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Overview

Comprehensive Description

Eudocimus albus, the white ibis, is a member of the Order Ciconiiformes (herons and storks). It is a long necked wading bird in which the sexes are similar in appearance. Major identifying characteristics include its long decurved bill, longer in males than in females; entirely white body coloration, pink bill, and legs, black tipped outer primary feathers, and distinctive bare face which ranges in shades from red to pink. The eyes of adults are blue.
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Distribution

occurs (regularly, as a native taxon) in multiple nations

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National Distribution

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

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Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) Resident from central Baja California, central Sinaloa, southern and eastern Texas, southern Louisiana, Florida, southeastern Georgia, and coastal North Carolina (rarely Virginia) south along coasts and through Greater Antilles to French Guiana and northwestern Peru. Wanders casually north. (AOU 1983). In the U.S., the highest winter densities occur in Florida and around the mouth of the Mississippi River (Root 1988).

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Range

Southern US to se Brazil, Bahamas and Greater Antilles.

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Geographic Range

The White Ibis breeds in the coastal region of Virginia, south and west to Louisiana, including inland S. Carolina through Florida. It also breeds along the entire coast of Mexico, Belize, Nicaragua, Cuba, Jamaica, Panama, and Costa Rica. Their non-breeding range extends only slightly deeper into inland Louisiana, Georgia, and S. Carolina.

Biogeographic Regions: nearctic (Native ); neotropical (Native )

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In the continental United States, Eudocimus albus, the white ibis, occurs from Virginia south along the Atlantic coast to the Gulf of Mexico. It is found on both coasts of Mexico, and ranges as far south as Columbia and Brazil. The white ibis is distributed throughout the Indian River Lagoon.
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Physical Description

Morphology

Physical Description

Resembling the Great Blue Heron in body shape, the White Ibis is a medium-sized wading bird. Its feathers are entirely white, except for its dark wing tips. The face of the ibis is bare and pink, blending into its long, curved bill, which is brown at the tip. It has long pink legs, which end with webbed toes. The adults eyes are light blue.

Average mass: 940 g.

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Size

Length: 64 cm

Weight: 1036 grams

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The white ibis is a medium sized wading bird that attains a height of 22 inches, with a wingspan of 38 inches. It may live as long as 16 years in the wild, and 20 years in captivity (Kushlan and Bildstein 1992).
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Diagnostic Description

No other North American bird of this size has both a long, slender, decurved, pink/scarlet bill and a white belly (bill of the much larger wood stork is yellow much thicker).

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Type Information

Type for Eudocimus albus
Catalog Number: USNM 525728
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Birds
Sex/Stage: Female;
Preparation: Skin: Whole
Collector(s): H. Bailey
Year Collected: 1924
Locality: Cape Sable, Monroe, Florida, United States, North America
  • Type: Bailey, H. H. April 1, 1930. Bailey Mus. Lib. Nat. Hist. Bull. 4.
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Ecology

Habitat

Moist Pacific Coast Mangroves Habitat

This taxon occurs in the Moist Pacific Coast mangroves, an ecoregion along the Pacific coast of Costa Rica with a considerable number of embayments that provide shelter from wind and waves, thus favouring mangrove establishment. Tidal fluctuations also directly influence the mangrove ecosystem health in this zone. The Moist Pacific Coast mangroves ecoregion has a mean tidal amplitude of three and one half metres,

Many of the streams and rivers, which help create this mangrove ecoregion, flow down from the Talamanca Mountain Range. Because of the resulting high mountain sediment loading, coral reefs are sparse along the Pacific coastal zone of Central America, and thus reef zones are chiefly found offshore near islands. In this region, coral reefs are associated with the mangroves at the Isla del Caño Biological Reserve, seventeen kilometres from the mainland coast near the Térraba-Sierpe Mangrove Reserve. The Térraba-Sierpe, found at the mouths of the Térraba and Sierpe Rivers, is considered a wetland of international importance.

Because of high moisture availability, the salinity gradient is more moderate than in the more northern ecoregion such as the Southern dry Pacific Coast ecoregion. Resulting mangrove vegetation is mixed with that of marshland species such as Dragonsblood Tree (Pterocarpus officinalis), Campnosperma panamensis, Guinea Bactris (Bactris guineensis), and is adjacent to Yolillo Palm (Raphia taedigera) swamp forest, which provides shelter for White-tailed Deer (Odocoileus virginianus) and Mantled Howler Monkeys (Alouatta palliata). Mangrove tree and shrub taxa include Red Mangrove (Rhizophora mangle), Mangle Caballero (R. harrisonii) R. racemosa (up to 45 metres in canopy height), Black Mangrove (Avicennia germinans) and Mangle Salado (A. bicolor), a mangrove tree restricted to the Pacific coastline of Mesoamerica.

Two endemic birds listed by IUCN as threatened in conservation status are found in the mangroves of this ecoregion, one being the Mangrove Hummingbird (Amazilia boucardi EN), whose favourite flower is the Tea Mangrove (Pelliciera rhizophorae), the sole mangrove plant pollinated by a vertebrate. Another endemic avain species to the ecoregion is the  Yellow-billed Cotinga (Carpodectes antoniae EN).  Other birds clearly associated with the mangrove habitat include Roseate Spoonbill (Ajaia ajaja), Gray-necked Wood Rail (Aramides cajanea), Rufous-necked Wood Rail (A. axillaris), Mangrove Black-hawk (Buteogallus anthracinus subtilis),Striated Heron (Butorides striata), Muscovy Duck (Cairina moschata), Boat-billed Heron (Cochlearius cochlearius), American White Ibis (Eudocimus albus), Amazon Kingfisher (Chloroceryle amazona), Mangrove Cuckoo (Coccyzus minor), Yellow Warbler (Setophaga petechia), and Black-necked Stilt (Himantopus mexicanus VU) among other avian taxa.

