Smith-supplement1
Supplementary data associated with: Evolution of body mass in the Pan-Alcidae (Aves,
2
Charadriiformes): the effects of combining neontological and paleontological data
3
N. Adam Smith1, 2, 3*
1
National Evolutionary Synthesis Center 2024 W. Main St., Suite A200, Durham, NC, USA;
2
3
4
North Carolina Museum of Natural Sciences, 11 W. Jones St., Raleigh, NC, USA;
The Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL, USA
*correspondence: adam_smith@utexas.edu
1
Smith-supplement5
Supplementary Appendix 1
6
Measurements of Sampled Taxa
7
2
Data Sources.—Body mass data for extant alcids are from Dunning (2008), with the
8
exception of C. carbo, which is from Gaston and Jones (1998). Body length data for extant alcids
9
are from del Hoyo et al. (1996). Measurement data for extant alcid eggs were assembled from the
10
following sources: (Baicich and Harrison 1997, Davie 1900, De Santo and Nelson 1995, del
11
Hoyo et al. 1996, Friesen et al. 1996, Gaston and Jones 1998, Hipfner et al. 2010, Hunter et al.
12
2002, Kaler et al. 2009, Konyukhov and Kitaysky 1995, Livezey 1988, Nelson and Hamer 1995).
13
Measurements of specimens representing extinct taxa were taken directly from holotype and
14
referred specimens. Skeletal measurements follow von den Driesch (1976), were taken using
15
digital calipers and were rounded to the nearest 0.1 millimeter. All masses are stated in grams (g)
16
and all length, breadth and width measurements are stated in millimeters (mm). With respect to
17
skeletal specimens representing extant species, a total of five or more adult specimens of each
18
species, assessed based upon degree of ossification (Chapman 1965), and including both sexes,
19
were evaluated whenever available. Skeletal specimens from multiple locations within the
20
geographic range of extant species (i.e., subspecies) were examined to account for potential
21
geographic variation within species. Sexual dimorphism in extant Alcidae is not significant with
22
respect to many life history traits, including plumage and body mass in some species (e.g.,
23
Common Murre Uria aalge; (Nettleship and Birkhead 1985, Storer 1952, Székely et al. 2000).
24
Measurement data for extant alcid eggs were assembled from the following sources: (Baicich
25
and Harrison 1997, Davie 1900, De Santo and Nelson 1995, del Hoyo et al. 1996, Friesen et al.
26
1996, Gaston and Jones 1998, Hipfner et al. 2010, Hunter et al. 2002, Kaler et al. 2009,
27
Konyukhov and Kitaysky 1995, Livezey 1988, Nelson and Hamer 1995). Additional oological
Smith-supplement-
3
28
data was collected from specimens housed in the collection of the Smithsonian National Museum
29
of Natural History (USNM). Body length data used for extant alcids are those compiled by del
30
Hoyo et al. (1996).
31
Associated skeletons of the Great Auk are not known (Fuller 1987, Livezey 1988).
32
Therefore, measurement data representing that species was collected from composite skeletons
33
and additional material from a series of disarticulated remains in the USNM collection that were
34
collected during the Lucas expedition to Funk Island (Lucas 1890). Despite numerous accounts
35
regarding the natural history of the Great Auk, only a single measurement of the mass of the
36
species is known. Feilden (1872) reported an individual weighing 9 Danish pounds, the
37
equivalent of ~4,500 g. The mass of the Great Auk has twice been independently estimated at
38
~5,000 g using a variety of bodily and oological dimensions (Coues 1868, Livezey 1988). An
39
intermediate value of 4,750 g is used herein. As with the lack of data on the body mass of the
40
Great Auk, measurements of fresh Great Auk eggs are not known. However, egg dimensions and
41
egg mass are strongly correlated across Aves (Anderson et al. 1979) and a previously generated
42
estimate of 327 g is used herein (Birkhead 1993). Additionally, newly collected measurements of
43
Great Auk body length and oological dimensions were included in an attempt to independently
44
estimate its body mass. The forelimbs (i.e., humerus, radius, ulna, and carpometacarpus) of
45
flightless pan-alcids (i.e., †Pinguinus and †Mancallinae) are relatively shorter than those of
46
volant pan-alcids (Howard 1971, Livezey 1988, Owen 1864, Raikow et al. 1988). Therefore,
47
non-forelimb skeletal variables (e.g., femoral dimensions) were evaluated to identify an
48
appropriate variable for estimating the body mass of flightless species. Measurements of the
49
mandible, vertebrae and pelvis were not evaluated because of the relative rarity of those elements
50
representing the flightless †Mancallinae lineage (Smith 2011b).
