ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by T. Worthy: 16 Mar. 2020; published: 7 May 2020 111
Zootaxa 4772 (1): 111–131
Copyright © 2020 Magnolia Press Article
Licensed under a Creative Commons Attribution 4.0 International License http://creativecommons.org/licenses/by/4.0/
The Irvingtonian Avifauna of Cumberland Bone Cave, Maryland
HELEN F. JAMES
Department of Vertebrate Zoology, National Museum of Natural History, Washington, DC, 20013. E-mail: email@example.com
The early and mid-Pleistocene avian communities of North America are best known from the Rocky Mountain region
and peninsular Florida. In the Appalachian Mountain region, only a small number of avian bones from mid-latitude cave
deposits have been attributed to this time period. Here, I enlarge this record by reporting on bird bones from Cumberland
Bone Cave in western Maryland, a well-known locality for large and small Irvingtonian mammals and other vertebrates.
The taxa identified encompass ground birds, waterfowl, a hawk, two eagles, a vulture, an owl, a jay, a flycatcher, a junco
or sparrow, and a finch. No purely boreal elements are confirmed as part of the avian assemblage, and all of the extant
species that are positively or tentatively identified in the assemblage still occur in the region today. An immature bone
referred to the Black Vulture (Coragyps atratus (Bechstein)) represents an Irvingtonian breeding record for the species
in Maryland. This record occurs at the northern limit of the current breeding range for the genus. Extinct species in the
assemblage include the Passenger Pigeon (Ectopistes migratorius (Linnaeus)), a large screech owl (Megascops guildayi
(Brodkorb & Mourer-Chauviré 1984)), and the large goose, Branta dickeyi Miller 1924. It can be argued that none of
these represent the extinction of a phyletic lineage during the Irvingtonian. Based on the broad habitat preferences of
modern counterparts of the birds in the assemblage, we can expect that Irvingtonian habitats near the site included mixed
forest with mast-producing hardwoods and both early and later successional stages represented. There must have been
fluvial, wetland, or lacustrine habitat suitable for waterbirds nearby, and probably also open woodland or grassy savannah
areas, suitable for vulture foraging, turkey nesting, and booming by Ruffed Grouse.
Key words: mid-Pleistocene, fossil birds, paleoecology, extinction, Branta dickeyi, Bonasa umbellus, Coragyps atratus,
Ectopistes migratorius, Megascops guildayi
The bone deposits at Cumberland Bone Cave in Alleghany County, Maryland, were discovered in 1912 during
dynamite blasting to create a roadbed for the Western Maryland Railway. Located in the Valley and Ridge Physio-
graphic Province, at the base of a limestone ridge near Corriganville (39.69 deg. N, 78.79 deg. W, 245 m asl), the
site soon became recognized as a major locality for Pleistocene mammals of the mid-latitude Appalachian Moun-
tains (Gidley 1914, 1920; Gidley & Gazin 1938). The diverse assemblage of mammals from the cave correlates
with the Irvingtonian North American Land Mammal Age, which covers the early to mid-Pleistocene from roughly
1.9 to 0.15 Ma (Bell et al. 2004; Cohen & Gibbard, 2011). Arvicoline biochronology suggests that the age of bone
accumulation can be further narrowed to a division of the Irvingtonian termed Irvingtonian II, estimated to range
from 0.85 to 0.4 Ma (Bell 2000; Bell et al., 2004). Coupled uranium-series and electron spin resonance dating of
two fossil peccary (Platygonus sp. Le Conte 1848) teeth offer more precise age estimates of 722±64 ka and 790±53
ka for the fossil accumulation (Withnell et al. in press). These dates are consistent with paleomagnetic evidence of
speleothem growth in the cave prior to the Brunhes-Matuyama reversal in the earth’s magnetic polarity (i.e., prior
to 780 ka; Withnell et al. in press).
Two previous papers reported on bird bones from the site: Wetmore (1927) identified a single avian bone as
a Ruffed Grouse (Bonasa umbellus (Linnaeus)), and Brodkorb & Mourer-Chauviré (1984) identified six bones as
representing six additional species of birds, notably including the extinct Passenger Pigeon (Ectopistes migratorius)
and a new species of owl (Megascops guildayi). Brodkorb & Mourer-Chauviré identified a tarsometatarsus from the
site as a Canada Jay, (Perisorius canadensis (Linnaeus)), and interpreted the presence of this species as evidence of
112 · Zootaxa 4772 (1) © 2020 Magnolia Press
a cooler climate compared with the present. Based on the presence of P. canadensis, Brodkorb & Mourer-Chauviré
thought it likely that the deposit accumulated during a glacial stage, most likely the Illinoian which extends from
about 130 ky to perhaps as long ago as 300 ky (Cohen & Gibbard 2011). However, as noted above, other data favor
an earlier age for the site.
I studied and catalogued a larger group of bird bones that were collected from Cumberland Bone Cave in the
1990s and 2000s. I also reexamined four of the seven bird bones reported from the site in previous literature. Here,
I present taxonomic identifications of the avian fossils from Cumberland Bone Cave and discuss the assemblage
in terms of inferred nearby habitats when the bones were accumulating and avifaunal turnover in the mid-latitude
Appalachians since then.
Materials and methods
All fossil specimens are housed at the Department of Paleobiology, National Museum of Natural History, Smith-
sonian Institution, Washington, DC (PAL USNM), except the six bones in Brodkorb & Mourer-Chauviré’s report,
which are at the Carnegie Museum of Natural History, Pittsburgh, PA (CM). The newly reported bones in this report
were collected from the site by Trent Spielman. Frederick V. Grady sorted the collection according to higher taxo-
nomic groups and freed some of the bones from breccia by acid etching at the Vertebrate Paleontology Preparation
Laboratory of NMNH.
The fossils were identified based on comparisons with the skeleton collection of the Bird Division, Department
of Vertebrate Zoology, NMNH. Avian body masses compiled by Dunning (2008) were useful for selecting species of
roughly similar body size to the fossil taxa, within families or orders, for further comparisons. Bone measurements
were taken with a digital caliper and rounded to the nearest 0.1 mm. Taxonomic nomenclature follows the American
Ornithological Society Checklist of North and Middle American Birds (Chesser et al. 2019). Taxonomic authorities
are given on first mention with dates and full references provided for the fossil taxa only. For extant taxa, dates and
taxonomic authorities can be found in prior published editions of and supplements to the Checklist cited above. The
letters l and r indicate left and right side of the body; f and m indicate female and male. In taxonomic attributions,
cf. indicates the specimen is referred to the named taxon, and aff. indicates only that the specimen has affinities
with the named taxon. Anatomical nomenclature follows Baumel & Witmer (1993), with many terms translated into
Statistics were calculated in R version 3.6.1 (2019-07-05) (R Core Team 2019), using the RStudio interface
(RStudio Team 2018) and the stats package. Principal components analysis was performed using the function
prcomp, with variables log-transformed and normalized (scaled to have unit variances and centered on zero). Graph-
ics were produced using the ggplot2 package (Wickham 2016). Box-and-whisker plots show the medians, lower
and upper quartiles, ranges, and outliers of the distributions. Plots of principal components were produced using the
function ggbiplot v0.55.
Comparative material examined
The following skeletons in the collection of the Bird Division at NMNH (catalog acronym USNM BIRDS), were
examined and/or measured: Coscoroba coscoroba (Molina) 345438 m, 291255 f; Cygnus buccinator Richardson
492496 m, 347734 f; C. columbianus (Ord) 488538, 499393; C. melanocoryphus (Molina) 428167 m (captive),
345227 m (captive); Anser albifrons (Scolopi) 488745 f; A. rossi Cassin 430474 f; A. canagicus (Sevastianov)
638783 m; A. caerulescens (Linnaeus) 501617 f; Branta canadensis (Linnaeus) 343185 m (captive), 343006 m
(captive), 488182 m, 488488 m, 488723 m, 489759 m, 610626 m, 488584 f, 555497 f (captive), 561860 f, 599464
f (captive); B. hutchinsii (Richardson) 630938 m, 430286 m; Oxyura jamaicensis (Gmelin) 492491 m, 499639,
623361; Anas crecca carolinensis Gmelin 610635 f, 431027 f, 610632 m, 638839; Spatula discors (Linnaeus)
502641 f, 502643 m, 502645 m, 562841 m; Bucephala albeola (Linnaeus) 492434 m, 632035 f, 644552; Meleagris
gallopavo Linnaeus 19634 m, 556311 m, 556360 m, 501019 f, 556334 f; Tympanuchus cupido (Linnaeus) 576677
f; T. c. cupido 319417; Bonasa umbellus 559643 m, 563661 m, 621367 m, 641459 m, 557528 f, 641448 f; Falci-
pennis canadensis (Linnaeus) 641435 m, 641460 m, 641432 m, 641444 m, 641446 f, 557527 f; Lagopus leucurus
THE AVIFAUNA OF CUMBERLAND BONE CAVE, MARYLAND Zootaxa 4772 (1) © 2020 Magnolia Press · 113
(Richardson) 640234; L. mutus (Montin) 641084 m, L. lagopus (Linnaeus) 622651 m; Cathartes aura (Linnaeus)
621554, 644556; Coragyps atratus 500989 m, 622507 m, 647497 m, 647499 m, 623354 f, 647498 f, 647500 f;
Haliaeetus leucocephalus (Linnaeus) 698, 2422, 8605, 19718, 611746 m, 615007 m, 611753 f, 611754 f imma-
ture, 611757 f immature; Circus cyaneus (Linnaeus) 291684 m, 291187 m immature, 489908 f; Accipiter cooperi
(Bonaparte) 636924 m, 614341 m imm., 553854 f, 612006 f, 623631 f; Accipiter gentilis (Linnaeus) 290342 m;
Buteo lineatus (Gmelin) 614338 m; B. platypterus (Viellot) 610745 m; B. lagopus (Pontoppidan) 499426 f; Aquila
chrysaetos (Linnaeus) 491476 m, 19394, 18194, 612086; Falco peregrinus Tunstall 553620 m; Tyto alba (Scopoli)
553887 m; Strix varia Barton 556919 f; Asio otus (Linnaeus) 554126 m, 555706 f, 553816 f; Asio flammeus (Pon-
toppidan) 499484 m, 614841 m, 499485 f, 610423 f; Megascops asio (Linnaeus) 502024 m, 556918 m, 610966 f;
M. kennecottii (Elliot) 621106 m, 637719 m, 637702 f, 641659 f; Aegolius funereus (Linnaeus) 623663 m; Surnia
ulula (Linnaeus) 622484 m, 610388 m; Athene cunicularia (Molina) 610970 m, 630744 m; Patagioenas fasciata
Say 499473 m, 499472 f; P. leucocephala (Linnaeus) 554982 m; P. cayennensis (Bonnaterre) 562537 sex unknown;
P. maculosa (Temminck) 630267 f; Ectopistes migratorius 18520 (complete), 292904 sex unknown (complete),
224320 m (partial), 224319 m (partial), 224322 m (partial); Zenaida asiatica (Linnaeus) 642168 m, 646526 f; Z.
