsh, and
    cord-grass prefers the wetter portions, growing to the edge of the
    water. The marsh-elder bushes mostly are restricted to the mounds
    of earth dug from the ditches. Cattails, in general, grow in a
    narrow band paralleling, but back a few yards from, the shoreline.
    Areas of mixed black grass and cord-grass occurred.


    REPTILES

    Diamond-backed terrapins (_Malaclemys terrapin_) were the only
    reptiles recorded from the study island. Several were taken on
    land, but the majority were seen in the waters about the marsh.

    On June 27 a black snake (_Coluber constrictor_) was seen in a
    bushy area bordering a marsh on the mainland side of Barnegat Bay.
    A few Sharp-tailed Sparrows were seen in the same locality and a
    singing male (G. E. W. 559) with testes 14 x 8 mm. and a female (G.
    E. W. 558) with a brood patch were collected.


    MAMMALS

    Only two species of mammals, both abundant, were present on the
    study island: the meadow vole (_Microtus pennsylvanicus_) and the
    muskrat (_Ondatra zibethicus_). The muskrats dug burrows beneath
    the level of the water into the banks of the island, used the
    ditches as routes to the interior of the marsh and built some small
    houses, mostly from cattail stems.


    PREDATORS

    Unless the above named mammals preyed on the sparrows, all of the
    enemies of the colony at Lavallette were avian. Both Crows (_Corvus
    brachyrhynchos_) and Fish Crows (_Corvus ossifragus_) visited the
    local marshes frequently as did a Marsh Hawk (_Circus cyaneus_). I
    watched the Marsh Hawk make many passes at what I thought were
    sparrows, but the only animal I ever saw caught by the hawk was a
    _Microtus_. The sparrows were alarmed when the hawk appeared,
    quickly and silently disappearing into the grass.

    At least two nests on the Lavallette Marsh were destroyed by
    predators in the course of the breeding season of 1955. One nest of
    the Seaside Sparrow was empty when I checked it on July 3; on July
    2 it had contained four young, three days old. On July 21 I found a
    dead Sharp-tailed Sparrow, approximately three days old, lying on a
    patch of matted grass. A hole was in the flank of the bird and
    blood was present about the bill. This nestling was not from a nest
    under observation.


    PASSERINE ASSOCIATES

    On the Lavallette marshes the only passerine associates of the two
    species of _Ammospiza_ were Song Sparrows (_Melospiza melodia
    atlantica_) and Long-billed Marsh Wrens (_Telmatodytes palustris
    palustris_). Two pairs of Song Sparrows and less than six pairs of
    marsh wrens nested on the study area. One Song Sparrow nest was
    found and is plotted on the map (pl. 1, fig. b); the other pair
    nested somewhere along the east shore of the island. The Song
    Sparrows at the east end of the island obtained most of their food
    from the grounds of the Lavallette Yacht Club across fifty yards of
    water to the east. The pair that nested in the western portion of
    the island fed along the sand fill or along the bases of the
    marsh-elder. Their nest was built in the most extensive area of
    these bushes; it was placed approximately one foot above the ground
    in a small dead bush and gained support and concealment from the
    surrounding black grass. Three of the four eggs hatched on June 30,
    and the young left the nest on July 11. Both parents fed the
    offspring.

    The marsh wrens fed and nested in the cattails. I never saw these
    wrens away from the cattails.


WINTER STATUS AND SPRING MIGRATION

Ocean County is ten miles south of the area treated in Cruickshank's
regional work (1942). He considers both species as rare to casual
winter residents. Concerning the spring migration of the Sharp-tailed
Sparrow he says (p. 456) "The first widespread wave never comes before
April 25, however, and most of the birds arrive in May." He mentions
that late May is the height of migration and that stragglers are
passing through up to the middle of June. The arrival of the Seaside
Sparrow in spring is similar (p. 458): the first widespread movement is
in early May, the peak is reached in the third week of the month, and
stragglers have been recorded through the second week in June.

I was in the field in Ocean County almost daily all spring and found no
Seaside Sparrows and only two Sharp-tailed Sparrows north of Barnegat
Inlet, Ocean County, before May 5. I waded through the marshes at
Chadwick, Lavallette, and Island Beach State Park on April 27 when high
tides covered all of the dense vegetation and saw no sparrows of the
Genus _Ammospiza_. If many had been present on this date I would have
seen them. On May 5 both species were plentiful on the Chadwick
marshes. Furthermore, the Seaside Sparrows were defending territories.
The absence of the two species the previous day indicates a large
nocturnal flight.

It was during the second and third weeks in May that the sparrows of
this genus were most abundant. In this period many unbanded Seaside
Sparrows were in the patches of cattails that were being defended by
the resident males from other territory-holders.

One _Ammospiza caudacuta subvirgata_ (G. E. W. 545) was taken in the
course of the study. It was a female (ovary: 7 x 5 mm.) weighing 15.3
grams ("moderate fat"), taken on June 8, 1955, on a marsh near the
mouth of the Manasquan River on the Monmouth-Ocean County line. This
marsh is decidedly less brackish than the Lavallette and Chadwick
marshes. The specimen was the only _Ammospiza_ seen there and was
probably a migrant, despite the late date; this subspecies is known to
occur late along the Atlantic Coast south of its breeding range.
Cruickshank (1942:454-455) considers the peak of spring migration for
this subspecies to be reached in late May.

To find _A. c. subvirgata_ in a marsh seemingly not saline enough for
the nominate race is not surprising. _A. c. subvirgata_ breeds in
marshes, along the coast of New England, which are almost fresh water
(Montagna, 1942b:256). _A. c. caudacuta_ is only casual away from salt
water.


TERRITORY

In a general treatise on the subject of territorialism, Nice (1933:98),
summarizing Howard, stated: "Territory implies in the male bird
isolation, advertisement, fixation, and intolerance." I concluded from
my observations that all four requirements are exhibited by male
Seaside Sparrows while none of them is well developed in male
Sharp-tailed Sparrows. This subject is discussed separately for the two
species.

Tomkins (1941:38-51) studied populations of _Ammospiza maritima
macgillivrayii_ near the mouth of the Savannah River in South Carolina
and Georgia and concluded that this form is not territorial. In support
of his conclusions, he quoted Nice (1933:90-91) as follows: "Territory
cannot mean just the nest spot when the adults feed in common; this may
be 'nest territory,' but it is a very different matter from a territory
in its strict sense to which parents confine themselves during the
breeding season. Again, the very essence of a territory lies in its
exclusiveness; if a bird's range is not defended, it is not a
territory."

The feeding and nesting grounds of breeding Seaside Sparrows are often
separated by a portion of the marsh which is not used by the birds.
This complicates study of the territorial habits of the species. It
does not mean, however, that the species is not territorial.

The birds studied by Tomkins had separate feeding and nesting grounds.
Concerning this, Tomkins (1941:43) states that "The Seaside Sparrows of
this locality [Savannah River area] often live where the two
requirements [adequate feeding grounds and suitable nesting cover] are
not always together or even meeting, but also where the feeding grounds
and the nesting place are separated by a short distance."

Six of the eight original pairs of breeding Seaside Sparrows of the
Lavallette colony fed in areas separate from those in which they
nested. I found the eight nests of the original residents and banded
and dyed all of the adults. The owners of two nests did not have
separate nesting and feeding areas. One nest was built within fifteen
feet of the south shore of the island, adjacent to the feeding area.
The other was built within a few feet of the north shore. The female of
this nest obtained food along the shore in the immediate vicinity of
the nest. Her mate was absent; in all probability it was the singing
male which I took from a nearby bush, before I found it advantageous to
use the island as a study area.