Mammals although not as numerous as birds, include species such as the Lowland Paca (Agouti paca), Mantled Howler Monkey (Alouatta palliata), White-throated Capuchin (Cebus capucinus), Silky Anteater (Cyclopes didactylus), Central American Otter (Lontra longicaudis annectens), White-tailed Deer (Odocoileus virginianus), feeds on leaves within A. bicolor and L. racemosa forests. Two raccoons: Northern Raccoon (Procyon lotor) and Crab-eating Raccoon (P. cancrivorus) can be found, both on the ground and in the canopy consuming crabs and mollusks. The Mexican Collared Anteater (Tamandua mexicana) is also found in the Moist Pacific Coast mangroves.

There are a number of amphibians in the ecoregion, including the anuran taxa: Almirante Robber Frog (Craugastor talamancae); Chiriqui Glass Frog (Cochranella pulverata); Forrer's Grass Frog (Lithobates forreri), who is found along the Pacific versant, and is at the southern limit of its range in this ecoregion. Example salamanders found in the ecoregion are the Colombian Worm Salamander (Oedipina parvipes) and the Gamboa Worm Salamander (Oedipina complex), a lowland organism that is found in the northern end of its range in the ecoregion. Reptiles including the Common Basilisk Lizard (Basiliscus basiliscus), Boa Constrictor (Boa constrictor), American Crocodile (Crocodilus acutus), Spectacled Caiman (Caiman crocodilus), Black Spiny-tailed Iguana (Ctenosaura similis) and Common Green Iguana (Iguana iguana) thrive in this mangrove ecoregion.

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Comments: Various salt water and freshwater habitats: marshes, mangroves, lagoons, lakes, marsh prairie, pasture, coastal swamps (AOU 1983, Kushlan 1979). Often perches in trees. Nests in trees or shrubs near water, especially in wooded swamps; also on matted clumps of JUNCUS (Frederick 1987) or other marsh vegetation. May show fidelity to nest area despite chronic nest loss due to tidal washover. Typically nests with smaller EGRETTA herons (Frederick 1987).

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Habitat and Ecology

Systems
  • Terrestrial
  • Freshwater
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Barriers, marshes, spoil islands on the coast, and islands in inland lakes are the preferred nesting sites for the White Ibis. These sites are in interior and coastal wetlands, in environments ranging from southeastern mixed forest to outer coastal plain forest, savanna, prairie parkland, and prairie bushland. Feeding habitats include sedge marshes, cypress swamps, salt marshes, and mangrove swamps.

Terrestrial Biomes: savanna or grassland ; forest

Aquatic Biomes: coastal

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Migration

Non-Migrant: Yes. At least some populations of this species do not make significant seasonal migrations. Juvenile dispersal is not considered a migration.

Locally Migrant: Yes. At least some populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).

Locally Migrant: No. No populations of this species make annual migrations of over 200 km.

Makes local seasonal movements along Gulf Coast.

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Trophic Strategy

Comments: Eats mainly crustaceans, also fishes, frogs, small snakes, slugs, snails, insects; probes into mud with bill or picks up food from surface among mangroves, along edges of ponds, in marsh prairie, fields, and in shallow estuaries (Palmer 1962). In South Carolina, dependent of crayfish availability in wetlands (Bildstein et al. 1990).

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Food Habits

Using its long decurved bill and long neck, the White Ibis probes the surface of its wetland habitat for aquatic crustaceans (such as crayfish and crabs) and insects. It washes the mud from its prey in the surrounding water, then swallows it with a quick, upward thrust of the neck and head. White Ibises tend to feed in large groups. They fly with this group to and from their feeding location.

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White ibis feed primarily on species of crustaceans (Kushlan and Bildstein 1992) from both saltmarsh and freshwater wetland habitats. They are tactile feeders that probe around soft bottom areas with their long bills in search of prawns, crayfish and fiddler crabs. Tactile foraging is somewhat more inefficient in capturing fish, so it is used rarely when foraging for fish (Kushlan 1979). Adult birds that are feeding chicks visit freshwater foraging sites more frequently than saltmarsh sites. However, once young have fledged, parental visits to saltmarsh feeding habitats double.Competitors: White ibises compete for food with other wading birds.Habitats: The white ibis has been described as a nomadic species, rather than a migratory species (Frederick et al. 1996). As such, it quickly colonizes wetlands having good food resources, and readily abandons areas where resources have become scarce. It utilizes both freshwater and estuarine wetlands such as mangrove and cypress swamps, bottomland hardwood, and marshes. In Florida studies, its preferred breeding habitat is in freshwater areas where winter and early spring water levels are low or receding (Smith and Collopy 1995).
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Associations

White ibises are highly gregarious and readily associate with other species of medium sized herons and egrets. They are known to nest in mixed flocks with cattle egrets (Kushlan and Bildstein 1992).
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Population Biology

Number of Occurrences

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: > 300

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Global Abundance

10,000 to >1,000,000 individuals

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Frederick et al. (1996) reported that the Florida Everglades population of white ibis is clearly in decline, with a more widespread overall decline in progress throughout the United States. Available records indicate there were approximately 125,000 breeding pairs in 1933; 170,000 breeding pairs in 1976; 51,000 pairs in 1991; and 43,000 pairs in 1992. Kushlan and Bildstein (1992) have suggested that these figures may not represent actual declines in population. Rather, they may indicate that range expansion of this species throughout the southeast, coupled with the nomadic lifestyle of the white ibis cause fluctuation in population numbers in any one area. These authors stress that population figures as a whole continue to be relatively high. One exception occurs in the Florida Everglades, where water management practices have altered the natural hydrologic regime of the system, causing a marked population decline (Kushlan and Bildstein 1992)(see Reproduction section below).Locomotion: White ibises walk at a rate of approximately 25 - 40 steps per minute. They fly with rapid wingbeats at the rate of 3.3 flaps per second. Flying is alternated with gliding for 60 - 100 m, sometimes as fast as 45 km/hour (Kushlan and Bildstein 1992).
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General Ecology

Highly gregarious. When not breeding, congregates at communal roosts; may move long distance between roost and feeding area (Hilty and Brown 1986).

Fish crow may prey on eggs but effect on ibis productivity was regarded as negligible in North Carolina (Shields and Parnell 1986).

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Life History and Behavior

Cyclicity

Comments: Forages during daylight (Powell 1987).

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Life Expectancy

Lifespan/Longevity

Average lifespan

Status: wild:
196 months.

