The Identity of Townsend’s Bunting

Mathew Louis

June 2, 2015

The name Emberiza townsendii was applied by Audubon to a single specimen collected by John Kirk Townsend in 1833 in New Garden, Pennsylvania. It is often referred as “Townsend’s Bunting,” and the details of the collection of this bird and its enigmatic status are well known.

A copy of Extinct Birds (Fuller, 2001; rev. ed.) in my library includes a color illustration of Audubon’s type (296) alongside a hypothetically-colored reconstruction by the artist (Julian Hume) of how that type may have appeared when first collected. While I had become familiar with Fuller’s book, and the illustration, since about the time of its publication, a notion occurred to me only recently which I feel is worth disseminating.

This is, very likely, a first-year or immature male Dickcissel x Lark Bunting hybrid (Spiza americana x Calamospiza melanocorys).

The illustration in Fuller is in copyright (but see www.extinction-website.com). Audubon figured the type (U.S. National Museum, #A10282) as plate 400, figure 4 in Birds of America [plate 157, 1841 edition.].

The most striking characteristic is the pale area along the folded wing, visible here in this, the decrepit specimen. The Lark Bunting’s tertiary and wing-coverts are white, and it is in Hume’s illustration that this character appears most obvious, whereas Audubon had not incorporated it in his portrait.

Comparing the above Townsend’s Bunting specimen to a male Lark Bunting specimen also conveys another peculiarity. Though the Lark Bunting’s undertail feathers are patterned with white panels on their outer extremities, the outer margins themselves are white. The image of the Townsend’s type likewise shows a margin to the outer right rectrix of a decidedly paler tone in contrast with the undertail itself. Audubon (Birds of America, 3: 62) had described the tail as “wood-brown, slightly edged with paler,…” This is quite different from the Dickcissel, which has a tail—an undertail—of a uniform brown without pale edges.

(Image source: www.miriameaglemon.com)

It is the Lark Bunting which, in immature or female plumages, has dark streaks along its crown (Byers, Sparrows and Buntings, pl. 17). As in our mystery bird, which is similarly marked (depictions of Audubon, Hume), the streaks appear as irregular flecks along the crown. The Dickcissel also bears a marked crown, but this is only true among immature or juvenile birds (Farrand, Western Birds (464)), and these marks tend to appear as evenly-arranged furrows. Though the streaks of the latter can also appear speckled if its crown is raised, there is a tendency for these rows of marks to be, nonetheless, nicely separated, and that aspect of this character also subtly distinguishes the Dickcissel from the other two. (See http://little-buffalo.com/tag/county/).

In the recent Extinct Birds (Hume & Walters, 2012, p. 346), the authors treat it as a hypothetical species combined in Spiza and provide these details of its description: “…eye-stripe, chin, throat, central line of underparts and edge of wing [emphasis mine] white; black-spotted line from lower corner of mandible down the side of throat, connecting with a crescent of streaks on upper edge of slate-blue breast.”

In ascribing the edge of wing as white, Hume and Walters are treating the type from a reconstructive perspective. Sharpe (Catalogue of the Birds in the British Museum, 12: 774 (1888)), likely working from American sources, described it in its actual state: “…the edge of the wing, and a gloss on the breast and the middle of belly, yellow.”

The Lark Bunting in immature or female plumages has discontinuous malar streaks, whereas in the Dickcissel the malar is a penciled line, with hardly an exception among individuals in the consistency of the marking. With respect to that character alone in the above image of the Townsend’s type, it may be an equivocal matter as to which species is represented by it.

The streaks on the breasts of the Townsend’s may actually be a correlating character of the Dickcissel, which has less streaking compared to the Lark Bunting, or limited streaking in younger birds.

Measurements [mm.] of Ridgway (Birds of North and Middle America, Part 1 (1901; 167, 171—175) on the Townsend’s and a series of the other two species also add to the argument.

Spiza townsendi [prevalent spelling of the Townsend’s]

Length: 146.05

Wing: 73.15

Tail: 53.59

Exposed culmen: 12.70

Depth of bill at base: 9.65

Tarsus: 20.07

Middle toe: 16.51

Ratio of exposed culmen to length—1: 11.50

Spiza americana

Length: 140.97—160.27 (148.08); female 139.70—145.80 (143.26)

Wing: 78.99—85.85

Tail: 55.12—61.47

Exposed culmen: 14.73—15.49 (14.99); female 12.70—14.22 (13.46)

Depth of bill at base: 10.41—11.43; female 9.91—10.67

Tarsus: 22.86—24.13

Middle toe: 16.76—18.03

Ratio of average female exposed culmen to average female length—1: 10.64

Calamospiza melanocorys

Length: 154.94—184.15 (163.32); female 144.78—165.10 (157.23)

Wing: 85.09—91.95

Tail: 65.53—71.12

Exposed culmen: 13.21—14.73 (13.97); female 12.70—13.21 (12.70)

Depth of bill at base: 10.67—12.19; female 10.16—11.94

Tarsus: 22.86—25.91

Middle toe: 16.76—18.03

Ratio of average female exposed culmen to average female length—1: 12.38

From this I can infer that the Townsend’s Bunting is appreciably larger than adult female Dickcissels, but as an immature or possibly adult individual (as the literature treats it as an adult), it averages much smaller than adult males of either species. For a (presumably) younger bird, it had already grown beyond the size of the largest female Dickcissels and in length is approaching that of the female Lark Bunting.

The most significant inference from the above is the comparative length of exposed culmen, the character where the Dickcissel averages greater than the Lark Bunting, as the former has a longer beak. The Townsend’s is on par with the smallest female Dickcissels, but this same figure is actually the mean of the female Lark Buntings. Even more compelling, culmen length is greatest in relation to body length in the Dickcissel, but the Townsend’s in this respect represents a significant departure from what would be expected if it was a pure Dickcissel, a figure midway between the two proposed parent species and even one approaching that of the Lark Bunting. [Female measurements are given as a comparative ratio on the basis of the small Townsend’s approaching females in size.]