Smith-supplement51
4
Institutional Abbreviations.—ANSP— Academy of Natural Sciences of Philadelphia, PA,
52
USA; GCVP—Georgia College Vertebrate Paleontology Collection, Milledgeville, GA, USA;
53
LACM—Natural History Museum of Los Angeles County, Los Angeles, CA., USA; NCSM—
54
North Carolina Museum of Natural Sciences, Raleigh, North Carolina, USA; SDSNH—San
55
Diego Natural History Museum, San Diego, CA, USA; UMMZ—University of Michigan
56
Museum of Zoology, Ann Arbor, MI, USA; UF/PB—Florida Museum of Natural History/Pierce
57
Brodkorb Collection, Gainesville, FL, USA; USNM—National Museum of Natural History,
58
Smithsonian Institution, Washington, D.C., USA.
59
60
Anatomical Abbreviations.—BM, body mass; BL, body length; EM, egg mass; EL, egg
61
length; ED, egg diameter; gbS. greatest breadth of skull; ghS, greatest height of skull; mlSt,
62
maximum length of sternum; dlSt, dorsal length of sternum; lcSt, length of sternal carina; sbRF,
63
smallest breadth between costal rib facets (on sternum); glC, greatest length of coracoid; mlC,
64
medial length of coracoid; bbC, basal breadth of coracoid; bfC, breadth of facies articularis
65
basalis of coracoid; diSc, diagonal of scapula; glH, greatest length of humerus; bpH, breadth of
66
proximal humerus; dpH, depth of proximal humerus; swH, shaft width of Humerus (at
67
midpoint); bdH, breadth of distal humerus; ddH, depth of distal humerus; glR, greatest length of
68
radius; bpR, breadth of proximal radius; swR = greatest width of radial shaft at midpoint; bdR,
69
breadth of distal radius; glU, greatest length of ulna; bpU, breadth of proximal ulna; swU, width
70
of ulnar shaft; bdU, breadth of distal ulna; ddU, diagonal of distal ulna; glF, greatest length of
71
femur; mlF, medial length of femur; bpF, breadth of proximal femur; dpF, depth of proximal
72
femur; swF, width of femoral shaft; bdF, breadth of distal femur; ddF, depth of distal femur; glT,
73
greatest length of tibiotarsus; laT, axial length of tibiotarsus; dpT, diagonal of proximal
Smith-supplement-
5
74
tibiotarsus; swT, width of tibial shaft; bdT, breadth of distal tibiotarsus; ddT, depth of distal
75
tibiotarsus; glTm, greatest length of tarsometatarsus; bpTm, breadth of proximal tarsometatarsus;
76
swTm, width of tarsometatarsal shaft; bdTm, breadth of distal tarsometatarsus.
77
78
79
Measurement Data.—See supplementary file "Appendix_1.xlsx" for raw values for all
sampled taxa. Dyrad.org DOI: 10.5061/dryad.3k7v7.
Smith-supplement80
Supplementary Appendix 2
81
82
83
84
Avian Body Mass Ranges for 65 Avian Clades
6
Values for clades with ranges exceeding that of Pan-Alcidae are bolded.
