About Tricolored Blackbirds

A summary of Tricolored Blackbird natural history, efforts to conserve the species, its legal status, and relevant literature.

Biological Profile

Distinguishing Characteristics

Appearance
Tricolored blackbirds look very much like the abundant and widespread red-winged blackbird (Agelaius phoeniceus), to which they are closely related. The plumage of adult male tricolors is glossy black and may sometimes show an iridescent blue-green sheen in bright sunlight. Adult females are brown dorsally and have brown streaks on their breasts that merge into a dark brown belly, unlike the belly of female red-winged which is streaked. Females may or may not have a brightly-colored epaulet. Juveniles are similar to females but paler (Pyle 1997). After-hatch year (1 year old) males are dark brown, not black, and have orange, not red, epaulets. Adult male epaulets are typically scarlet red, unlike the epaulet of the male red-wing which is orange-red. The band below the epaulet is bright white, not yellow or buffy as in the red-winged blackbird. Breeding males average 65-68 grams and breeding females average 40-45 grams. Total body length is 18-24 cm.

 

Male tricolored blackbird
Displaying male tricolored blackbird.

The bills of both sexes are long, narrow, round, and pointed (Beedy and Hamilton 1999). Both sexes have narrow, pointed wings with primary 9 longer than primary 6 (Mailliard 1910, Pyle 1997), giving the bird a flight profile that has more pointed wings than does the red-winged blackbird.  These different flight profiles can readily be seen by experienced observers (Mailliard 1910, Orians 1961). Flight typically undulating, with powered down-strokes followed by upward glides.  Foraging birds fly out from and return to the breeding colony head-to-tail, "follow the leader", and form long flightlines that may persist and allow the birds to be seen at relatively great distances.  Foraging flocks appear to roll across the landscape as birds in the rear of foraging flocks fly up to the front.  Foraging flocks may consist of hundreds or, in rare cases, even thousands of birds and be seen from some distance, although foraging flocks consisting of thousands of birds have not been reported since prior to 2020 as the number of breeding birds, even in large colonies, has declined. 

Vocalizations
Males may be highly vocal, females silent except at breeding colonies when they may chatter conspicuously or when disturbed, when they may shriek in brief, high-pitched single notes. Males may be silent in flight but when leaving breeding colonies may respond en masse to a conspicuous high-pitched, unmusical call as they depart.  Occasionally males produce a distinctive "wuk" call (Collier 1968). During settlement and early incubation, males typically utter a strange, cat-like mewing call that is very unlike the sounds produced by other blackbirds. Males sing while courting females, and in larger colonies this produces a very loud song chorus that may be heard at least a kilometer away.

Subspecies

There are no identified subspecies, although the tricolored blackbird has two fairly distinct population segments: a Southern California population that occurs in southern California south of the Tehachapi Mountains and into northern Baja California, and a Central Valley population segment that occurs throughout the Central Valley. Banding studies involving in excess of 100,000 birds have documented a very small rate of movement of individuals between these population segments, with a total (in early 2021) of only 5 birds known to have moved from one population segment to the other.

Distribution

Historical Breeding Distribution
Formerly abundant both in coastal California, where it was considered the most abundant bird from Los Angeles to San Diego in the mid-19th Century (Cooper cited in Baird and Cooper 1859), and in the Central Valley of California through interior southern California to northern Baja California (Grinnell 1915). Reported to breed in small numbers in southern Oregon (Neff 1937) and at scattered sites in eastern Washington.

Current Breeding Distribution
Since 1980, breeding has occurred in 46 California counties (Beedy and Hamilton 1999), although the species has essentially been extirpated as a breeder in coastal locations (Kelsey 2008, Meese 2017) and persists in very small numbers (12,000 in 2017; Meese 2017) at widely scattered sites in southern California (Unitt 2004). Gaps also occur in the breeding distribution in the Central Valley (e.g., Meese 2017). Small colonies of tens of individuals persist in northern Baja California (Erickson et al. 2007, 2008, Erickson in Meese 2017), a single colony of fewer than 30 individuals breeds in western Nevada (Ammon and Woods 2006), formerly up to two thousand birds breed at several scattered sites in Oregon (Marshall et al. 2003), and since 1998 a few tens of birds have bred in a few locations near Walla Walla, Washington (Wahl et al. 2005).

Non-breeding Distribution
Resident in Washington, Oregon and Baja California, absent in Nevada. May occur in small numbers along coastal regions and portions of northern California outside of the Central Valley. Wintering populations concentrate in the Sacramento-San Joaquin Delta and formerly along the central coast in Monterey, Marin, and Sonoma counties, where they occur in multispecies flocks with other blackbirds, brown-headed cowbirds, and European starlings (Beedy and Hamilton 1999). Wintering tricolors are associated with open rangeland in the Sacramento-San Joaquin Delta and with dairies along the central California Coast. From mid-October to February flocks to 25,000 frequented dairies on the outer Point Reyes Peninsula in Marin County (Beedy and Hamilton 1999), but few birds (< 1,000) have been seen during winter there since 2012 (Evens, pers. obs., Meese, pers. obs.). A wintering population estimated at 50,000 birds was observed by Ted Beedy and Bob Meese outside Birds Landing, Solano County in late October, 2007 but only 1,500 birds were observed wintering in the same location in winter 2020/2021 and only 800 birds during the winter of 2023-2024 (Meese, pers. obs.).  Small numbers (< 100) of wintering birds have been observed in western Idaho in 2022-2023 and 2023-2024.

The precise locations of and habitats utilized by wintering tricolors are poorly known and there is great need for extensive study of the distribution and abundance of this species throughout the winter.

Ecology

Territoriality
Often described as nonterritorial, but both males and females maintain breeding territories held for the duration of a single nesting effort. Territories include only nesting space, not foraging areas, and range in size from less than one to several square meters, with most territories between 2 and 6 square meters (Beedy and Hamilton 1999). In some instances, especially in especially high nest densities, tricolor territories may be vertically stacked (e.g., Hamilton and Meese 2005).