Both the Lark Bunting and Dickcissel breed in the Great Plains region of the United States, the former a rare bird in Pennsylvania and the latter having been known to breed in the southern region of the state. Both are represented in rare records from Prince Edward Point National Wildlife Area (Prince Edward County, Ontario). From my own observations of the Lark Bunting, it is an irregular species. It will sometimes occur singly among flocks of other species (as I have observed one or small numbers among gambelii White-crowned Sparrows, Brewer’s Sparrows, or House Finches). The Dickcissel, too, is often an irruptive species and one which will turn up in single or small numbers among flocks of other species, notably House Sparrows. Given consideration of these arguments, it is plausible to argue that perhaps once in a great while the two may interbreed. Hybridization, despite the two species being in different respective subfamilies (Emberizinae, Cardinalinae) of the Emberizidae, these being treated in more recent works as separate families.

I am not aware if the literature has discussed hybridization between these species and assume no formal record has been published. Thus, I am arguing that each indiviual in question—Audubon’s type and the bird photographed in Ontario—represent legitimate hybrids and that further discussion on these two individuals should appropriate such a designation. Implying that there is but a singular instance of known hybridization of the Dickcissel, Denis Lepage (see link below for “Open Mic: The Townsend’s Bunting Story”) refers to a Dickcissel x Blue Grosbeak hybrid “with the name Spiza townsendii also attached to it.”

The discovery last year of a Dickcissel similar to the famed Townsend’s Bunting and having aberrant plumage characters deserves attention —

http://www.kyleblaney.com/photo-blog/

http://blog.aba.org/2014/09/open-mic-the-townsends-bunting-story.html

Reflection on this case gives me reason to present the arguments in this essay as more than speculation. This new bird not only adds to the discussion, it makes it very likely that it represents an instance—another—of hybridization. The superb photographs by Kyle Blaney appear in his blog, where he, and later Denis Lepage of the American Birding Association, argue for it being an aberrant color variant of the Dickcissel, and suggest the same is true for the Townsend’s. This argument I reject, and in a major irony, it is these very images which more than reinforce my own arguments, not only of this individual but of the original Townsend’s Bunting.

These photographs reveal a bird that is similar to the Townsend’s Bunting, but with obvious differences. No streaks on crown, a white mark below the eye, brown tertiaries only edged white, a continuous malar streak, gray of lower back the same color as the head. The tail appears to be of a uniform brown color, though the images do not reveal the acute detail of the edge of the underside.

These images also reveal a number of stunning aspects about the bird which show a definite correlation to Lark Bunting characters—

*While the tertiaries are brown in this bird, they are edged pale whitish, typical of immature Lark Buntings. To clarify, the Lark Bunting has white panels at the outer edge of the tertiaries which extend along the bend of the wing to the outer wing-coverts and alula feathers. The inner tertiaries are brown in young or female birds, edged paler.

*The wing-coverts of this individual are white with black chevrons in their centers—this an obvious character of immature Lark Buntings. The last of the images in sequence shows how this white area appears as panels on the outstretched wing.

*The beak is dark blackish above, and of a weak bluish tone on the lower mandible. This beak appears bicolored, and that tends to favor suggesting the Lark Bunting, as it is distinctively bicolored in all stages, each mandible a separate hue. While the shape of this bird seems lengthy and pointed like the Dickcissel’s, the immature Dickcissel has a more pale or flesh-colored beak and the adult’s is not truly bicolored—coloration is similar to the Lark Bunting but the blue almost always leads beyond the lower mandible to the base of the upper, with only the area at and approaching the culmen dark or blackish.

The lower eye is marked white with a faint whitish suffusion leading along the lores, and this is a character of the Dickcissel, along with the gray color of the back.

Reinforcing these arguments also is a belatedly published description of the Townsend’s Bunting type by John Kirk Townsend himself. The record of Townsend’s original notes were not published until 1909 (Auk, 26: 262—272). He carefully described something approaching a Lark Bunting beak: “[u]pper mandible black, middle edge white, lower light blue with a longitudinal line of black extending from the point half way to the base;…” He described it having “shoulders yellowish white,” a reference to the aforementioned panels.

Its song was compared to that of an Indigo Bunting Passerina cyanea (“a succession of lively notes somewhat resembling…but louder and more varied”), which is significant as the Dickcissel has a distinct harsh call from which its vernacular name is derived. A melodic bird, as conveyed by Townsend, would be an Indigo Bunting, but the characters of that species, as also with the Blue Grosbeak as Denis Lepage has argued, do not at all allow for a correlation to the characters of the mystery birds in question. So it is to the vocalizations of the Lark Bunting which consideration is afforded, and it is plausible to argue that Townsend may have been hearing the whistled phrases of one in song. Just years later, he would be first to describe the Lark Bunting as Fringilla bicolor, but in describing it (from flocks he and Thomas Nuttall watched) no recollection or comparison to the earlier specimen, particularly its vocalizations, was made. In his notes, he compared the earlier mystery in its plumage to the Gray-headed Sparrow Passer griseus, a species which was originally and erroneously believed to have been first collected in the United States.

Another character deserving note, if only that, are the stipples below the eye in Julian Hume’s reconstructive illustration of the type. Though Audubon’s portrait does not show this, he having it fresh, it is another trait common in the Lark Bunting. The specimen may be too worn to be able to examine this point further.

I feel that a hybrid Dickcissel x Lark Bunting is a better argument than that of a simple aberrantly-plumaged individual for both cases. Each bird represents, in its plumage characters, examples of both species, and for the sake of my arguments, it is the Lark Bunting which is adequately represented as a parent and most likely the one species from which crosses such as these would originate. Despite the assertions made, a Blue Grosbeak hybrid is unlikely as that species does not adequately betray characters in these mystery individuals, and an aberrantly-plumaged individual is unlikely as this is a type of bird with remarkable proportions unlike those of a pure Dickcissel. It is possible that the Ontario bird may have different proportions (or even be an adult), but in regards to an actual identity of what these individuals are, I feel that what holds in the identity in the original Townsend’s Bunting holds for this more recent example, one and the same. A hybrid argument is also reinforced in that there is considerable variation between both individuals in question. A DNA analysis of the Townsend’s Bunting type would ultimately rectify this mystery, and this essay presupposes a course of further action.