Taxon
Alcidae (n = 34)
(extant alcids)
Alcidae (n = 35)
(including P. impennis)
Pan-Alcidae (n = 60)
(pan-alcids)
Stercorariidae (n = 16)
(skuas)
Laridae (n = 99)
(gulls)
Sternidae (n = 60)
(terns)
Turnicidae (n = 24)
(buttonquail)
Scolopacidae (n = 154)
(snipes and sandpipers)
Jacanidae (n = 13)
(jacanas)
Haematopididae
(oystercatchers)(n = 15)
Recurvirostridae
(stilts) (n = 13)
Burhinidae (n = 10)
(thick-knees)
Glareolidae (n = 16)
(coursers & pratincoles)
Charadriidae (n = 77)
(plovers & lapwings)
Spheniscidae (n = 40)
(penguins)
Gaviidae (n = 7)
(loons)
Podicipedidae (n = 45)
(grebes)
Diomedeidae (n = 40)
(albatross)
Procellariidae (n = 98)
(petrels & shearwaters)
Hydrobatidae (n = 30)
(storm petrels)
Minimum
(g)
84
(Aethia pusilla)
84
(Aethia pusilla)
84
(Aethia pusilla)
270
(Stercorarius longicaudus)
118
(Larus minutus)
42
(Sternula saundersi)
18
(Ortyxelos meiffrenii)
21
(Caladris minuta)
41
(Microparra capensis)
517
(Haematopus finschi)
161
(Himantopus himantopus)
320
(Burhinus vermiculatus)
37
(Glareola cinerea)
12
(Charadrius thoracicus)
842
(Eudyptula minor)
1486
(Gavia stellata)
116
(Tachybaptus dominicus)
2,060
(Thalassarche chlororhynchos)
99
(Bulweria bulwerii)
17
(Oceanites gracilis)
Maximum
(g)
992
(Uria aalge)
4750
(Pinguinus impennis)
5,363
(Miomancalla howardae)
1,935
(Stercorarius antarcticus)
1,855
(Larus hyperboreus)
655
(Hydroprogne caspia)
110
(Turnix ocellatus)
869
(Numenius arquata)
261
(Actophilornis albinucha)
819
(Haematopus fuliginosus)
361
(Recurvirostra andina)
1032
(Esacus magnirostris)
150
(Rhinoptilus chalcopterus)
387
(Vanellus miles)
38,200
(Aptenodytes forsteri)
5,460
(Gavia immer)
1,646
(Podiceps major)
10,300
(Diomedea epomorpha)
4,940
(Macronectes giganteus)
86
(Oceanodroma tristrami)
Range
(g)
908
4,666
5,279
1,665
1,737
613
92
848
220
302
200
712
113
375
37,358
3,974
1,530
8,240
4,841
69
Smith-supplementPelecanoididae (n = 6)
(diving petrels)
Pelecanidae (n = 14)
(pelicans)
Sulidae (n = 30)
(gannets & boobies)
Phalacrocoracidae
(n = 79)(cormorants)
Anhingidae (n = 6)
(darters)
Ardeidae (n = 78)
(herons)
Ciconiidae (n = 21)
(storks)
Threskiornithidae
(ibis) (n = 32)
Anatidae (n = 320)
(ducks & geese)
Cathartidae (n = 12)
(vultures)
Accipitridae (n = 348)
(hawks & eagles)
Falconidae (n = 116)
(falcons)
Megapodiidae (n = 29)
(mound-builders)
Cracidae (n = 58)
(curassows & guans)
Pteroclidae (n = 12)
(sandgrouse)
Tetraonidae (n = 43)
(grouse)
Odontophoridae
(quail) (n = 47)
Phasianidae (n = 180)
(pheasants)
Gruidae (n = 27)
(cranes)
Rallidae (n = 173)
(rails & crakes)
Otididae (n = 30)
(bustards)
Apodidae (n = 105)
(swifts)
Trochilidae (n = 372)
(hummingbirds)
Trogonidae (n = 38)
(trogons)
Alcedinidae (n = 94)
(kingfishers)
121
(Pelecanoides georgicus)
3,174
(Pelecanus occidentalis)
857
(Sula sula)
427
(Phalacrocorax niger)
1235
(Anhinga anhinga)
80
(Ixobrychus involucris)
1,081
(Anastomus lamelligerus)
511
(Plegadis ridgwayi)
266
(Nettapus auritus)
935
(Cathartes burrovianus)
74
(Accipiter superciliosus)
41
(Microhierax latifrons)
308
(Megapodius laperouse)
439
(Ortalis leucogastra)
160
(Pterocles quadricinctus)
257
(Bonasa sewerzowi)
115
(Colinus leucopogon)
45
(Coturnix adansonii)
2,417
(Anthropoides virgo)
24
(Porzana flaviventer)
585
(Eupodotis senegalensis)
5
(Collocalia esculenta)
2