In contrast, the closely-related red-winged blackbird is highly territorial and during the breeding season males defend territories that include two to several females as well as some foraging habitat. Thus, red-winged blackbirds are quite sparsely distributed across the landscape during the breeding season while tricolored blackbirds tend to form large, dense colonies during the breeding season and it is this difference in behavior that most clearly distinguishes the two species for most people.

Breeding
Males may arrive on the breeding sites 1-3 days before the females, typically in mid-to-late March in southern California and the San Joaquin Valley (DeHaven 1975b) although the onset of breeding has occurred as early as late February since 2014 and has been shown to be advancing (Holyoak et al., unpub.). The tricolor typically nests twice, and the second breeding attempt often occurs in a different, more northerly location (Hamilton 1998) for Central Valley breeders, although when breeding conditions permit, second breeding attempts may occur in the same or immediately adjacent locations (Meese 2006, 2007, 2008). Comparable movements have not been reported in southern California, where the species is believed to be resident.

In most cases, one male breeds with two females in each breeding attempt. Pair bond persistence from for one breeding attempt to another has not been studied. Tricolors may be site-faithful, and if landscape conditions permit, will return to breed in the same locations year after year (Meese, unpub. data), but much within-year between-colony movement has been documented (Wilson, Meese, and Wyckoff, 2016, Meese, unpub.).

Nesting Substrate
The tricolor builds its nests on a variety of substrates, although most of these are either flooded or armored. Historically, most colonies were established in freshwater marshes dominated by cattails (Typha spp.) and bulrushes (Scirpus or Schoenoplectus spp.), with willows (Salix spp.) and nettles (Urtica spp.) also common (Neff 1937). The introduced mustards (Brassica spp.), Himalayan blackberry (Rubus armeniacus), thistles (Circium spp.), and cheeseweed mallow (Malva parviflora) have been commonly used for several decades.

Since the 1980's, the largest colonies have typically formed in grain (almost exclusively triticale, a hearty wheat x rye hybrid) fields in the San Joaquin Valley, especially so-called "dirty fields" that have relatively large amounts of invasive mustards or mallows.

Colony in triticale adjacent to dairy (Kern County).
Colony in triticale, Kern County

Nest
The tricolor builds an open cup nest out of long leaves of a variety of plants. In most cases, the leaves are collected dry, then taken to a nearby water source, placed in the water for several seconds until thoroughly wet, and then taken to the nest site (Meese, pers. obs.). The leaves are woven around stout, upright stems of many plant species, at heights of only a few cm to 2 meters above ground or water level, with ca. 1 meter being typical. The female builds the nest alone, with males only very rarely providing nest materials to the females. Nest-building takes about 3 days. Coarse materials are woven around the stems to produce a platform, then similar materials are used to create the sides, a mud bottom is added, and when this dries, very fine materials are placed on top of the mud to provide a soft substrate for the eggs.

Eggs
Typically, four eggs are laid, with 3-egg clutches also common (the most common number for second clutches in a season), and 5-egg clutches rare. The eggs are oval, typically light blue but occasionally light green, with dark reddish-brown splotches concentrated on the blunt end. Egg length averages 24-25 mm, and egg weight averages 2.5-5 grams. Egg laying typically begins the day after nest completion, and one egg is laid per day (Beedy and Hamilton 1999).

Incubation
Incubation of eggs is exclusively by the females and begins with the laying of the first egg (Beedy and Hamilton 1999). Eggs are incubated for approximately 12 days, exceptionally to 14 days (Payne 1969).

Nestling Growth and Development
The young are born naked and blind. Young weigh 2-3 grams at birth (Beedy and Hamilton 1999). Nestlings typically fed for 11-12 days; feeding by both parents. Young fed almost exclusively animal prey from birth until day 9, after which a mixture of both plant and animal foods are provided (Crase and DeHaven 1977; Skorupa, Hothem, and DeHaven 1980; Meese pers. obs.).  Tricolors are brood-reducers, meaning that the females remove "surplus" newly-hatched eggs and young from nests in environmental conditions which they assess to be insufficient to enable them to find enough insect food to rear all of the young from the eggs that hatch.  Carcasses of these discarded young can sometimes be quite conspicuous and easily found around the perimeters of nesting substrates.

brood-reduced young
Carcass of brood-reduced young not yet fully hatched from egg.

Fledging
Young typically fledge at day 10-14. Fledglings remain in or on periphery of colony for several days post-fledging, actively and quite vocally soliciting food from adults. Both parents feed fledglings. In many cases, adults lead fledglings away from the colony by returning to the fledgling with food, but instead of feeding the fledgling, fly away from the colony with the fledgling in pursuit.

Sue Liberto photo.
Adult female and fledgling.  Sue Liberto photo.

Diet and Foraging
Tricolored blackbirds, like other blackbird species, are fundamentally granivores (Orians 1980); however, tricolors consume a wide variety of plant and animal foods (Skorupa, Hothem, and DeHaven 1980), and respond opportunistically to the most abundant, readily available food resource (Payne 1969, Meese 2013). In the San Joaquin Valley, most tricolors feed on stored grains associated with dairies, with additional animal foods taken from both aquatic and terrestrial habitats up to 9 km from the colony. Birds may forage over a wide area, but females forming eggs and adults feeding nestlings typically concentrate foraging efforts on small, highly productive habitats, including shrublands (often excellent sources of caterpillars), pasturelands (many types of insects, but especially grasshoppers during grasshopper "breakout" years), wetlands (aquatic insect larvae), and rice paddies (aquatic insect larvae).