macroura (Linnaeus) 490801 f, 491239 m, 622529 f; Z. aurita (Temminck) 554975 m; Sayornis phoebe (Latham)
489807 m; Cyanocitta stelleri (Gmelin) 614422 f; Cyanocitta cristata (Linnaeus) 499266 m, 614424 f hatch year
based on skull ossification, 614427 m, hatch year based on skull ossification; Perisoreus canadensis 622673 f,
489846 m; Aphelocoma coerulescens (Bosc) 614082 m; A. californica (Vigors) 489825 m, 641219 f; A. wood-
houseii (Baird) 614433 m, 346480 f; Riparia riparia (Linnaeus) 499691 m, 490504 f; Tachyineta bicolor (Viellot)
499694 m; Toxostoma rufum (Linnaeus) 17710; Coccothraustes vespertinus (Cooper) 490800 f; Pinicola enuclea-
tor (Linnaeus) 489737 m, 502668 m; Molothrus ater (Boddaert) 634899 m; Quiscalus quiscula (Linnaeus) 502026
m 502008 f; Pheucticus ludovicianus (Linnaeus) 553159 m; Cardinalis cardinalis (Linnaeus) 611386 m; Passerina
caerulea (Linnaeus) 642184 m, 499098 m, 642137 m. Specimens examined are adults unless otherwise noted; sex
is given if known. Additional skeletons in the Bird Division’s collection were briefly consulted and/or measured to
narrow down identifications and evaluate intraspecific variability.
Below, I present a taxonomic list with descriptions of identification criteria and remarks about the fossil bird bones
from the cave, including those previously identified by Wetmore (1927) and Brodkorb & Mourer-Chauviré (1984).
Branta dickeyi Miller, 1924
Material. USNM PAL 641972, r carpometacarpus: proximal end with damage to trochlea carpalis, collected July
2002 (Fig. 1F).
Description. A large anserine carpometacarpus with a long extensor process of the alular metacarpal that is
perpendicular to the long axis of the bone and narrows distinctly towards the tip. In contrast, the extensor process
is shorter in Chen and Anser; and shorter and wider at the tip in Coscoroba and Cygnus. The fossil agrees well in
morphology with the largest comparative skeletons of Branta canadensis but exceeds them in size (at least if skel-
etons of captive-reared individuals are excluded from comparisons). It is distinctly larger than other North American
species of Anserinae. The extensor process resembles large individuals of B. canadensis in being long and robust.
Rough-surfaced exostoses are present at the tip of the extensor process; these tend to be present in large males of B.
canadensis but can also occur in other Anseriformes.
114 · Zootaxa 4772 (1) © 2020 Magnolia Press
Measurements. In the fossil, the maximum depth from the carpal trochlea through the extensor process is 28.6
mm. This exceeds the range for 12 modern carpometacarpi of B. canadensis measured by Emslie (1995; range
21.8-25.8) and nine measured by me (range 19.8–25.3, including two from South Dakota, within the original range
of the largest subspecies of Canada Goose, B. canadensis maxima Delacour). It closely matches the measurements
recorded by Emslie (1995) for two Irvingtonian fossil carpometacarpi of the fossil species Branta dickeyi, one from
Florida and one from Oregon, both of which measured 28.7 mm.
Remarks. The large fossil goose B. dickeyi has the osteological characteristics of Branta but is roughly the size
of a Tundra Swan (C. columbianus). It has previously been reported from three disparate localities in North America:
the Rancholabrean McKittrick tar seeps of California (Miller 1924), a Blancan locality in Malheur County, Oregon
(Miller 1944), and an early Irvingtonian locality in Florida (Leisey Shell Pit, Emslie 1995). There are few complete
bones among these fossils, and not many bones from each locality. I have ascribed the large but fragmentary carpo-
metacarpus of Branta from Cumberland Bone Cave to the species, although this does require the assumption that
incomplete remains from widely separated localities represent a single species.
In the Oregon and Florida sites, B. dickeyi co-occurs with smaller Branta fossils that match B. canadensis in
size. This supports the view that the fossil species represents an extinct phyletic lineage rather than a chronospecies
of its modern relative. However, if we consider the modern species B. canadensis as a potential modern analog,
interpreting the fossils becomes more complex. B. canadensis is migratory within North America and exhibits con-
siderable geographic variation in body size across its broad range (Aldrich 1946; Delacour 1951). Populations that
breed at high latitudes in northern Canada and Alaska tend to migrate farther than, and to be smaller in body size
than, those that breed at mid-latitudes (Mowbray et al. 2002). (This is independent of the human-mediated expan-
sion in the distribution of large-bodied, resident Canada Geese that began in the 1960s due to captive propagation
and release (Ankey 1996; Mowbray et al. 2002.)) Such a pattern of size variation and migration could theoretically
cause individuals of the same species, but quite different body sizes, to be present in the same fossil site.
The largest modern subspecies of Canada Goose, B. canadensis maxima Delacour, bred in the Great Plains
well west of Cumberland Bone Cave and was thought to be extinct when it was described (Delacour 1951). A small
population of these birds was discovered in 1962 and taken into captive propagation. Releases of the captive-reared
birds into the wild after habituation to human-modified habitats enabled them to expand in geographic distribution,
become resident year-round in many regions, and increase exponentially in population size (Ankey 1996; Mowbray
et al. 2002). Some captive-reared individuals of this subspecies in the USNM collection approach B. dickeyi in
skeletal size, although it is unclear whether birds with no history of captivity attain the same body size. In ascribing
the Cumberland Bone Cave fossil to B. dickeyi, I have followed the lead of prior authors and have left unresolved
the question of whether the fossils of this species represent larger-bodied populations that are ancestral to modern
Anatinae, aff. Anas crecca carolinensis/Spatula discors
Material. CM 34027, l carpometacarpus: proximal end lacking most of the alular metacarpal.
Description. Smaller in size than all but the smallest group of anatids (Table 1). Larger than Oxyura jamaicen-
sis and with a deeper fossa distal to the pisiform process and the carpal trochlea less narrow and blade-like. Larger
than Bucephala albeola, with the shaft of the major metacarpal wider and with a different configuration of rugosities
in the supratrochlear fossa. Surface area of the supratrochlear fossa generally greater than, and the fossa generally
wider distally than, in Spatula discors. Surface area of supratrochlear fossa greater than in A. c. carolinensis. Shaft
depth across the major and minor metacarpals greater than observed in the modern comparative taxa.
Measurements. See Table 1.
Remarks. Fragmentary anatine postcranial bones can be undiagnosable to species. This particular bone was
previously attributed to A. c. carolinensis by Brodkorb & Mourer-Chauviré (1984), and I agree that that is the likely
identification. However, the bone differs in at least minor ways from all comparative skeletons and I cannot exclude
the possibility that it should be referred to S. discors or an extinct species.
THE AVIFAUNA OF CUMBERLAND BONE CAVE, MARYLAND Zootaxa 4772 (1) © 2020 Magnolia Press · 115
TABLE 1. Measurements (mm) of the proximal end of the carpometacarpus in small North American taxa of Anatidae,
in comparison with the Cumberland Bone Cave fossil CM 34027. Depth = shaft depth through major and minor meta-
carpals. Width = width of proximal end excluding os metacarpale alulare (measurement taken from the ventral surface of
Taxon Specimen Depth Width
Fossil CM 34027 4.8 5.0
Anas crecca carolinensis USNM BIRDS 610632 4.0 4.8
USNM BIRDS 610635 4.0 4.7
USNM BIRDS 638839 3.5 4.5
Bucephala albeola USNM BIRDS 492434 3.6 4.8
USNM BIRDS 644552 3.3 4.7
USNM BIRDS 632035 3.2 4.5
Oxyura jamaicensis USNM BIRDS 492491 3.7 4.7
USNM BIRDS 499639 3.9 4.6
USNM BIRDS 623361 3.6 4.3
Spatula discors USNM BIRDS 502641 4.2 4.8
USNM BIRDS 502645 4.2 5.0
USNM BIRDS 502643 4.0 5.0
USNM BIRDS 562841 3.9 5.1
Meleagris sp. Linnaeus
Material. USNM PAL 641967, l humerus: proximal half missing the deltopectoral and bicipital crests and a portion
of the shaft below the head, collected July 2002 (Fig 1C). The bone appears to be from a full adult.
USNM PAL 641969, l humerus: nearly complete shaft missing articular surfaces of both ends and the crista
bicipitalis, collected October 8, 1999 (Fig. 1A). On the caudal surface of the shaft, near the distal end, there are
four distinct circular perforations and several similar lesions that do not fully perforate the bone (Fig. 1A). These
are likely to be carnivore tooth punctures, resulting from an act of predation or scavenging. They show no sign of
healing so they likely occurred at the time of death or afterwards.
