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Carolina wren

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Carolina wren
Carolina Wren1.jpg
Recorded in Cape May, New Jersey, US
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Passeriformes
Family: Troglodytidae
Genus: Thryothorus (but see text)
Vieillot, 1816[2]
Species: T. ludovicianus
Binomial name
Thryothorus ludovicianus
(Latham, 1790)
Subspecies [3]
Thryothorus ludovicianus species distribution map.svg
Range of the Carolina wren (T. ludovicianus)
Synonyms[4]

Sylvia ludoviciana
Latham, 1790

The Carolina wren (Thryothorus ludovicianus) is a common species of wren that is a resident in the eastern half of the United States of America, the extreme south of Ontario, Canada, and the extreme northeast of Mexico. Severe winters restrict the northern limits of their range while favorable weather conditions lead to a northward extension of their breeding range. Their preferred habitat is in dense cover in forests, farm edges and suburban areas. This wren is the state bird of South Carolina.

There are seven recognized subspecies across the range of these wrens and they differ slightly in song and appearance. The birds are generally inconspicuous, avoiding the open for extended periods of time. When out in the open, they investigate their surroundings and are rarely stationary. After finding a mate, pairs maintain a territory and stay together for several years. Both sexes give out alarm calls, but only males sing to advertize territory. Carolina wrens raise multiple broods during the summer breeding season, but can fall victim to brood parasitism by brown-headed cowbirds, among other species. Some populations have been affected by mercury contamination.

Taxonomy[edit]

The Carolina wren was first described under the name of Sylvia ludoviciana by John Latham in 1790.[3][note 1] Louis Jean Pierre Vieillot considered all wrens under the genus Troglodytes and called the Carolina wren Troglodytes arundinaceus but placed it subsequently in a separate genus Thryothorus (initially misspelled Thriothorus [2]) that he created in 1816.[7] Thryothorus is of Greek origin from the combination of thryon (rush, reed) and thouros (derivative of verb throskein to leap up, spring, jump at) which means 'reed jumper'; its specific name ludovicianus is a post-classical Latin term for Ludovicus (derivative from Louis XIV) that means 'of Louisiana' that identifies the locality of the specimen collected near New Orleans.[8][9]

There are seven recognized subspecies of the Carolina wren:[3][10]

T. l. ludovicianus (Latham, 1790) – Southeast Canada (Southern Ontario, irregularly in Eastern and Southern Quebec) and the eastern United States (Southern Wisconsin and New England southward to Texas and northern Florida).

T. l. miamensis Florida wren (Ridgway, 1875) – Florida from approximately 30 degrees (Gainesville) region southward through the rest of the state.

T. l. nesophilus (Stevenson, 1973) – Dog Island in Northwestern Florida.

T. l. burleighiBurleigh's Carolina wren (Lowery, 1940) Offshore islands off of the Mississippi coast: Cat Island, Ship Island (Mississippi), and Horn Island.

T. l. lomitensisLomita wren (Sennett, 1890) southern Texas to the extreme northeast of Mexico (Tamaulipas).

T. l. berlandieriBerlandier's wren (S. F. Baird, 1858) Northeastern Mexico (eastern Coahuila, Nuevo Leon, and southwestern Tamaulipas)

T. l. tropicalis – Northeastern Mexico (eastern San Luis Potosi and southern Tamaulipas).

T. ludovicianus is traditionally placed within its own genus as its only representative of North America, but recent DNA work suggests it is closely allied with the Bewick's wren.[11] A distinct population in the Yucatan Peninsula of Mexico, Belize, Nicaragua, and in Guatemala is treated as a separate species, either known as Cabot's wren or white-browed wren (Thryothorus albinucha).[3][12] It is considered a subspecies of T. ludovicianus by some authors, however.[10]





Campylorhynchus megalopterus



Campylorhynchus brunneicapillus





Thryomanes bewicki




Thryothorus ludovicianus ludivicianus



T. l. albinucha








Cinnycerthia peruana




Thryothorus guarayanus



Thryothorus leucotis










Henicorhina leucosticta



Henicorhina leucophrys




Uropsila leucogastra




Cyphorhinus arada





Thryothorus maculipectus



Thryothorus coraya





Thryothorus sinaloa





Description[edit]