The remaining six pairs flew to the south or west shores of the island
in order to feed. None of these six nests was more than 100 yards from
the feeding grounds (pl. 1, fig. b).

It was comparatively easy to see that the males defended an area
surrounding the nest. It was more difficult to see that the pairs fed
on separate plots of shoreline, each defended by the male, but I am
convinced that this was the case.

The nest area was defended by the males through singing and chasing. I
saw no instances of a female entering into territorial disputes;
nevertheless, I did see a female chase a Sharp-tailed Sparrow away from
the vicinity of her nest.

Tomkins (1941:46) did not consider the song of _A. m. macgillivrayii_
to be "a declaration warning other birds away." After observing the
behavior of males of _A. m. maritima_ I am convinced that advertisement
of intolerance is the primary purpose of song in this species. An
account of the activities of a male Seaside Sparrow on May 6 on the
marsh at Chadwick demonstrates this point. In an hour (6:01-7:01 a.m.)
the bird sang 395 times, an average of 6.6 times per minute. He faced
his nearest singing competitor when singing, which in the course of
this hour was usually a male approximately 250 feet away across a
creek. The two competitors almost always alternated their songs and
frequently the singing of one seemed to stimulate the other bird to
sing. Although the song of the Seaside Sparrow is short and unmusical
it is loud, especially when compared with the song of the Sharp-tailed
Sparrow. Elevated perches such as the tallest cattail stems or isolated
bushes were used as singing and observation perches.

The chase of the Seaside Sparrow is not vigorous, but in all cases the
intruder was seen to give way to the defender. I saw no physical fights
between Seaside Sparrows. Chase by a defending bird was close to the
ground and directly toward the intruder. Sometimes the attacking male
emitted chipping notes when first sighting or flying towards his
adversary.

In the hour of observation mentioned above, no other Seaside Sparrows
entered the bird's territory, which consisted of a strip of cattail and
shoreline, 250 feet long and no more than 25 feet wide. At other times
Seaside Sparrows did enter this male's territory, and he drove them out
as soon as he saw them. Savannah and Swamp Sparrows, which for a few
weeks migrated through the area, were not chased, but Sharp-tailed
Sparrows were chased at times.

Several times I flushed a particular male Seaside Sparrow from the
northwest tip of the Lavallette study island so that it flew to the
island to the north. Seaside Sparrows of this north island immediately
made themselves conspicuous by chipping and then drove the non-resident
individual back to its own territory.

The first time I heard what is described below as the social call of
the Seaside Sparrow was on June 30 when an unbanded sparrow alighted in
a marsh-elder bush near a nest. The individual called twice as it came
near. The sound immediately aroused the owners of the nest and the male
flew directly toward the strange bird. The intruder quickly and
silently flew away.

My field notes refer to many other instances of territorial defense of
the nesting area; it seems superfluous to cite them here.

Additional proof of territorialism in Seaside Sparrows was obtained by
identifying and plotting the location of all the marked individuals,
which I saw each day while systematically traversing all the available
habitat on the island. Surprisingly, I did not once record a resident
Seaside Sparrow in what I considered another male's territory in the
month and a half (June 15-August 1) that I worked on the island at
Lavallette.

The fact that the adult Seaside Sparrows did not search for food
communally, or that different pairs did not utilize one particular area
at different times was most apparent when the pairs were feeding young.
From the blinds I first noted that the adults from any given nesting
territory always flew in the same direction towards the shore. After
moving a blind closer to the shore I noted that once an adult arrived
at the open or sparsely vegetated shoreline, that adult restricted
itself to a certain portion of the shoreline. These shoreline
territories were plotted on field maps and appear on the map in plate
1, figure b. One method used to ascertain the boundaries of these
shoreline feeding territories was a census taken from a boat. Many
times I circled the island in a skiff identifying the marked sparrows
as they appeared along the shore. The feeding sparrows were always
found in the same areas around the island. Straight lines can be drawn
between the nest sites and feeding areas of each pair of Seaside
Sparrows without having any lines cross (pl. 1, fig. b). These lines
correspond to the flyways used by each pair to go to and return from
the feeding area. I consider the area defended about the nest, the
segment of shoreline used by a pair of Seaside Sparrows and the
connecting flyway to constitute the territory of a male Seaside
Sparrow. If the flyways of any of the pairs had crossed, a situation of
mutual exclusiveness would not have existed and a territory could not
have been defined for the species.

It is generally agreed that territorial species engage in a minimum of
fighting. Song and display have been evolved to substitute for actual
combat which demands a greater amount of energy. Additionally, the mere
presence of an individual in an area previously established as its
territory probably serves to keep birds of nearby territories away. I
think that male Seaside Sparrows defend the feeding area and flyway as
a part of their territory by advertisement through use of these areas.
The birds at Lavallette rarely sang on the feeding grounds and I noted
only a few chases originating there. The sparrows rarely landed along
the flyways. The constant use of these areas probably served as
territorial defense, however. This supposition is supported by the fact
that feeding areas and flyways of different pairs were mutually
exclusive.

Nice (1941:457) divided territory into six categories. Type A (mating,
nesting, and feeding ground for young) is the type exhibited by the
Seaside Sparrow. The territory of a male Seaside Sparrow must contain
an area of open mud and/or sparse vegetation where food can be obtained
and also enough suitable cover to conceal the nest. I suspect in the
case of the few males studied on the marshes at Chadwick that the
territories the males established (strips of cattails and adjacent
shoreline) did not have suitable nesting cover, because these males
were unmated on June 15 when I left this study area because of human
interference. Suitable nesting cover and feeding areas were separated
by short distances of unusable marsh for most of the sparrows on the
Lavallette study area. This fact caused the adults to commute from one
site to the other. Photographs of shoreline habitat suitable for
feeding by Seaside Sparrows appear in plate 2.

The area defended about the nest tended to follow the rows of
marsh-elder bushes (pl. 3, fig. a), probably because these bushes
supplied suitable song and observation perches. The segments of
shoreline used by each pair were less than 75 yards in length and
scarcely 20 feet wide. I never recorded Seaside Sparrows foraging in
the interior of the marsh.

Sharp-tailed Sparrows were more abundant than Seaside Sparrows on the
marshes at Chadwick and Lavallette. Sharp-tailed Sparrows were the more
difficult to net because of the peculiar organization of the colonies.
This organization, described below, also made nests of that species the
more difficult to find. Only intensive netting at both localities
produced enough marked individuals for me to study the breeding
behavior of the species.

At Chadwick, where I netted most of the 85 Sharp-tailed Sparrows that I
banded, my efforts were concentrated on one segment of the marsh.
Marking made it evident that the males were not territorial, although
they did confine themselves to what might appropriately be called a
breeding home range, the area to which an individual confines itself in
the course of one nesting attempt. Observations of marked birds also
indicated that there was considerable overlap of the breeding home
ranges of individual males.

I recorded a few marked Sharp-tailed Sparrows often enough and over a
long enough period (more than one month) to gain a good idea of the
size of the breeding home range of the males, which I estimate to be
three to four acres. This estimate was made at Chadwick, where large
areas of suitable uniform habitat occur. Females are more secretive
than males, but seem to restrict themselves to areas considerably
smaller than those of the males. My observations of two females that
were feeding young indicated that each female restricted herself to an
area of less than one acre. Female Sharp-tailed Sparrows possibly are
territorial, although I recorded no disputes that would substantiate
this possibility.