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Lifespan, longevity, and ageing

Maximum longevity: 27.6 years (captivity) Observations: These animals can live up to 27.6 years in captivity (Brouwer et al. 1994).
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Reproduction

Clutch size is 3-4 in the north, usually 2 in the south (Central and South America). Incubation lasts about 21- 23 days, by both sexes. Young leave nest at about three weeks, fly at about five weeks. Captive birds first bred at 2 years (Terres 1980, Palmer 1962). Largest colonies in coastal U.S. comprise about 5000-6200 birds on Atlantic coast (in Carolinas), 20,000 at Cedar Keys, Florida, and 60,000 just north of Lake Maurepas, Louisiana, on Gulf Coast (Spendelow and Patton 1988). In Florida, nesting success and high nesting numbers were associated with rapid water drying rate in spring (Frederick and Collopy 1989).

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Pair formation in ibises tends to depend on the amount of rain, light, and available food rather than occurring at a set time every year. Nest building, however, tends to fall between May and early June. Colonies begin with the roost sites of males, who form a sort of daytime bachelor party. The females then come and build the nests nearby. Nests are built in live or dead woody plants, usually in branch crotches. On average, 2-3 eggs are laid. Both the male and female incubate, and the eggs hatch about 21 days after incubation begins. The nestlings are born with their eyes closed and cannot stand, preferring to sleep for the first week of life. They are easily overheated, so both parents make sure to keep them guarded from the sun with their wings. After about 40-50 days of care from the parents, the fledgling ibises first leave the colony. They do not leave permanently until they are at least 79% the adult mass, which takes about two years to achieve.

Average time to hatching: 22 days.

Average eggs per season: 3.

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White ibises are highly social breeders and nest in large mixed colonies numbering from hundreds to ten thousands of nests. They begin nesting in their third summers, and will nest yearly under optimal conditions. In Florida, nesting begins as early as mid-March and April (Smith and Collopy 1995; Smith 1997). Further to the south, in Costa Rica, nesting takes place between May and July (Leber 1980). Nest success is highly correlated to low and receding water levels in freshwater areas, as nestling are highly susceptible to salt stress. Between 2 - 5 eggs are laid per clutch, and 1 brood is raised per year. Renesting often occurs following early-season nest failure.White Ibises shift breeding locations in response to rainfall patterns, with breeding success shown to increase in years with high rainfall (Bildstein 1990). In south Florida, it has been observed that 35 times as many breeding birds are present in wet years than in dry years (Kushlan 1976). In dry years, clutch size is decreased, nests are more likely to be abandoned, and chicks are more likely to die from starvation (Bildstein et al. 1990). In Florida, the active management of water levels in wetland areas such as Lake Okeechobee and the Florida everglades may also contribute to nest success (Smith and Collopy 1995). It has been suggested that in order to enhance nest success and later fledgling success, water levels in managed wetlands should be maintained at low or receding levels throughout the late winter to early spring nesting season. Low water levels help to concentrate prey in small, shallow areas, thereby making it simpler for parent birds to feed nestlings. It also contributes to the foraging success of fledglings (Smith and Collopy 1995).Nest takeovers and nest parasitism are sometimes problems in ibis breeding colonies. A pair that is nesting, but without eggs will occasionally attack a nesting female with eggs and force her from her nest. If eggs are present, the intruding female will peck at eggs to destroy them, and then eject them from the nest. Nesting material is then rearranged and the intruding pair will begin copulation. The ejected pair will often return to the nest and chase off the intruders; however, in all cases where this behavior was observed, nest failure resulted (Frederick 1986). Conspecific egg dumping, or nest parasitism also occurs during nesting, where non-nesting females will deposit eggs into a nesting female's nest and leave the nesting female to raise chicks that are not her own (Kushlan and Bildstein 1992).White ibises from wild populations in Venezuela and Florida are known to hybridize with the scarlet ibis, Eudocimus ruber. Hybridization with the scarlet ibis also readily occurs in captivity (Ramo and Busto 1987).
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Growth

Eggs are incubated 21 - 23 days. Hatching is asynchronous and occurs over several days. The oldest chick quickly gains experience at food handling and aggressive behaviors toward its siblings, and thus grows at a faster rate than its nestmates. Nestling mortality is greatest in the first 20 days of life (Kushlan and Bildstein 1992).At hatching, the skin, legs and feet of chicks are flesh colored. Blue-gray down covers the body by the third day, with black down covering the head and neck. Feathers begin to emerge around day 5. The eyes open after 1 - 3 days, with chicks becoming fully alert by day 9. Chicks are mobile at 8 days of age, and by day 15 often begin leaving the nest to join crèches (social groups) of similarly aged birds. Chicks in crèches continue to be fed by adults. Fledging occurs at 28 - 35 days.
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Molecular Biology and Genetics

Molecular Biology

Barcode data: Eudocimus albus

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 6 barcode sequences available from BOLD and GenBank.

Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.

See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

NNNNNNNNNNNNNNTTGGCGCATGAGCTGGTATAGTTGGAACTGCCCTTAGCTTACTTATCCGCGCAGAACTAGGTCAACCAGGAACACTCCTGGGAGACGACCAAATCTACAATGTAATCGTCACTGCCCATGCCTTCGTAATAATCTTCTTCATAGTCATACCAATCATGATTGGCGGATTTGGCAACTGACTAGTGCCCCTTATAATCGGTGCCCCCGACATAGCCTTTCCCCGCATAAACAATATAAGCTTCTGACTACTACCTCCATCCTTCCTACTCCTTCTAGCCTCCTCCACAGTAGAAGCAGGAGCAGGCACAGGATGAACTGTATACCCACCTCTAGCTGGCAACCTCGCCCATGCTGGAGCCTCAGTAGACCTAGCCATCTTCTCCCTTCACCTAGCAGGGGTATCTTCCATTCTAGGAGCTATTAACTTCATCACAACTGCCATTAACATAAAACCTCCAGCCCTCTCACAATATCAAACACCACTATTCGTTTGATCCGTCCTAATCACTGCCGTTTTACTACTACTCTCTCTACCAGTCCTCGCTGCCGGCATCACCATGCTGCTAACAGATCGAAACCTGAACACTACATTCTTCGACCCTGCAGGAGGAGGAGATNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNNN
-- end --

Download FASTA File

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Statistics of barcoding coverage: Eudocimus albus

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 6
Specimens with Barcodes: 6
Species With Barcodes: 1
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Conservation

Conservation Status

National NatureServe Conservation Status

United States

Rounded National Status Rank: N5B,N5N : N5B: Secure - Breeding, N5N: Secure - Nonbreeding

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NatureServe Conservation Status

Rounded Global Status Rank: G5 - Secure

Reasons: Still common in many areas of the large range. Large declines have occurred in Florida in recent decades.