At 12.5 to 14 cm (4.9 to 5.5 in) long, with a 29 cm (11 in) wingspan and a weight of about 18 to 23 g (0.63 to 0.81 oz), the Carolina wren is a fairly large wren; the second largest in the United States species after the cactus wren. Among standard measurements, the wing chord is 5.4 to 6.4 cm (2.1 to 2.5 in), the tail is 4.5 to 5.6 cm (1.8 to 2.2 in), the culmen is 1.4 to 1.8 cm (0.55 to 0.71 in) and the tarsus is 2 to 2.3 cm (0.79 to 0.91 in).[3] Sexual dimorphism is slight with males being larger than their mates. A study indicated that out of 42 mated pairs, all but one of male was larger than the female of the pair. The males were on average 11 percent heavier along with having longer wing chords.[13]

There are several differences among the subspecies. For T. l. ludovicianus, the crown is rich brown that appears more chestnut-colored on its rump and uppertail-coverts. Shoulders and greater coverts are a rich brown, with a series of small white dots on the lesser primary coverts. The secondary coverts are rich brown with a darker brown barring on both webs; the bars on the primaries are on the outerwebs only, but darker and more noticeable. The retrices are brown with 18 to 20 bars that span across the tail. The white supercilious streak borders thinly with a black above and below, and extends above and beyond its shoulders. The ear coverts are speckled gray and grayish-black. Its chin and throat are grey that becomes buff on its chest, flank and belly, though the latter two are of a warmer color. The underwing coverts sport a grayish buff color. Its iris is reddish-brown, the upper mandible is lemon-colored and paler at the base and lower mandible. The legs are flesh-colored.[3]

As for the other subspecies in contrast to T. l. ludovicianus, T. l. berlandieri is of a slightly smaller build, but possesses a larger bill, the upperparts are duller brown with deeper colored underparts, T. l. lomitensis is of a duller color (than either ludovicianus or berlandieri) with its underparts either pale or almost white, T. l. miamensis contains darker rusty chestnut upperparts and deeper colored below. T. l. burleighi is duller and sootier with less distinct tail markings, T. l. mesophilus has paler underparts and a whiter supercilium, and T. l. tropicalis is darker than all races, and contains heavier bars than T. l. berlandieri.[3]

Plumage[edit]

The juvenile T.l. ludovicianus is similar in appearance, but the plumage is generally paler with a softer texture with buff-tipped wing coverts, a superciliary streak is less white, a fluffy vent and crissum (flanks) without bars.[3][14] In August and September, the partial plumage molt for the post-juvenile wrens is darker in color and affects the contour plumage, wing coverts, tail and develops a whiter superciliary stripe. The post-nuptial molt for adults in the same time period is more pronounced in color than the spring molt, with both sexes similar in appearance.[14]

Life span[edit]

Sketches of Thryothorus ludovicianus, T.l. lomitensis, and Thryomanes bewickii (Bewick's wren) and one of its subspecies.

Survival rates differ by region. A male captured in Arkansas lived to be at least 73 months old, and in Alabama, the oldest female and male captured were six and ten years old, respectively. A mark-and-recapture analysis of the wrens analyzing survival probability within the Southeastern United States captured was monitored from 1992 to 2003. Roughly 90 percent of the banded wrens died within 10 years.[15]

Similar species[edit]

The easiest species to confuse with the Carolina wren is Bewick's wren,[16] which differs in being smaller but with a longer tail, grayer-brown above and whiter below. The Carolina and white-browed wrens differ from the house wren in being larger, with a decidedly longer bill and hind toe; their culmen has a notch behind the tip.[17]

Habitat and distribution[edit]

These birds are largely resident, and will only disperse beyond their range after mild winters.[3] Carolina wrens sporadically breed as far north as Maine and Quebec after mild winters.[3][18] In certain parts of their range, such as most of Iowa, prolonged periods of snow can curtail potential expansion.[19] Permanent breeding locations range from eastern Nebraska, southern Michigan, southeast Ontario and the New England states to Mexican states such as Coahuila, Nuevo León, San Luis Potosí and Tamaulipas and the Gulf Coast of the United States.[15] Local occurrences with infrequent and likely breeding locations include southeast South Dakota, central Kansas, eastern Colorado, western Oklahoma and Texas as far as Maine and New Brunswick.[15] There have been occasional vagrants spotted in Colorado, New Mexico, and Arizona, Wyoming, South Dakota, Manitoba, Nova Scotia, and the Gulf of St. Lawrence.[3][18]