(Thaumastura cora)
42
(Trogon violaceus)
10
(Ispidina lecontei)
239
(Pelecanoides garnotii)
11,450
(Pelecanus onocrotalus)
3,067
(Morus capensis)
3,500
(Phalacrocorax harrisi
1,700
(Anhinga melanogaster)
4,468
(Ardea goliath)
6,892
(Jabiru mycteria)
3,515
(Pseudibis gigantea)
11,900
(Cygnus buccinator)
12,500
(Vultur gryphus)
8177
(Gyps coprotheres)
1,752
(Falco rusticolus)
2,520
(Alectura lathami)
4133
(Crax rubra)
428
(Pterocles orientalis)
4,100
(Tetrao urogallus)
457
(Odontophorus capueira)
4,766
(Pavo cristatus)
8,786
(Grus japonensis)
2,700
(Fulica gigantea)
11,975
(Otis tarda)
180
(Hirundapus celebensis)
20
(Patagona gigas)
206
(Pharomachrus mocinno)
356
(Dacelo novaeguineae)
7
118
8,276
2,210
3,073
465
4,388
5,811
3,004
11,634
11,565
8,103
1,711
2,212
3,694
268
3,843
342
4,721
6,369
2,676
11,390
175
18
164
346
Smith-supplementMeropidae (n = 21)
(bee eaters)
Coraciidae (n = 12)
(rollers)
Bucerotidae (n = 83)
(hornbills)
Galbulidae (n = 12)
(jacamars)
Bucconidae (n = 29)
(puffbirds)
Capitonidae (n = 94)
(barbets)
Ramphastidae (n = 68)
(toucans & aracaris)
Indicatoridae (n = 24)
(honeyguides)
Picidae (n = 247)
(woodpeckers)
Columbidae (n = 296)
(doves & pigeons)
Cacatuidae (n = 29)
(cockatoos)
Psittacidae (n = 321)
(parrots)
Musophagidae (n = 23)
(turacos)
Cuculidae (n = 189)
(cuckoos & allies)
Tytonidae (n = 15)
(barn owls)
Strigidae (n = 249)
(true owls)
Caprimulgidae (n = 84)
(nightjars & allies)
Passeriformes (n = 6,593)
(oscines & suboscines)
Cinclidae (n = 5)
(dippers)
Tinamidae (n = 59)
(tinamous)
15
(Merops orientalis)
96
(Eurystomas gularis)
97
(Tockus camurus)
16
(Brachygalba lugubris)
16
(Nonnula frontalis)
9
(Pogoniulus simplex)
121
(Pteroglossus inscriptus)
10
(Prodotiscus zambesiae)
9
(Picumnus aurifrons)
29
(Columbina passerina)
92
(Nymphicus hollandicus)
12
(Loriculus tener)
198
(Corythaixoides leucogaster)
18
(Chrysococcyx meyeri)
195
(Phodilus prigoginei)
41
(Micrathene whitneyi)
25
(Nyctiphrynus yucatanicus)
4
(Phylloscopus sichuanensis)
37
(Cinclus shulzi)
43
(Taoniscus nanus)
60
(Merops nubicoides)
171
(Coracias temminckii)
4,191
(Bucorvus leadbeateri)
63
(Jacamerops aureus)
106
(Monasa morphoeus)
202
(Megalaima virens)
709
(Ramphastos swainsonii)
54
(Melichneutes robustus)
516
(Campephilus principalis)
2,384
(Goura victoria)
841
(Probosciger aterrimus)
2000
(Strigops habroptila)
965
(Corythaeola cristata)
769
(Centropus milo)
887
(Tyto multipunctata)
2,992
(Bubo bubo)
174
(Eurostopodus mystacalis)
1,135
(Corvus crassirostris)
88
(Cinclus pallasii)
1,636
(Tinamus solitarius)
8
45
75
4,094
47
90
193
588
44
507
2,355
749
1,988
767
751
692
2,951
149
1,132
51
1,593
85
86
Supplementary Appendix 2 Footnote: Number of taxa sampled in each clade (excluding Pan-
87
Alcidae which includes extinct taxa) reflects the number of data points provided by Dunning
88
(2008; i.e., includes males, females and sub-species). Taxonomy follows Dunning (2008) and all
89
values are means from Dunning (2008) rounded to nearest 1.0 gram. Despite small sample sizes,
Smith-supplement-
9
90
Gaviidae (n = 7), Pelecanoididae (n = 6), Pelecanidae (n = 14) and Anhingidae (n = 6) were
91
included to sample as many diving taxa as possible. Despite small sample sizes, clades such as
92
Burhinidae (n = 10) and Jacanidae (n = 13) were sampled to include as broad a range of