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IUCN Red List Assessment


Red List Category
LC
Least Concern

Red List Criteria

Version
3.1

Year Assessed
2012

Assessor/s
BirdLife International

Reviewer/s
Butchart, S. & Symes, A.

Contributor/s

Justification
This species has an extremely large range, and hence does not approach the thresholds for Vulnerable under the range size criterion (Extent of Occurrence <20,000 km2 combined with a declining or fluctuating range size, habitat extent/quality, or population size and a small number of locations or severe fragmentation). The population trend appears to be stable, and hence the species does not approach the thresholds for Vulnerable under the population trend criterion (>30% decline over ten years or three generations). The population size is very large, and hence does not approach the thresholds for Vulnerable under the population size criterion (<10,000 mature individuals with a continuing decline estimated to be >10% in ten years or three generations, or with a specified population structure). For these reasons the species is evaluated as Least Concern.
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The main threat to white ibis populations is the destruction of colony sites and wetland foraging environments by humans. The management of these areas is a problem, because ibises tend to change sites frequently. As a whole, however, the population is not experiencing large decreases in number and no special status has been issued for this species. In Florida white ibises are a species of special concern.

US Migratory Bird Act: protected

US Federal List: no special status

CITES: no special status

State of Michigan List: no special status

IUCN Red List of Threatened Species: least concern

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Global Short Term Trend: Decline of 10-30%

Comments: In southern Florida, declined 90% from 1940 to 1974, 80% from 1975 to the late 1980s; 50% decline statewide in the past decade (BWD Skimmer, April 1993). Populations in the south-central U.S. may be benefiting from crayfish aquaculture; bird population increases may be related to favorable foraging opportunities afforded by expanding crayfish aquaculture (Fleury and Sherry 1995).

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Population

Population Trend
Stable
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Threats

The white ibis is listed as a Species of Special Concern (SSC) in Florida. It is not federally listed as threatened or endangered.
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Relevance to Humans and Ecosystems

Benefits

Economic Importance for Humans: Negative

In Louisiana, White Ibises are considered a threat to commercial crayfish farmers and are sometimes shot as vermin.

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Economic Importance for Humans: Positive

White Ibis was hunted as game throughout its range. Its appealing taste is thought to originate from its crustacean diet.

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Benefit in IRL: In contrast with migratory birds that commonly utilize the same habitats from year to year, the white ibis is nomadic, readily abandoning habitats that can no longer support it. Its presence in wetlands is thus a good indicator of overall environmental health and habitat quality (Frederick et al. 1996).Cost in IRL: As a nomadic species, the white ibis is thus unlikely to be insular and easily protected because any conservation effort must address habitat quality over extensive areas. Should more evidence of a declining U.S. population continue to accumulate, the continued survival of this species will depend both regional planning and a continued commitment to maintain high quality habitat areas (Frederick et al. 1996). Though not of economic value, in some regions throughout its range, the white ibis continues to be hunted for food. Their appealing taste is perhaps due to the ibis' dietary preference for crabs and other crustaceans (Kushlan and Bildstein 1992).
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Wikipedia

American white ibis

This article is about the American white ibis. For the Australian bird, see Australian white ibis.

The American white ibis (Eudocimus albus) is a species of bird in the ibis family, Threskiornithidae. It is found from the mid-Atlantic and Gulf Coast of the United States south through most of the coastal New World tropics.[2] This particular ibis is a medium-sized bird with an overall white plumage, bright red-orange down-curved bill and long legs, and black wing tips that are usually only visible in flight. Males are larger and have longer bills than females. The breeding range runs along the Gulf and Atlantic Coast, and the coasts of Mexico and Central America. Outside the breeding period, the range extends further inland in North America and also includes the Caribbean. It is also found along the northwestern South American coastline in Colombia and Venezuela. Populations in central Venezuela overlap and interbreed with the scarlet ibis. The two have been classified by some authorities as a single species.

Their diet consists primarily of small aquatic prey, such as insects and small fishes. Crayfish are its preferred food in most regions, but it can adjust its diet according to the habitat and prey abundance. It is a tactile, non-visual forager, whose main foraging behavior is probing with its beak at the bottom of shallow water to feel form, and to capture its prey.

During the breeding season, the American white ibis gathers in huge colonies near water. Pairs are predominantly monogamous and both parents care for the young, although males tend to engage in extra-pair copulation with other females to increase their reproductive success. Males have also been found to pirate food from unmated females and juveniles during the breeding season.

Human pollution has affected the behavior of the American white ibis via an increase in the concentrations of methylmercury, which is released into the environment from untreated waste. Exposure to methylmercury alters the hormone levels of American white ibis, affecting their mating and nesting behavior and leading to lower reproduction rates.