Carolina wren at feeder

The range of the wrens increased northward and westward in several regions over the past few centuries. In Massachusetts, the wrens had expanded westward and northeastward from its former southeastern location in approximately 35 years, in New York the population increased three-fold in roughly 25 years, while in midwest states such as Ohio and Michigan have seen numbers of the birds increase since the mid-1800s and early 1900s, respectively.[15] Expansion around Ontario occurred since early reports in 1890 and 1905. Explanations given include infrequent winter storms in the 20th century, expanded forest habitats, and the wrens taking advantage of urban areas containing feeders, especially in winter.[15]

Carolina wrens adapt to various habitats. Natural habitats include various types of woodland such as oak hardwoods and mixed oak-pine woodlands, ash and elmwoods, hickory-oak woodlands with a healthy amount of tangled undergrowth.[3][20] The preferred habitats are riparian forests, brushy edges, swamps, overgrown farmlands, and suburban yards with abundant thick shrubs and trees, and parks.[3][20] It has an affinity for dilapidated buildings and unkempt yards in man-made areas.[20] Subspecies burleighi and neophilus inhabit slash pine and palmettos.[3][20]

Behavior[edit]

Song and calls[edit]

Carolina wrens sing year round and at any point during the daytime, with the exception of performing during the most harsh weather conditions.[20] The birds are also the only species in the Certhiidae family that neither sings in duet nor has their song control regions affect repertoire size.[21] Males alone sing, and have a repertoire of at least twenty different phrase patterns and on average, thirty two.[21][22] One of these patterns is repeated for several minutes, and although the male's song can be repeated up to twelve times, the general amount of songs range from three to five times in repetition. While singing, the tail of the birds is pointed downward. Some general vocalizations have been transcribed as teakettle-teakettle-teakettle and cheery-cheery-cheery.[20][14] Various descriptions of the teakettle song include whee-udel, whee-udel, whee-udel, che-wortel, che-wortel and túrtee-túrtee-túrtee and familiar names and phrases such as sweet heart, sweet heart, come to me, come to me, sweet William, and Richelieu, Richelieu.[14]

Males are capable of increasing their repertoire through song learning, but due to their sedentary nature and territorial defense habits, the song learning must occur within the first three months of life.[23] Geographic barriers affect song repertoire size from male wrens, as one study indicated that distances separated as close as 3 kilometres (1.9 mi) by water barriers can have the same effect as that of a distance of 145 kilometres (90 mi) in the mainland with no barriers.[23]

Female Carolina wrens possess song control regions that would appear to make them capable of singing with repertoires like the male. Due to vocalizations that they occasionally make with the male, it has been suggested that song perception plays a role and is of behavioral relevance.[21]

Carolina wren on Rutland Township Forest Preserve

Different subspecies have variations in songs and calls, such as miamensis having a more rapid song that contains more notes than the races that are further north.[3]

Their songs can be confused with the Kentucky warbler. The song patterns are similar, but is of a different quality, as the warbler's songs is described as richer, with more ringing and a hurried pace.[20] Other bird species that have their songs described as akin to the wrens are the flicker, Baltimore oriole, gray catbird, and more specifically the peto, peto, peto calls of the tufted titmouse, and whistles of the northern cardinal.[14] Occasionally, the wrens mimic other species, and in Pennsylvania it has led for it to be also known as the 'Mocking wren'.[3]

Sexual selection[edit]

A 2006 study suggested that the correlation of tail length and body size in males, wing length in females, and lifespan for both sexes were signs of individual quality, and the wrens of high quality tend to mate with like individuals. The courting and antagonistic encounters that involve the tail fanning and wing drooping was suggested to be a possible signaling use. Age and life experience are not thought of as significant for potential mates due to their relatively short lifespan and sedentary lifestyle. Due to the large size of male wrens and the male's vigor in defending its territory, intrasexual selection was given as a possible explanation for the sexual dimorphism.[13]