93
charadriiforms as possible. Based on previous phylogenetic results (Baker et al., 2007),
94
Ptercolidae is not considered a part of Charadriiformes herein. The following taxa were not
95
included owing to small sample sizes (i.e., relatively few extant species in comparison with
96
Alcidae): Struthionidae; Rheidae; Casuariidae; Dromaiidae; Apterygidae; Phaethontidae;
97
Fregatidae; Scopidae; Balaenicipididae, Phoenicopteridae, Anhimidae; Pandionidae;
98
Sagittariidae; Meleagrididae; Numididae; Opisthocomidae; Mesitornithidae, Aramidae;
99
Psophiidae; Heliornithidae; Rhynochetidae; Eurypygidae; Cariamidae; Hemiprocnidae; Coliidae;
100
Todidae; Momotidae; Brachypteraciidae; Leptosomatidae; Upupidae; Phoeniculidae;
101
Rostratulidae; Dromadidae; Ibiorhynchidae; Pluvianellidae; Pedionomidae; Thincoridae;
102
Chionidae; Rynchopidae; Steatornithidae; Podargidae; Nyctibiidae.
Smith-supplement103
104
Supplementary Figure 1. Bar graph of avian body mass ranges.
10
Smith-supplement105
11
Supplementary Appendix 3
106
107
108
Supplementary Body Mass Estimation Methods and Results
Additional Details of Body Mass Estimation for Flightless Pan-Alcidae.—As stated above,
109
forelimb measurements of the Great Auk were excluded from initial PGLS analyses because of
110
the markedly different allometric relationship between those skeletal elements and body mass in
111
flightless pan-alcids (i.e., flightless pan-alcids have relatively small wings). Attempts to
112
independently estimate the body mass of the Great Auk were not successful. No individual, non-
113
forelimb variable met all three criteria (see methods) evaluated to assess the predictive power of
114
variables in relation to body mass. This included explorations of non-skeletal variables (e.g.,
115
body length, egg volume). Therefore, as stated above, a conservative body mass estimate of
116
4,750 g was used for the Great Auk. The body mass estimate for the Pliocene sister taxon of the
117
Great Auk, †Pinguinus alfrednewtoni (Olson 1977), was generated by assessing the ratio of body
118
mass to humeral length in the Great Auk (4,750 g: 104.1 mm = 45.63 g/mm) and extrapolating to
119
produce an estimate for †P. alfrednewtoni (101.0 mm x 45.63 g/mm = 4,609 g; Table 2). This
120
estimate is consistent with the overall smaller size of †P. alfrednewtoni relative to †P. impennis
121
(based on greatest length of coracoids, humeri and femora; see Appendix 1). If the body mass of
122
†P. impennis is assumed to be 5,000 g (sensu Bédard 1969, Birkhead 1993, Coues 1868, Livezey
123
1988), then the estimate for †P. alfrednewtoni increases to 4851 g, a difference of only 242 g
124
(i.e., < 5.3 %).
125
Estimation of body mass for species of the flightless †Mancallinae clade (Lucas Auks)
126
was complicated owing to the lack of complete skeletons representing the 6 included species.
127
Moreover, as in †Pinguinus, the forelimbs of Lucas auks are relatively small in comparison to
Smith-supplement-
12
128
their overall body mass, putatively based on the relative dimensions of other skeletal elements
129
(Howard 1970). The best non-forelimb predictor of body mass among extant alcids (including
130
the Great Auk) was the greatest length of the femur (glF; r2 = 0.95, AICc = -62.1, λ = 0.9; Fig.