If I am correct in my estimates of size of breeding home range in
Sharp-tailed Sparrows (males, three to four acres; females,
approximately one acre), certain observations made by Montagna and me
are readily explainable.

My netting operations indicated a surplus of male Sharp-tailed Sparrows
in a given area. At Chadwick, I netted as many Sharp-tailed Sparrows as
I could, without regard to sex. Here I captured 39 males and 16 females
(six individuals remained unsexed). On the Lavallette study island,
netting was more selective; here I attempted to net the females of the
nests I found. The sex ratio at Lavallette was 15 males to eight
females (one juvenile was not sexed). Three of the eight females were
netted at their nests.

Montagna (1940:196) decided from collecting and observations that male
Sharp-tailed Sparrows either outnumbered the females or were
polygamous. The results I obtained from netting seemed to indicate a
surplus of males. Banding, however, showed that in the breeding season
males range over a larger area than do females. With this knowledge,
the discrepancy between the number of males and females captured is
explainable without an unbalanced sex ratio. If the males range over an
area four times as large as that of the females, theoretically, four
times as many males should be caught at every placement of the net
provided the net remained in place long enough to capture all the birds
using the area. In practice, this is essentially what occurred.

Other behaviorisms of this species indicate that it is not territorial.
The song of the male is not loud and does not seem to be an
advertisement to other birds. In fact, the song of this species is so
quiet and lengthy when compared to that of the Seaside Sparrow that I
at first thought I was hearing "whisper" or "practice" songs. These
qualities of the song seem to indicate that the "advertising" function
of song of territorial species is lacking or unimportant in
Sharp-tailed Sparrows.

I suspect that male Sharp-tailed Sparrows do not even know where nests
are. On July 18 at 7:00 a.m. I was watching a nest from a nearby blind
when an unbanded male (I saw the individual sing later) appeared. As
the bird foraged through the black grass, it headed directly toward the
nest. When the male was almost one foot from the nest the incubating
female left. She ran from the tussock and flew a short distance away to
a cattail stem. From here she watched the male, which seemingly
oblivious continued foraging, coming within inches of the nest. As the
male walked away from the nest the female returned. At 8:00 p.m. the
same day I was in the blind again. The female was out searching for
food when a different, banded male appeared. In his foraging, the male
walked up on the grass stems over the nest. The male apparently saw the
young (two had hatched on July 17 and one on July 18) for he turned his
head and seemed to peer down under the stems. The female appeared (with
food) as he was doing this; she flew directly toward him and he flew
away. The male was not seen near the nest in later observations.

On July 1 (6:50 a.m.) I was in a blind near another nest as the female
approached with food for the young. At this moment a male appeared and
the female immediately flew away. The male perched on a tussock within
two feet of the nest, sang, and then flew off. The female reappeared in
a few seconds without the food. She searched through a clump of black
grass four feet from the nest, caught a small, pale green insect, fed
it to her one young (there were also two eggs in the nest) and began
brooding.


VOICE


_Song_

Only males of the two species sing. The normal song of the Seaside
Sparrow lasts just under two seconds, the buzzing final note
constituting three-quarters of the song. Saunders (1951:257-258)
describes this song as short, and buzzlike, beginning with two or three
short, rather faint notes and ending in a trill at first loud but
fading away toward the end. The introductory notes are followed by a
higher-pitched, loud, strongly accented, but buzzy note. This note is
usually higher than the final trill and connected with it. The song has
been written as _tup tup ZEE reeeeeeeeee_ and _tup TEE tle reeeeeeeeeee_
(Saunders, 1951:257), _cutcut_, _zhé-eeeeeeeee_ (Peterson, 1947:232)
and _che-zheéeege_, _che-zhée_, _che-wéege_, _chur-zhée_ and
_too-szheée_ (Stone, 1937:910). My field notes contain the following:
_CHUR-er eeeee_, _CHUR eeeeee_ and _oka-CHE weeeee_. These variations
in the phonetic representation of the songs are attributable mostly to
the birds. Not only is there variation among individuals, but also
individuals vary their songs. Birds that I heard giving a characteristic
song suddenly sang a different type for awhile, and then reverted to
the original. The bill is elevated and opens considerably with each
note; the head bobs with the loud note. Typical singing postures are
shown by Tomkins (1941: pl. 3).

The song of the Sharp-tailed Sparrow, as described by Saunders
(1951:256-257), is short and insectlike, introduced by one or two short
notes; the remainder is a somewhat wheezy trill, growing fainter
towards the end. Sometimes there are two trills on different pitches,
and occasionally a final short, low note. The quality is as though the
sound _sh_ ran through all but the introductory notes. Saunders writes
these trills as: _tsup tsup shreeeeeeeee_ and _tip tish eeeeee
shaaaaaaay_. The bill is opened slightly with each note, as I saw when
I watched a singing bird with the sun directly behind it. Montagna
(1942a:116) noted that _A. c. caudacuta_ sang less often than the more
northern _A. c. subvirgata_.

Both species have specialized flight songs, but in the birds that I
studied these songs were infrequent and seemingly unimportant. The
flight song of the Seaside Sparrows consisted of a double version of
the normal song. Although I heard it only a few times, the flight song
of the Sharp-tailed Sparrow seemed slightly louder than the normal
song. This song is given by both species as the bird flutters upward
ten or 20 feet and glides back down.

Singing begins at daylight and decreases at 9 or 10:00 a.m. when the
temperature rises. On cloudy days singing seemed to last longer.
Towards dusk singing again increases, but not to the frequency of the
morning peak.

The major differences between the songs of the two species are in
loudness, length, and frequency. The fact that the Seaside Sparrow
sings louder than the Sharp-tailed Sparrow is mentioned by Stone
(1937:906). On windless days I heard singing Seaside Sparrows more than
200 yards away; Sharp-tailed Sparrows were inaudible at distances of
more than 40 yards. The song of a Seaside Sparrow is rarely longer than
two seconds; the song of a Sharp-tailed Sparrow usually lasts for
almost 20 seconds and consists of a variable number of phrases like
those described by Saunders. A Seaside Sparrow that I watched for one
hour sang 395 times or 6.6 times per minute. I doubt that any of the
Sharp-tailed Sparrows sang more than 20 times per hour, although I made
no comparable count.

Additionally, Seaside Sparrows sing from exposed perches such as tall
cattail stems and tall or isolated marsh-elder bushes. Sharp-tailed
Sparrows do not often use conspicuous perches for singing. They sing
while on the ground or when in flight. They do use exposed perches as
observation posts and occasionally sing from them.

Seaside Sparrows often face their nearest neighbor when singing and
alternate songs with this bird. The one time Sharp-tailed Sparrows
almost always sing is when they are involved in fighting. In such a
case the several birds sing simultaneously.

Seaside Sparrows began singing the morning after their nocturnal
arrival. For resident birds, singing is at its maximum at this time and
is maintained at a high level throughout incubation. At hatching of the
eggs, singing declines sharply; males then are busy aiding in care of
the young. Males that have successfully reared a brood rarely sing
after the young leave the nest.

Sharp-tailed Sparrows sang infrequently when they first arrived, and
singing did not reach its peak until late May. By August singing had
almost ceased in this species.