Territorial and predator defense[edit]

Both sexes are involved in defending the territory. One aspect of territorial defense involves identifying the proximity of the threat based on the loudness of bird song as well as the level of degradation of the calls. In experiments involving playback, the wrens are capable of discriminating between degraded and undegraded songs, as well as degraded songs in the same acoustic conditions, and can detect changes of acoustic properties within their territories, such as songs under foliage.[24] Song degradation can also be used to determine the proximity of potential intruders. If the song of a bird appears to be degraded, the wrens will assume that the threat is distant not respond; if the song is not degraded, they respond by attacking.[25] Not all birds within their territory are potential enemies. Some species of birds that are neighbors are designated as 'dear-enemies' by the wrens, and the responses to neighbors and intruders in their territories differ by the season. In spring, the wrens respond more aggressively toward neighbors, though in the fall, no major discrepancy in responses is shown.[26] When protecting their nest, alarm calls are the general response. The wrens judge the size of the potential threat, such as a blue jay and avoid the risk of injury when attacking.[27] Countersinging produced by intruder birds is more likely to be taken as an aggressive threat to male Carolina wrens.[28]

Both males and females utilize calls in alarm situations, especially in territorial disputes and encounters with predators. Males alone produce the cheer call, which can sound indistinct. In southern regions of their range, the sound males use in alarm disputes is a ringing pink or p'dink sound. Females are the only ones that can perform the paired dit-dit or chatter sounds. The former can be used in territorial disputes with predators, and with at least northern populations the songs are used in alternation with the males cheer chant. The chatter is used exclusively with territorial encounters with male song, and the song can either follow or overlap her mate's song.[21][29]

Feeding[edit]

Carolina wrens spend the majority of their time on or near the ground searching for food, or in tangles of vegetation and vines. They also probe bark crevices on lower tree levels, or pick up leaf-litter in order to search for prey. Their diet consists of invertebrates, such as beetles, true bugs, grasshoppers, katydids, spiders, ants, bees, and wasps. Small lizards and tree frogs also make up the carnivorous portion of their diet. Vegetable matter makes up a small percentage of their diets, such as fruit pulp and various seeds. In the northern portion of their range, they frequent bird feeders.[3][20]

Movement[edit]

Carolina wrens are wary, and are more often heard than seen. When on the ground, they move in jerky hops pillaging through various objects, whether man-made or natural.[20] While moving abruptly, they pause momentarily for chattering or singing.[14] When stationary, they move in twitched motions, jerking their breast around.[20] They also sun- or sand-bathe.[30] The wrens also displays a skittish behavior when encountered by humans, as they can be seen thrusting off into cover slowly if approaching is detected. However, they occasionally seek out humans that are near, so long as there is no movement from them.[14] Other movements involve being capable of crawling like a creeper and hanging upside-down like a nuthatch.[20]

Flights are generally of short duration, rapid, low-leveled, and wavelike. The wings during flight are flapped rapidly, and are frequently used during foraging. They are also capable of flying vertically from the base of a tree to the top in a single wing assisted bound.[3][20]

Breeding[edit]

Carolina wren nesting in a duck nestbox.

Carolina wrens are both genetically and socially monogamous and will usually mate for life. Mate changing is rare,[13] and there has been one possible instance of polygamy.[3] During the winter season, males are more responsible for guarding the territory. Females vary in succeeding to maintain winter territories without a mate.[3] It has been suggested that the possibility of desertion and decline in care-taking from males along with the need for security in resources year-round prevent extra pair copulations from females, as the mortality rate for Carolina wrens peaks during the winter.[31] Along with thermoregulatory benefits, roosting is thought to reinforce pair-bonding and prevent divorce between mates.[32]

The nests are arch-shaped structures with a side entrance and built of dried plants or strips of bark, as well as horsehair, string, wool and snake sloughs. Males obtain nesting materials while the females remains at the site to construct the nest. Nests are located in fragmented or complete cavities in trees, or in man-made structures such as bird-boxes, buildings, tin cans, mailboxes or unorthodox places such as pockets of hanging jackets in sheds or in a tractor in everyday use.[14][3] Nests are from 1–3 m (3.3–9.8 ft) from the ground and are rarely higher. They occasionally can be built in sloping locations or at ground level.[3]