Sources of Mortality
Exposure to severe weather can cause massive mortality to adult females, eggs, and nestlings (Beedy and Hamilton 1999, Meese pers. obs.). Intense rainfall and strong winds are the two primary weather-related mortality sources, but colony desertion may occur when daytime temperatures exceed 38C (100F) for three or more days in a row.  These colony abandonment events used to occur in mid-summer, but the recent trend has been for earlier heat-related abandonment events and in 2021, a 3-day heat wave began in late May that resulted in the abandonment of a settlement of 20,000 birds at Colusa National Wildlife Refuge in Colusa County, CA (Isola, Carpenter, and Meese, pers. obs.). The tricolor has a large number of predators, and predation has been implicated in colony desertion (Beedy and Hayworth 1992, Hamilton 1998, Hamilton and Meese 2005). Serious tricolored blackbird predators include black-crowned night heron (Nycticorax nycticorax), white-faced ibis (Plegadis chihi), raccoon (Procyon lotor) and river otter (Lontra canadensis) in wetland colonies and coyote (Canis latrans), cattle egret (Bubulcus ibis; Meese 2011), and great blue heron (Ardea herodias) in upland and silage (triticale) colonies. Additional predators include red-tailed hawk (Buteo jamaicensis), Swainson's hawk (Buteo swainsonii), peregrine falcon (Falco peregrinus), northern harrier (Circus cyaneus), great horned owl (Bubo virginianus), northern raven (Corvus corax), Cooper's hawk (Accipiter cooperii; Beedy and Hamilton 1999), and California king snake (Lampropeltis getula).

Lifespan
Demographic information is only now becoming available as until recently relatively few banded birds have been recovered to make generalizations. Work by Neff (1942) and DeHaven and Neff (1973) suggests a maximum lifespan of at least 12 years, while Kennard (1975) suggests a 13 year lifespan. These are most likely to maxima for males, as they are about 50% larger than females and likely live longer. Band returns analyzed by Meese (unpub. data) suggest an average annual adult survivorship of 60%.

Females are believed to breed at one year while males are believed to defer breeding until their second year (Orians 1961; Payne 1969).

Conservation

Conservation

The tricolored blackbird has been the subject of intensive study for several decades due to observed declines and absences from portions of their range, and the Tricolored Blackbird Working Group produced a Conservation Plan in 2007 to guide conservation efforts.

Decline in Abundance

The original, pre-Columbian abundance of the tricolored blackbird is unknown and likely unknowable. The tricolor was described as the most abundant bird species in coastal southern California, in the region from Santa Barbara through San Diego, at the end of the 19th Century (Cooper in Baird and Cooper 1859, Unitt 2004). The first intensive studies dedicated to estimating tricolored blackbird abundance began in the 1930's in response to criticism of the Bureau of Biological Survey (the forerunner of the U.S. Fish & Wildlife Service) for failure to respond to perceived widespread reductions in abundance prior to 1931 (Neff 1937). Neff conducted six years of field work across large portions of the species' range, from the northern Sacramento Valley to southern California, and his abundance estimates were derived from counts of nests during or following the breeding season. Neff (1937) and his assistants studied breeding tricolored blackbirds and found single colonies consisting of up to 282,000 nests, with a maximum number 491,000 nests from a sample of 8 colonies in 1934. Extrapolating from Neff's estimates, one may conservatively estimate the number of tricolored blackbirds as in the range of 2-3 million birds in the 1930's.

DeHaven and colleagues looked at the abundance of tricolors in the early 1970s and concluded that the species had declined by at least 50% in the 35 years since Neff's work. Subsequent field workers tried to estimate the statewide population of tricolors via a volunteer-based survey originally conceived and coordinated by Ted Beedy and Bill Hamilton. These so-called Tricolored Blackbird Statewide Surveys attempted to detect and estimate the sizes of all colonies across the entire state during a 3-day interval in April, when the birds are most reliably counted. These statewide surveys began in 1994 and from 1994 to 2005 produced estimates of numbers of tricolors in the range of 250,000 birds (i.e. fewer birds than had been estimated from a single colony by Neff in the 1930's). There was much variation in the level of effort and in geographic completeness among the 1994-2005 Statewide Surveys, however, and trying to deduce a population trend by comparing the results of these surveys is difficult.

The 2008 triennial Statewide Survey, which differed from previous Statewide Surveys by having 1) county coordinators, 2) data management, and 3) data entry via this Tricolored Blackbird Portal, estimated the statewide population at approximately 400,000 birds.  The 2011 Statewide Survey estimated that only 258,000 birds remained, the 2014 the statewide population estimate was only 145,000 birds, but the 2017 statewide population estimate went up to 177,600 birds (Meese, 2017) and the 2022 Statewide Survey statewide population estimate went up to 218,000.

Reasons for the Decline

The reasons for the decline of tricolors are many, but include:

  1. habitat losses due to water diversions and draining of wetlands
  2. habitat losses due to conversion to agriculture
  3. habitat losses due to urbanization
  4. deliberate shooting for sale at markets ("market-hunting")
  5. destruction of breeding colonies in grain fields, most often adjacent to dairies, during normal agricultural operations
  6. deliberate shooting and poisoning as agricultural pests

Taken together, these factors have reduced the available breeding and foraging habitats by transforming the landscapes in which the birds evolved, and reduced the numbers of adults by shooting and poisoning during and following the breeding season. Although deliberate shooting and poisoning were prohibited in the 1970's and 1980's, habitat losses continue.

More recently, and continuing to the present, are three additional factors that affect productivity of tricolors:

  1. replacement of annual and perennial crops (primarily alfalfa and sunflowers) and shrublands, which often support large insect populations and therefore provide excellent food sources, by woody perennial crops (vineyards and nut [almond, walnut, and pistachio] orchards), which are insect-poor and do not provide foraging opportunities; this process severely restricts essential foraging habitats surrounding potential nesting sites and may prevent the utilization of otherwise suitable nesting substrate;
  2. predation by large numbers (e.g., 125 or more in a single colony) of cattle egrets, a voracious predator that was unknown in tricolor colonies until first observed in 2006, but that has subsequently become a serious threat to large, potentially highly productive colonies in the southern San Joaquin Valley;
  3. harvest of the grain fields, or chopping of weedy fields, used as nesting substrate while eggs or young are still in the nests; this causes catastrophic failure of entire breeding efforts that may affect thousands to tens of thousands of birds.