Song of the Seaside Sparrow functions importantly in the establishment
and maintenance of its territory. Newly-arrived males sing vigorously.
In the Sharp-tailed Sparrow I think song is merely an expression of
sexual excitement because song does not reach maximum frequency until
the females arrive and become receptive to the males.

Differences in song correspond to differences in territorial behavior.
The distinct, loud song, sung often and from exposed perches, which is
frequently alternated with that of the nearest competitor, is given by
the Seaside Sparrow, a territorial species. The indistinct, quiet song,
sung infrequently and often from unexposed places belongs to the
Sharp-tailed Sparrow, a non-territorial species.


_Calls_

Seaside Sparrows give a soft, lisping call note, probably the one
referred to by Saunders as a squeaky _tseep_ (1951:258), that functions
as a social call. When migrants were numerous on the marshes at
Chadwick I heard this note often. At Lavallette I did not hear it
until June 30 (work began there on June 16) and then it was from an
unbanded, non-resident bird. In late July and in August the number of
non-resident sparrows increased and the social call was heard often. I
never heard a resident bird give this call. On December 29, 1955, on a
marsh at the mouth of the Manasquan River on the Monmouth-Ocean County
line, a group of wintering Seaside Sparrows frequently used this call.
I do not know whether the Sharp-tailed Sparrow has a comparable call.

Both species emit alarm notes. Although variable, the Seaside Sparrow
has two general types. One, recorded by me as a short _chip_ or _tick_
was given by both sexes whenever I was near a nest. The other type, a
high, sharp _tsip_, is indicative of a higher degree of excitement.
When I captured young already out of the nest, or when I investigated
nests containing young old enough to depart, the adults gave this call.
The tail is jerked downward each time this note is given.

The alarm call of the Sharp-tailed Sparrow is not so loud as that of
the Seaside Sparrow and it is not given so often. I described it as a
short _tsick_ or _tsuck_. Females emitted such calls when I was at
their nests or when male Sharp-tailed Sparrows came near their nests.
Males may have a similar call, but I never recorded it. Montagna
(1942a:116) remarks on the quietness of this species. This is
especially evident when one compares Sharp-tailed Sparrows with Seaside
Sparrows.


PLATE 1

[Illustration: FIG. _a_. An aerial photograph of the Lavallette study
island. One inch equals approximately 375 feet. The area covered by
sand has been extended since this photograph was taken. This is
indicated in figure _b_ of this plate.]

[Illustration: FIG. _b_. Map of the Lavallette study island. All
fringillid nests that I found are indicated and the territorial
boundaries of the Seaside Sparrows are shown.

  [triangle]--Seaside Sparrow
  [dot]--Sharp-tailed Sparrow
  [star]--Song Sparrow]


PLATE 2

[Illustration: FIG. _a_. The south shore of the Lavallette study island
showing the two major driftlines and the sparsely vegetated areas. This
is the feeding habitat of the Seaside Sparrow.]

[Illustration: FIG. _b_. A close-up view of a segment of the shoreline.
Note the spacing of the clumps of cord-grass (_Spartina alterniflora_).
A six inch ruler propped against a 12 inch stick is included to
indicate the size and spacing of the plants.]


PLATE 3

[Illustration: FIG. _a_. The inner portion of the marsh on the
Lavallette study island showing the rows of marsh-elder bushes (_Iva
frutescens_) and the extensive areas of black grass (_Juncus gerardi_).
Areas of mixed black grass and cord-grass appear in the foreground. All
the nests of Sharp-tailed Sparrows were found in the areas of black
grass. Four of the eight nests of Seaside Sparrows were in the
marsh-elder. One of the blinds that I used is shown in this
photograph.]

[Illustration: FIG. _b_. A mated, banded pair of Seaside Sparrows in a
dead marsh-elder bush near their nest. Note the abdomen of a moth
protruding from the bill of the female on the right.]


PLATE 4

[Illustration: FIG. _a_. A female Sharp-tailed Sparrow at the entrance
to her nest. The throat on this bird is dark because of dye applied by
me.]

[Illustration: FIG. _b_. The nest of a Sharp-tailed Sparrow viewed from
above. Stems of black grass were parted to take the picture. The outer
rim of this nest (lower right) is made of living stems of black grass.]


COPULATION

In late June at the Lavallette area there was an influx of unbanded
Seaside Sparrows. Certain of these new arrivals established territories
in areas unoccupied by the remaining original residents. These new
residents were birds that probably had unsuccessful nestings elsewhere.
Because of tropical storms that almost covered the island with water in
August, I doubt that any of these late nestings were successful. On
July 7 at 8:30 a.m., while watching a pair of these new arrivals, I
recorded my only observation of copulation in the Seaside Sparrow. The
female seemed to be searching for a nest site when copulation occurred.
The female crawled about in a marsh-elder bush seemingly testing the
various forks in the branches for size. The male followed her,
remaining a few inches above and behind. Several times the two birds
disappeared in the lower branches and were hidden by the surrounding
black grass. Finally, while the female squatted on a branch the male
mounted. He fluttered his wings before mounting and continued to do so
as coition took place.

I began observations at Lavalette on June 16, too late to observe
copulation of the early residents. All the nests contained eggs by that
time. At Chadwick, pair formation seemingly never occurred, at least
with the males I was studying. The territories established by males at
Chadwick contained few marsh-elder bushes. Possibly females, finding no
suitable nest sites, refused to accept these territories.

Copulation in the Sharp-tailed Sparrow was observed several times. It
occurs most frequently in the course of, or immediately following, a
fight between several males. I do not know what instigates the
gathering of several males into these groups; it may be a certain
behaviorism of a female, or possibly, merely the appearance of a
female. Montagna (1942a:117) was convinced that females of _A. c.
subvirgata_ were present in these fights. On the other hand, in two
instances with _A. c. diversa_ where he collected all the birds in the
group, no females were present. Twice, at Chadwick, my observations
indicated that females of _A. c. caudacuta_ were not always involved in
these groups. In these instances all the birds in the group had
previously been banded and diagnosed as males. Possibly a female was
the original stimulus of these groups, and she may have disappeared
while the males were fighting with each other. I found it difficult to
distinguish fighting males from a copulating pair. On June 3, however,
a banded pair was observed. Copulation occurred on the ground. The male
fluttered his wings as he mounted and the female remained motionless.
Copulation lasted approximately three seconds; immediately thereafter
the male flew to a nearby cattail stem and the female climbed a tussock
of grass and chipped quietly. This same female was seen to copulate
with other males, and males were observed copulating with several
females.

_A. m. maritima_ is monogamous, the pair-bond being maintained
throughout a breeding cycle. _A. c. caudacuta_ is promiscuous,
relations between the sexes being limited to copulation. For _A. c.
subvirgata_ a relationship other than promiscuity has been intimated
(Lewis, 1920:587-589). Concerning observations of the nest he found at
Yarmouth, Nova Scotia, Lewis wrote: "The nest was found after I had
quietly watched the parent Sparrows for about an hour, while they were
bringing food to their young.... The male sang from time to time from a
piece of driftwood on the marsh about 30 feet distant from the nest.
When I was examining the nest and the young birds, the parents made no
demonstration for some minutes, but later they came near and uttered
chip's, much like those of Savannah Sparrows."