USNM PAL 641971, r carpometacarpus: severely abraded proximal end and the proximal 1/3 of the shaft, miss-
ing the minor metacarpal, collected October 8, 1999.
CM 34028, l coracoid: dorsal portion, reported by Brodkorb & Mourer-Chauviré (1984).
USNM PAL 641968, r femur: the shaft and the abraded head are preserved; collected October 8, 1999, by Trent
Spielman. The bone is immature as indicated by pits and striations on the shaft.
Measurements. The following bone measurements correspond with measurements reported by Steadman (1980).
I have repeated Steadman’s letter designations and wording for ease of comparison with his extensive statistical
tables. USNM PAL 641969, humerus measurement C (width of midshaft) 12.5, humerus measurement D (depth
of midshaft) 10.4. USNM PAL 641971, carpometacarpus measurement B (proximal depth) 15.9, measurement C
(length of metacarpal one) 10.3. USNM PAL 641968, femur measurement D (width of midshaft) 9.4, measurement
E (depth of midshaft) 7.3.
Description. Bones of the genus Meleagris are distinguished by very large size in the context of Phasianidae.
The bones listed above are assigned to Meleagris based on size and morphological agreement. The bone measure-
ments from fossils can be compared with Steadman’s summary statistics for approximately 70 specimens of modern
116 · Zootaxa 4772 (1) © 2020 Magnolia Press
M. gallopavo. Measurements of humerus USNM PAL 641969 fall below the reported range for males of M. gal-
lopavo. The width of midshaft of the humerus is near the mean for M. gallopavo females, whereas the depth of mid-
shaft is 0.1 mm greater than the range for females. The proximal depth of the carpometacarpus, USNM PAL 641971,
falls below the reported range for males and females of M. gallopavo by 0.8 mm, and its length of metacarpal one
falls within the range for both male and female M. gallopavo. Measurements of the femur USNM PAL 641968, an
immature bone, fall below the range for males of M. gallopavo. Its width of midshaft is within the range for females,
and depth of midshaft is below the reported range for females by 0.8 mm. Published measurements for the coracoid,
CM 34028, are within the range for females of M. gallopavo.
Remarks. The comparative osteology and fossil record of Meleagris have been extensively studied (e.g., Stead-
man 1980; Bocheński & Campbell 2006). The two extant species of Meleagris show few fixed osteological differ-
ences and considerable intra-specific variation, including pronounced sexual size dimorphism. Only Pleistocene
sites with large numbers of Meleagris bones provide an adequate basis for taxonomic assessment taking into ac-
count variability. The detailed study by Bocheński & Campbell confirmed that the abundant bones from the Rancho
La Brea tar pits (Los Angeles, California) can be diagnosed as a distinct species, M. californica (Miller 1909a),
closely related to M. gallopavo. Steadman studied the Pleistocene fossil record before the Rancholabrean land
mammal age, including larger samples from an Irvingtonian (Coleman 2A, University of Florida Vertebrate Fossil
Locality SM001) and a late Blancan (Inglis 1A, University of California Vertebrate Fossil Locality CI001) site in
Florida. The Irvingtonian turkeys tended to be larger than those from the late Blancan, and the late Blancan birds
lacked a pneumatized scapula which is present in later time periods. Steadman considered it likely that one or more
successive fossil species of Meleagris had a very broad distribution in North America (similar to that of modern M.
gallopavo) and had geographically variable morphology during the Early and Middle Pleistocene. He posited that
most known Blancan and Irvingtonian fossil turkeys represent archaic forms of M. gallopavo.
Steadman (1980) and Bocheński & Campbell (2006) both reported osteological character states that differ at
least in prevalence among various named taxa and local fossil assemblages of Meleagris, but unfortunately, none
of them can be observed in the fossils from Cumberland Bone Cave. Considering their lack of diagnostic species-
level traits, I have refrained from assigning the fossils a species-level identification, while recognizing that the
combined evidence of morphology and biogeography suggests that they represent an archaic form of M. gallopavo,
as envisioned by Steadman. This small collection of mid-Pleistocene turkey bones provides no evidence to refute
Steadman’s finding that Meleagris in North America has increased in body size since the late Blancan.
Material. USNM PAL 641979, l tarsometatarsus: shaft lacking the trochleae and proximal articular surfaces, col-
lected by Trent Spielman, probably on October 2, 1996 (Fig. 1I).
USNM PAL 769092, l tarsometatarsus: shaft lacking most of the plantar surface, collected by Trent Spielman
in the 1990s.
Description. All of the fossils of Tetraoninae from Cumberland Bone Cave fall within or close to the size range
of modern B. umbellus. Among New World Tetraoninae, Bonasa umbellus is much larger than Lagopus leucura but
considerably smaller than all others except Falcipennis canadensis, Lagopus lagopus, and L. muta. Bonasa differs
from Lagopus and Falcipennis in having a proportionately long and slender tarsometatarsus. The two tarsometatarsi
listed above are referable to B. umbellus on that basis. Note that USNM PAL 641979 appears to be from a full adult
and is large compared with some B. umbellus skeletons, but is a good match for USNM BIRDS 641459, a male
from Alberta, Canada.
Remarks. See below.
THE AVIFAUNA OF CUMBERLAND BONE CAVE, MARYLAND Zootaxa 4772 (1) © 2020 Magnolia Press · 117
FIGURE 1. Fossils from Cumberland Bone Cave compared with modern taxa. A) fossil left humerus of Meleagris sp., USNM
PAL 641969, caudal aspect; B) left humerus of Meleagris gallopavo, USNM BIRDS 556334 f, caudal aspect; C) fossil left
humerus of Meleagris sp., USNM PAL 641067, caudal aspect; D) fossil right femur of Coragyps atratus, USNM PAL 641790,
cranial aspect; E) right femur of C. atratus, USNM BIRDS 647499 m, cranial aspect; F) fossil proximal right carpometacarpus
of fossil Branta dickeyi, USNM PAL 641972, ventral aspect; G) proximal right carpometacarpus of B. canadensis, USNM
BIRDS 488182 m, ventral aspect; H) left tarsometatarsus of Falcipennis canadensis, USNM BIRDS 557527 f, plantar aspect;
I) fossil left tarsometatarsus of Bonasa umbellus, USNM PAL 641979, plantar aspect; J) left tarsometatarsus of B. umbellus,
USNM BIRDS 600360 f imm., plantar aspect; K) fossil left tarsometatarsus of Ectopistes migratorius, USNM PAL 769090,
plantar aspect; L) tarsometatarsus of E. migratorius, USNM BIRDS 292904 (captive), plantar aspect; M) fossil left femur of
Megascops guildayi, USNM PAL 769089, caudal aspect; N) left femur of Megascops asio, USNM 623630 f, caudal aspect; O)
fossil right ulna of M. guildayi, USNM PAL 641984, ventral aspect; P) right ulna of M. asio, USNM BIRDS 623630 f, ventral
aspect; Q) right tarsometatarsus of Perisoreus canadensis, USNM BIRDS 639077 f, dorsal aspect; R) fossil right tarsometa-
tarsus of a species of jay (Corvidae, aff. Perisoreus/Cyanocitta), CM 24274, dorsal aspect; S) fossil right tarsometatarsus of a
species of jay (Corvidae, aff. Cyanocitta/Perisoreus), USNM PAL 641989, dorsal aspect; T) right tarsometatarsus of Cyanocitta
cyanea, USNM BIRDS 499504 f, dorsal aspect. Scale bar 1, for images A–G, = 2 cm. Scale bar 2, for images H–N, = 1 cm.
118 · Zootaxa 4772 (1) © 2020 Magnolia Press
Material and Descriptions. USNM PAL 641987, l coracoid: sternal half lacking the lateral process and with much
of the dorsal surface of the bone damaged, collected in the 1990s. The bone agrees in morphological details with
the four species under consideration but does not preserve enough morphological detail to allow for diagnosis at
the species level. It is moderately larger than comparative skeletons of B. umbellus. The bone has been gnawed by
a small mammal, likely a rodent.
USNM PAL 641975, l coracoid: shaft missing sternal end and part of processus acrocoracoideus, collected May
14, 1999. In galliforms, the ventral surface of the shaft typically has a blunt crest running from the acrocoracoid
process nearly to the sternal end of the bone. The fossil differs from L. lagopus, L. mutus, and F. canadensis, and
agrees with B. umbellus, in having this crest more prominent, so that the ventral surface attains a more acute angle.
It differs from B. umbellus and F. canadensis, and agrees more with Lagopus, in having a wider facies articularis
humeralis. This combination of attributes leaves the generic assignment of the bone uncertain.
USNM PAL 641976, l scapula: cranial one-third only, collected on October 8, 1999. The bone is immature,
with its shaft heavily striated and articular surfaces pitted. It is smaller than comparative adult Bonasa umbellus
USNM PAL 641974, l humerus: shaft only, preserving the proximal tip of the brachial depression, collected in
the 1990s. The bone appears adult. The size of the bone and the stout, curvy shaft agree well with B. umbellus and
USNM PAL 11690, l humerus: distal half, referred to B. umbellus by Wetmore (1927). A well-preserved speci-
men that agrees in fine detail with B. umbellus but also closely resembles L. lagopus and F. canadensis. In the com-
parative series examined, B. umbellus tends to have a more distinct olecranon fossa than the other two species. The
fossil tends to agree with B. umbellus in this regard, but the trait is variable and is insufficient grounds for positive
USNM PAL 641988, abraded ulnare collected in the 1990s. An uncatalogued radiale, stored with this one and
with the same provenience data, could well be the same species.
USNM PAL 769091, ulnare: complete, collected on May 14, 1999.
USNM PAL 641978, left carpometacarpus: proximal end only, collected by Trent Spielman probably on Octo-
ber 2, 1996 (illegible).
USNM PAL 641977, l femur: shaft only, collected in 2009. Agrees with B. umbellus and F. canadensis in having
the caudal intramuscular lines less wide-set than in Lagopus. The shaft circumference is greater than in comparative
skeletons of B. umbellus and F. canadensis, but still much smaller than in Tympanuchus.