This video was filmed in Merced County in 2006 and shows the destruction of a Tricolored Blackbird colony during the harvest of a grain field.

 

Conservation Actions

Several conservation actions have been undertaken to stem the decline of the tricolored blackbird and to rebuild its numbers to sustain the species well into the future; these include:

  1. annual detection and monitoring of colonies in the Central Valley; this effort provides for the detection of the largest colonies in the San Joaquin and Sacramento Valleys and near-real-time monitoring of their fates
  2. conservation of the largest "silage colony" (i.e. nesting colony established in a triticale field in the San Joaquin Valley) through voluntary sale by the landowner and purchase by the state or federal government or non-governmental organization; a process known as a "silage buy-out" that was phased out due to cost in 2010
  3. conservation of "silage colonies" by harvest delay; this procedure involves paying a willing landowner to delay the harvest of his grain field occupied by breeding birds until the birds have finished breeding and have left the site; this procedure is desirable because it reimburses landowners for costs associated with the harvest delay (and consequent reduction in value of the silage) while allowing them to harvest their crop; this process replaced silage buy-outs, which were far more expensive
  4. provision and maintenance of attractive nesting substrate on National Wildlife Refuges and other protected areas and on Tricolored Blackbird Conservation Easements on private ranches; the provision and proper maintenance of secure nesting substrate is an essential component of the long-term strategy to conserve the birds; nesting birds forage widely around the colony, on both protected land and private property
  5. assisting private property owners to provide nesting substrate
  6. provision of "lure grains", typically rolled corn, on the periphery of colonies to provide grains for birds that would otherwise consume grains from nearby dairies
  7. triennial statewide survey; every 3 years, a volunteer statewide survey is performed to comprehensively monitor tricolor distribution and abundance
  8. education and outreach; brochures have been produced and distributed which educate landowners and agency personnel to the status of the tricolor and to efforts being made to conserve it; in addition, good working relations have been established with many landowners who permit monitoring of colonies on their properties; presentations to local Audubon chapters and similar conservation organizations provide additional education and outreach opportunities
  9. banding; by summer 2022, over 106,000 tricolors have been banded by Bob Meese and his assistants since beginning to band the birds in 2007; these efforts have documented spatial and temporal patterns of movements and provided estimates of life history parameters (publications available in Tricolored Blackbird Literature under the About menu)
  10. web portal; this web portal provides current, documented information on tricolors and access to data and reports provided by tricolor researchers, as well as web-based data entry capabilities for field workers and participants in the triennial Statewide Surveys

Legal Status

Federal Status

The tricolored blackbird is federally protected under the Migratory Bird Treaty Act, or MBTA (https://www.fws.gov/migratorybirds/RegulationsPolicies/mbta/mbtintro.html). Formerly, the tricolored blackbird was exempt from the protections offered to many North American birds by the Migratory Bird Treaty Act, as most blackbirds and relatives ("yellow-headed red-winged, rusty, and Brewer's blackbirds, cowbirds, all grackles, crows, and magpies") are considered potential agricultural pests and exceptions to protection under the MBTA are possible in cases where birds are about to cause depredations "to ornamental or shade trees, agricultural crops, livestock, or wildlife, or when concentrated in such numbers and manner as to constitute a health hazard or other nuisance" (50 CFR 21.43). However, in 1985, the tricolored blackbird was removed from the list of exempted species and now is fully protected under the MBTA (50 CFR Part 10 13708).

Currently, the tricolored blackbird is considered a Species of Conservation Concern (http://www.fws.gov/migratorybirds/reports/BCC2002.pdf).  On August 15, 2019, a listing under the Endangered Species Act was found to be unwarranted due to "high nesting success in both small and large colonies" and existing regulatory mechanisms such as the California Endangered Species Act that "are currently acting to ameliorate the severity of some existing threats" (https://federalregister.gov/d/2019-17536).

State Status

The tricolored blackbird was given emergency Endangered status under the California Endangered Species Act in December, 2014. This listing provided temporary (6 month) protection but was allowed to expire in June, 2015. The California Fish and Game Commission voted to list the tricolored blackbird as Threatened under CESA at its April 19, 2018 meeting in Ventura and was formally listed as Threatened under CESA on March 18, 2019.

See the list of State and Federally Listed Endangered and Threatened Animals of California at: http://www.dfg.ca.gov/biogeodata/cnddb/pdfs/TEAnimals.pdf.

BLM

The Bureau of Land Management designates the tricolored blackbird a Sensitive Species (BLM 2003; http://www.blm.gov/nhp/efoia/nv/ib/2003/nvib2003-097a.pdf).

IUCN

Not a regulatory agency, the World Conservation Union, or IUCN (http://www.iucn.org/), maintains lists of threatened and endangered species world-wide. The tricolored blackbird was placed on the IUCN Red List as Endangered in 2006 (http://www.iucnredlist.org/details/22724196/0).

