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diff --git a/36285-0.txt b/36285-0.txt new file mode 100644 index 0000000..16fb004 --- /dev/null +++ b/36285-0.txt @@ -0,0 +1,1907 @@ +The Project Gutenberg EBook of Comparative Breeding Behavior of Ammospiza +caudacuta and A. maritima, by Glen E. Woolfenden + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Comparative Breeding Behavior of Ammospiza caudacuta and A. maritima + +Author: Glen E. Woolfenden + +Release Date: May 30, 2011 [EBook #36285] + +Language: English + +Character set encoding: UTF-8 + +*** START OF THIS PROJECT GUTENBERG EBOOK COMPARATIVE BREEDING *** + + + + +Produced by Chris Curnow, Joseph Cooper and the Online +Distributed Proofreading Team at http://www.pgdp.net + + + + + + +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._ + + + + + + +End of the Project Gutenberg EBook of Comparative Breeding Behavior of +Ammospiza caudacuta and A. marit, by Glen E. Woolfenden + +*** END OF THIS PROJECT GUTENBERG EBOOK COMPARATIVE BREEDING *** + +***** This file should be named 36285-0.txt or 36285-0.zip ***** +This and all associated files of various formats will be found in: + http://www.gutenberg.org/3/6/2/8/36285/ + +Produced by Chris Curnow, Joseph Cooper and the Online +Distributed Proofreading Team at http://www.pgdp.net + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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