NESTS

I found the nests of all eight pairs of Seaside Sparrows which nested
on the Lavallette study island in 1955. Four nests were supported by
marsh-elder bushes, three of which were dead. These nests were placed
low enough to be hidden by numerous stems of black grass, as were the
other four nests. Of the remaining four nests, three were placed in
tussocks of black grass and the fourth one gained support mostly from
cord-grass stems. The eight nests ranged from 9 to 11 inches (9.6 inch
average) from the rim to the ground, the four nests in the bushes being
the highest. The outside diameters of the nests ranged from 3 to 4.5
inches (3.9 inch average) and the outside depth varied between 2 and
3.5 inches (2.7 inch average). Seven of the nests had an inside depth
of 1.5 inches; the other one was only an inch from the rim to the
floor. The inside diameter of the cup varied between 2 and 2.5 inches.

As mentioned above all eight nests were shielded by stems of black
grass. Stems were not woven over the nests by the birds; rather it was
the choice of the nest sites that resulted in the concealment. The only
plant used for nest material was black grass.

In all cases the black grass limited the directions from which the
nests could be entered. Six of the nests were approached from a
direction varying between northeast and southeast. The prevailing winds
of spring and summer are from the south and southwest; the black grass
consequently leans in the opposite direction. The remaining two nests
were entered from the northwest. These were nests built in marsh-elder
bushes where the grass stems were held upright by the branches of the
bushes.

One nest, built in a small dead marsh-elder bush, was tilted by the
growth of stems of black grass which were used for support on one side.
This tilting did not cause the contents to spill, but, I judged, did
cause the adults to desert the nest.

Seven nests of the Sharp-tailed Sparrow were found; two of these were
old nests. Four of the five nests used in the breeding season of 1955
were found on the Lavallette marsh study area, the other one I
discovered on the Chadwick marshes. Two young Sharp-tailed Sparrows
that I saw at Lavallette were not from nests I found, nor were they
from the same nest. Therefore, a minimum of six Sharp-tailed Sparrows
bred on the Lavallette island. Measurements were taken of only the five
nests that were used in 1955. The Sharp-tailed Sparrow builds its nest
closer to the ground than does the Seaside Sparrow. The five nests were
five to six inches off the ground; the two nests of a previous year
appeared to have been no higher. The Sharp-tailed Sparrow nests were
built in areas where black grass was the predominant plant, and the
nests were constructed entirely from this grass. The outside diameters
varied from 3 to 4.25 inches (3.4 inch average). The outside depth of
the nests varied from 2 to 3.5 inches (2.8 inch average). The inside
depth was 1.5 inches in all nests and the inside diameter ranged from 2
to 2.5 inches (2.1 inch average).

Harrison F. Lewis (1920:587) studied a nest of _A. c. subvirgata_ in a
small salt marsh near Bunker's Island at the southern end of Yarmouth
Harbor, Yarmouth, Nova Scotia, which he found on June 12, 1920. For
details of this nest I quote Dr. Lewis. "The nest proper was a neat,
round cup of fine, dry, dead grass, with some horsehair in the lining.
Its foundation consisted of some small masses of 'eel-grass' and roots.
Its dimensions were: inside diameter, 2.5 in.; outside diameter, 4.5
in.; inside depth, 1.5 in.; outside depth 2.375 in. It was elevated
above the general surface of the marsh by being placed on the top of a
low, grassy ridge, about fourteen inches high, formed from material
thrown up when a ditch was dug across the marsh, many years before.
During some storm a mat of dead 'eel-grass' had been left on top of
this ridge, and this had later been lifted by the growing marsh grass,
leaving several inches between it and the ground. The nest was placed
on the northwest edge of this mat, about half of the nest being under
it, while the other side was sheltered and concealed by grass about six
inches high. The nest was not sunk in the ground at all."

Two of the nests found were entered from the north-northeast, the other
three from the east-southeast. All five nests were sheltered above by
stems of black grass. Three of the nests were beneath a layer of dead
black grass where a clump of erect living stems parted the mat. One
nest (pl. 4, fig. a) was situated where cattail stubs held the black
grass somewhat erect. Green stalks as well as dead stalks were woven
into a canopy over this nest. Another nest was constructed on a mat of
black grass under and among numerous horizontal living stems, some of
which were woven into the outer lining of the nest (pl. 4, fig. b).

Nests of both species were found in tussocks of black grass. The
locations of these sites differed. The Sharp-tailed Sparrow prefers the
higher and therefore dryer portions of the marsh where black grass is
the characteristic plant. Contrastingly the Seaside Sparrow almost
always chooses the wetter portions of the marsh (Cruickshank, 1942:45;
Forbush and May, 1939:514; Stone, 1937:906; personal observations)
where several species of plants are abundant. In areas that have been
ditched, as have almost all marshes in New Jersey, the mound of
excavated muck is ideal for the growth of marsh-elder. Rows of these
bushes are present on many of the marshes of New Jersey (pl. 3, fig.
a). The location of four of the eight Seaside Sparrow nests in these
"hedgerows" indicates that they provide suitable, if not preferred,
sites for the species.


EGGS AND INCUBATION

I found no nests of either species before they contained a complete
complement of eggs and therefore was unable to ascertain the incubation
period for these species. Brood patches were evident on female
Sharp-tailed Sparrows by June 1, probably indicating that laying began
near this date. Cruickshank (1942:456) lists egg dates of the
Sharp-tailed Sparrow as concentrated in early June, with extremes of
May 19 and August 4. He thinks the species probably has two broods. For
the Seaside Sparrow, Cruickshank (1942:458) states there is probably
but one brood and that egg dates are concentrated in early June, with
extremes May 23 and July 2. Stone (1937:907, 911) considers four eggs a
normal clutch for both species, but cites instances where three and
five eggs were thought to be complete sets. Four of the eight Seaside
Sparrow nests I found contained at least three eggs, and four contained
at least four eggs. Four of the five Sharp-tailed Sparrow nests I found
contained at least three eggs and one contained four eggs.

Female Seaside Sparrows do all of the incubation. The male, while the
female is on the nest, remains a short distance away. He sings often
and gives alarm notes when there is a local disturbance. These chipping
notes bring the female off the nest, and then they both chip at the
intruder. The male accompanies the female to the feeding grounds and
normally they return together.

As previously mentioned, male Sharp-tailed Sparrows take no part in the
nesting activities.


YOUNG

I studied growth and changes in behavior of the young. Since I could
see no behavioral differences between the nestlings of the two species,
this subject will be discussed jointly for the two forms.


_Growth_

The color of the natal downs of both species is similar. Dwight
(1900:190), who saw newly hatched nestlings only of the Sharp-tailed
Sparrow, described the color as grayish wood-brown. A series of white
neossoptiles is present at the posterior end of the ventral tract in
both species. These feathers are more numerous in the Seaside Sparrow.
Dwight (1900:98) saw no neossoptiles on the underparts of any of the
passerines he examined. Seaside Sparrows have a mid-dorsal row of downs
in the dorsal tract near the uropygium. These feathers are lacking in
the Sharp-tailed Sparrow and constitute the major difference, in this
plumage, between the two species. The neossoptiles of three Seaside
Sparrows and one Sharp-tailed Sparrow were counted. These counts were
checked on the young birds studied in the field. The number and
placement of these feathers appear in plate five. There appears to be a
consistently greater number of natal downs in Seaside Sparrows, when
compared with Sharp-tailed Sparrows.