"The Sage Sparrow, like the Lark Bunting and other related sparrows, usually reveals a panel of white wing-coverts with dark chevrons in the center of each. Comparison from this mystery bird to the Sage seems plausible from that, but it is important to note the distinction between the panels of the Sage and Lark Buntings, which also reduces the question of the former representing a hybrid parent. In Blaney's bird, these markings are unequivocally black, as night/day against the white areas--exactly like that of the Lark Bunting; whereas, in the Sage Sparrow, the dark markings on its wing-coverts are never as black, but a dark grayish-brown or blackish-brown and somewhat dull. There is proportionately a much lesser amount of white."

Having run search queries on the net of “Emberiza townsendi” in conjunction with the following terms: hybrid, Calamospiza, “Fringilla bicolor,” I was unable to find precedent on the Lark Bunting being associated with the discussion on this form.

(Also see Audubon Society Master Guide to Birding, 3: 214—215, 246 and Threatened Birds (ICBP, 2000).

Record Status of Ancient Avifauna Database

Additional records are still being added to the ancient avifauna database because of research at Criss Library at the University of Nebraska at Omaha. Efforts continue to focus upon searching the variety of online historic newspapers, locating a pertinent article, and then printing it nearby the cubicle of the day. New website options and search methods have also meant a repeat of searches within known record sources, especially since it's now easier to evaluate, to an extent, whether found content is relevant or can be ignored.

Numerous unique records have been recently added that indicate significant details on occurrence and distribution of many species. Articles with interesting anecdotes pertaining to the history of ornithology have been documented. Key details embedded within this one-of-a-kind database as of the end of January include:

• more than 148,045 distinct occurrence records for more than 1700 species, according to International Ornithological Council taxonomy of 2013, and from Canada, all of the continental states, including the District of Columbia, along with Mexico and southward to the Panama Canal;
• a bibliography of 8215 items, with more than 500 newspaper articles or poems now posted online in their entirety;
• items from at least 900 different newspapers, which have, thus far, contributed more than 5,300 records, primarily from the continental United States; and
• 14,412 sites, though not all are distinct since some sites are repeated to allow a comparison of temporal differences.

These are the top-ten years for which records are currently available:

» 1882: 5547 records

» 1881: 5192

» 1883: 5144

» 1878: 4757

» 1880: 4659

» 1884: 4644

» 1875: 4330

» 1877: 4141

» 1876: 4036

» 1868: 3819

There are also more than 14,000 items in the game market prices datatable. Each item is separately documented. Additional details for this distinctive aspect of bird history are pending further research.

It is astounding to realize the extensive results of this endeavor. Years ago there was a sense that the records available at some particular time was the culmination of what could be achieved. This perspective quickly changed. In January 2008, the record count was about 25,000, which was supposedly thought to be significant at that time. By September of the same year, the tally was 50,000. The number of records continued to steadily increase, by increments of one record at a time, individually considered and documented. The tally was 75,000 records in February 2009, with 100,000 by November. It was 125,000 distinctive records in January, 2011 when there were only 3,000 items in the bibliography.

The focus on newspaper sources blossomed started in March, 2012 after an initial review of online sources. The bibliography was up to about 4,000. By June, 1000 newspaper articles had been found and dealt with in a documentary manner. By August of 2012, the tally was 2,000.

This quest continues unabated, thou sometimes stymied. There are still other records to be found from sources still unknown that will contribute to the tally of what could readily be 150,000 records. The effort will never be completed because there is no single-source to evaluate any record. It is rather a disparate bunch of stuff which does not allow comparison, evaluation and contribution. The wonderful variety of records associated with historic ornithology has not yet been embraced in an inclusive manner. Only through a cooperative effort will the vibrant old-time history of northern American birds be realized to its truest extent, with appreciation for every contributor.

Family contributions of some dollars have been essential to deal with project expenses, essentially to ensure having photocopies. The cost for printing at UNO recently increased to 7 cents per page from 5 cents, dramatically decreasing the value of a dollar.

The extent of boxes in the basement with copies continues to increase.

Others of the clan have helped with funds to get notebooks and sheet protectors to organize copies with a similar topic. Pencils have also been bought. This is a digital-based effort, but it could not have been done without a sharp-tipped pencil to indicate one-at-a-time, particulars upon a piece of paper. These are the details so essential to documenting the history of wildbirds in northern America.

Significant Milestones in Historic Ornithology Compilation

While adding records of historical notations for birds in northern America, there were two significant milestones achieved in latter December.

Because of references found on internet archive sites, further records of bird specimens for 1880 and later were discovered. Once one particular reference was evaluated, there were an additional 496 records added to the database of records. In order to suitably add the observations, more than 75 new distinct site names were also added.

Both of these items increased the number of records and extent of coverage for this compilation of birds records for the region of America from Panama northward. Nearly every record is also based upon an observation made more than 125 years before present.

Specimen Catalogue

The most recent publication reviewed was a catalogue of birds, eggs and nests in the collection of the Greene Smith Museum, of Peterboro, New York, as published in 1880. Someone compiled the contents within 48 cases of material.

Smith was the son of a "New York reformer, philanthropist and abolitionist Gerrit Smith," and "was an avid sportsman and early conservationist who established a distinguished legacy as a self-taught expert ornithologist and taxidermist."

There were 486 usable records representing more than 320 species, which included an identification, and very importantly, the date and location for the record. Each record was entered into a database and designated to a particular site.

Information on eggs and nests was not usable since no dates were given.

This publication was found online at the Biodiversity Heritage Library. Resources found here and at other places are essential in any effort to evaluate historic ornithology since they provide sources which can be found by careful searching. Many of the older publications noted by such an effort would have never been located using the bibliographic options available even five years ago.

Interesting Details

An essential aspect of the specimen records are the many different localities indicated as the source of the collection's material. The catalogue has items from the Baja Peninsula (lower California) and elsewhere in Mexico, the Bahama Islands, Disco Island along the coast of western Greenland. There are 26 states represented, with the larger number of records from these states:

Illinois, including Calumet, Cook County and six other localities: 95 records

New York (14 localities): 87

Texas (11 localities: 66

Florida (12 localities): 61

Some of the records have a vague location designated (i.e., Atlantic Ocean, North Pacific Ocean), but nonetheless are a part of the historic record.

What is interesting is that some of the dates for specimens match notations from other references, and are useful in indicating a possible source of specimens, during an era when correspondents exchanged material in their individual effort to establish their personal collection. Specimens were also sold, as shown in ads in the early ornithological publications.