131
1B; Table 1). Femur length was also identified as a reliable predictor of body mass in a larger
132
sample of charadriiforms [(Field et al. 2013), figure 6, r2 = 0.935)]. Although femoral length is
133
potentially linked with phylogeny (λ = 0.9) and the percent predicted error associated with
134
estimates based on this regression is relatively high in some instances (PPE: range = 1.1-66.9%;
135
average = 25.3%; median = 22.9%), more precise estimates will only be possible as additional
136
fossil material is referred to the species level in †Mancallinae (i.e., specimens that would
137
facilitate a multi-step regression). Comparisons with previous estimates for these taxa are not
138
feasible given the methodological issues with the only previous analysis of body mass of
139
mancalline auks (Livezey, 1988; discussed below). Estimates for †Miomancalla howardae,
140
†Mancalla cedrosensis and †Mancalla lucasi were generated using femoral measurements from
141
the holotypes of those species. However, femora representing †Miomancalla wetmorei,
142
†Mancalla vegrandis and †Mancalla californiensis are not known (Smith 2011b). Therefore, to
143
derive a body mass estimate for those †Mancallinae species lacking femora, the ratio of humerus
144
length to femoral length in †M. howardae (0.774), †M. cedrosensis (0.779), and †M. lucasi
145
(0.752) was calculated and then the average ratio (glH:glF = 0.74) was used to estimate femoral
146
lengths for †M. wetmorei, †M. vegrandis and †M. californiensis (Table 2). Body mass estimates
147
for †M. wetmorei, †M. vegrandis and †M. californiensis were then generated using the same
148
regression equation based on the relationship between greatest length of the femur and body
149
mass in extant alcids (Table 2).
150
Smith-supplement151
13
Comparisons with Previous Studies.—Previous body mass estimates for extinct pan-
152
alcids are restricted to a single volant species, the large Pliocene auk †Alca stewarti and some
153
flightless species including the Great Auk and selected members of the Alcidae stem lineage
154
Mancallinae (Lucas Auks; Livezey 1988, Martin et al. 2001). Body length was not strongly
155
correlated with body mass regardless of whether the Great Auk was included (Table 1) or
156
excluded (r2 = 0.86, AICc = -29.9, λ = 0.0). In contrast, body length contributed significantly to
157
the multi-step regression used by Livezey (1988) to estimate a 4,999 g body mass for the Great
158
Auk. However, Livezey (1988) also included measures of culmen length, which varies with diet
159
(relatively shorter beaks in planktivorous species versus longer beaks in piscivorous species) and
160
tail length. Tail length variability in Alcidae has not been studied in detail. However, differences
161
in locomotor style (both underwater and aerial) and incubation posture (i.e., upright as in
162
Cepphus versus pronated as in Alca; see Birkhead 1993, figure 9) are likely factors influencing
163
variability in alcid tail length. Body length may prove to be a better estimator in avian clades
164
other than Alcidae, clades with more homogeneous ecologies and ethologies.
165
Flightless pan-alcids and penguins possess pachyostotic limb bones that are relatively
166
greater in mass than those of comparably sized volant pan-alcids (Ksepka 2007, Smith and
167
Clarke 2014). However, previous comparison of body mass with skeletal mass across a broad
168
sample of Aves did not identify penguins as an outlier and skeletal mass is a minor contributor to
169
overall mass in birds (Prange et al. 1979). Therefore, while it remains possible that body mass
170
estimates based on regressions generated using data from extant, volant alcids with non-
171
pachyostotic limb bones may underestimate the mass of flightless pan-alcids, it seems unlikely
172
that pachyostotic limb bones would introduce significant bias into these estimates.
Smith-supplement-
14
173
It is not surprising that egg mass, egg length and egg diameter were not strongly
174
correlated with body mass in Alcidae as Charadriiformes in general, and alcids in particular, are
175
characterized by a wider range of developmental strategies than most other avian clades (i.e.,
176
semi-precocial, intermediate, precoccial (De Santo and Nelson 1995). Developmental strategy
177
has been correlated with a variety of life history traits including egg mass and associated egg
178
dimensions (Ricklefs and Starck 1998). Exclusion of potentially inaccurate estimates for the
179
body mass and egg mass of the Great Auk did not result in stronger support for these non-
180
skeletal variables as good predictors of body mass in Alcidae (results not shown).
181
Body mass estimates for extinct pan-alcids were previously restricted to †A. stewarti, †P.