Remarks. I have referred only two of the ten bones of Tetraoninae in the assemblage to B. umbellus and have
left the rest unidentified to genus, including the distal humerus referred to the species by Wetmore (1927). As Stead-
man (2005) observed, postcranial skeletal elements of B. umbellus and F. canadensis, other than the tarsometatarsus,
generally cannot be distinguished. However, I note that the other eight bones provide no clear evidence to refute
the supposition that B. umbellus is the only species of Tetraoninae present. Note too that although I referred the two
tarsometatarsi to the living species, it is possible that they instead represent a chronospecies of B. umbellus or an
extinct species of Bonasa.
Cumberland Bone Cave provides the sole Irvingtonian fossil record of B. umbellus and the oldest fossil record
of the genus and species. The species has been reported from late Pleistocene and Holocene sites in North America,
particularly in the southeast (e.g., Brodkorb 1959; Wetmore 1962; Steadman 2005). Bonasa umbellus is the only
extant species of grouse that occurs in the mid-latitude Appalachians or in the mixed hardwood forest habitat that is
characteristic of the region at present (Rusch et al. 2000).
USNM PAL 641970, r femur: proximal end and most of the shaft, collected October 8, 1999 (Fig. 1D).
THE AVIFAUNA OF CUMBERLAND BONE CAVE, MARYLAND Zootaxa 4772 (1) © 2020 Magnolia Press · 119
Description. The bone displays adult features including a distinct crista and fossa trochanteris, articular facet
for the antitrochanter, and femoral head with the neck and the fovea ligamentum capitis developed, but it is clearly
immature in that the exposed shaft is densely covered with short, fine striations, the fossa trochanteris is spongy,
and the head retains dense, fine pores. It agrees remarkably in size, form and fine morphological details with Cora-
gyps atratus, including the shaft curvature, proportions of the proximal end including the thick neck, the sharp and
moderately elevated form of the crista trochanteris, and the proximal placement of the pneumatic foramen on the
cranial surface. Larger than Cathartes aura and with the femur shaft distinctly bowed rather than straight as in that
species. An immature individual of H. leucocephalus with similar bone surface textures (USNM BIRDS 611757 f)
has a much longer femur (length from trochanter to the prominent nutrient foramen on the caudal aspect of the shaft,
57.7 mm in H. leucocephalus vs. 37.0 in the fossil) but is less advanced towards adult morphology (e.g., femoral
head much less produced, crista trochanteris not distinct, no fossa trochanteris or fovea ligamentum capitis, articular
facet for the antitrochanter bulbous). Likewise, the fossil is much smaller than the extant Aquila chrysaetos, and
the adult features that the fossil has attained differ in form from the two eagles (crista trochanteris less elevated
proximally above the proximal articular surface than in H. leucocephalus, more elevated and more distinct than in
A. chrysaetos). Amplibuteo concordatus Emslie and Czaplewski 1999, an extinct eagle known from Blancan and
early Irvingtonian sites in Florida, was not directly compared but is unlikely to be relevant due to its eagle-like rather
than Coragyps-like osteology.
Measurements. Howard (1968) summarized measurements from a large series of Coragyps occidentalis (Miller
1909b) bones from the Rancho La Brea tar pits, of which only the least transverse breadth of the shaft (8.9 mm) can
be observed in the Cumberland Bone Cave femur. For this measurement, Howard reported a mean of 9.1 mm in
modern C. atratus and a minimum of 9.0 mm in 65 femora of C. occidentalis.
Remarks. The late Pleistocene (Rancholabrean) species C. occidentalis, is known from multiple localities in
southwestern North America and is believed to have become extinct along with the Pleistocene megafauna (Brasso
& Emslie 2006). Bones of this species are larger on average than in modern C. atratus (Howard 1962, 1968; Frailey
1972). Because the fossil femur has not attained adult surface textures, we cannot exclude the possibility that its full
adult size would be above the range of modern C. atratus and within the range of the larger late Pleistocene species.
It is notable that C. atratus has not moved into most of the southwestern range of C. occidentalis in the time since
the extinction of the latter, roughly ten thousand years ago. This suggests that C. occidentalis occupied arid habitats
that are not suitable for C. atratus.
The immature bone of C. atratus provides evidence that the species was breeding in or near the cave. We can
rule out the possibility that the bird had flown from elsewhere to reach Cumberland Bone Cave by analogy with a
growth series of California Condor (G. californianus) skeletons. The developmental stage of the fossil most closely
matches that of a condor that died at 136 days old (USNM BIRDS 658188), in which the femur is likewise at or
close to full adult size and form, has a finely striated shaft, dense pores on the femoral head, and the fossa trochan-
teris still spongy. The fossil femur is more advanced developmentally than a bird that died at 80 days old (USNM
BIRDS 658191), which has not attained adult size and has the articular ends not fully formed or ossified. In two
older condor chicks that died ten or fifteen days after the usual age of fledging (AMNH 32153, 184 days, and USNM
BIRDS 658221, 187 days), the surface of the femoral shaft has attained the smooth surface texture and rugose to-
pography seen in adult birds, although both femora retain some porous structure on the proximal articular surface.
The G. californianus skeleton with the femur at a similar stage of development to the fossil has a relatively un-
derdeveloped forelimb, with the major wing bones still spongy and growing at the articular surfaces and the carpals
and metacarpals unfused, and has the sternum still entirely porous and spongy (see USNM BIRDS 658188). It is no
surprise, then, that California Condor chicks typically fledge at an older age (about 30 to 40 days older) than that of
the individual that best matched the fossil (Snyder & Snyder 2000). Assuming the developmental pattern is similar
in Coragyps, the fossil vulture from Cumberland Bone Cave would not yet have been able to fly.
120 · Zootaxa 4772 (1) © 2020 Magnolia Press
Aquila chrysaetos (Linnaeus)/A. bivia Emslie & Czaplewski 1999
Material. CM 34018, r ungual phalanx with tip and base missing.
Description. Corresponds in size and shape with the terminal phalanx of digits one (the hind toe) and two of
an eagle. Brodkorb & Mourer-Chauviré (1984) referred it to A. chrysaetos rather than H. leucocephalus based on
large overall size and more gentle distal tapering of the claw core (plantar view), implying a longer claw. Based
on these criteria, I found it to be larger than all comparative specimens of H. leucocephalus and most comparative
skeletons of A. chrysaetos. It agrees well in size and shape with one comparative skeleton of A. chrysaetos (USNM
BIRDS 19724). However, the possibility that it belongs to the larger early Irvingtonian species Aquila bivia cannot
Remarks. This large ungual phalanx appears to be attributable either to the Golden Eagle (A. chrysaetos) or to
a larger, early Irvingtonian species of Aquila described from Florida and Arizona (Emslie & Czaplewski 1999). The
modern distribution of the Golden Eagle in North America is primarily in the west, but the species does have a rarely
observed wintering population in the region of Cumberland Bone Cave (Brodeur et al. 1996).
Material. USNM PAL 641980, l coracoid: dorsal one-third, collected October 8, 1999.
Description. The bone falls in the size range of large males and small females of modern A. cooperi. I compared
it with modern species of North American Accipitriformes in roughly the same size class. The fossil is much more
slender and less pneumatic than in Elanoides forficatus or Rostrhamus sociabilis. The pneumatic foramen in the
sulcus m. supracoracoidei is smaller and the acrocoracoid process is less inflated than in Circus cyaneus. The fossil
resembles A. cooperi as opposed to Buteo lineatus and B. platypterus in having a relatively slender facies articularis
humeralis (glenoid facet) and a less extended procoracoid process.
Remarks. This species still occurs in the region of the cave. It was previously reported from other Irvingtonian
fossil localities and from a late Blancan locality in Florida (Emslie 1998).
Accipitridae, aff. Accipiter cooperi
Material. USNM PAL 641996, pedal phalanx, entire, with the articular cotyla slightly crushed; collected in the
USNM PAL 641983, ungual phalanx, entire; collected Sept 22, 1994.
Description. Neither bone is diagnosable at the species level but both compare well with A. cooperi in size and
Remarks. See above.
Material. USNM PAL 641982, ungual phalanx: proximal 1/3, collected July 2002.
Description. Agrees in size with the third or fourth ungual phalanx in H. leucocephalus and A. chrysaetos. Its
small size and narrow articular cotyla suggest it is from the fourth digit. Although there is morphological overlap
between the two species in this element, the fossil can be assigned to H. leucocephalus as opposed to A. chrysaetos
based on the following combination of traits: articular cotyla narrower (proximal view) and with a tighter curve
(lateral view), flexor tubercle bulbous and distended as opposed to crested and elongated.
Remarks. The assignment of this fossil to H. leucocephalus is based on morphological agreement but it comes
with the caveat that comparisons with the Plio-Pleistocene fossil eagle from Florida and Arizona, Amplibuteo con-
cordatus Emslie & Czaplewski 1999, were not possible. However, based on the published measurements of its long
bones, A. concordatus was smaller in body size than H. leucocephalus or A. chrysaetos. Haliaeetus leucocephalus
currently occurs in the region of the cave.
THE AVIFAUNA OF CUMBERLAND BONE CAVE, MARYLAND Zootaxa 4772 (1) © 2020 Magnolia Press · 121
Material. CM 34020, r coracoid: cranial portion lacking procoracoid process, referred to E. migratorius by Brod-
korb & Mourer-Chauviré (1984). Not reexamined for this study.
USNM PAL 769090, l tarsometatarsus: shaft missing the proximal end and the trochleae, collected Sept 22,
1994 (Fig. 1K).