Interesting Species

Species represented the most in the entered records are the Song Sparrow, Greater Prairie-Chicken, Northern Bobwhite, Ruffed Grouse, Horned Lark and Song Sparrow.

Among the distinctive records from the catalogue notations for four rare or now extinct species.

Carolina Parakeet: specimen from 26 Feb 1877 at Bayport, Florida

Eskimo Curlew: specimen from 19 Mar 1878 at Gainesville, Texas

Ivory-billed Woodpecker: specimen from 17 Mar 1876 at Hernando Co., Fla.

Whooping Crane: specimen 21 Mar 1871 from Champaign, Ills.; this record is perplexing due to the northerly locality in early spring. Though this would not be expected, based on more contemporary information, having a specimen adds to the credibility of the record.

These records are just a few of the interesting details derived from this particular source. Add in the whole set of information and there is information available but not yet obvious on the occurrence and distribution during a period of ca. 10,000 years.

Another similar source awaits analysis and data entry is another catalogue, this one from Europe, which lists the 6006 specimens in the Hugh Edwin Strickland collection. A preliminary review indicates there are records from Carlisle, Pennsylvania as provided by S.F. Baird, and items from Jamaica and Guatemala.

Among the specimens in the Oxford Museum of Natural History, are some from Northern America, and which are available to review and consider in any evaluation of species records for the continent.

None of these three sources have been noted as being previously evaluated for their contribution to the continental avifauna. They certainly have essential details of occurrence.

Bone Studies Reveal Avifauna of North America in Past Millenniums

Relicts of bone from dead birds that have been carefully removed at different types of excavations provide subtle clues to identify the avifauna present during past millenniums in North America. This type of ornithological research is an interesting and intriguing investigation of fragile remnants of excavated bird skeletons, and a subsequent, detailed comparison to known bones to determine - if possible - the particular species.

Published finding present clues of the birds present at a particular locale at a place and time long before primitive peoples had any means of recording history, and hundreds to many thousands of years prior to the first written record of a bird on the continent.

Prehistoric Law's Diving-Goose. Image courtesy of Wikipedia.

A wide variety of published findings are available. Each scholarly article may provide details of only a few species, but combined results from the wide range of anthropology, ornithology, archeology and related sources do provide a set of occurrence records for a wonderfully representative variety of avifauna extant during eleven millennia. In this case, starting with ca. 12,000 years before the common era (B.C.E.) through two thousand years ago.

A list of known species was determined by extracting bird information from more than 500 published sources. Information was entered into a database, and included the given bird species name, and the particular locale including country, state or province if applicable and county if available. Chronology was determined from radio carbon dating for the site as given with the research findings. In some cases, when several date ranges were provided, an average was used to designate an approximate, but accurate time of occurrence. Also, in some instances, when avian remains occurred in different strata, distinct names for a particular set of records were designated to separate the species into a time frame with a greater degree of precision.

Ancient Avifauna

More than 1850 distinct records provide the details to document more about 305 recognized species for North America prior to 2000 years B.C.E. (also commonly referred to as Y.B.P., or Years Before Present). For the purposes of this comparison, the base year used is A.D. 2000.

The information given is the species in modern taxonomic sequence, the thousand year period of time before common era, and the number of records within each designated time period. If a record is included within a particular millennia, the site chronology was correlated within that period of a thousand years. For example, if a site was dated to approximately 7750 years B.C.E., it would be within the 7000 millennia period.

Notable are several currently extinct species which were a short time ago in the past - biologically speaking - were an integral part of the continental avifauna.

The value given indicates the number of records for the particular period.