Egg laying dates and clutch size vary by region; in Texas the time period is from late February to late August, in Iowa it ranges from late April to June.[12][19] The clutch size is generally 3 to 6 eggs, but can reach as high as seven in Texas.[3][12] The eggs are creamy white with brown or reddish-brown spots, and are more heavily marked at the broad end.[3] The eggs are incubated by the female for 12–16 days. After the young hatch, they are fed exclusively on invertebrates and they fledge in 12–14 days. As many as three broods may be raised by a pair in a single breeding season.[3] In one study, three of the 70 fledglings remained or defended territory adjacent to the natal area.[15]

Male and females are involved in the process of provisioning at similar rates throughout most nest stages, with the males providing slightly more in the nestling stages. Both sexes increase their provision rates as the nestlings grow in age.[33]

Predation and threats[edit]

Brood parasitism by brown-headed cowbirds is common, with up to 25% of Carolina wren nests being affected in certain regions such as Oklahoma and Alabama.[3][15] Cowbird parasitism peaks in April at 41%, and is as low as 8% and 0% in July and August, respectively. Female cowbirds sometimes eject Carolina wren eggs before laying their own, and even if host eggs are retained, the size of cowbird eggs negatively affect the hatching success of wren eggs. As a result, cowbirds may have a significant impact on the reproductive success of wrens.[3][15] The feeding rate for cowbird nestlings is higher than wren feeding rates, and some have been raised to independence.[15] This also can be detrimental to the survival of wren nestlings.[15] A rare instance of brood-parasitism by a House finch has been recorded.[15][34] The rate of brood parasitism is thought to be lower in more natural and concealed nesting locations.[15] Body parasites such as the larvae of blowflies feed on nestlings and the blood loss weakens nestlings.[15] Fellow species of wren such as Bewick's wren and the winter wren compete for nesting locations and food, respectively.[15]

In Virginia, some Carolina wrens populations show high levels of mercury in their blood and this is acquired from feeding all-year-round on spiders.[35][note 2] Spiders being at a higher trophic levels contain a higher concentrations of mercury (through biomagnification) than herbivorous invertebrates. As these wrens are year-round residents, they are at a higher risk than other species to acquire mercury in its blood. Nest abandonment and failure to raise young were more common with higher mercury content.[35] Exposure, and prolonged periods of cold, ice, and snow is thought to affect the wren nestling and adult populations, respectively.[15] Wrens that outlast those winters reside in sheltered areas during the season.[14]

Among the top predators of adult Carolina wrens are domestic cats, and snakes such as the canebrake rattlesnake.[36][37] Racoons and black rat snakes also feed on wren eggs and nestlings.[15]

In culture[edit]

South Carolina state quarter

In 1930, the South Carolina Federated Women's club adopted the Carolina wren as the unofficial state bird over the eastern mourning dove and pushed for its official state adoption until 1939, when the South Carolina Legislature named the northern mockingbird as the state bird. In 1948, the legislature repealed their previous decision, and the wren became the official state bird.[38]

In 2000, the Carolina wren was featured on the back of the South Carolina edition of the 50 State Quarters.[39]

Notes[edit]

  1. ^ Johann Friedrich Gmelin wrongly associated the Carolina wren with the European wrens under the name Motacilla troglodytes (originally coined by Carl Linnaeus in Systema Naturae)[5] in 1788.[3][6]
  2. ^ The North Folk Holston and South Rivers were used in the study and were both contaminated by industrial activity.