Description. The tarsometatarsus is much larger than in Zenaida and closer in size to those of males of Patagio-
enas fasciata. Proportionately long and slender compared with P. fasciata, P. leucocephala, and P. cayanensis. The
median crest on the plantar surface increases in depth as it extends proximally to join the hypotarsus in Patagioenas
but is shallower along its full length in the fossil. The fossil shares a shallow median crest with those of Ectopistes
and Zenaida. This crest is less deep proximally in P. leucocephala than in the other Patagioenas, but still more deep
than in Ecotopistes and the fossil. Only two of the five comparative skeletons of Ectopistes include the tarsometa-
tarsus, and in both cases the bone is distinctly smaller than the fossil. Three additional partial comparative skeletons
are known to be males, and these three have considerably larger skeletons, suggesting that the fossil is within the
size range of male Ectopistes.
Remarks. The Cumberland Bone Cave bones referred to E. migratorius appear to be the oldest documented
fossil record of the species, which is also known from late Pleistocene (Rancholabrean) sites including some west-
ern localities outside the species’ historical range (Hargrave & Emslie 1980; Chandler 1982). The cave locality falls
within the primary breeding range of the species during the 19th century (Blockstein 2002).
cf. Ectopistes migratorius
Material. USNM PAL 641973, r coracoid, cranial half missing part of the acrocoracoid process, collected Sept. 22,
Description. Agrees in size and morphological detail with a male comparative skeleton of E. migratorius
(USNM BIRDS 224320), including in having a slender procoracoid process.
Remarks. See above.
Megascops guildayi (Brodkorb and Mourer-Chauviré 1984)
Material. USNM PAL 641984, r ulna: shaft and proximal end, collected in 1991 (Fig. 1O).
USNM PAL 641985, r ulna: distal articular end only, collected October 2, 1996.
USNM PAL 769089, l femur: proximal 2/3, collected Sept. 22, 1994 (Fig. 1M).
CM 8040 holotype, l tarsometatarsus lacking proximal end. Not reexamined for this study.
USNM PAL 769088, pedal phalanx: entire, collected Sept. 22, 1994.
USNM PAL 641981, ungual phalanx: entire, collected Sept. 22, 1994.
Description. The bones of a strigid owl listed above are referred to M. guildayi based primarily on the criteria of
size (smaller than Asio otus, larger than Megascops asio and M. kennicottii) and agreement in morphological details
with modern Megascops. The proximal half of the ulna agrees with those of Megascops, Athene, and Aegolius in
being more curved than in Asio, Strix, or Surnia. The proximal articular surfaces of the femur are broader in relation-
ship to depth than in Athene or Aegolius, agreeing in this trait with Megascops and larger owls. No complete long
bones of the species are known, and the larger size of the fossil species compared with other Megascops is evident
122 · Zootaxa 4772 (1) © 2020 Magnolia Press
mainly from the thicker shafts of the ulna and femur. The tarsometatarsus, described and figured by Brodkorb &
Mourer-Chauviré (1984: Figure 1), is referable to Megascops based on size and narrow, slender form, though pro-
portionately stouter and shorter than in Athene cunicularia. The tarsometatarsus is not as stout as in Asio brevipes
Ford & Murray 1967 from the Upper Pliocene of Idaho nor as slender as in Athene megalospeza (Ford 1966) from
the Upper Pliocene of Kansas and Idaho.
Measurements. USNM PAL 641984 (ulna), greatest width distal to cotylae, 5.2 mm. USNM PAL 769089 (fe-
mur), proximal lateromedial width from head through trochanter, 7.0 mm; width of head, 3.3 mm.
Remarks. The only previously known material of this species is the type specimen. The new material bolsters
the evidence for a large extinct species of Megascops in the Irvingtonian of Maryland. It is odd that the species has
no other fossil record, but perhaps some of the Pliocene fossils referred to Otus (=Megascops) are relevant to it (i.e.,
two bones from the Hagerman local fauna of Idaho (Ford & Murray 1967) and one from the Rexroad Formation of
Kansas (Ford 1966)).
Material. USNM PAL 641986, r femur: distal end only, collected May 14, 1999.
Description. Smaller than, and with the fibular trochlea wider than in, Tyto alba. Among extant North American
owls, closest to Asio otus and males of Surnia ulula in size. Differs from S. ulula and agrees with A. otus in having
the medial condyle less bulbous and projecting less far mediad, the popliteal fossa and the intercondylar sulcus shal-
lower, and the juncture of the tibiofibular crest with the shaft forming a tight curve as opposed to a shallow slope.
Measurements. The maximum distal breadth in the fossil (8.4 mm) falls just below the range reported by Emslie
(1982) for A. otus (8.5-10.2 mm, n=10).
Remarks. The fossil compares favorably with Asio, however, the possibility that it represents a large female of
Megascops guildayi cannot be ruled out. I consider the bone to be undiagnostic at the level of the genus.
Material. USNM PAL 641993, r tarsometatarsus: distal extremity lacking trochlea metatarsi IV, with shaft extend-
ing to just proximal of fossa metatarsi I, collected in the 1990s. The bone appears fully adult.
Description. The fossil closely resembles but is somewhat more gracile than Sayornis phoebe. I consider it to
be undiagnostic at the genus and species level.
Corvidae, aff. Cyanocitta/Perisoreus
Material. CM 24274, r tarsometatarsus with damage to the hypotarsus and repair to the shaft (Fig. 1Q). Referred to
Perisoreus canadensis by Brodkorb & Mourer-Chauviré (1984).
USNM PAL 641989, r tarsometatarsus: entire, collected in the 1990s (Fig 1s). A break mid-shaft has been re-
USNM PAL 641990, l tarsometatarsus: distal three-fourths, collected July 2002.
USNM PAL 641991, r tibiotarsus: distal half, collected Sept. 22, 1994.
USNM PAL 641994, l carpometacarpus lacking proximal end, collected July 2002.
Description. These five passerine bones are recognizable as New World jays, family Corvidae, based on size
and morphological agreement. They represent either a single species or multiple species of similar size. They cannot
THE AVIFAUNA OF CUMBERLAND BONE CAVE, MARYLAND Zootaxa 4772 (1) © 2020 Magnolia Press · 123
be assigned to any of several fossil species of North American jays of late Blancan and Irvingtonian ages (Protocitta
dixi Brodkorb 1957, Protocitta ajax Brodkorb 1972, Henocitta brodkorbi Holman 1959), which are considerably
larger in body size. One of the fossil tarsometatarsi (CM 24274) has a particularly narrow shaft distally, and the
tibiotarsus also has a narrow distal shaft in relationship to condyle breadth. These traits are not diagnostic to species
according to the morphometric analysis described below.
Measurements. See Appendix 1 and Figures 2-4.
Morphometric Analysis. I used a morphometric approach to compare the fossils with modern species of jays
from continental North America north of Mexico (Canada Jay P. canadensis, Blue Jay Cyanocitta cristata, three
species of scrub jays, Aphelocoma coerulescens, woodhouseii, and californica, and the larger Steller’s Jay Cya-
nocitta stelleri). Bone lengths could only be taken from the fossil carpometacarpus and the two tarsometatarsi. Box-
plots for these measurements illustrate differences in intermembral proportions among the taxa, with Aphelocoma
having a long tarsometatarsus relative to carpometacarpus, and Perisoreus and even more so Cyanocitta cristata
having a short tarsometatarsus relative to carpometacarpus (Fig. 2). For the fossils, even with the small sample size,
it can be seen that if only one species of jay is present, its body proportions are not consistent with assignment to
Aphelocoma. The fossil tarsometatarsi are also distinctly smaller than in C. stelleri.
FIGURE 2. Box and whisker plots summarizing long bone lengths in North American jays. A) Carpometacarpus, length of
major metacarpal (mm); B) tarsometatarsus length (mm). Data from Appendix 1.
To evaluate the fossil tibiotarsus, width of the distal shaft and width across the distal condyles were taken. The
ratio of these measurements shows that the fossil tibiotarsus has an unusually slender shaft (Fig. 3). It agrees with
P. canadensis in this trait although it is also within the range of A. woodhouseii.
Brodkorb & Mourer-Chauviré (1984) referred the tarsometatarsus CM 24274 to P. canadensis based on its
slender shaft. To evaluate the intramembral proportions of the two fossil tarsometatarsi, I measured tarsometatar-
sus length and four width measurements covering proximal and distal articular and shaft widths (Appendix 1). I
performed a principal components analysis using four variables: 1) length from intercotylar area to the trochlea of
the third metatarsal, 2) proximal width across articular facets, 3) proximal shaft width at level of proximal vascular
foramina, 4) distal shaft width at level of distal vascular foramen (Fig. 4). I also ran this analysis with five variables
(distal width across trochleae included), with very similar results (not shown). In the four-variable analysis, for the
first principal component, the variables’ loadings had the same sign and a moderate range of values (variable 1 =
-0.52, variable 2 = -0.55, variable 3 = -0.49, variable 4 = -0.44). This component, weighted most heavily on length
and the proximal widths, reflects size and also some shape variance, as clearly shown by the reversal of the posi-
tions of C. cristata and P. canadensis relative to their tarsometatarsus lengths (compare Figs. 2b and 4). The second
principal component contrasted tarsometatarsus distal shaft width (component loading -0.80) with proximal shaft
width (loading +0.48) and total length (loading +0.33). This component tended to distinguish A. californica and
A. woodhouseii from the other taxa but otherwise was ineffective at separating the species. The two Cumberland
Bone Cave fossils are centrally placed in the principal components biplot (Fig. 4). CM 24274 falls just outside the
probability ellipse for P. canadensis, which supports Brodkorb & Mourer-Chauviré’s opinion of the bone; however,
it also falls within the probability ellipse for C. cristata and just outside the ellipses of the three species of Aphelo-
124 · Zootaxa 4772 (1) © 2020 Magnolia Press
coma. USNM PAL 641989 falls near the center of the plot, within the ellipses for C. cristata and A. coerulescens.
Considering that these are merely 68% probability ellipses, representing one standard deviation, I conclude that the
tarsometatarsus data are indecisive with regard to the genus or species of jay(s) represented.
FIGURE 3. Box and whisker plot summarizing the ratio of shaft width to distal condylar width in North American jays. Data
from Appendix 1.