List of Recorded Bird Species

Common Name	12000	11000	10000	9000	8000	7000	6000	5000	4000	3000	2000
Black-bellied Whistling-Duck	-	-	-	-	-	-	-	-	-	1	-
Greater White-fronted Goose	-	-	-	-	-	-	-	2	-	-	2
Snow Goose	-	2	2	1	-	4	1	2	2	4	6
Ross's Goose	-	-	-	-	-	-	-	1	-	-	1
Brant	-	1	-	-	-	-	-	1	-	-	1
Canada Goose	2	2	3	3	-	6	1	8	6	7	6
Trumpeter Swan	-	-	2	-	-	1	-	2	1	3	2
Tundra Swan	-	-	1	1	-	-	1	3	3	1	4
Muscovy Duck	-	-	-	-	-	-	-	-	-	1	-
Wood Duck	-	-	2	1	1	-	1	1	2	1	-
Gadwall	-	-	-	1	-	1	-	2	2	-	2
American Wigeon	2	-	1	1	-	1	-	1	3	-	3
American Black Duck	-	-	-	1	-	-	-	-	2	1	-
Mallard	3	3	5	2	-	2	1	7	5	4	6
Blue-winged Teal	2	-	2	1	-	2	1	1	3	-	2
Cinnamon Teal	-	-	-	1	-	1	-	3	1	-	1
Northern Shoveler	1	1	-	1	-	1	-	3	1	1	2
Northern Pintail	1	-	3	1	-	1	2	4	4	1	5
Green-winged Teal	3	1	4	2	2	4	-	3	3	1	5
Canvasback	1	-	1	1	-	-	-	3	5	2	2
Redhead	1	-	-	1	-	-	-	1	2	-	1
Ring-necked Duck	-	-	1	-	-	-	1	1	3	1	1
Greater Scaup	1	-	-	1	-	-	-	1	1	1	3
Lesser Scaup	1	-	2	2	-	-	-	3	4	3	1
Law's Diving-Goose	-	-	1	-	1	4	2	2	3	3	1
King Eider	-	-	-	-	-	-	-	-	-	1	-
Common Eider	-	1	-	-	-	-	-	-	-	-	1
Harlequin Duck	-	-	-	-	-	-	-	-	-	-	1
Surf Scoter	-	1	-	-	-	-	-	1	-	1	4
White-winged Scoter	-	1	-	-	-	-	-	1	-	1	3
Black Scoter	-	-	-	-	-	-	-	1	-	-	1
Long-tailed Duck	-	1	-	-	-	-	-	1	1	1	1
Bufflehead	1	1	1	2	-	1	-	3	5	2	3
Common Goldeneye	-	1	-	1	-	1	-	1	-	-	2
Hooded Merganser	1	-	2	2	1	1	-	-	4	2	2
Common Merganser	1	-	-	1	-	1	-	3	1	3	4
Red-breasted Merganser	-	-	-	-	-	-	-	2	1	1	2
Ruddy Duck	2	-	1	1	-	2	1	3	7	2	3
Plain Chachalaca	-	-	-	-	-	-	-	-	-	1	1
White-bellied Chachalaca	-	-	-	-	-	-	-	-	-	1	-
Ring-necked Pheasant	-	-	-	-	-	-	-	-	-	-	2
Ruffed Grouse	3	2	3	3	1	2	1	6	-	1	3
Greater Sage-Grouse	3	2	4	2	-	4	-	4	-	4	-
Spruce Grouse	1	1	-	-	-	-	-	-	-	-	-
Willow Ptarmigan	-	-	-	-	-	-	-	-	-	1	-
Dusky Grouse	1	1	3	-	-	3	1	4	-	1	2
Sharp-tailed Grouse	3	1	4	-	-	1	-	3	-	2	-
Greater Prairie-Chicken	3	-	5	1	-	3	-	1	2	3	-
Lesser Prairie-Chicken	-	-	-	-	-	-	-	-	-	-	1
Wild Turkey	3	2	10	3	4	2	3	10	11	10	20
Ocellated Turkey	-	-	-	-	-	-	-	-	-	-	2
Mountain Quail	-	-	2	-	-	1	-	-	-	-	-
Scaled Quail	1	-	-	-	-	-	-	-	-	-	-
Northern Bobwhite	3	-	4	2	2	1	1	1	3	3	1
Spotted Wood-Quail	-	-	-	-	-	-	-	-	-	1	-
Ocellated Quail	-	-	-	-	-	-	-	-	-	1	-
Red-throated Loon	-	-	-	-	-	-	-	2	-	-	3
Arctic Loon	-	-	-	-	-	-	-	-	-	1	1
Pacific Loon	-	1	-	-	-	-	-	1	-	-	2
Common Loon	1	1	-	-	1	1	-	1	3	6	6
Least Grebe	-	-	-	-	-	-	-	-	-	1	-
Pied-billed Grebe	3	-	-	1	-	1	-	3	10	7	2
Horned Grebe	1	-	1	2	1	1	1	2	2	3	4
Red-necked Grebe	-	-	-	-	-	-	-	1	2	1	2
Eared Grebe	-	-	1	1	-	5	-	5	2	4	2
Western Grebe	1	-	-	1	-	-	-	5	3	2	4
Black-footed Albatross	-	-	-	-	-	-	-	1	1	-	-
Short-tailed Albatross	-	-	-	-	-	-	-	1	1	1	2
Northern Fulmar	-	-	-	-	-	-	-	1	1	2	1
Bermuda Petrel	-	-	-	-	-	-	-	-	-	-	1
Pink-footed Shearwater	-	-	-	-	-	-	-	1	1	-	-
Buller's Shearwater	-	-	-	-	-	-	-	-	1	-	-
Sooty Shearwater	-	-	-	-	-	1	-	1	-	-	1
Short-tailed Shearwater	-	-	-	-	-	-	-	1	1	-	-
Manx Shearwater	-	-	-	-	-	-	-	-	-	-	3
Audubon's Shearwater	-	-	-	-	-	-	-	-	-	-	2
Leach's Storm-Petrel	-	-	-	-	-	-	-	1	-	-	2
Ashy Storm-Petrel	-	-	-	-	-	-	-	-	-	-	2
Black Storm-Petrel	-	-	-	-	-	-	-	-	-	-	1
Brown Booby	-	-	-	-	-	-	-	-	-	-	1
Northern Gannet	-	-	-	-	-	-	-	-	-	1	-
American White Pelican	1	-	-	-	-	2	-	2	3	1	3
Brown Pelican	-	-	-	-	-	-	-	1	1	2	2
Brandt's Cormorant	-	-	-	-	-	-	-	1	-	1	3
Neotropic Cormorant	-	-	-	-	-	-	-	-	-	1	-
Double-crested Cormorant	1	1	-	-	1	2	-	4	7	5	4
Pelagic Cormorant	-	1	-	-	-	-	-	1	-	1	2
Anhinga	-	-	-	-	-	-	-	-	1	-	-
American Bittern	1	-	1	-	-	1	-	1	1	2	2
Least Bittern	-	-	-	-	-	-	-	-	-	-	1
Bare-throated Tiger-Heron	-	-	-	-	-	-	-	-	-	1	-
Great Blue Heron	-	-	-	1	-	3	-	4	3	5	3
Great Egret	-	-	-	-	-	1	-	1	3	2	-
Snowy Egret	-	-	-	-	-	-	-	2	-	1	1
Little Blue Heron	-	-	-	-	-	1	-	-	-	-	-
Green Heron	-	-	-	-	-	-	-	1	1	1	1
Black-crowned Night-Heron	-	-	1	-	-	2	-	-	2	2	1
Yellow-crowned Night-Heron	-	-	-	-	-	-	-	-	-	1	1
White Ibis	-	-	-	-	-	-	-	-	-	-	2