References[edit]

  1. ^ BirdLife International (2012). "Thryothorus ludovicianus". IUCN Red List of Threatened Species. Version 2013.2. International Union for Conservation of Nature. Retrieved 26 November 2013. 
  2. ^ a b David, Normand; & Dubois, Alain (2011). "The original spellings of Thryothorus Vieillot, 1816 (Vertebrata, Aves): a correction" (PDF). Zootaxa 2918: 68–68. 
  3. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab Brewer, David (2001). Wrens, Dippers, and Thrashers. Yale University Press. pp. 132–4. ISBN 978-0-300-09059-8. 
  4. ^ Mayr, E. and Greenway, J.C. Jr., eds. (1960). Check-list of birds of the World. Volume IX. Cambridge, Mass.: Museum of Comparative Zoology. pp. 409–410. 
  5. ^ Linnaeus, C. (1758). Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Tomus I. Editio Decima, Reformata (in Latin). Holmiae: Laurentius Salvius. p. 188. 
  6. ^ Harding, J. (1824). Journal of the Academy of the Sciences of Philadelphia: Volume IV, Part I. Philadelphia: Academy of Sciences of Philadelphia. pp. 28–29. 
  7. ^ Oberholser, Harry C. (1902). "A synopsis of the genus commonly called Anorthura". Auk: 175–181. JSTOR 4069309. 
  8. ^ Sandrock, James (2014). The Scientific Nomenclature of Birds in the Upper Midwest. University of Iowa Press. p. 144. ISBN 978-1609382254. 
  9. ^ Coues, Elliot (1882). Coues check list of North American Birds (2 ed.). Boston: Estes and Lauriat. p. 31. 
  10. ^ a b "Carolina or White-browed Wren (Thryothorus [ludovicianus or albinucha])". AviBase. Retrieved 16 February 2015. 
  11. ^ Mann, Nigel I.; Barker, F. Keith; Graves, Jeff A.; Dingess-Mann, Kimberly A. & Slater, Peter J.B. (2006). "Molecular data delineate four genera of "Thryothorus" wrens" (PDF). Molecular Phylogenetics and Evolution 40 (3): 750–9. doi:10.1016/j.ympev.2006.04.014. PMID 16750640. 
  12. ^ a b c "Thryothorus ludovicianus". Texas A&M AgriLifeExtension. Texas A&M University. 2006. Retrieved 31 January 2015. 
  13. ^ a b c Haggerty, Thomas M. (2006). "Sexual size dimorphism and assortative mating in Carolina Wrens" (PDF). Journey of Field Ornithology 77 (3): 259–65. doi:10.1111/j.1557-9263.2006.00051.x. 
  14. ^ a b c d e f g h i Bent, Arthur C. (1948). "Life Histories of North American Nuthatches, Wrens, Thrashers, and their Allies: Order Passeriformes". Bulletin of the United States National Museum: 205–16. doi:10.5479/si.03629236.195.1. 
  15. ^ a b c d e f g h i j k l m n o p "The Birds of North America Online: Carolina Wren". 
  16. ^ Martínez Gómez, Juan E.; Barber, Bruian R. & Peterson, A. Townsend (2005). "Phylogenetic position and generic placement of the Socorro Wren (Thryomanes sissonii)" (PDF). The Auk 122: 50. doi:10.1642/0004-8038(2005)122[0050:PPAGPO]2.0.CO;2. 
  17. ^ Brattstrom, Bayard H. & Howell, Thomas R. (1956). "The Birds of the Revilla Gigedo Islands, Mexico" (PDF). Condor 58 (2): 107–120. doi:10.2307/1364977. 
  18. ^ a b Dunn, John Lloyd; Alderfer, Jonathan K. (2004). National Geographic Illustrated Birds of North America. National Geographic Books. p. 342. ISBN 978-1426205255. 
  19. ^ a b Jackson, Laura Press (1996). The Iowa Breeding Bird Atlas. University of Iowa Press. pp. 266–7. ISBN 978-1426205255. 
  20. ^ a b c d e f g h i j k l Dunne, Pete (2006). Pete Dunne's Essential Field Guide Companion: A Comprehensive Resource for Identifying North American Birds. Houghton Mifflin Harcourt. p. 479. ISBN 978-0-300-09059-8. 