182
impennis and some species of †Mancallinae (Livezey, 1988; Martin et al., 2001). The previously
183
generated ~1,900 g estimate for the large volant auk A. stewarti (Dyke and Walker, 2001) is
184
relatively consistent with the ~2,100 g estimate generated herein (Table 2). However, further
185
comparisons with previous studies that estimated the body mass of pan-alcids are limited
186
because those previous studies (Livezey, 1988; Martin et al., 2001) did not report the
187
measurement data on which their regressions and resulting estimates were based, did not provide
188
the catalog numbers of specimens from which measurements were taken, and did not consider
189
the potentially biasing effect of phylogeny (Harvey and Pagel, 1991). Moreover, both previous
190
studies that estimated extinct pan-alcid body masses (Livezey, 1988; Martin et al., 2001) relied
191
on measurement data from fossil specimens that were previously attributed to species on the
192
basis of size or provenance, and that may not have been representative of the taxa they were
193
referred to. For example, both previous studies reportedly considered measurement data for
194
humeri and ulnae of †Mancalla milleri. Martin et al. (2001) utilized measurements reported in
195
the original description of M. milleri (Howard, 1970). In contrast, no sources of data or specimen
Smith-supplement-
15
196
numbers were reported by Livezey (1988), and it is unclear from what specimens data were
197
collected. Furthermore, the holotype specimen of †M. milleri is a femur (LACM 2185) and
198
associated specimens that would allow for referral of elements other than femora to that taxon
199
are not known (Smith 2011b). Moreover, the holotype specimen of †M. milleri is now
200
considered Pan-Alcidae incertae sedis and some material in the size range of †M. milleri is now
201
referred to †Mancalla vegrandis (Smith, 2011b). Data reported for †Mancalla cedrosensis,
202
†Mancalla emlongi and †Mancalla diegensis by Livezey (1988) also lack documentation of
203
specimen numbers or citation of sources and include elements not comparable to the holotypes of
204
those taxa. Finally, it is unclear what specimens Livezey (1988) included in '†Praemancalla spp.'.
205
The lack of documentation, repeatability and phylogenetic context, in combination with the
206
inclusion of specimens of uncertain taxonomic referral call into question the reliability of those
207
previous estimates.
208
Smith-supplement-
16
209
Supplementary Table A3.1: Mean and 95% confidence interval of maximum likelihood
210
estimates of ancestral body mass for nodes in extant Alcidae (lettered nodes correspond to those
211
in Figure 3).
Node
A
B
C
D
E
F
G
H
I
J
K
L
M
N
O
P
Q
R
S
T
U
V
212
213
Estimate
415.31
411.94
222.71
194.53
200.57
208.06
550.86
627.11
607.58
421.17
434.66
442.64
538.70
556.71
868.81
236.85
218.88
167.05
462.08
549.25
325.79
269.74
Lower CI
208.03
204.03
56.90
59.98
73.42
80.33
371.09
482.96
497.38
243.12
261.14
267.70
356.42
370.51
710.78
62.12
76.96
87.13
298.44
429.42
122.45
91.70
Upper CI
622.60
619.84
388.53
329.07
327.71
335.79
730.64
771.25
717.78
599.22
608.18
617.57
720.98
742.92
1026.84
411.58
360.80
246.98
625.73
669.07
529.12
447.79
Smith-supplement-
17
214
Supplementary Table A3.2: Mean and 95% confidence interval of maximum likelihood
215
estimates of ancestral body mass for nodes in Pan-Alcidae (lettered nodes correspond to those in
216
Figure 2).