TABLE 1--DAILY WEIGHT IN GRAMS OF NESTLING SEASIDE SPARROWS AND
SHARP-TAILED SPARROWS FROM LAVALLETTE, OCEAN CO., NEW JERSEY.

  ======+================================================================+
    Day |                   _Ammospiza maritima_               | Average |
  ------+---------+------+---------+------+------+------+------+---------+
   0    |     ... |  2.2 |     2.3 |  ... |  ... |  2.2 |  1.8 |    2.1  |
   1    |     3.0 |  3.3 |     3.5 |  3.7 |  3.3 |  2.6 |  2.1 |    3.1  |
   2    |     4.6 |  5.0 |     5.2 |  5.0 |  4.8 |  3.7 |  3.0 |    4.5  |
   3    |     7.0 |  5.5 |     7.2 |  6.9 |  6.9 |  5.9 |  4.7 |    6.3  |
   4    |     9.4 |  8.1 |    10.6 |  9.1 |  9.1 |  7.6 |  6.4 |    8.6  |
   5    |    12.5 | 11.1 |    12.3 | 11.4 | 11.1 |  9.9 |  8.7 |   11.0  |
   6    |    14.6 | 13.1 | [1]11.1 | 13.9 | 13.7 | 12.6 |  9.6 |   13.0  |
   7    | [1]11.6 | 13.9 |    12.1 | 15.1 | 14.8 | 14.3 | 11.8 |   13.7  |
   8    |    14.9 | 15.5 |    13.4 | 14.9 | 14.8 | 14.6 | 12.4 |   14.4  |
   9    |    15.2 | 15.8 |    13.8 | 16.2 | 16.1 | 16.0 | 14.4 |   15.4  |
  10    |     ... |  ... |     ... |  ... | 15.9 | 15.5 | 14.3 |   15.2  |
  ------+---------+------+---------+------+------+------+------+---------+

  ------+---------+------+---------+------+------+---------+
    Day |         _Ammospiza caudacuta_          | Average |
  ------+---------+------+---------+------+------+---------+
   0    |   1.6   |  ... |    1.8  |  1.7 |  1.7 |    1.7  |
   1    |   1.8   |  2.3 |    2.5  |  3.0 |  2.1 |    2.3  |
   2    |   3.3   |  2.7 |    4.1  |  4.6 |  3.1 |    3.6  |
   3    |   5.0   |  3.7 |    5.9  |  6.4 |  4.7 |    5.1  |
   4    |   6.8   |  5.4 |    8.4  |  9.1 |  6.7 |    7.3  |
   5    |   8.6   |  6.9 |   10.7  | 11.2 |  9.5 |    9.4  |
   6    |  10.2   |  8.9 |   12.8  | 13.0 | 10.9 |   11.2  |
    7   |  12.1   | 11.4 |   14.5  | 13.6 | 12.3 |   12.8  |
   8    |  13.5   | 12.9 |   15.3  | 14.5 | 13.3 |   13.9  |
   9    |  12.2   | 13.4 |   15.9  | 14.9 | 13.6 |   14.0  |
  10    |  12.7   | 14.0 |   15.5  | 15.0 | 14.0 |   14.2  |
  11    |   ...   |  ... |   ...   | 15.1 | 14.4 |    ...  |
  ------+---------+------+---------+------+------+---------+

      [1] These weights are not figured in the averages; see text.

Seven nestling Seaside Sparrows and five nestling Sharp-tailed Sparrows
were weighed at 24-hour intervals until they left their nests. The
birds were weighed in early morning before they had received much food.
Weights of these individuals, and daily averages for each species are
shown in Table 1. The weights in the zero column were of nestlings that
had not been fed. The weight of one hatchling (1.9 gm.), which does not
appear in the table, is included in the average for the zero column.
Two young Seaside Sparrows, approximately a week old, fell out of a
nest between 9:30 a.m. July 6 and 5:30 a.m. July 7. When I found them
below the nest, at the latter time, their temperatures were far below
normal, and they had lost a considerable amount of weight. These
abnormally low weights were not figured in the averages. The weights of
Sharp-tailed Sparrows 11 days old were obtained by confining the birds
to the vicinity of the nest with a screen.


[Illustration: FIG. 1. The development of the young of Seaside Sparrows
(solid line) and Sharp-tailed Sparrows (dotted line) as evidenced by
four linear measurements taken at 24 hour intervals.]


At hatching and throughout nestling life and post nestling life Seaside
Sparrows average heavier than Sharp-tailed Sparrows of comparable age
(Table 1). Weights of adults of the two species that were netted or
collected between May 6 and June 27, 1955, within two miles of
Chadwick, Ocean County, New Jersey, follow: Fourteen males of
_Ammospiza maritima_ averaged 24.2 gm. (21.9-27.4 gm.); three females
averaged 22.3 gm. (19.8-24.4 gm.). Thirty-three males of _A.
caudacuta_ averaged 20.7 gm. (18.0-23.1, 25.8 gm.); 14 females
averaged 17.8 gm. (15.3-19.0 gm.), 2.9 gm. less than the males. One
female Sharp-tailed Sparrow, weighing 23.1 gm., was not included in the
averages because it had an egg with shell in the oviduct.

Montagna (1940:195-196) weighed a series of breeding Sharp-tailed
Sparrows (21 males; 5 females) from Popham Beach, Maine, and found the
males to be only 0.2 gm. heavier than the females, but he stated that
the small number of females weighed, and the high percentage of these
that contained eggs, probably lessened the difference in weight found
at other seasons.

The four linear measurements that I took on the same series of adults
confirmed the size difference: Seaside Sparrows average larger than
Sharp-tailed Sparrows, and males average larger than females in both
species. The average and range for each measurement taken on the
sparrows is presented in Table 2.

Four linear measurements were also taken daily on the young sparrows. A
summary of these data appears in Figure 1.


_Behavior_

The first indication of hatching is a crack in the side of the egg
along the line of greatest circumference. The crack is extended along
this line by the egg tooth, and then contraction of muscles of the neck
by the embryo separates the shell into two pieces. Extension of the
legs frees the bird from the shell. I held the eggs of two Seaside
Sparrows in my hand and watched this procedure. In each instance the
young bird defecated in the shell before freeing itself. A barely
audible "peep" note was heard from one hatchling Sharp-tailed Sparrow
when I held it near my ear. When free from the shell, the young birds
rest on their tarsi, abdomen and forehead; their down dries in a few
minutes, and their skin becomes noticeably darker. One sparrow gaped
five minutes after hatching and all the young gaped later the same day.
The abdomen of the young becomes distended when they are fed by the
parents.


[Illustration: PLATE 5

_Ammospiza caudacuta_

_Ammospiza maritima_

Drawings of the nestlings of the two species of _Ammospiza_
approximately three days of age showing the variation in the amount and
placement of the neossoptiles in the two species. Abbreviations for
feather tracts in which downs were found: ca, capital; h, humeral; a,
alar; d, d´, dorsal; cr, crural; v, ventral.]


[Illustration: PLATE 6

An aerial view of the marshes at Chadwick (upper left) and Lavallette
(lower left). The Atlantic Ocean appears in the upper right of this
photograph.]


TABLE 2--LINEAR MEASUREMENTS IN MILLIMETERS OF ADULT SEASIDE SPARROWS
AND SHARP-TAILED SPARROWS CAPTURED OR COLLECTED WITHIN TWO MILES OF
CHADWICK, OCEAN COUNTY, NEW JERSEY, BETWEEN MAY 6 AND JUNE 27, 1955.