Taxonomy[edit]

The American white ibis was one of the many bird species originally described by Carl Linnaeus in the 1758 10th edition of his Systema Naturae, where it was given the binomial name of Scolopax albus.[3] The species name is the Latin adjective albus "white".[4] Alternative common names that have been used include Spanish curlew and white curlew.[5] English naturalist Mark Catesby mistook immature birds for a separate species, which he called the brown curlew.[6] Local creole names in Louisiana include bec croche and petit flaman.[7]

Johann Georg Wagler gave the species its current binomial name in 1832 when he erected the new genus Eudocimus, whose only other species is the scarlet ibis (E. ruber). There has long been debate on whether the two should be considered subspecies or closely related species,[8] and the American Ornithologists' Union considers the two to be a superspecies as they are parapatric.[9] The lack of observed hybrids was a large factor in the view that the species were separate.[8]

However, in a field study published in 1987, researchers Cristina Ramo and Benjamin Busto found evidence of interbreeding in a population where the ranges of the scarlet and white ibises overlap along the coast and in the Llanos region of Colombia and Venezuela. They observed individuals of the two species mating and pairing, as well as hybrid ibises with pale orange plumage, or white plumage with occasional orange feathers; their proposal that these birds be classified as a single species,[9] has been followed by least one field guide.[5] Hybrid ibises have also been recorded in Florida, where the scarlet ibis has been introduced into wild populations of American white ibis. Birds of intermediate to red plumage have persisted for generations.[8]

Ornithologists James Hancock and Jim Kushlan also consider the two to be a single species, with the differences in plumage, size, skin coloration and degree of bill darkening during breeding season forming the diagnostic characters. They have proposed the populations recontacted in northwestern South America after a period of separation, and that the color difference is likely due to the presence of an enzyme that allows uptake of pigment in the diet. They have questioned whether white-plumaged birds of South America are in fact part of the ruber rather than the albus taxon, and acknowledge that more investigation is needed to determine this.[8]

Description[edit]

Adult flying in Huntington Beach State Park, South Carolina, United States

The white plumage and pink facial skin of adult American white ibises are distinctive.[10] Adults have black wingtips that are usually only visible in flight.[11] In non-breeding condition the long downcurved bill and long legs are bright red-orange.[12] During the first ten days of the breeding season, the skin darkens to a deep pink on the bill and an almost purple-tinted red on the legs. It then fades to a paler pink, and the tip of the bill becomes blackish.[13] It is difficult to determine the sex of an adult American white ibis from its external appearance, since the sexes have similar plumage.[14] However, there is sexual dimorphism in size and proportion as males are significantly larger and heavier than females and have longer and stouter bills.[15] A study of the American white ibis in southern Florida yielded weight ranges of 872.9 to 1,261 g (1.924 to 2.780 lb) for males and 592.7 to 861.3 g (1.307 to 1.899 lb) for females, with average weights of 1,036.4 g (2.285 lb) for males and 764.5 g (1.685 lb) for females.[15][16] The length of adult female and male birds ranges from 53 to 70 cm (21 to 28 in) with a 90 to 105 cm (35 to 41 in) wingspan.[17][18] Among standard measurements, American white ibis measure 20.5–31 cm (8.1–12.2 in) along each wing, have a tail measurement of 9.3–12.2 cm (3.7–4.8 in), a tarsus of 6.75–11.3 cm (2.66–4.45 in) and a culmen of 11–16.9 cm (4.3–6.7 in).[8]

The newly hatched American white ibis is covered with purple down feathers, deepening to dark brown or black on the head and wings. The chest is often bare and there can be a white tuft on the head. The irises are brown. The exposed skin is pinkish initially, apart from the tip of the bill which is dark gray, but turns gray within a few days of hatching.[8] The bill is short and straight at birth and has an egg tooth which falls off between days five and nine,[19] and develops three black rings from around day six, before turning gray by around six weeks of age. The gray to sandy gray brown juvenile plumage appears between weeks two and six, and face and bill become pink a few weeks later, while the legs remain gray. The irises have turned slate-gray by this stage.[8] Once fledged, the juvenile American white ibis has largely brown plumage and only the rump, underwing and underparts are white.[11] The legs become light orange. As it matures, white feathers begin appearing on the back and it undergoes a gradual molt to obtain the white adult plumage.[11] This is mostly complete by the end of the second year, although some brown feathers persist on the head and neck until the end of the third year. Juvenile birds take around two years to reach adult size and weight.[8]

Like other species of ibis, the American white ibis flies with neck and legs outstretched, often in long loose lines or V formations—a 1986 field study in North Carolina noted over 80% of adult ibis doing so, while juveniles rapidly took up the practice over the course of the summer. The resulting improvement in aerodynamics may lower energy expenditure.[20] These lines fly in an undulating pattern as they alternately flap and glide. Soaring in a circular pattern is also seen.[21] Heights of 500 to 1,000 m (1,600 to 3,300 ft) may be reached as birds glide over flights of 20 km (12 mi) or more. More commonly, birds fly between 60 and 100 m (200 and 330 ft) above the ground, gliding or flapping at a rate of around 3.3 wingbeats a second.[22]

The main call of the American white ibis is a honking sound, transcribed as urnk, urnk,[2] or hunk, hunk.[8] The call is used in flight, courtship or when disturbed. Birds also utter a muted huu-huu-huu call while foraging, and make a squealing call in courtship. Young in the nest give a high-pitched zziu as a begging call.[8]

Similar species[edit]

Immature American white and scarlet ibises are very difficult to tell apart, although scarlet ibises tend to have darker legs and bare skin around the face.[23] An immature American white ibis could be mistaken for an immature glossy ibis, but the latter is wholly dark brown and lacks the white belly and rump. The adult is distinguishable from the wood stork, which is much larger and its wings have more black on them.[21]

Distribution and habitat[edit]

An American white ibis at Riverside Park, Jacksonville, Florida
Adults in shallow water at Merritt Island National Wildlife Refuge near the Atlantic coast of Florida

The American white ibis is most common in Florida, where over 30,000 have been counted in a single breeding colony. It also occurs throughout the Caribbean, on both coasts of Mexico (from Baja California southwards) and Central America, and as far south as Columbia and Venezuela. The non-breeding range extends further inland, reaching north to Virginia, and west to eastern Texas.

Adult American white ibis on pavement outside of Orlando, FL.