FIGURE 4. Plot of the first two principal components for tarsometatarsus dimensions in North American jays. Data from Ap-
Remarks. The fossil jays received extra scrutiny because Brodkorb & Mourer-Chauviré cited their identifica-
tion of P. canadensis as evidence that the Cumberland Bone Cave assemblage accumulated during a cold climate
phase. Although the osteometric analysis eliminated C. stelleri on size and provisionally eliminated Aphelocoma on
body proportions (assuming only one species is represented by the fossils), it was otherwise indecisive as to the ge-
nus and species represented. Perisoreus canadensis has also been reported as a fossil in another southerly location,
THE AVIFAUNA OF CUMBERLAND BONE CAVE, MARYLAND Zootaxa 4772 (1) © 2020 Magnolia Press · 125
Cheek Bend Cave in Tennessee (Parmalee & Klippel 1982). The bird and mammal assemblage from Cheek Bend
Cave includes a number of boreal species and was interpreted as dating to the Last Glacial Maximum. In the case
of Cumberland Bone Cave, Perisoreus, if present, would be the only purely boreal element in the avian assemblage
Material. USNM PAL 641992, l mandibular ramus lacking symphyseal part and medial process. Collected Oct. 7,
Description. Similar to but somewhat larger than in Junco hyemalis and somewhat smaller than in Zonotrichia
Material. USNM PAL 641995, l quadrate lacking orbital process. Collected in the 1990s.
Description. Agrees in general form with species in the Fringillidae and Cardinalidae that have finch-like bill
shapes. The limited extent of pneumatization of the caudal surface of the otic process suggests that the quadrate is
assignable to Fringillidae not Cardinalidae. In size, it is somewhat smaller than Cardinalis cardinalis.
North American birdlife during the Irvingtonian land mammal age is best known from the southern Rocky Moun-
tains (Porcupine Cave, Emslie 2004) and from several localities in peninsular Florida (Emslie 1995, 1998). The as-
semblage from Cumberland Bone Cave comprises at least 14 species (Table 2) and helps to fill the geographic gaps
in distribution of avian fossils from this time period. The ecological types represented as fossils include ground-
feeding birds (a turkey, the Ruffed Grouse, the Passenger Pigeon), a scavenger (Black Vulture), a number of rapto-
rial species (Bald Eagle, Coopers Hawk, Golden Eagle or a close relative, a species of screech owl), a few aquatic
species (a large goose, a small duck), and several perching birds (a flycatcher, a jay, a junco or sparrow, and a finch).
Immature bones in the assemblage document breeding records for turkeys, grouse, and the Black Vulture. The fos-
sils include new anatomical parts for the large extinct screech owl that is known only from this locality and appear to
provide the oldest fossil record of Passenger Pigeon, extending the age range of the species to the mid-Pleistocene,
approximately 700 to 800 kya. The large goose, B. dickeyi, has a broad geographic and long temporal distribution, as
it is now reported from the Blancan of Oregon, the early Irvingtonian of Florida, the Irvingtonian of Maryland, and
the Rancholabrean of California (although see the species account for a caveat about this poorly known species).
The proportion of extinct species in the avifauna is 20% or higher (minimally, Branta dickeyi, Ectopistes migra-
torius, and Megascops guildayi, plus there is uncertainty about several other taxa such as Meleagris sp. and Aquila
chrysaetos/bivia). Whether any of these represent extinctions of a phyletic lineage by the end of the Irvingtonian is
debatable. The Passenger Pigeon became extinct only a little more than a century ago. Branta dickeyi may repre-
sent the extinction of a phyletic lineage, but it’s taxonomic status is uncertain and its fossil record extends into the
Rancholabrean. In his comprehensive study of Blancan and Irvingtonian fossil turkeys, Steadman (1980) concluded
that the modern Wild Turkey is a lineal descendant of fossil turkeys that have been geographically widespread in
North America since Blancan and Irvingtonian times. Similarly, Brodkorb and Mourer-Chauviré (1984) considered
it likely that their new species M. guildayi represents a larger Pleistocene form of the Eastern Screech Owl, M. asio,
rather than an extinct lineage. From this perspective, it is possible to interpret the avifauna of the cave as containing
no phyletic extinctions that were restricted to the Irvingtonian.
126 · Zootaxa 4772 (1) © 2020 Magnolia Press
TABLE 2. Taxonomic list of birds identified in the Cumberland Bone Cave Local Fauna. NISP, Number of Identified Specimens.
Taxon Common Name or Informal Description NISP
Branta dickeyi large extinct goose 1
Anatinae, aff. Anas crecca carolinensis/Spatula discors similar to Green- or Blue-winged Teal 1
Meleagris sp. turkey 5
Bonasa umbellus Ruffed Grouse 2
Tetraoninae sp. similar to Ruffed Grouse 9
Coragyps atratus Turkey Vulture 1
Aquila chrysaetos/bivia Golden Eagle or larger extinct eagle 1
Accipiter cooperi Cooper’s Hawk 1
Accipitridae, aff. Accipiter cooperi similar to Cooper’s Hawk 2
Haliaeetus leucocephalus Bald Eagle 1
Ectopistes migratorius Passenger Pigeon 2
cf. Ectopistes migratorius similar to Passenger Pigeon 1
Megascops guildayi large extinct screech owl 6
Strigidae sp. Medium-sized owl 1
Tyrannidae sp. similar to Eastern Phoebe 1
Corvidae, aff. Cyanocitta/Perisoreus similar to Blue or Canada Jay 5
Passerellidae sp. a junco or sparrow 1
Fringillidae or Cardinalidae
Fringillidae/Cardinalidae sp. a finch 1
For the sake of paleoecological interpretation, it was important to scrutinize the five bones attributable to New
World jays because one of them was previously identified as a northern boreal and alpine species, the Canada Jay.
This brought disharmony into the faunal list, considering that the modern distributions of Canada Jay and the Black
Vulture are non-overlapping. The Black Vulture fossil from Cumberland Bone Cave occurs near the northern limit
of the species’ current breeding range (Buckley 1999), and the species historically had an even more southerly dis-
tribution, with the first breeding record in Maryland not occurring until 1922 (Robbins & Blom 1996). Canada Jays
have a northern boreal distribution in the eastern half of North America and extend farther south only in western
montane regions (Strickland & Ouellet 2018). Modern Wild Turkeys, Coopers Hawks, and screech owls also have
essentially non-overlapping ranges with the Canada Jay if both geographic distribution and habitat type are taken
into account. Considering that my osteometric analysis of the jay fossils neither confirmed nor excluded the pres-
ence of Canada Jay in the avifauna, the most parsimonious interpretation is that the species was absent. Under this
assumption, there are no purely boreal elements in the Cumberland Bone Cave avifauna.
Setting aside the Canada Jay, all of the extant or potentially extant species identified still occur in the region
of the cave today (as would the Passenger Pigeon if not forced into extinction). The avifauna thus paints a picture
of only modest avifaunal turnover between the mid-Pleistocene and the late Holocene in the mid-latitude Appala-
chians. This is not to say that habitats were the same, especially considering the importance of human-modified
habitats to modern populations of species like the Wild Turkey, Black Vulture, and Cooper’s Hawk.
THE AVIFAUNA OF CUMBERLAND BONE CAVE, MARYLAND Zootaxa 4772 (1) © 2020 Magnolia Press · 127
To gain insight into Irvingtonian habitats, it is worth considering the habitats used by modern counterparts of the
fossil taxa. Canada Geese make use of diverse habitats, from lakes and rivers to marshes, meadows, and agricultural
and urban habitats. They are not a woodland species except in the presence of wetlands, lakes, or rivers. Wild Turkey
use a wide variety of mixed woodlands and grassy savannah habitats across their extensive range in North America.
They use trees for roosting and fruit, seed, and mast for food. Hardwoods are often an important forest component
in their habitat, and the presence of some more open habitat such as small pastures is preferred, especially for breed-
ing. They are absent from high mountain regions and part of the Great Plains, and barely extend northward into
Canada. Ruffed Grouse have a largely boreal distribution but descend to Tennessee in the Appalachians; they are
year-round residents over most of their range. Aspen woodlands and boreal forest are primary habitat types but in
the southeastern part of their range they associate with mixed deciduous-coniferous forest (oaks, hickories, pines).
Males seek logs or other prominences for drumming, preferably located in protected understory clearings in early
successional forest (Rusch et al. 2000). Black Vultures often seek the protection of forest for nesting and roosting;
their distribution in forest tends to be riverine. They frequently lay their eggs on the ground in crevices or cavities
(Robbins & Blom 1996), which may explain the presence of an unfledged chick in the cave site. As visual foragers,
Black Vultures prefer to forage in more open habitat, and they require large home ranges during breeding (Coleman
& Fraser 1989). Bald Eagles favor larger rivers, lakes, and shorelines but also occur along smaller rivers, including
in the region of the cave locality. They prefer large trees near water for nesting and roosting but will nest on the
ground if trees are absent. Golden Eagles are primarily birds of open habitat across their cosmopolitan distribution;
however, birds that breed near Hudson Bay have been found to secretively over-winter in montane forest in the mid-
latitude Appalachians, not far from the cave site. Cooper’s Hawks are broadly distributed in deciduous, mixed, and
coniferous forest and in small woodlots in North America. They tend to forage in wooded habitats more than open
ones. Passenger Pigeons fed primarily on mast of beeches, oaks, and to a lesser extent, chestnuts, as well as grain
and fruit. They required large expanses of forest for feeding. Most species of screech owls are neotropical; the two
species with mainly temperate distributions occur in a tremendous variety of treed habitats but most commonly in
riverine forest. They generally avoid high montane regions, and when they do enter these regions, they are often
restricted to valley bottoms.
Taken together, these observations suggest that in the mid-Pleistocene, roughly 700 to 800 kya, habitat near
Cumberland Bone Cave included mixed forest with mast-producing deciduous hardwoods like beeches and oaks.