  =========================================================
                    _Ammospiza maritima_
  --------------+-------------------+---------------------+
                |     14 males      |      3 females      |
  --------------+-------------------+---------------------+
  wing (chord)  |  64.14 (60-66)    |  58.33 (58-59)      |
  tail          |  55.28 (54-59)    |  51.00 (49-53)      |
  tarsus        |  23.00 (22-25)    |  22.17 (21-23)      |
  culmen        |  15.18 (15-15.5)  |  14.50 (13.5-15.0)  |
  ---------------------------------------------------------
                    _Ammospiza caudacuta_
  --------------+-------------------+---------------------+
                |     33 males      |     15 females      |
  --------------+-------------------+---------------------+
  wing (chord)  |  58.79 (55-61)    |  55.67 (54-58)      |
  tail          |  49.48 (46-53)    |  46.93 (45-50)      |
  tarsus        |  20.91 (20-22)    |  20.30 (20-21)      |
  culmen        |  13.67 (13-14)    |  13.23 (12.5-14.0)  |
  --------------+-------------------+---------------------+

In the first 24-hour period after hatching the soft "peep" note is
heard frequently. The young are better able to right themselves, and
many feather papillae show distinctly through the skin.

On the second day young are capable of moving short distances by using
their wings and feet. A thick ridge of tissue forms over the eyeball
where the eyelids later delaminate. The call is now a double version of
the "peep" note described above.

When the young are three days old the eyelids open, but only slightly.
In the next three days the young become better co-ordinated and the
eyes open fully. The egg tooth was last seen on a young bird on the
sixth day. All incoming feathers remain sheathed until the seventh day.

On the seventh day young show the first signs of cowering. Previously,
they all begged when I came to the nest. The remiges remain sheathed,
but the body feathers emerge from the tips of the sheaths. A quiet
reedy call replaces the "peep" note. A quiet, but squealing distress
call was also first noted on the seventh day, when the young were
handled.

On the eighth day the remigial sheaths become gray (previously they
were dark blue) and begin to slough off. When removed from the nest,
the young attempt to escape. Begging is less frequent and cowering is
the predominant attitude towards intruders.

The first young of both species left the nest on the ninth day. It must
be remembered, however, that this remark, and succeeding remarks,
concerning departure of young from nests pertains to young that were
disturbed daily by me. The others climbed to the edge of the nest when
they were left alone, but remained in the nest when they were all
replaced. Gaping was recorded once on the ninth day. Stuart W. John
watched two Sharp-tailed Sparrows on my study area leave a nest. They
climbed out and immediately hid in a tussock of grass a few inches
behind the nest.

On the tenth day when I parted the branches over one Seaside Sparrow
nest, the four young jumped from the nest and scattered in the grass.
One of these birds gave a chipping note similar to the distress call of
adults. No bird remained in a nest longer than ten days. Four young
left the nest after nine days, seven young left on the tenth day. When
the young leave the nest they are able to run rapidly through the dense
grass. The young are fed by the parents for approximately 20 days after
they leave the nest. Twenty-three days after one young Seaside Sparrow
left the nest it was netted at the opposite end of the island, 300
yards from the territory of its parents.

Young Seaside Sparrows fly in an uncertain but characteristic manner
when they are flushed from the grass. They dive clumsily into the grass
after a short flight, making it easy to identify them as birds of the
year.

The plumage of sparrows of the Genus _Ammospiza_ serves to conceal them
in their habitat. In juvenal and adult plumage, the Sharp-tailed
Sparrow is a brown-backed, streaked bird, the color and pattern
blending with the matted grasses (Allen, 1925:67) where the species
feeds and nests. The Seaside Sparrow, as an adult, is olive-gray. Its
color corresponds to that of the substratum where the species forages.
The juvenal plumage of the Seaside Sparrow resembles that of the
Sharp-tailed Sparrow. I believe that young Seaside Sparrows have this
brown, streaked plumage because they spend most of their time in the
dense grass. In the Seaside Sparrow a complete post-juvenal molt begins
in late August. The resulting plumage resembles that which is acquired
by the adults when they complete their post-nuptial molt (Dwight,
1900:192-193).


FOOD, FEEDING, AND CARE OF THE YOUNG

The food habits of Seaside Sparrows and Sharp-tailed Sparrows have been
studied by Judd (1901:64-66), who concluded that both species are
highly insectivorous. In 51 stomachs of Sharp-tailed Sparrows 81 per
cent of the contents was animal. The results of investigation of
stomachs of Seaside Sparrows were similar. In each of the two species
the bill is more elongated and less conical than in other sparrows. For
the two species studied, the shape of the bill seems to be an
adaptation for feeding on insects.

When searching for food, Sharp-tailed Sparrows walk through the dense
black grass, deftly brushing stems aside with their bill as they go.
Open areas are generally traversed by rapid running. I never noticed
either species hopping. They stop to investigate openings in the matted
understory of grass, often sticking their heads into the holes. Many
times I saw these sparrows stretch or jump to pick insects from stems.
Many droppings, almost certainly those of Sharp-tailed Sparrows, were
present in areas of damp, matted grass. Females, when feeding young,
obtain most of the food near the nest; several times I saw birds catch
insects when they were within inches of their nest. Sharp-tailed
Sparrows feed also along the banks of pools and creeks, and along the
perimeters of marshes. Sharp-tailed Sparrows seem to be less restricted
in the types of feeding habitats they can use than are Seaside
Sparrows.

Seaside Sparrows always returned to the edge of the marsh to procure
food, according to my observations. The birds at Lavallette fed
extensively on noctuid moths. In the feeding territories of two pairs
of Seaside Sparrows, along the strip of washed-up eel grass, I found at
least 40 wings of these moths. In several instances the four wings of
one moth were lying close together in the same relative position in
which they had been on the animal. Legs and pieces of thorax were also
discarded occasionally. I watched adults take these moths from the
stems of the smooth cord-grass and snip the wings off with their bills.
Moth wings were present in the other feeding territories, but not in so
great a quantity. Once I saw a female return to the nest with a spider
in her bill. Spiders were abundant throughout the marsh.

Dwight (1900:193) was surprised that the two species living in the same
environment, and therefore suffering equally from abrasion from the
coarse marsh grasses, should have a different number of molts per year.
The Sharp-tailed Sparrow has a complete pre-nuptial, as well as a
complete postnuptial, molt. The Seaside Sparrow has only a postnuptial
molt, the nuptial plumage being acquired by wear. My observations of
the feeding habits of the two species indicate that they do not live in
precisely the same environment. The Sharp-tailed Sparrow, which has two
complete molts annually, generally forages in dense, abrasive
vegetation. The Seaside Sparrow, which has but one molt each year,
forages in relatively open areas.

Several times I saw adult Seaside Sparrows fly from their nests toward
the feeding territories with fecal sacs in their bills. On the feeding
grounds, I found several of these sacs discarded near the moth wings. I
saw also female Sharp-tailed Sparrows leave their nests with fecal
sacs. I did not see sparrows of either species swallow fecal sacs.