The species is known to wander, and has been sighted, sometimes in small flocks, in states far out of its usual range.[5][11]

In North America, breeding takes place along the Atlantic coast, from the Carolinas south to Florida and thence west along the Gulf Coast.[2] Laguna Cuyutlán is an isolated and regionally important wetland in the state of Colima on México's west coast where a breeding colony has been recorded.[24] American white ibises are not faithful to the sites where they breed,[25] and large breeding colonies composed of ten thousand birds or more can congregate and disband in one or two breeding seasons.[26] Breeding populations across its range have fluctuated greatly with wholesale movement between states.[27] Until the 1940s, the species only bred in large numbers in Florida, mostly within the Everglades.[26] Drought conditions elsewhere in the United States led to over 400,000 American white ibis breeding there in the 1930s.[27] In the 1950s and 1960s, large colonies appeared in Alabama, Louisiana, and then North and South Carolina and the Gulf Coast of Florida, and finally Texas in the 1970s. Then, between the 1970s and early 1990s, breeding colonies declined and disappeared in South Carolina and Florida, and greatly increased in Louisiana,[26] and North Carolina.[27] Colonies last between one and seventeen years, their longevity related to size and quality of nearby wetlands. The longest-lasting are associated with wetlands over 800 km2 (310 sq mi) in size. Degradation of wetland or breeding sites are reasons for abandonment.[26] The population of American white ibises in a colony at Pumpkinseed Island in Georgetown County, South Carolina dropped from 10,000 to zero between 1989 and 1990 as Hurricane Hugo had inundated nearby freshwater foraging areas with salt water.[28]

The American white ibis is found in a variety of habitats, although shallow coastal marshes, wetlands and mangrove swamps are preferred.[29] It is also commonly found in muddy pools, on mudflats and even wet lawns. Populations that are away from the coast and shoreline, particularly in southern Florida, often reside in other forms of wetlands such as marshes, ponds and flooded fields.[2][11] In summer, these move to more coastal and estuarine habitats as inland waterways become flooded with summer rains and the ibis find the water levels too deep to forage effectively.[29]

Fossil record[edit]

Remains similar to the American white ibis have been found in Middle Pliocene deposits of the Bone Valley formation in central Florida, and Lower Pliocene deposits of the Yorktown Formation at Lee Creek in North Carolina.[30] Two species, one living and one extinct, have been recovered from the Talara Tar Seeps in northern coastal Peru. Eudocimus peruvianus was described from a tarsometatarsus that differed slightly from E. albus, whose remains were also found there. Remains of neither species are common in the beds. The tar seeps have been dated at 13,900 years old. The American white ibis is still found in Peru.[31]

Behavior[edit]

American white ibis in a neighborhood pond in Tampa Bay, Florida

A field study late in the Florida nesting season revealed that on an average day, adult American white ibis spent 10.25 hours looking for food, 0.75 hours flying, 13 hours resting, roosting, and attending to their nests.[32] Much of the time roosting is spent preening, biting and working their feathers with their long bills, as well as rubbing the oil glands on the sides of their heads on back plumage. American white ibis generally only preen themselves, not engaging in allopreening unless part of courtship behavior. Bathing often takes place before preening; ibis squat in water 5–20 cm (2.0–7.9 in) deep and flick water over themselves with each wing in succession. Hundreds of birds may bathe together around the time of courtship.[22]

The American white ibis is territorial, defending the nesting and display sites against intruders. Agonistic or threat displays include lunging forward with the bill in a horizontal posture, and standing upright and snapping the bill opposite another bird engaging in the same display. Birds also lunge and bite, often holding onto an opponent's head or wings.[22]

Breeding and lifespan[edit]

Birds roosting in a tree near to St. Johns River, Florida

The American white ibis pairs up in spring and breeds in huge colonies, often with other waterbird species. Nesting begins as soon as suitable foraging and nesting habitat is available. The female selects the site, usually in the branches of a tree or shrub, which is often over water, and builds the nest, and males assist by bringing nest material.[12] Anywhere from one to five eggs are typically laid, with two or three being the most common. The eggs are matt pale blue-green in color with brown splotches and measure 5.8 cm × 3.9 cm (2.3 in × 1.5 in) and weigh on average 50.8 g (1.79 oz).[5] Clutch sizes are usually lower in coastal colonies as compared to inland colonies, although there are no statistically significant differences in the fledging rate of both colonies.[32] Throughout the mating and incubation period, the male undergoes a period of starvation to stay close to the nest and aggressively defend his nest and mate from both predators and other ibises in preference to foraging for food.[33] In the 2006 breeding season, a non-breeding adult female was observed to be tending to multiple nests that belonged to other American white ibises—the first time the behavior has been documented for this species.[34]

Although the American white ibis is predominantly monogamous and both sexes provide parental care to their young, the male often flies off to engage in extra pair copulation with other nesting females after mating with its primary female partner. These extra-pair copulations are usually done after the within-pair copulations,[35] and make up about 45% of all total matings, although only about 15% of all extra-pair copulations are successful.[35] By not restricting the number of females it copulates with, the male is able to increase its reproductive success considerably. Although females are receptive towards extra-pair copulations, male mate-guarding greatly reduces the rate of successful female involvement in attempted extra-pair copulations by other males.[33]

Juvenile in Everglades National Park. Some of its brown feathers have molted and have been replaced with white feathers.

The breeding success of the American white ibis is sensitive to the hydrological conditions of the ecosystem such as rainfall and water levels. Low and decreasing water levels predict good prey accessibility. Water level reversals, where levels rise in the breeding season, disperse prey and impact on foraging success. Nest numbers and average clutch sizes are smaller in periods of reduced prey availability.[36] The success rate of parents raising one or more young to 20 days of age ranges widely from 5 to 70% of nests, and varies greatly between nearby colonies.[27] American white ibis parents have been known to supplement their chicks' diet with items such as cockroaches and rotting food from human garbage in poorer years, when fish and crayfish are more scarce.[37] Studies have also shown that years with higher nesting numbers had significantly faster spring drying rates of water bodies than years with low nesting numbers. This is because faster drying rates means that there are fewer fish and increased available area where crayfish can be hunted.[38] This highlights the fact that American white ibises do not use probability of nesting failure as a key factor in determining nesting sites but instead, rely on other criteria such as prey availability and nest-site predation rates.[39] The draining of wetlands in south Florida has also impacted on species that forage in shallow water such as the American white ibis, and its increase in numbers is a key indicator of restoration of habitat within the Everglades.[40]