Tricolored Blackbird Literature

Airola, DA et al. 2015.  Tricolored blackbird breeding status in 2015 in the Foothill Grasslands of the Sierra Nevada, California.  CVBC Bulletin 18: 96-114.  Airola et al. 2015.pdf

Airola, DA, Meese, RJ, Krolick DE. 2015. Tricolored blackbird conservation status and opportunities in the Sierra Nevada Foothills of California.  CVBC Bulletin 17: 57-78.  Airola, Meese, and Krolick 2015.pdf

Airola, DA et al. 2016.  Breeding status of the Tricolored Blackbird in the grassland-dominated region of the Sierra Nevada, California in 2016.  CVBC Bulletin 19: 82-109.  Airola  et al. 2016.pdf

Airola, DA et al. 2018.  Breeding status of the Tricolored Blackbird in the foothill grasslands of the Sierra Nevada, California, in 2017.  CVBC Bulletin 21: 1-24.  Airola et al. 2018.pdf

Airola, DA, Harris, J, Ross D. 2018.  Tricolored blackbird breeding status in the Central Sierra Nevada Foothills, California, in 2018.  CVBC Bulletin 21: 66-74.  Airola, Harris, and Ross 2018.pdf

Airola, DA, Young, L. 2015. Protecting nesting habitat for the Tricolored Blackbird on private rangelands in the foothill grassland region of the Sierra Nevada. CVBC Bulletin 17: 116-121.  Airola and Young 2015.pdf

Ammon, EA, Woods J.  2008.  Status of Tricolored Blackbirds in Nevada. Great Basin Birds 10: 63-66.  Ammon and Woods 2008.pdf

Anderson, DW, Hamilton, WJIII, Bowen, R, Cahill, TM, Herzog, SK. undated.  Large, recent colonies of nesting tricolored blackbirds in Northern Baja California, Mexico.  Anderson et al. undated.

Anonymous.  1991. Tricolored Blackbird Statewide Survey, 1991 (unpublished).

Anonymous.  1994.  Tricolored Blackbird Statewide Survey, 1994 (unpublished).

Anonymous.  1997.  Tricolored Blackbird Statewide Survey, 1997 (unpublished).

Anonymous.  1999.  Tricolored Blackbird Statewide Survey, 1999 (unpublished).

Avery, ML, DeHaven RW.  1982.  Bird damage to sunflowers in the Sacramento Valley, California. Tenth Vertebrate Pest Conference: 197-200.  Avery and DeHaven 1982.pdf

Barr, K. 2023. Expanding genomics toolkits to facilitate the conservation and recovery of two protected species, the Tricolored Blackbird and Burrowing Owl. Dissertation, UCLA. Barr 2023.pdf

Barr, K, Beichman, AC, Kalhori, P, Rajbhandary, J, Bay, RA, Ruegg, K, Smith, TB. 2020.  Persistent panmixia despite extreme habitat loss and population decline in the threatened tricolored blackbird (Agelaius tricolor). Evolutionary Applications 2020;00:1–11.  Barr et al 2020.pdf

Beedy, EC, Hamilton WJ.  1997.  Tricolored Blackbird status update and management guidelines.  Beedy and Hamilton 1997.pdf

Beedy, EC, Hamilton WJ.  1999.  Tricolored blackbird (Agelaius tricolor). The Birds of North America. 423

Beedy, EC, Hamilton, WJ III, Meese, RJ, Airola, DA, Pyle, P. 2018.  Tricolored Blackbird (Agelaius tricolor) version 3.1 In: The Birds of North America (P. G. Rodewald, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America: https://birdsna.org/Species-Account/bna/species/tribla

Beedy, EC, Hayworth A.  1992.  Tricolored blackbird nesting failures in the Central Valley of California: general trends or isolated phenomena? Endangered and sensitive species of the San Joaquin Valley, California: 33-46.  Beedy and Hayworth 1992.pdf

Beedy, EC, Sanders SD, Bloom D.  1991.  Breeding status, distribution, and habitat associations of the Tricolored Blackbird (Agelaius tricolor) 1859-1989.  Beedy, Sanders, and Bloom 1991.pdf

Beedy, EC.  1989.  Draft habitat suitability index model, Tricolored blackbird (Agelaius tricolor).

Bent, AC.  1958.  Life histories of North American blackbirds, orioles, tanagers, and allies.

Berg, EC, Pollinger, JP, Smith TB. 2010. Population structure of the Tricolored Blackbird (Agelaius tricolor) in California: Are northern and southern populations genetically distinct? Berg Pollinger and Smith 2010.pdf

California Department of Fish and Wildlife.  2018.  Report to the Fish and Game Commission.  A Status Review of the Tricolored Blackbird (Agelaius tricolor) in California.  CDFW Tricolored Blackbird Status Review.pdf

California Farm Bureau Federation. 2020.  Annual Report: Tricolored Blackbird Voluntary Local Program 2019.  California Farm Bureau Federation 2020.pdf

Center for Diversity. 2004.  Petition to list Tricolored Blackbird under the State and Federal Endangered Species Acts and Request for Emergency Action to Protect the Species.

Churchwell, R, Geupel GR, Hamilton WJ, Schlafmann D.  2005.  Current monitoring and management of tricolored blackbirds. USDA Forest Service: 169-173.  Churchwell, Geupel, Hamilton, and Schlafmann 2005.pdf

Clark, KB, Rideout, B, Garrett, KL, Unitt, P, OConnor, B. 2019.  Historical and geographical patterns in Knemidokoptes mite infestations in Southern California birds.  Western Birds 50: 26-36.  Clark et al. 2019.pdf

Colibri Ecological Consulting, LLC. 2022.  2022 Tricolored Blackbird Statewide Survey.  Fresno, CA.  Prepared for California Department of Fish and Wildlife, Sacramento, CA. 2022 Tricolored Blackbird Statewide Survey Final Report.pdf

Collier, G.  1968.  Annual cycle and behavioral relationships in the Red-Winged and Tricolored Blackbirds of southern California. Ph.D. Dissertation.  Collier 1968.pdf

Cook, L.  1996.  Nesting adaptations of Tricolored Blackbirds (Agelaius tricolor). M. S. Thesis.  U.C. Davis, Division of Environmental Studies.  Cook 1996.pdf

Cook, L.  1999.  Tricolored Blackbird (Agelaius tricolor) conservation plan for southern Sacramento County.