One nest, that of a Seaside Sparrow containing four young, became
fouled with excrement when the young were nine to ten days old. It is
interesting that these young were cared for only by a male, at least
for the last four days of nest life, and that one of the young birds
died two days before the others left the nest. This male's mate was
probably a female that I banded on June 18 (the young left the nest on
June 23) and never saw again. A female, whose mate was probably killed
by me on June 15, continued to incubate the three eggs until they
hatched on June 29, but deserted the nest when the young were two days
old. This female was seen again on August 1 more than 500 yards from
her nest site on the island immediately north of the study area.

Devotion of parent passerine birds to the young typically increases
with the growth of the young (Nice, 1943:245). This may explain why the
mateless female deserted its nest when the young were only two days
old, whereas a mateless male continued to care for his six-day-old
young. The death of one nestling, and the eventual fouling of his nest
may indicate that the job was too much for one adult to perform. The
correlation of increasing devotion of the parents with increasing age
of the young was further illustrated by the distraction display, noted
by me, on the part of a pair of Seaside Sparrows on the day their young
left the nest. As I lifted the four nine-day-old young from the nest
for weighing, they began to give the distress call. This attracted the
parents from the feeding area approximately 60 yards away. The two
adults ran around on the ground within ten feet of me giving the
_tsip_ note and fluttering their wings. Several times the adults
flew within a few feet of me, making a vibrating sound with their
wings. Although I realized the function of this display, it was
distracting nevertheless.


    ACKNOWLEDGMENTS

    I am indebted to Assistant Professor Harrison B. Tordoff for
    comments and suggestions throughout the preparation of this
    manuscript, and to Mr. Stuart W. John who photographed the birds
    and the habitat scenes. The aerial photograph of the study island
    is reproduced with the permission of Fairchild Aerial Surveys,
    Inc., and the photograph of Lavallette and Chadwick through the
    kindness of the Lavallette Yacht Club. The drawings were made by
    Mr. John R. Beeder. Additionally I wish to express my gratitude to
    Dr. Dean Amadon, Mr. H. Lyman Sindle, and Mr. Lester B. Woolfenden
    for help and advice in certain aspects of the field work.


SUMMARY

A comparative study of the breeding behavior of the Seaside Sparrow and
Sharp-tailed Sparrow was made in New Jersey in 1955.

Observations of marked individuals indicate that the Seaside Sparrow is
monogamous and territorial, whereas the Sharp-tailed Sparrow is
promiscuous, and at least the male is non-territorial. The male Seaside
Sparrow defends its territory by chasing and singing. The male
Sharp-tailed Sparrow confines itself to a breeding home range. This
range is not a territory; it is inhabited by several males. Female
Sharp-tailed Sparrows may be territorial; this is not certainly known.

The Seaside Sparrow sings louder, more distinctly, more often, and from
more exposed perches than does the Sharp-tailed Sparrow. These
characteristics seem to be correlated with territorial habits. Other
calls are described and their functions are discussed.

The Seaside Sparrow nests in marsh-elder bushes, or in areas of mixed
vegetation. The Sharp-tailed Sparrow prefers the inner, drier areas of
a marsh, where black grass is dominant. The Seaside Sparrow places its
nest farther above the ground than does the Sharp-tailed Sparrow. Both
species used only black grass in constructing the nest.

Copulation is described. The incubation period was not determined for
either species. Three or four eggs seem to be a normal clutch. Females
do all of the incubating.

The young remained in the nests nine to ten days. These nests, of
course were disturbed, for I visited them at least daily. The nestlings
of the Seaside Sparrow are fed by both parents. Male Sharp-tailed
Sparrows seem not to know the location of the nests and take no part in
rearing the young at least up to time of fledging. The natal down of
both species is described. Data on growth and behavior of the young are
presented.

Seaside Sparrows obtained most of their food from the shoreline of the
marsh, in areas of open mud and smooth cord-grass. The plumage of the
adult matches, in color, this mud. The Sharp-tailed Sparrow feeds
everywhere in the marsh, but mostly in areas of dense and matted black
grass. The plumage on the dorsum of this species is brown and streaked
resembling the dead grass. Juvenal Seaside Sparrows, which spend most
of their time concealed in the dense grass, resemble adult and juvenal
Sharp-tailed Sparrows in plumage. Sharp-tailed Sparrows molt completely
twice per year. The Seaside Sparrow molts but once per year. The
difference in number of molts, too, is correlated with habitat
preference, since the grassy forage habitat of the Sharp-tailed Sparrow
must result in greater abrasion of the plumage than does the open
feeding habitat of the Seaside Sparrow.


LITERATURE CITED

ALLEN, GLOVER M.

1925. Birds and their attributes. Marshall Jones Co., Boston,
Massachusetts. xiii + 338 pp., frontispiece, 45 pls., 6 figs.

CRUICKSHANK, ALLAN D.

1942. Birds around New York City, where and when to find them. The
American Museum of Natural History, Handbook Series, No. 13, New York.
xvii + 489 pp., frontispiece, 35 pls.

DWIGHT, JONATHAN

1900. The sequence of plumages and moults of the passerine birds of New
York. Annals N.Y. Acad. Sci., Vol. XIII, pp. 73-360, 7 pls.

FORBUSH, EDWARD HOWE and MAY, JOHN BICHARD

1939. Natural history of the birds of eastern and central North
America. Riverside Press, Cambridge, Massachusetts. xxv + 553 pp., 97
pls.

GRISCOM, LUDLOW

1944. A second revision of the Seaside Sparrows. Occ. papers of the
Museum of Zool., L.S.U., No. 19.

JUDD, SYLVESTER D.

1901. The relation of sparrows to agriculture. U.S.D.A. Bull. No. 15,
98 pp., 4 pls., 19 figs.

LEWIS, HARRISON F.

1920. Notes on the Acadian Sharp-tailed Sparrow (_Passerherbulus
nelsoni subvirgata_). Auk, 37:587.

MONTAGNA, WILLIAM

1940. The Acadian Sharp-tailed Sparrows of Popham Beach, Maine. Wilson
Bull., 52:191-197, 2 figs., 1 table.

1942a. The Sharp-tailed Sparrows of the Atlantic coast. Wilson Bull.,
54:107-120, 4 figs.

1942b. Additional notes on Atlantic coast Sharp-tailed Sparrows. Wilson
Bull., 54:256.

NICE, MARGARET MORSE

1933. The theory of territorialism and its development. Fifty years'
progress of American ornithology 1883-1933. A.O.U., Lancaster,
Pennsylvania, pp. 89-100.

1941. The role of territory in bird life. Amer. Mid. Nat., 26:441-487.

1943. Studies in the life history of the Song Sparrow II. Trans. Linn.
Soc. N.Y. Vol. VI, viii + 329 pp., frontispiece, 6 figs.

PETERSON, ROGER TORY

1947. A field guide to the birds. Houghton Mifflin Co., Boston,
Massachusetts. xxiv + 290 pp., 60 pls.

SALT, W. RAY

1954. The structure of the cloacal protuberance of the Vesper Sparrow
(_Pooecetes gramineus_) and certain other passerine birds. Auk,
71:64-73, 5 figs.

SAUNDERS, ARETAS A.

1951. A guide to bird songs. Doubleday and Co., Garden City, New York.
xiv + 307 pp., 201 figs.

STONE, WITMER

1937. Bird studies at Old Cape May. D.V.O.C. Philadelphia, Vol. II, pp.
521-941.

TOMKINS, IVAN R.

1941. Notes on Macgillivray's Seaside Sparrow. Auk, 58:38-51, pls. 2,
3.


_Transmitted June 14, 1956._