The main cause of nest failure among the species is due to nest abandonment,[38] the leading cause of which is inundation from extremely high tides. Parents abandoned 61% of all nesting starts either during or immediately after extremely high tides. The eggs float out of the flooded nests, or get washed out into the sea by wave action. Incubating parents usually abandon the nest when the water or tidal levels reaches 3 to 8 cm (1.2 to 3.1 in) above the bottom of the nest cup. Nevertheless, there have been instances where the parents have been observed to transport their eggs to another nest in an attempt to salvage some eggs. However, despite the fact that some nesting sites face high chance of tidal damage every breeding season, American white ibises still continue to nest in these areas because of other favorable conditions such as abundant nearby food sources and low egg predation rates.[39]

Juvenile in Florida

The eggs hatch after about three weeks and the young are attended by both parents. Males are present around the nest for most of the day, and females most of the night. The parents exchange nest duties in the morning and in the evening. Most of the feeding of the chicks occurs during the period where they swap nesting duties. Little feeding is done in the period of the day that is between the two duty swaps and no feeding is done between midnight and 6 a.m.[41] Chick mortality is highest in the first twenty days post hatching, with anywhere from 37 to 83% of hatchlings surviving to three weeks of age in the Everglades.[27] During periods of food limitations and starvation events, the American white ibis tends to exhibit sex-dependent pre-fledgling mortality.[42] For many bird species that have sexually dimorphic nestlings, mortality rates are higher for larger-sized male nestlings as a result of the parents' inability to meet its greater nutritional needs.[43] However, in the case of the American white ibis, the male nestlings actually have a lower mortality rate as compared to the females despite being on average 15% greater in mass as compared to its female counterparts.[42] Although current research has yet to discover the underlying factors to why the males tend to have better survival rates under such conditions, it is suspected that the parents tend to feed the larger male nestlings first because they are either perceived by the parents to have a higher chance of survival, or, being generally larger, the male nestlings simply out-compete the small females for food.[43]

Bird predators may seize anywhere from 7% to 75% of the progeny in a breeding colony.[22] The fish crow (Corvus ossifragus) is common raider of American white ibis nests, accounting for up to 44% of egg loss in a field study at Battery Island, North Carolina.[44] Other predators of eggs and young include the boat-tailed grackle (Quiscalus major), black-crowned night heron (Nycticorax nycticorax), gulls, and possibly vultures, as well as the common opossum (Didelphis marsupialis), raccoon (Procyon lotor), and rat snakes (Elaphe species).[22] Egg predation rates of the American white ibis decline with nest age owing to increased nest attentiveness by the parents, especially during the last week of incubation. High nest densities and reduced synchrony increase egg predation rates because of the increased opportunities afforded by the longer incubation times, as well as the greater availability of nests available for predation.[44]

The American white ibis begins breeding in its third summer, although birds in captivity may breed as young as nine or ten months of age.[27] The oldest member of the species recorded in captivity was over 20 years of age, and a wild bird has been picked up 16 years and 4 months after being banded.[5]

Diet[edit]

Adult eating a fish

The American white ibis prefers to eat crayfish and other crustaceans, but also takes aquatic insects and small fishes.[12][45] Outside the nesting season, the diet is highly variable, as abundance and types of prey depend on the both region and habitat. In Los Llanos, located on the border of Columbia and Venezuela, the most frequent prey are insects, such as fly larvae and beetles. Generally in North America the main prey are crustaceans, mostly crayfish.[46] In the Everglades and cypress swamps, the diet is primarily made up of crayfish, while those that feed in willow ponds eat predominantly fish. American white ibises that feed in mangrove swamps focus on crabs.[29] The tactile nature of the ibis's probing for food in mud means that it catches prey that are too slow to evade the ibis once located by its bill. In the Everglades, this means that crayfish make up a large part of the diet, but a more diverse array of invertebrates are taken in coastal areas.[47] Although crayfish are sought by foraging ibises, prey switching to fish does occur if fish are found in great abundance. It is unclear whether the fish are more easily caught if overcrowded, or whether sheer numbers of fish mean that ibises are catching them instead of crayfish—normally, fish are more agile than crayfish and hence elude the ibis's bill more easily. Fish are a more energy-rich source of food for the American white ibis.[29] In the breeding season, American white ibises in a colony at Pumpkinseed Island travelled further to forage in freshwater wetlands and catch crayfish, than nearby saltwater areas where fiddler crabs predominated, indicating their benefit was worth the extra energy expended in fetching them for their young.[48] This travel results in the wholesale transport of nutrients across the landscape by the colony; in a successful breeding year the colony at Pumpkinseed Island was estimated to have contributed a third as much phosphorus to the neighboring estuary as other environmental processes.[49]

The American white ibis is found in mixed-species foraging flocks with the glossy ibis (Plegadis falcinellus) in flooded fields, and the two species select different food items with little overlap; the former foraging for crabs and aquatic insects and the latter feeding mainly on grain.[5] The Wood Stork is also found in the same habitat in Florida, but hunts larger prey and a higher percentage of fish, so there is little overlap.[47] In the Llanos, where American white ibis coexist with the scarlet ibis, their diets differ, the former consuming more bugs, fish and crustaceans, while the latter eat a much higher proportion of beetles.[46] The willet (Tringa semipalmata) has been observed trailing American white ibis and catching prey disturbed by them, and even kleptoparasitizing (stealing) from them, in J. N. "Ding" Darling National Wildlife Refuge on Sanibel Island in Florida.[50] An isolated event of intraspecific predation in juvenile American white ibis has been observed, where a juvenile attacked and consumed a chick from another nest.[51]

Foraging[edit]

Video of adults foraging on Bonita Beach, Bonita Springs, Florida, United States

During the summer, the American white ibis roams along the coast of tidal flats and mangrove swamps as the inland marshes are usually flooded. However, as the water level recedes in the fall, populations at the coast shift their foraging area inland, to inland marshes and swamplands.[29] It has become more common in urban landscapes in Florida since the late 1990s,[52] and is one of a number of wetland-dependent bird species which forages in man-made ponds on golf courses in the southwest of the state.[53]