Cook, L, Toft C. 2005.  Dynamics of extinction: population decline in the colonially-nesting tricolored blackbird, Agelaius tricolor.  Bird Conservation International 15: 73-88.  Cook and Toft 2005.pdf

Cook, R. 2010.  Recent history and current status of the Tricolored Blackbird in Southern California.  Western Riverside County MSHCP Biological Monitoring Program  Cook 2010.pdf

Cooper, JG.  1870.  Ornithology of California.

Cousens, BF, Meese, RJ, van Vliet, HEJ, Shapiro, AM, Forister, ML, Thorne, JH, Waetjen, DP, Casner, KL, and Airola, DA. 2016.  Declines in Butterflies and Insectivorous Birds in Areas of High Insecticide Use in California.  Poster presented at North American Ornithological Conference, Washington, DC.  Cousens et al. 2016.pdf

Crase, FT.  1975.  Analysis of rice field test of methiocarb for reducing blackbird damage to ripening rice in the Sacramento Valley, California.

Crase, FT, DeHaven RW.  1977.  Food of nestling tricolored blackbirds. Condor 79: 265-269.  Crase and DeHaven 1977.pdf

Crase, FT, DeHaven RW.  1978.  Food selection by five sympatric California blackbird species. California Fish & Game. 64: 255-267.  Crase and DeHaven 1978.pdf

DeHaven, RW.  1971.  Blackbirds and the California Rice Crop. The Rice Journal. :7-14.

DeHaven, RW.  1974.  Bird damage appraisal methods in some agricultural crops. 6th Vertebrate Pest Conference.

DeHaven, RW.  1975.  Plumages of the tricolored blackbird. Western Bird Bander 50: 59-61.

DeHaven, RW.  2000.  Breeding Tricolored Blackbirds in the Central Valley, California: A Quarter-Century Perspective.  DeHaven 2000.pdf

DeHaven R. W., Schafer J.  1971.  Methiocarb for repelling blackbirds from ripening rice. International Rice Commission Newsletter: 25-29.

DeHaven, RW, Crase FT, Woronecki PD.  1975.  Breeding status of the tricolored blackbird, 1969-1972. California Fish & Game 61: 166-180.  DeHaven, Crase, and Woronecki 1975.pdf

DeHaven, RW, Crase FT, Woronecki PD.  1975.  Movements of Tricolored Blackbirds banded in the Central Valley of California, 1965-1972.  Bird-banding 46: 220-229.  DeHaven, Crase, and Woronecki 1975.pdf

DeHaven, RW, Neff JA.  1973.  Recoveries and returns of tricolored blackbirds, 1941-1964. Western Bird Bander. 48: 10-11.  DeHaven and Neff 1973.pdf

Emlen, JT.  1941.  An experimental analysis of the breeding cycle of the tricolored red-wing. Condor 43: 209-219.  Emlen 1941.pdf

Emlen, JT. 1985.  Morphological Correlates of Synchronized Nesting in a Tricolored Blackbird Colony.  Auk 102: 882-884.  Emlen 1985.pdf

Erickson, RA, de la Cueva, H.  2008.  Nesting Tricolored Blackbird Survey: Baja California 2008.  Erickson and de la Cueva 2008.

Erickson, RA, de la Cueva H, Billings MJ.  2007.  Nesting tricolored blackbird survey: Baja California 2007.  Erickson, de la Cueva, and Billings 2007.pdf

Erickson, RA, de la Cueva, H, Feenstra, JS, Zamora-Hernández, ED.  2016. On the edge of extinction: Can the Tricolored Blackbird (Agelaius tricolor) persist in Mexico?  Poster presented at North American Ornithological Conference.  Erickson et al. 2016 NAOC Poster.

Erickson, RA, de la Cueva, H, Zamora-Hernández, ED. 2021. Requiem for the Tricolored Blackbird in Mexico? Western Birds 52: 207-221; doi 10.21199/WB52.3.2.  Erickson, de la Cueva and Zamora-Hernandez 2021.pdf

Evermann, BW.  1919.  A colony of tricolored blackbirds. The Gull 1: 2-3.

Frayer, WE, Peters DD, Pywell HR.  1989.  Wetlands of the California Central Valley: status and trends 1939 to the mid-1980's.  U.S. Fish & Wildlife Service.

Graves, E, Holyoak, M, Kelsey, TR, Meese, RJ. 2013.  Understanding the contribution of habitats and regional variation to long-term population trends in tricolored blackbirds.  Ecology and Evolution 3: 2845-2858.  Graves, Holyoak, Kelsey, and Meese 2013.pdf

Graves, E, Meese, RJ, Holyoak, M. 2022.  Neonicotinoid exposure in Tricolored Blackbirds (Agelaius tricolor).  Environ. Sci. and Pollution Res.  https://doi.org/10.1007/s11356-022-23290-4.  Graves Meese and Holyoak 2022.pdf

Green, M, Edson L.  2004.  The 2004 Tricolored Blackbird April Survey. Central Valley Bird Club Bulletin 7(2 and 3): 23-31. 

Grinnell, J.  1915.  A distributional list of the birds of California. Pacific Coast Avifauna 11.

Grinnell, J, Miller AH.  1944.  The distribution of the birds of California. Pacific Coast Avifauna 27.

Hamilton, WJ III, Cook L, Grey R.  1995.  Tricolored blackbird project 1994.  Hamilton, Cook, and Grey 1995.pdf

Hamilton, WJ III, Meese RJ.  2005.  Habitat and population characteristics of tricolored blackbird colonies in California.  Hamilton and Meese 2005.pdf

Hamilton, WJ III.  1998.  Tricolored blackbird itinerant breeding in California. Condor 100: 218-226.  Hamilton 1998.pdf

Hamilton, WJ III. 2004.  Tricolored blackbird management recommendations and 2005 Survey priorities.  Hamilton 2004.pdf

Hamilton, WJ III. 2000.  Tricolored Blackbird 2000 Breeding Season Census and Survey - Observations and Recommendations.  Hamilton 2000.pdf

Harrison, C.  1978.  A field guide to the nests, eggs, and nestlings of North American birds.