Both early and later successional stages were probably represented. A nearby river, lake or wetland would have pro-
vided habitat for Canada Goose, Bald Eagle, a screech owl and a small duck. Wills Creek currently provides such
habitat near the site, and the North Branch of the Potomac River is close enough for eagle foraging. There are no
clear boreal elements in the avifauna, and the habitats near the cave may not have differed greatly from those of the
late Holocene. The exception may be that the region surrounding the cave apparently included enough open habitat
for Black Vulture foraging during the breeding season, well before human influence.
I am grateful to Trent Spielman for donating the collection to the USNM and to Frederick V. Grady and Ralph
Eshelman for encouraging me to study it. I thank Ami Henrici of the Carnegie Museum of Natural History for the
loan of fossils, Mark Florence in the USNM Paleobiology Department for assistance with registration of the fossils,
Brian Schmidt for creating Figure 1, and Sorilis Ruiz-Escobar for insights on skeletal ontogeny of condors. I thank
Steven Emslie for discussion of Irvingtonian fossil birds and Ralph Eshelman and Frederick Grady for comments
on the manuscript.
Aldrich, J.W. (1946) Speciation in the White-cheeked Geese. The Wilson Bulletin, 58 (2), 94–103.
Ankey, C.D. (1996) An embarrassment of riches: Too many geese. Journal of Wildlife Management, 60 (2), 217–223.
Baumel, J.J. & Witmer, L.M. (1993) Osteologia. In: Baumel, J.J., King, A.S., Breazile, J.E., Evans, H.E. & Vanden Berge, J.C.
(Eds.), Handbook of Avian Anatomy: Nomina Anatomica Avium. 2nd Edition. Publications of the Nuttall Ornithological
128 · Zootaxa 4772 (1) © 2020 Magnolia Press
Club No. 23. Nuttall Ornithological Club, Cambridge, Massachusetts, pp. 45–132.
Bell, C.J. (2000) Biochronology of North American microtine rodents. In: Noller, J.S., Sowers, J.M. & Lettis, W.R. (Eds.), Qua-
ternary Geochronology, Methods and Applications. AGU Reference Shelf 4, American Geophysical Union, Washington,
D.C., pp. 379–406.
Bell, C.J., Lundelius, E.L. Jr., Barnosky, A.D., Graham, R.W., Lindsay E.H., Ruez, D.R. Jr., Semken, H.A. Jr., Webb, S.D. &
Zakrzewski, R.J. (2004) The Blancan, Irvingtonian, and Rancholabrean Mammal Ages. In: Woodburne, M.O. (Ed.), Late
Cretaceous and Cenozoic Mammals of North America: Biostratigraphy and Geochronology. Columbia University Press,
New York, New York, pp. 232–314.
Blockstein, D.E. (2002) Passenger Pigeon (Ectopistes migratorius). Version 2.0. In: Poole, A.F. & Gill, F.B. (Eds.), The Birds
of North America. Cornell Lab of Ornithology, Ithaca, New York. Available from: https://birdsna.org/Species-Account/bna/
home (accessed 30 January 2020)
Bocheński, Z.M. & Campbell, K.E. (2006) The extinct California turkey, Meleagris californica, from Rancho La Brea: Com-
parative Osteology and Systematics. Contributions in Science of the Natural History Museum of Los Angeles County, 509,
Brasso, R.L. & Emslie, S.D. (2006) Two new Late Pleistocene avifaunas from New Mexico. The Condor, 108, 721–730.
Brodeur, S., Décarie, R., Bird, D.M. & Fuller, M. (1996) Complete migration cycle of Golden Eagles breeding in northern
Quebec. The Condor, 98, 293–299.
Brodkorb, P. (1957) New passerine birds from the Pleistocene of Reddick, Florida. Journal of Paleontology, 31 (1), 129–138.
Brodkorb, P. (1959) The Pleistocene avifauna of Arredondo, Florida. Bulletin of the Florida State Museum, Biological Sciences,
4 (9), 271–291.
Brodkorb, P. (1972) Neogene fossil jays from the Great Plains. The Condor, 74 (3), 347.
Brodkorb, P. & Mourer-Chauviré, C. (1984) Pleistocene Birds from Cumberland Cave, Maryland. In: Genoways, H.H. & Daw-
son, M.R. (Eds.), Contributions in Quaternary Vertebrate Paleontology: A Volume in Memorial to John E. Guilday. Carn-
egie Museum of Natural History, Special Publication No. 8, pp. 39–43.
Buckley, N.J. (1999) Black Vulture (Coragyps atratus). Version 2.0. In: Poole, A. F. & Gill, F. B. (Eds.), The Birds of North
America. Cornell Lab of Ornithology, Ithaca, New York. Available from https://birdsna.org/Species-Account/bna/home
(accessed 30 January 2020)
Chandler, R.M. (1982) A second record of Passenger Pigeon from California. The Condor, 84, 242.
Chesser, R.T., Burns, K.J., Cicero, C., Dunn, J.L,. Kratter, A.W., Lovette, I.J., Rasmussen, P.C., Remsen, J.V. Jr., Stotz, D.F. &
Winker, K. (2019) Check-list of North American Birds. Online. American Ornithological Society. Available from http://
checklist.aou.org/taxa (accessed 21 February 2020)
Cohen, K.M. & Gibbard, P. (2011) Global chronostratigraphical correlation table for the last 2.7 million years. Subcommission
on Quaternary Stratigraphy (International Commission on Stratigraphy), Cambridge. Available from: http://quaternary.
stratigraphy.org/charts/ (accessed 3 March 2020)
Coleman, J.S. & Fraser, J.D. (1989) Habitat use and home ranges of Black and Turkey Vultures. Journal of Wildlife Manage-
ment, 53, 782–792.
Delacour, J. (1951) Preliminary note on the taxonomy of Canada Geese, Branta canadensis. American Museum Novitates, 1537,
Dunning, J.B. Jr. (2008) CRC Handbook of Avian Body Masses. 2nd Edition. Taylor & Francis, Boca Raton, Florida, vii + 655
Emslie, S.D. (1982) Osteological identification of Long-Eared and Short-Eared Owls. American Antiquity, 47 (1), 155–157.
Emslie, S.D. (1995) An early Irvingtonian avifauna from Leisey Shell Pit, Florida. Bulletin of the Florida Museum of Natural
History, 37, Part I (110), 299–344.
Emslie, S.D. (1998) Avian community, climate, and sea-level changes in the Plio-Pleistocene of the Florida Peninsula. Ornitho-
logical Monographs, No. 50, 1–113.
Emslie, S.D. (2004) The Early and Middle Pleistocene Avifauna from Porcupine Cave. In: Barnosky, A.D. (Ed.), Biodiversity
response to climate change in the Middle Pleistocene: The Porcupine Cave Fauna from Colorado. University of California
Press, Berkeley, California, pp. 127–140.
Emslie, S.D. & Czaplewski, N.J. (1999) Two new fossil eagles from the late Pliocene (late Blancan) of Florida and Arizona and
THE AVIFAUNA OF CUMBERLAND BONE CAVE, MARYLAND Zootaxa 4772 (1) © 2020 Magnolia Press · 129
their biogeographic implications. Smithsonian Contributions to Paleobiology, 89, 185–198.
Ford, N.L. (1966) Fossil owls from the Rexroad Fauna of the Upper Pliocene of Kansas. The Condor, 68, 472–475.
Ford, N.L. & Murray, B.G. Jr. (1967) Fossil owls from the Hagerman local fauna (Upper Pliocene) of Idaho. The Auk, 84,
Frailey, C.D. (1972) Additions to the Pleistocene avifauna of Arredondo, Florida. Quarterly Journal of the Florida Academy of
Sciences, 35 (1), 53–54.
Gidley, J.W. (1914) Preliminary report of a recently discovered Pleistocene cave deposit near Cumberland, Maryland. Proceed-
ings of the National Museum, 46 (2014), 93–102.
Gidley, J.W. (1920) Pleistocene peccaries from the Cumberland Cave Deposit. Proceedings of the United States National Mu-
seum, 57, 651–678.
Gidley, J.W. & Gazin, C.L. (1938) The Pleistocene vertebrate fauna from Cumberland Cave, Maryland. Bulletin of the United
States National Museum, 171, 1–199.
Hargrave, L.L. & Emslie, S.D. (1980) Passenger Pigeon bones from archaeological sites in New Mexico. Contributions in Sci-
ence of the Natural History Museum of Los Angeles County, 330, 257–260.
Holman, J.A. (1959) Birds and mammals from the Pleistocene of Williston, Florida. Bulletin of the Florida State Museum,
Biological Sciences, 5 (1), 1–24.
Howard, H. (1962) Bird remains from a prehistoric cave deposit in Grant County, New Mexico. The Condor, 64 (3), 241–242.
Howard, H. (1968) Limb measurements of the extinct vulture, Coragyps occidentalis, with a description of a new subspecies.
Papers of the Archaeological Society of New Mexico, 1, 115–128.
Le Conte, J.L. (1848) On Platygonus compressus: a new fossil pachyderm. Memoirs of the American Academy of Arts and Sci-
ences, New Series, 3, 257–274.
Miller, L.H. (1909a) Pavo californicus, a fossil peacock from the Quaternary asphalt beds of Rancho La Brea. University of
California Publications, Bulletin of the Department of Geology, 5 (19), 285–289.
Miller, L.H. (1909b) Teratornis, a new avian genus from Rancho La Brea. University of California Publications, Bulletin of the
Department of Geology, 5 (21), 305–317.
Miller, L.H. (1924) Branta dickey from the McKittrick Pleistocene. The Condor, 26, 178–180.
Miller, L.H. (1944) Some Pliocene birds from Oregon and Idaho. The Condor, 46, 25–32.