White-faced Ibis	-	-	-	-	-	-	-	-	-	-	1
Wood Stork	1	-	-	-	-	-	-	-	-	-	-
Black Vulture	2	1	1	-	1	-	-	-	-	1	-
Turkey Vulture	3	1	-	2	1	-	-	4	3	3	3
California Condor	6	-	2	-	1	-	-	1	-	-	3
Osprey	-	-	-	-	-	-	-	3	-	1	2
Swallow-tailed Kite	-	-	1	-	-	-	-	-	1	-	-
Mississippi Kite	-	-	1	-	-	-	-	-	1	-	-
Bald Eagle	1	1	2	-	1	1	1	4	4	4	5
Northern Harrier	1	1	-	1	-	1	-	1	1	2	3
Sharp-shinned Hawk	3	-	-	1	-	-	-	-	-	2	1
Cooper's Hawk	2	-	-	-	-	-	-	3	1	-	1
Northern Goshawk	-	-	1	-	1	-	-	1	-	-	-
Common Black-Hawk	-	-	-	1	-	-	-	-	-	-	-
Roadside Hawk	-	-	-	-	-	-	-	-	-	1	-
Red-shouldered Hawk	2	-	2	-	2	1	-	1	2	1	1
Broad-winged Hawk	1	-	1	-	-	-	-	-	-	-	1
Swainson's Hawk	4	-	-	-	-	-	-	1	-	-	3
Red-tailed Hawk	5	1	5	1	1	-	-	3	6	2	4
Ferruginous Hawk	-	-	-	1	-	-	-	-	-	-	-
Rough-legged Hawk	1	-	-	-	-	-	-	-	-	-	4
Harpy Eagle	-	-	-	-	-	-	-	-	-	1	-
Golden Eagle	6	1	1	2	-	1	-	7	2	3	2
Crested Caracara	-	-	-	-	-	-	-	-	-	1	-
Bahama's Caracara	-	-	-	-	-	-	-	-	-	-	1
American Kestrel	6	2	2	2	1	1	2	4	2	-	7
Merlin	-	1	1	-	-	1	-	1	-	-	-
Aplomado Falcon	1	-	-	-	-	-	-	-	-	-	-
Peregrine Falcon	1	-	1	-	-	1	-	1	1	-	-
Prairie Falcon	2	2	2	-	1	-	-	3	1	1	1
Yellow Rail	1	-	-	-	-	-	-	-	-	-	-
Gray-breasted Crake	-	1	-	-	-	-	-	-	-	-	-
Clapper Rail	-	-	-	-	-	-	-	1	-	-	-
King Rail	-	-	-	1	-	-	-	-	1	-	-
Virginia Rail	-	1	1	-	-	1	-	1	-	1	1
Sora	-	1	1	1	-	2	-	1	2	-	1
Purple Gallinule	-	-	-	-	-	-	-	-	1	-	-
Common Moorhen	-	-	-	2	-	-	-	-	3	1	1
American Coot	3	1	3	2	1	4	-	7	11	5	4
Sandhill Crane	-	-	2	-	-	1	1	1	5	2	4
Whooping Crane	1	-	-	-	-	-	-	-	-	-	-
Black-bellied Plover	-	-	-	-	-	-	-	1	-	-	1
Semipalmated Plover	-	-	-	-	-	-	-	-	-	1	-
Killdeer	1	-	-	-	-	-	-	-	1	-	-
Mountain Plover	-	1	-	-	-	-	-	-	-	-	-
Black-necked Stilt	1	-	-	-	-	-	-	-	-	1	1
American Avocet	2	-	1	1	-	2	-	1	1	-	1
Spotted Sandpiper	-	-	-	-	1	-	-	-	-	-	1
Wandering Tattler	-	-	-	-	-	-	-	-	-	-	1
Willet	3	-	-	-	-	-	-	1	1	1	2
Upland Sandpiper	2	-	-	-	-	-	-	-	1	-	1
Whimbrel	-	-	-	-	-	-	1	-	1	1	-
Long-billed Curlew	1	-	2	1	-	2	-	-	1	-	1
Hudsonian Godwit	-	1	-	-	-	-	-	-	-	-	-
Marbled Godwit	-	1	-	-	-	-	-	1	-	1	-
Ruddy Turnstone	-	-	-	-	1	-	-	-	-	-	-
Sanderling	-	-	-	-	-	-	-	-	-	-	1
Least Sandpiper	1	-	1	-	-	-	-	-	-	1	1
Short-billed Dowitcher	-	-	1	-	-	-	-	-	-	-	-
Wilson's Snipe	2	-	-	1	1	-	-	2	1	-	2
American Woodcock	3	-	1	1	1	-	-	1	-	-	1
Red-necked Phalarope	-	-	-	-	-	-	-	-	-	-	1
Red Phalarope	-	-	-	-	-	-	-	-	-	-	1
Bonaparte's Gull	-	1	-	-	-	-	-	-	1	-	1
Heermann's Gull	-	-	-	-	-	-	-	1	1	1	-
Mew Gull	-	1	-	-	-	-	-	1	-	-	-
Ring-billed Gull	-	-	-	-	-	-	-	1	-	-	-
California Gull	-	-	-	-	-	1	-	2	1	-	3
Herring Gull	-	-	-	-	-	-	-	1	-	1	1
Western Gull	-	-	-	-	-	-	-	1	-	-	2
Glaucous-winged Gull	-	1	-	-	-	-	-	1	-	-	-
Glaucous Gull	-	-	-	-	-	-	-	1	-	-	-
Black-legged Kittiwake	-	-	-	-	-	-	-	1	-	-	3
Sooty Tern	-	-	-	-	-	-	-	-	-	-	1
Caspian Tern	-	-	-	-	-	1	-	1	-	-	-
Black Tern	-	-	-	-	-	-	-	-	-	1	-
Common Tern	-	-	-	-	-	-	-	-	-	-	1
Arctic Tern	-	-	-	-	-	-	-	1	-	-	-
Sandwich Tern	-	-	-	-	-	-	-	-	-	1	-
Parasitic Jaeger	-	-	-	-	-	1	-	-	-	-	-
Common Murre	-	1	-	-	-	-	-	2	-	1	5
Razorbill	-	-	-	-	-	-	-	-	-	-	1
Great Auk	-	-	-	-	-	-	-	-	-	2	-
Pigeon Guillemot	-	1	-	-	-	-	-	1	-	1	1
Marbled Murrelet	-	1	-	-	-	-	-	-	-	-	1
Xantus's Murrelet	-	-	-	-	-	-	-	-	-	-	2
Cassin's Auklet	-	1	-	-	-	-	-	1	1	1	1
Parakeet Auklet	-	1	-	-	-	-	-	-	-	-	-
Least Auklet	-	1	-	-	-	-	-	-	-	-	-
Rhinoceros Auklet	-	1	-	-	-	-	-	1	1	1	3
Tufted Puffin	-	1	-	-	-	-	-	1	-	1	-
Scaled Pigeon	-	-	-	-	-	-	-	-	-	-	1
White-crowned Pigeon	-	-	-	-	-	-	-	-	-	-	2
White-winged Dove	-	-	-	-	-	-	-	-	1	-	-
Zenaida Dove	-	-	-	-	-	-	-	-	-	-	1
Mourning Dove	3	-	1	2	-	2	-	-	2	1	2
Passenger Pigeon	4	-	4	3	4	2	2	4	5	5	4
Inca Dove	-	-	-	-	-	-	-	-	-	1	-
Blue Ground-Dove	-	-	-	-	-	-	-	-	-	1	-
White-tipped Dove	-	-	-	-	-	-	-	-	-	1	-
Key West Quail-Dove	-	-	-	-	-	-	-	-	-	-	1
Bridled Quail-dove	-	-	-	-	-	-	-	-	-	-	1
Ruddy Quail-Dove	-	-	-	-	-	-	-	-	-	1	-
Orange-fronted Parakeet	-	-	-	-	-	-	-	-	-	1	-
Military Macaw	-	-	-	-	-	-	-	-	-	1	-