  21. ^ a b c d Nealen, Paul M.; Perkel, David J. (2000). "Sexual Dimorphism in the Song System of the Carolina Wren (Thryothorus ludovicianus)". The Journal of Comparative Neurology 418 (4): 346–360. doi:10.1002/(sici)1096-9861(20000313)418:3<346::aid-cne8>3.3.co;2-b. PMID 10701831. 
  22. ^ Elliott, Lang (1999). Music of the Birds: A Celebration of Bird Song. Houghton Mifflin Harcourt. p. 30. ISBN 978-0618006977. 
  23. ^ a b Morton, Eugene S. (2005). "The Effects of Distance and Isolation on Song-type Sharing in the Carolina Wren" (PDF). Wilson Bulletin 99 (4): 601–10. 
  24. ^ Naguib, Marc (1995). "Ranging by Song in Carolina Wrens (Thryothorus ludovicianus): Effects of Environmental Acoustics and Strength of Song Degradation" (PDF). Animal Behaviour 133: 541–559. doi:10.1163/156853996x00206. 
  25. ^ Richards, Douglas G. (1981). "Estimation of the Distance of Singing Conspecifics by the Carolina Wren" (PDF). The Auk 98: 127–33. JSTOR 4085615. 
  26. ^ Hyman, Jeremy (2005). "Seasonal Variation in Response to Neighbors and Strangers by a Territorial Songbird" (PDF). Ethology 111: 951–61. doi:10.1111/j.1439-0310.2005.01104.x. 
  27. ^ D'Orazio, Kelly; Neudorf, Diane L. H. (2008). "Nest Defense by Carolina Wrens". The Wilson Journal of Ornithology 120 (3): 467–72. doi:10.2307/20456180. 
  28. ^ Hyman, Jeremy (2003). "Countersinging as a signal of aggression in a territorial songbird" (PDF). Animal Behaviour 65: 1179–85. doi:10.1006/anbe.2003.2175. 
  29. ^ Elliott, Lang (2004). Know Your Bird Sounds: Songs and calls of yard, garden, and city birds. Stackpole Books. p. 29. ISBN 978-0811729635. 
  30. ^ Hauser, Doris C. (1957). "Some Obversations of Sun-bathing in Birds" (PDF). The Wilson Bulletin 69 (1): 259–65. 
  31. ^ Haggerty, Thomas M.; Morton, Eugene M.; Fleischer, Robert C. (2001). "Genetic Monogamy in Carolina Wrens (Thryothorus ludovicianus)" (PDF). The Auk 118 (1): 215–19. doi:10.1642/0004-8038. JSTOR 4089770. 
  32. ^ Labinsky, Ronald F.; Arnett Jr., John E. (2006). "Pair Roosting of Carolina Wrens (Thryothorus ludovicianus)". The Wilson Journal of Ornithology 118 (4): 566–569. doi:10.1676/05-109.1. JSTOR 20455927. 
  33. ^ Neudorf, Diane L. H.; Broddick, Mallory J.; Cureton II, James C. (2013). "Parental Provisioning by Carolina Wrens". The Wilson Journal of Ornithology 125 (1): 179–184. doi:10.1676/12-009.1. 
  34. ^ Wood, Douglas R. & William A. Carter (2006). "Carolina Wren Nest Successfully Parasitized by House Finch". The Wilson Journal of Ornithology 118 (3): 413–415. doi:10.1676/05-102.1. 
  35. ^ a b Jackson, Alison K.; Evers, David C.; Etterson, Matthew A.; Condon, Anne M.; Folsom, Sarah B.; Detweiler, Jennifer; Schmerfeld, John; Cristol, Daniel A. (2011). "Mercury Exposure Affects the Reproductive Success of a Free-Living Terrestrial Songbird, The Carolina Wren (Thryothorus ludovicianus)". The Auk 128 (4): 759–769. doi:10.1525/auk.2011.11106. 
  36. ^ Mitchell, Joseph C.; Beck, Ruth A. (1992). "Free-Ranging Domestic Cat Predation on Native Vertebrates in Rural and Native Virginia" (PDF). Virginia Journal of Science 43 (1B): 197–208. 
  37. ^ Parmley, Dennis; Parmley, Amanda M. (2001). "Food Habits of the Canebrake Rattlesnake (Crotalus horridus atricaudatus) in Central Georgia" (PDF). Georgia Journal of Science 59 (4): 172–80. 
  38. ^ "South Carolina State Bird – Thryrothorus ludovicianus". NetState. Retrieved 31 January 2015. 
  39. ^ "The Official South Carolina State Quarter". TheUS50. Retrieved 31 January 2015. 

External links[edit]