Mowbray, T.B., Ely, C.R., Sedinger, J.S. & Trost, R.E. (2002) Canada Goose (Branta canadensis). Version 2.0. In: Poole, A.F. &
Gill, F.B. (Eds.), The Birds of North America. Cornell Lab of Ornithology, Ithaca, New York. Available from https://birdsna.
org/Species-Account/bna/home (accessed 30 January 2020)
Parmalee, P.W. & Klippel, W.E. (1982) Evidence of a boreal avifauna in middle Tennessee during the late Pleistocene. The Auk,
99 (2), 365–368.
R Core Team. (2019) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna.
URL https://www.R-project.org/ (accessed 16 March 2020)
Robbins, C.S. & Blom, E.A.T. (1996) Atlas of the breeding birds of Maryland and the District of Columbia. University of Pitts-
burgh Press, Pittsburgh, 479 pp.
RStudio Team. (2018) RStudio: Integrated Development for R. RStudio, Inc., Boston, MA. Available from http://www.rstudio.
com/ (accessed 1 November 2019).
Rusch, D.H., Destefano, S., Reynolds, M.C. & Lauten, D. (2000). Ruffed Grouse (Bonasa umbellus). Version 2.0. Available
from https://birdsna.org/Species-Account/bna/home (accessed 29 January 2020)
Snyder, N. & Snyder, H. (2000) The California Condor: A saga of natural history and conservation. Academic Press, London,
xxi + 410 pp.
Steadman, D.W. (1980) A review of the osteology and paleontology of turkeys (Aves: Meleagrinidae). Contributions in Science
of the Natural History Museum of Los Angeles County, 330, 131–207.
Steadman, D.W. (2005) Late Pleistocene birds from Kingston Saltpeter Cave, southern Appalachian Mountains, Georgia. Bul-
letin of the Florida Museum of Natural History, 45 (4), 231–248.
Strickland, D. & Ouellet, H.R. (2018) Canada Jay (Perisoreus canadensis). Version 2.1. In: Rodewald, P.G. (Ed.), The Birds
of North America. Cornell Lab of Ornithology, Ithaca, New York. Available from https://birdsna.org/Species-Account/bna/
home (accessed 15 November 2020)
Wetmore, A. (1927) A record of the Ruffed Grouse from the Pleistocene of Maryland. The Auk, 44, 561.
130 · Zootaxa 4772 (1) © 2020 Magnolia Press
Wetmore, A. (1962) Notes on fossil and subfossil birds. Smithsonian Miscellaneous Collections, 145 (2), 1–17.
Wickham, H. (2016) ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag, New York. ISBN 978-3-319-24277-4.
Available from: https://ggplot2.tidyverse.org (accessed 9 March 2020)
Withnell, C.B., Joannes-Boyau, R. & Bell, C.J. (2020) A Reassessment of the Age of the Fauna from Cumberland Cave, Mary-
land (Early-Middle Pleistocene) using coupled U-series and Electron Spin Resonance Dating (ESR). Quaternary Research.
APPENDIX 1. Measurements (mm) of North American jays (Corvidae) in comparison with the Cumberland Bone Cave
fossils. 1) Carpometacarpus, dorsal aspect, length of minor metacarpal from point just distal to facies articularis ulnocar-
palis to distal end. 2) Tibiotarsus, cranial aspect, greatest width across condyles. 3) Tibiotarsus, cranial aspect, width of
shaft across pons supratendineus. 4) Tarsometatarsus length from intercotylar area to trochlea metatarsi III. 5) Tarsometa-
tarsus, proximal width across articular facets. 6) Tarsometatarsus, proximal shaft width at level of foramina vascularia
proximalia. 7) Tarsometatarsus, distal shaft width at level of foramen vasculare distale. 8) Tarsometatarsus, width across
Species Catalog Number Sex 1 2 3 4 5 6 7 8
Fossils CM 24274 36.9 4.8 2.8 2.5
USNM PAL 641989 38.3 5.3 2.7 2.6 3.4
USNM PAL 641990 2.7 3.5
USNM PAL 641991 4.8 2.7
USNM PAL 641994 15.5
Aphelocoma californica USNM BIRDS 611085 f 16 5 3.1 42.2 5.4 3.2 2.6 3.8
USNM BIRDS 556692 f 13.4 4.1 2.8 36.2 4.4 2.9 2.3 3.2
USNM BIRDS 641219 f 14.9 4.6 2.9 39.1 5 3.2 2.8 3.2
USNM BIRDS 489825 m 14.8 4.6 3 42.8 4.9 3.3 2.3 3.3
USNM BIRDS 556687 m 15.7 4.9 2.9 41 5 2.9 2.4 3.4
USNM BIRDS 556689 m 14.5 4.7 3.1 40.4 4.9 3.2 2.7 3.5
USNM BIRDS 556691 m 14.5 4.6 2.9 40.2 4.8 3.2 2.2 3.3
Aphelocoma coerulescens USNM BIRDS 560527 f 13.9 4.4 3.1 38 4.7 3.4 2.5 3.3
USNM BIRDS 611083 m 14.3 4.7 2.8 39.1 5 2.9 2.8 3.6
USNM BIRDS 489957 m 13.7 4.6 3 37 5 3.1 2.6 3.6
USNM BIRDS 611082 m 13.6 4.7 3.2 36 4.8 2.9 2.7 3.5
USNM BIRDS 611081 m 14.1 4.4 2.7 37.4 4.8 2.8 2.7 3.3
Aphelocoma woodhouseii USNM BIRDS 555357 f 13.7 4.1 2.7 38 4.6 3 2.2 3
USNM BIRDS 346480 f 4.4 2.4 39.5 4.8 3.1 2.6 3.2
USNM BIRDS 611086 f 14.9 4.4 2.6 40 4.7 2.6 2.3 3.2
USNM BIRDS 614433 m 14.2 4.6 3 39.7 4.8 3.1 2.7 3.3
USNM BIRDS 611084 m 15.4 4.8 3 41.7 5.1 3 2.5 3.3
Cyanocitta cristata USNM BIRDS 614422 f 16.3 4.9 2.9 41 5.1 3 2.8 3.5
USNM BIRDS 556922 f 15.4 4.4 3.1 33.8 4.6 2.7 2.6 3.3
USNM BIRDS 15486 f 15.9 4.7 2.9 35.1 4.8 2.8 2.4 3.2
USNM BIRDS 553644 f 15.8 4.5 3 33.9 4.7 2.7 2.2 3.5
USNM BIRDS 554885 f 15 4.3 2.9 32.9 4.5 2.7 2.4 3.2
USNM BIRDS 614425 f 15.8 4.5 2.8 34.7 4.7 2.9 2.6 3.5
USNM BIRDS 499199 m 15.8 4.6 2.8 34.9 4.9 2.9 2.4 3.4
USNM BIRDS 244063 m 16.3 4.5 3 35.9 4.7 2.9 2.4 3.2
USNM BIRDS 612793 m 15.9 4.4 3 35.3 4.8 3 2.9 3.5
......continued on the next page
THE AVIFAUNA OF CUMBERLAND BONE CAVE, MARYLAND Zootaxa 4772 (1) © 2020 Magnolia Press · 131
APPENDIX 1. (Continued)
Species Catalog Number Sex 1 2 3 4 5 6 7 8
USNM BIRDS 614429 m 15.6 4.9 3.1 34.9 5 2.9 2.9 3.7
USNM BIRDS 501365 m 16.3 4.6 2.7 35.1 4.9 2.7 2.4 3.2
USNM BIRDS 554017 m 16.1 4.7 2.8 36.8 4.9 2.7 2.5 3.5
Cyanocitta stelleri USNM BIRDS 557597 ? 17.5 5.1 3.1 43.9 5.5 3.2 2.8 3.9
USNM BIRDS 639067 f 17.5 5.2 3.4 42.9 5.7 3 3 3.9
USNM BIRDS 614423 f 16.9 4.9 3.2 43.2 5.4 3.3 2.7 3.7
USNM BIRDS 621170 f 18 5.6 3.5 46.2 5.9 3.4 3.1 4.2
USNM BIRDS 554249 f 16.1 5 3 42.5 5.2 3.3 2.7 3.5
USNM BIRDS 614412 f 17 5.1 3.1 42.7 5.3 3.4 2.5 3.5
USNM BIRDS 637677 m 17.3 5 3.3 43.7 5.7 3.3 2.9 3.7
USNM BIRDS 614420 m 16.8 4.7 2.8 42.1 5.1 3 2.8 3.5
USNM BIRDS 634952 m 15.8 5.3 3.4 42 5.6 3.1 3 3.6
USNM BIRDS 554248 m 16.9 5.4 3.5 44.6 5.7 3.3 2.7 3.4
USNM BIRDS 611073 m 17 5.2 3.3 41.8 5.6 2.9 2.8 3.7
Perisoreus canadensis USNM BIRDS 622673 f 15.4 4.4 2.3 34.3 4.7 2.3 2.6 3
USNM BIRDS 638977 f 15.3 4.1 2.3 34.3 4.4 2.5 2.5 3.1
USNM BIRDS 611071 f 15.9 3.9 2.5 34.5 4.2 2.4 2.3 2.9
USNM BIRDS 489636 f 15.2 4.1 2.6 35.6 4.5 2.4 2.6 3.1
USNM BIRDS 622714 f 15.9 4.2 2.6 36.5 4.4 2.5 2.3 3.1
USNM BIRDS 622715 f 15 4.1 2.6 34.5 4.5 2.6 2.4 3.2
USNM BIRDS 489629 f 15.5 4 2.4 35.1 4.5 2.4 2.4 3
USNM BIRDS 489846 m 16.4 4 2.2 36.4 4.4 2.2 2.3 3
USNM BIRDS 613826 m 16.3 4.1 2.4 36.3 4.5 2.4 2.5 3.2
USNM BIRDS 639078 m 16 4.5 2.7 36.7 4.9 2.7 2.5 3.3
USNM BIRDS 639078 m 15.9 4.4 2.7 36.6 4.8 2.7 2.4 3.3
USNM BIRDS 489632 m 16.4 4.7 2.8 37.3 5 2.6 2.6 3.4