White-fronted Parrot	-	-	-	-	-	-	-	-	-	1	-
Cuban Parrot	-	-	-	-	-	-	-	-	-	-	2
Lesser Ground-Cuckoo	-	-	-	-	-	-	-	-	-	1	-
Greater Roadrunner	-	-	-	-	-	-	-	-	1	-	-
Groove-billed Ani	-	-	-	-	-	-	-	-	-	1	-
Barn Owl	2	-	1	-	-	-	-	1	2	1	5
Western Screech-Owl	-	-	-	-	1	-	-	-	-	-	1
Eastern Screech-Owl	1	1	1	2	1	1	-	2	1	1	-
Great Horned Owl	5	1	2	1	1	5	1	5	4	1	3
Snowy Owl	-	-	-	-	-	-	-	-	-	1	2
Northern Hawk Owl	1	-	-	-	-	-	1	-	-	-	-
Northern Pygmy-Owl	1	-	-	-	-	-	-	-	-	-	-
Elf Owl	1	-	-	-	-	-	-	-	-	-	-
Burrowing Owl	-	1	-	-	-	1	-	1	-	-	4
Spotted Owl	1	-	-	-	-	-	-	1	-	-	-
Barred Owl	2	-	1	1	2	-	-	2	1	2	1
Long-eared Owl	2	-	4	-	-	1	-	2	-	-	1
Short-eared Owl	3	1	2	1	-	2	-	2	-	1	2
Boreal Owl	2	-	-	-	-	-	-	-	-	-	-
Northern Saw-whet Owl	3	-	-	-	-	-	-	-	-	-	-
Common Nighthawk	1	-	-	1	1	1	-	-	-	-	-
Chuck-will's-widow	-	-	-	-	-	-	-	-	1	-	-
Chimney Swift	1	-	-	-	1	-	-	-	-	-	-
White-throated Swift	1	-	-	-	-	-	-	-	1	-	-
Blue-crowned Motmot	-	-	-	-	-	-	-	-	-	1	-
Ringed Kingfisher	-	-	-	-	-	-	-	-	-	1	-
Belted Kingfisher	1	-	-	-	1	1	-	-	1	-	-
Emerald Toucanet	-	-	-	-	-	-	-	-	-	1	-
Red-headed Woodpecker	1	-	-	1	-	-	-	1	-	-	1
Red-bellied Woodpecker	3	-	-	1	-	-	-	1	2	-	1
West Indian Woodpecker	-	-	-	-	-	-	-	-	-	-	1
Downy Woodpecker	1	-	-	-	-	-	-	1	-	-	-
Hairy Woodpecker	2	-	-	-	-	-	-	-	-	-	-
White-headed Woodpecker	-	-	-	-	-	-	-	1	-	-	-
Northern Flicker	4	3	3	1	1	4	-	6	4	2	4
Pileated Woodpecker	-	1	-	1	1	-	-	-	1	-	-
Ivory-billed Woodpecker	-	-	-	-	-	-	-	-	1	-	-
Western Wood-Pewee	-	-	-	1	-	-	-	-	-	-	-
Eastern Wood-Pewee	1	-	-	-	-	-	-	-	-	-	-
Cuban Pewee	-	-	-	-	-	-	-	-	-	-	1
Eastern Phoebe	3	-	-	-	-	-	-	-	1	-	-
Say's Phoebe	-	-	-	1	-	-	-	-	-	-	-
Sulphur-bellied Flycatcher	-	-	-	-	-	-	-	-	-	1	-
Rose-throated Becard	-	-	-	-	-	-	-	-	-	1	-
Northern Shrike	-	-	-	1	-	-	-	-	-	-	-
Gray Jay	2	-	-	-	-	-	-	-	-	-	-
Steller's Jay	1	-	-	-	-	-	-	-	-	-	-
Blue Jay	3	-	-	1	1	-	-	1	2	-	1
Florida Scrub-Jay	-	-	-	1	-	-	-	-	-	-	-
Pinyon Jay	-	-	-	-	-	-	-	-	-	1	-
Clark's Nutcracker	2	1	-	-	-	-	-	-	-	-	-
Black-billed Magpie	5	-	4	1	1	2	-	1	-	2	-
American Crow	1	-	4	1	3	-	1	3	2	-	4
Cuban Crow	-	-	-	-	-	-	-	-	-	-	2
Fish Crow	-	-	-	-	-	-	-	-	2	-	1
Common Raven	8	3	5	1	2	6	-	3	4	4	6
Horned Lark	2	1	1	-	-	-	-	1	-	1	1
Cliff Swallow	1	-	1	-	2	-	-	-	-	-	1
Cave Swallow	-	-	-	-	-	-	-	-	-	-	1
Carolina Chickadee	-	-	-	-	-	-	-	-	-	1	-
Red-breasted Nuthatch	2	-	-	-	-	-	-	-	1	-	-
Rock Wren	-	1	-	1	-	-	-	-	-	-	-
Canyon Wren	-	-	-	1	-	-	-	-	-	-	-
Western Bluebird	-	-	-	-	-	-	-	-	-	-	1
Mountain Bluebird	-	-	-	1	-	-	-	-	-	-	-
Townsend's Solitaire	-	-	-	1	-	-	-	-	-	-	-
American Robin	4	-	2	1	1	1	-	-	-	-	1
Red-legged Thrush	-	-	-	-	-	-	-	-	-	-	1
Gray Catbird	1	-	-	-	-	-	-	1	-	-	-
Northern Mockingbird	-	1	-	-	-	-	-	-	-	-	1
Bahama Mockingbird	-	-	-	-	-	-	-	-	-	-	1
Brown Thrasher	1	1	-	-	-	-	-	-	-	-	-
Pearly-eyed Thrasher	-	-	-	-	-	-	-	-	-	-	2
European Starling	-	-	-	1	-	-	-	-	-	-	1
Bohemian Waxwing	-	-	-	1	-	-	-	-	-	-	-
Yellow-rumped Warbler	-	-	-	1	-	-	-	-	-	-	-
Black-and-White Warbler	-	-	-	-	-	1	-	-	-	-	-
Bahama Yellowthroat	-	-	-	-	-	-	-	-	-	-	1
Scarlet Tanager	1	-	-	-	-	-	-	-	-	-	-
Black-faced Grassquit	-	-	-	-	-	-	-	-	-	-	1
Eastern Towhee	1	-	-	1	-	-	-	-	-	-	-
Chipping Sparrow	-	-	-	-	-	-	-	-	-	-	1
Vesper Sparrow	-	1	-	-	-	-	-	-	-	-	-
Fox Sparrow	1	1	-	-	-	-	-	-	-	-	3
Song Sparrow	1	-	-	-	-	-	-	-	-	-	-
White-throated Sparrow	2	-	-	-	-	-	-	-	-	-	-
Northern Cardinal	-	-	-	-	-	-	-	1	1	-	-
Dickcissel	-	-	-	1	-	-	-	-	-	-	-
Red-winged Blackbird	2	1	1	-	1	-	1	1	-	-	1
Eastern Meadowlark	-	-	-	-	-	-	-	-	-	-	1
Western Meadowlark	-	-	1	-	-	4	-	-	-	-	3
Yellow-headed Blackbird	-	-	-	-	-	-	-	-	-	1	1
Brewer's Blackbird	-	-	-	-	-	-	-	-	-	-	1
Common Grackle	1	-	3	1	-	1	-	-	1	1	-
Great-tailed Grackle	-	-	-	-	-	-	-	-	-	1	-
Brown-headed Cowbird	1	-	-	-	-	-	-	-	-	-	1
Gray-crowned Rosy-Finch	-	-	-	-	-	-	-	1	-	1	-
Pine Grosbeak	1	-	-	-	-	-	-	-	-	-	-
Purple Finch	1	-	-	-	-	-	-	-	1	-	-
White-winged Crossbill	-	-	-	-	-	-	-	-	1	-	-
Pine Siskin	1	-	-	-	-	-	-	-	-	-	-