Holyoak, M, Meese, RJ, Graves, E. 2014.  Combining site occupancy, breeding population sizes and reproductive success to calculate time-averaged reproductive output of different habitat types: an application to tricolored blackbirds.  PLOS One 9: e96980.  Holyoak, Meese, and Graves 2014.pdf

Hosea, R.  1986.  A population census of the tricolored blackbird, Agelaius tricolor (Audubon), in four counties in the northern Central Valley of California. M.S. Thesis, C.S.U.S. Department of Biological Sciences.

Jaramillo, A, Burke P.  1999.  New World Blackbirds: The Icterids.

Kelsey, R. 2008.  Results of the 2008 Tricolored Blackbird Census.  Sacramento: Audubon California.  Kelsey 2008.pdf

Kennard, JH.  1975.  Longevity records of North American birds. Bird-banding 46: 55-73.

Kyle, K, Kelsey R. 2011.  Results of the 2011 Tricolored Blackbird Statewide Survey.  Sacramento: Audubon California.  Kyle and Kelsey 2011.pdf

Lack, D and Emlen JT. 1939.  Observations on breeding behavior of tricolored red-wings.  Condor 16: 225-230.  Lack and Emlen 1939.pdf

Lanyon, SM. 1994.  Polyphyly of the Blackbird Genus Agelaius and the importance of assumptions of monophyly in comparative studies.  Evolution 48: 679-693.  Lanyon 1994.pdf

Liu, IA and Meese RJ.  Low genetic diversity in a population of Tricolored Blackbird (Agelaius tricolor), a species pending Endangered status.  Uploaded to bioRxiv preprint doi: https://doi.org/10.1101/2020.07.13.201574.  Liu and Meese 2020.pdf

Mailliard, J. 1900.  Breeding of Agelaius tricolor in Madera Co., Cal. Condor 2: 122-124.  Mailliard 1900.pdf

Mailliard, JW. 1910.  For the better determination of Agelaius tricolor.  Condor 12: 39-41.  Mailliard 1910.pdf

Maillard, J.  1910.  The status of the California bi-colored blackbird. Condor 12: 63-70.

Maillard, J.  1914.  Notes on a colony of tricolored red-wings. Condor 16: 204-207.

Marshall, DB, Hunter MG, Contreras AL.  2003.  Birds of Oregon: A General Reference.

McCabe, TT.  1932.  Wholesale poison for the red-wings. Condor 34: 49-50.

McCormick, AJ. 2022.  The tricolored blackbird in Washington State: First Assessment of Status and Distribution.  Western Birds 53: 186-198.  McCormick 2022.pdf

Meehan, T, Arthur, S. Michel, N, Wilsey, CB, Langham, GM. 2019.  Trends in tricolored blackbird colony size: 2008-2017.  J. Wildlife Management 1-7.  Meehan, Arthur, Michel, Wilsey, and Langham 2019.pdf

Meese, RJ.  2006.  Settlement and breeding colony characteristics of tricolored blackbirds in 2006 in the Central Valley of California.  Meese 2006.pdf

Meese, RJ.  2007.  Settlement, breeding, productivity and color-banding of tricolored blackbirds in 2007 in the Central Valley of California.  Meese 2007.pdf

Meese, RJ. 2008.  Detection, monitoring, and fates of Tricolored Blackbird Colonies in 2008 in the Central Valley of California.  Calif. Dept. of Fish and Game and U.S. Fish and Wildlife Service.  Meese 2008.pdf

Meese, R. J. 2009a.  Detection, monitoring, and fates of Tricolored Blackbird colonies in 2009 in the Central Valley of California.  Final Report submitted to California Dept. of Fish & Game, Sacramento, CA and U.S. Fish and Wildlife Service, Sacramento, CA. Meese 2009a.pdf

Meese, RJ. 2009b.  Contribution of the conservation of silage colonies to the conservation of tricolored blackbirds from 2005-2009.  Report submitted to U.S. Fish and Wildlife Service.  Meese 2009b.pdf

Meese, R. J. 2010.  Detection, monitoring, and fates of Tricolored Blackbird colonies in 2010 in the Central Valley of California.  Final Report submitted to California Dept. of Fish & Game, Sacramento, CA and U.S. Fish and Wildlife Service, Sacramento, CA. Meese 2010.pdf

Meese, R. J. 2011.  Reproductive success of Tricolored Blackbird colonies in 2011 in the Central Valley of California.  Final Report submitted to California Dept. of Fish & Game, Sacramento, CA. Meese 2011.pdf

Meese, RJ. 2012.  Cattle egret predation causing reproductive failures of nesting tricolored blackbirds.  Calif. Fish and Game 98: 47-50.  Meese 2012.pdf

Meese, RJ. 2013.  Chronic low reproductive success of the colonial tricolored blackbird.  Western Birds 44: 98-113.  Meese 2013.pdf

Meese, RJ. 2014. Results of the 2014 Tricolored Blackbird Statewide Survey.  Meese 2014.pdf

Meese, RJ. 2015.  Efforts to assess the status of the Tricolored Blackbird from 1931 to 2014.  CVBC Bulletin 17: 37-50.  Meese 2015.pdf

Meese, RJ. 2016 Detection, monitoring, and fates of Tricolored Blackbird colonies in California in 2016. Meese 2016.pdf

Meese, RJ.  2017.  Results of the 2017 Tricolored Blackbird Statewide Survey.  Meese 2017.pdf

Meese, RJ, Airola, D, Beedy, EC, Cook R. 2015.  Recovering the Tricolored Blackbird in California.  CVBC Bulletin 17: 97-109.  Meese, Airola, Beedy, and Cook 2015.pdf