The American white ibis is a tactile, nonvisual forager, which limits its ability to choose from a wide variety of prey.[54] For the most part, the American white ibis forages for food by tactile probing. It wades slowly through shallow water and sticks its long, downcurved bill into the substrate of the water body with its bill held at around 1 to 2 cm (0.39 to 0.79 in) agape at the tip, and sweeps its long bill back and forth across the bottom to pick out suitable food items.[2] Birds may also probe when standing still. Groping with a wide open bill is a technique used by ibis in deeper water when alone, as is head swinging, in which the ibis swings its wide open bill widely in open water. Others copy this type of foraging if they see one ibis doing it. On land, the American white ibis locates prey by sight and pecks, and does not have to insert its bill into the substrate.[54] The American white ibis seeks small prey when other birds are around, as it needs time to break up larger food items into smaller pieces to eat, and other predators such as herons and egrets often take the opportunity to rob the ibis of its catch.[29][55] Along with the scarlet ibis, the species coexists with another five species of ibis in the Llanos in Venezuela. American white ibis males are aggressive to and take prey items from smaller ibises, but the smaller females are more often the victims of this behavior.[56]

Adults foraging for food in a front garden in Florida

Juveniles have lower foraging efficiency compared to adults and in most feeding flocks, the juveniles are usually outnumbered by the adults. They usually tend to stay close to one another and forage for food together at the peripheral region of the group.[57] During the breeding season, adult male ibises have been recorded raiding other parent ibises who are feeding their young in the colony. The raiders force their bill down the throat of the victim—either the parent about to disgorge their food or recently fed young—and extract the ball of food.[58] This behavior allows the otherwise starving adult males to obtain food without having to spend long periods of time away from the nest, and prevent its female mate copulating with another male ibis, which would reduce its own reproductive success. Females and juveniles almost never try to drive off the larger and more aggressive pirating males, but instead try to avoid or move away from them.[58] This pirating behavior is less common between two male ibises as the males will actively fight off the pirate.[58] The explanation of the species' sexual dimorphism of body size is unclear, because no differences between the sexes in feeding success rates or the foraging behavior have been observed and, as males are larger, they need more food than females.[59]

Parasites and mortality[edit]

Causes of death of adult ibis are not well known. Alligators could feasibly prey on nesting ibises but there has been little research in the area.[27] A flock of fifty adult American white ibis were killed in a fire in the Everglades. The corpses were found in a dense swathe of cattail (Typha angustifolia), which suggested they had taken shelter there. It is unclear why they had not been able to fly away from the fire, but one hypothesis was that they had been foraging for insects disturbed by the fire.[60]

A total of 51 species of parasitic worm have been recovered from the American white ibis, predominantly from the gastrointestinal system and particularly the small intestine. These include Cestoda (tapeworms), Acanthocephala (thorny headed worms), Nematoda (roundworms), Digenea and Spirurida. Several roundworm and spirurid species have been found in the lining of the gizzard.[61] Nematodes are more prevalent in American white ibis from freshwater habitats, and cestodes more frequent in those from saltwater areas. One nematode found in adult birds, Skrjabinoclavia thapari, is borne in the fiddler crab as an intermediate host, while the thorny headed worm species Southwellina dimorpha is carried in crayfish and infests both adult and juvenile ibis.[27]

Parasitic protozoa of the genus Sarcocystis have been recovered from the smooth muscles of adult American white ibis,[62] and another species, Haemoproteus plataleae, has been recovered from the blood of adults and nestlings, and can hence be transmitted before the young leave the nest.[63][64] The larvae of two species of mite of the family Hypoderidae, Phalacrodectes whartoni and Neoattialges eudocimae, have been recovered from under the skin.[65] Two species of the louse suborder Mallophaga, Plegadiphilus eudocimus and Ardeicola robusta, also parasitise the bird.[63]

Status[edit]

The American white ibis is classed as Least Concern on the IUCN Red List.[1] The population consists of 150,000 mature adults, and is stable, although some populations have unknown trends. A partial survey of under 50% of the North American population published in 2007 found an almost six-fold increase in the last four decades. The estimated breeding range is huge, at 1,200,000 km2 (460,000 sq mi).[66] Fluctuating breeding populations and high mobility of colonies make estimating the population difficult. Attempted censuses of breeding colonies across Texas, Louisiana, Florida, and the Carolinas yielded a count of 166,000 breeding birds in 2001, and 209,000 in 2004.[27] The conservation status has been listed in two states—it is a Species of Special Concern in Florida, and a species of Moderate Conservation Concern in Alabama. The preservation of colony sites and freshwater foraging areas is important to maintaining populations; however, the highly mobile nature of breeding colonies makes this challenging.[27]

Human impact[edit]

John James Audubon reported that the American white ibis was hunted and sold in Louisiana, and mainly eaten by American Indians. It had orange flesh and a strong fishy taste.[7] Elsewhere, the flesh has been described as appealing on account of the crayfish diet, and both members of the genus Eudocimus have been hunted, which has been responsible for decline, across their range.[8] Crawfish farmers in Louisiana have also shot them for foraging in crawfish ponds. Overall, the impact of hunting is not thought to be major.[27]

The pollutant methylmercury is a globally distributed neurotoxin and an endocrine system disruptor. In the Everglades ecosystem, human pollution has led to increased concentrations of methylmercury, which have impacted the behaviors of the American white ibis.[67] Hormone levels in males are affected, leading to a decrease in the rates of key courtship behavior, and fewer approaches by females during the mating season.[68] In addition, methylmercury also increased male-male pairing behaviors by 55%. Both the chemically induced "homosexual" behavior and the diminished ability to attract females by males have reduced reproduction rates in affected populations.[68] Exposure of American white ibises to methylmercury causes reduced foraging efficiency[69] and it also makes them more likely to abandon nests owing to the disruptive effect of the pollutant on the bird's hormone systems, which in turn affects parental care behavior.[67] Tests on captive birds have not shown a decreased survival of American white ibis exposed to methylmercury.[70]

In culture[edit]

Native American folklore held that the bird was the last to seek shelter before a hurricane, and the first to emerge afterwards. The bird was thus a symbol for danger and optimism.[71]

The University of Miami adopted the American white ibis as its official athletics mascot in 1926,[72] and the yearbook was known as The Ibis from that year.[71] The mascot was initially known as Ibis before adopting the name Sebastian in 1957. It was named after San Sebastian Hall, a residence hall on campus, which sponsored an Ibis entry in the college's homecoming celebration.[73]

References[edit]

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