There are additional records for these expansive time periods which refer to generic species (i.e., goose, duck, hawk, gull, woodpecker, etc.) but these are not included in the list. Neither are bone remains identified to a group of birds such as unidentified waterfowl, or in some cases, simply as unidentified birds.

Grouping the records into distinct millennia shows how the analysis of species varies during a period of time. Larger species - thus larger bones - are better represented in the older periods, with smaller species occurring more often in the more recent periods since the bone remnants have not disintegrated.

There is a large amount of information which could be determined if the records were geo-mapped using the current geographic systems. This would be an extremely useful endeavor to determine species biogeography and to evaluate changes in distribution through time.

The largest number of records are from the United States:

Country - Number of Records

U.S.A. - 2254

Canada - 126

Mexico - 63

Bahama Islands - 49

Belize - 3

This table has a larger number of records since it includes all records and is not limited to those records particular to a valid species.

The number of records from the different states or provinces can also be derived:

State or Province - Country - Number of Records

California - U.S.A. - 406 Nevada - U.S.A. - 262 Utah - U.S.A. - 202 Alabama - U.S.A. - 201 Illinois - U.S.A. - 173 New Mexico - U.S.A. - 101 British Columbia - Canada - 84 Arizona - U.S.A. - 77 Washington - U.S.A. - 77 Tennessee - U.S.A. - 73 Texas - U.S.A. - 71 Idaho - U.S.A. - 69 Florida - U.S.A. - 62 Alaska - U.S.A. - 61 Chiapas - Mexico - 60 Missouri - U.S.A. - 55 Nebraska - U.S.A. - 55 Wisconsin - U.S.A. - 43 Virginia - U.S.A. - 40 Kansas - U.S.A. - 36 Ontario - Canada - 30 Pennsylvania - U.S.A. - 25 Ohio - U.S.A. - 25

New York - U.S.A. - 21 Minnesota - U.S.A. - 21 Michigan - U.S.A. - 20 Oregon - U.S.A. - 17 North Carolina - U.S.A. - 15 Colorado - U.S.A. - 9 South Dakota - U.S.A. - 9 New Hampshire - U.S.A. - 8 Montana - U.S.A. - 7 Northwest Territories - Canada - 7 South Carolina - U.S.A. - 4 Wyoming - U.S.A. - 4 Quebec - Canada - 3 Rhode Island - U.S.A. - 3 Alberta - Canada - 2 Maine - U.S.A. - 1 Mexico - Mexico - 1 Yucatan - Mexico - 1 Oaxaca - Mexico - 1 New Jersey - U.S.A. - 1

The total does not match other tallies since localities where the state or province is not known are not included (i.e., the Bahama Islands and Belize).

One factor which limits the list derived from bone studies is information given for Arizona locales. Many of the studies evaluated do not include site chronology, so the material cannot be attributed to a particular time period, so the records could not be included in the analysis.

It should also be noted that there are certainly additional sources that should be included in an analysis of ancient avifauna. Many of the citations are included in the literature cited for a reviewed source, but are obscure reports that are not generally available and could note be accessed, yet alone get decent consideration. There are valuable additions that would help increase the extent of the base data to evaluate, and help provide a broader view of species occurrence during a particular millennia of former time.

Disclaimer: This is a preliminary analysis subject to change that may occur as a result of further analysis and study, and if additional informational details become available. Further resources - which are not currently available - will also be needed to provide complete geo-coding for each locale that would allow accurate mapping. Currently, there is no foreseeable time when this can be done for the 31,800+ records available overall in the Ancient Avifauna database which includes a diverse bunch of records from excavation and narrative sources through the 1850s.