Meese, RJ, Beedy EC. 2015.  Managing nesting and foraging habitats to benefit breeding tricolored blackbirds.  CVCB Bulletin 17: 79-96.  Meese and Beedy 2015.pdf

Meese, RJ, Simmons S. 2010.  Safe and effective for trapping and color banding tricolored blackbirds in the Central Valley of California.  Calif. Fish and Game 96: 23-35.  Meese and Simmons 2010.pdf

Meese, RJ, Yee, JL, and Holyoak, M.  2015.  Sampling to estimate population size and detect trends in Tricolored Blackbirds.  CVBC Bulletin 17: 51-56.  Meese, Yee, and Holyoak 2015.pdf

Neff, JA.  1937.  Nesting distribution of the tricolored red-wing. Condor 39: 61-81.  Neff 1937.pdf

Neff, JA.  1942.  Migration of the tricolored red-wing in central California. Condor 44: 45-53.  Neff 1942.pdf

Ohlendorf, HM, Hoffman DJ, Saiki MK, Aldrich TW.  1986.  Embryonic mortality and abnormalities of aquatic birds. Science of the Total Environment 52:49-63.

Orians, GH, Collier G.  1963.  Competition and blackbird social systems. Evolution 17: 449-459.  Orians and Collier 1963.pdf

Orians, GH.  1960.  Autumnal breeding in the tricolored blackbird. Auk 77: 379-398.  Orians 1960.pdf

Orians, GH. 1961.  The ecology of blackbird (Agelaius) social systems.  Ecological Monographs 31: 285-312.  Orians 1961.pdf

Orians, GH.  1961.  Social stimulation within blackbird colonies. Condor 63: 330-337.  Orians 1961.pdf

Orians, GH.  1963.  Notes on fall-hatched tricolored blackbirds. Auk 80: 552-553.  Orians 1963.pdf

Orians, GH.  1980.  Some adaptations of marsh-nesting blackbirds. Monographs in Population Biology 14.

Orians, GH, Christman GM.  1968.  A Comparative Study of the Behavior of Red-winged, Tricolored, and Yellow-headed Blackbirds.

Payne, R.  1969.  Breeding seasons and reproductive physiology of tricolored blackbirds and red-winged blackbirds. University of California Publications in Zoology 90: 1-137.

Payne, R, Landolt M. 1970.  Thyroid histology of Tricolored Blackbirds (Agelaius tricolor) in the annual cycle, breeding, and molt.  Condor 72: 445-451.  Payne and Landolt 1970.pdf

Phillips, R, McCluskey, J, Bradshaw, A, Higgins, P. 2015.  Calero Reservoir Tricolored Blackbird Nest Monitoring 2015 Report.  Report submitted to Santa Clara County Parks Department.  Phillips et al. 2015.pdf

Phillips, R, Bradshaw, A, Smith, K, Ackland, C. 2019.  Santa Clara County Tricolored Blackbird Nesting and Foraging Monitoring Project 2017-2018.  Final Report prepared for California Dept. of Fish and Wildlife and Santa Clara Habitat Agency.  Phillips et al. 2019.pdf

Robinson, O, Ruiz-Gutierrez, V, Fink, D, Meese, RJ, Holyoak, M, Cooch, EG. 2018.  Using citizen science data in integrated population models to inform conservation.  Biological Conservation 227: 361-368.  Robinson et al. 2018.pdf

Robinson, O, Ruiz-Gutierrez, V, Fink, D. 2017.  Correcting for bias in distribution modelling for rare species using citizen science data.  Diversity and Distributions 1-13.  Robinson, Ruiz-Gutierrez, and Fink 2017.pdf

Robinson, O, Ruiz-Gutierrez, V, Meese, RJ, Graves, EE, Holyoak, M, Wilson, CR, Wyckoff, AC, Merriell, BD, Snyder, C, Cooch, EG. 2021. Multi-scale demographic analysis reveals range contraction via pseudo-source and sink population structure. Ecosphere 12(5):e03521. 10.1002/ecs2.3521.  Robinson et al. 2021. pdf

Schackwitz, W., Airola, DA, Greene, A, Schackwitz, M Woodruff, J. 2022.  Bioacoustic monitoring reveals details of tricolored blackbird breeding phenology.  J. Fish & Wild. Mgt. 11: 518-530.  Schackwitz et al 2020.pdf

Skorupa, JP, Hothem RL, DeHaven RW.  1980.  Foods of breeding tricolored blackbirds in agricultural areas of Merced County, California. Condor 82: 465-467.  Skorupa, Hothem, and DeHaven 1980.pdf

Sloat, T.  2005.  South Sacramento Habitat Conservation Plan (2005): Appendix A.

Tricolored Blackbird Working Group. 2009.  Conservation Plan for the Tricolored Blackbird (Agelaius tricolor).  2.0 Update.  Conservation Plan 2009 2.0 update.pdf

Tyler, J.  1907.  A colony of tri-colored blackbirds. Condor 9: 177-178.  Tyler 1907.pdf

U.S. Fish & Wildlife Service. 1995.  Migratory nongame birds of management concern in the United States.

U.S. Fish & Wildlife Service. 2002.  Birds of conservation concern 2002.

Unitt, P.  2004.  San Diego County Bird Atlas. Proceedings of the San Diego Society of Natural History 39.

Wahl, TR, Tweit B, Mlodinow SG., Editors.  2005.  Birds of Washington: Status and Distribution.  Oregon State University Press, Corvallis, OR.

Weintraub, K, George TL, Dinsmore SJ. 2016.  Nest survival of Tricolored Blackbirds in California's Central Valley.  Condor 118: 850-861.  Weintraub, George, and Dinsmore 2016.pdf

Wilson, C, Meese, RJ, Wyckoff, AC. 2016.  Breeding chronology, movements, and life history observations of tricolored blackbirds in the California Central Coast.  Calif. Fish and Game 102: 162-174.  Wilson, Meese, and Wyckoff 2016.pdf