Node
A
B
C
D
E
F
G
H
I
J
K
L
M
N
O
P
Q
R
S
T
U
V
W
X
Y
Z
a
b
c
d
e
f
g
h
i
j
k
l
m
Estimate
1236.50
1041.71
966.58
994.78
1106.61
1382.39
1809.68
1678.35
1672.33
1613.24
1515.77
1374.35
850.95
4399.68
817.67
925.18
960.39
558.66
287.65
570.62
346.48
240.01
180.17
236.74
607.08
544.68
541.42
544.14
393.53
331.87
282.94
259.65
486.91
359.49
280.42
227.21
224.82
198.78
501.74
LowerCI
1136.67
955.44
886.93
919.63
1034.54
1313.38
1743.06
1617.313
1610.374
1551.747
1457.527
1322.036
808.6849
4354.418
759.4156
872.8599
918.1272
508.3691
245.6618
528.6617
283.5357
186.57
137.75
192.78
536.85
483.76
488.47
501.79
322.35
270.65
229.92
217.29
414.87
291.41
216.53
168.26
172.34
156.49
438.20
UpperCI
1336.33
1127.97
1046.22
1069.94
1178.68
1451.40
1876.31
1739.39
1734.29
1674.74
1574.01
1426.67
893.21
4444.96
875.93
977.50
1002.66
608.95
329.64
612.64
409.43
293.45
222.59
280.70
677.31
605.60
594.37
586.49
464.71
393.10
335.96
302.01
558.95
427.57
344.30
286.16
277.31
241.06
565.27
Smith-supplementn
o
p
q
r
s
t
u
v
w
217
218
554.35
619.26
602.59
416.12
474.37
2670.90
2430.07
1902.55
1826.85
3898.58
495.50
566.80
560.30
361.62
431.80
2599.44
2368.75
1849.51
1784.48
3853.81
613.19
671.71
644.87
470.63
516.94
2742.36
2491.38
1955.59
1869.22
3943.35
18
Smith-supplement219
19
Supplementary Literature Cited
220
221
222
223
224
Anderson, J. F., H. Rahn, and H. D. Prange. 1979. Scaling of Supportive Tissue Mass. Quarterly
Review of Biology 54:139-148.
Baicich, P. J., and C. J. O. Harrison. 1997. A Guide to the Nests, Eggs, and Nestlings of North
American Birds. Academic Press, San Diego, CA.
225
Baker, A. J., S. L. Pereira, and T. A. Paton. 2007. Phylogenetic relationships and divergence
226
times of Charadriiformes genera: multigene evidence for the Cretaceous origin of 14
227
clades of shorebirds. Biology Letters 3:205-209.
228
Bédard, J. 1969. Adaptive radiation in the Alcidae. Ibis 111:189-198.
229
Birkhead, T. R. 1993. Great Auk Islands : a field biologist in the Arctic. Poyser, London.
230
Chapman, W. L. 1965. Appearance of ossification centers and epiphyseal closures as determined
231
by radiographic techniques. Journal of the American Veterinary Medical Association
232
147:138-141.
233
234
Coues, E. 1868. A monograph of the Alcidae. Proceedings of the Academy of Natural Sciences
of Philadelphia 20:2-81.
235
Davie, O. 1900. Nests and Eggs of North American Birds. David McKay, Philadelphia.
236
De Santo, T. L., and S. K. Nelson. 1995. Comparative reproductive ecology of the auks(Family
237
Alcidae) with emphasis on the Marbled Murrelet. Pp. 33-47. USDA Forest Service
238
General Technical Report PSW-152.
239
240
241
del Hoyo, J., A. Elliott, and J. Sargatal. 1996. Handbook of the Birds of the World Vol. 3.
Hoatzin to Auks. Lynx Edicions, Barcelona.
Dunning, J. B. J. 2008. CRC Handbook of Avian Body Masses, 2nd Ed. CRC Press, Boca Raton.
Smith-supplement-
20
242
Feilden, H. W. 1872. The birds of the Faroe Islands. The Zoologist 7 (Ser. 2):3277-3294.
243
Field, D. J., C. Lynner, C. Brown, and S. A. F. Darroch. 2013. Skeletal correlates for body mass
244
estimation in modern and fossil flying birds. Plos One 8(11):E82000.
245
doi:10.1371/journal.pone.0082000.
246
Friesen, V., P. Piatt, and A. J. Baker. 1996. Evidence from cytochrome b sequences and
247
allozymes for a new species of alcid: the Long-Billed Murrelet (Brachyramphus perdix).
248
Condor 98:681-690.
249
Fuller, E. 1987. Extinct birds. Cornell University Press, Ithaca NY.
250
Gaston, A. J., and I. L. Jones. 1998. The auks :Alcidae. Oxford University Press, Oxford ; New
251
252
York.
Hipfner, J. M., K. B. Gorman, R. A. Vos, and J. B. Joy. 2010. Evolution of embryonic
253
developmental period in the marine bird families Alcidae and Spheniscidae: roles for
254
nutrition and predation? BMC Evolutionary Biology 10:179.
255
http://www.biomedcentral.com/1471-2148/10/179.
256
257
258
259
260
Howard, H. 1970. A review of the extinct avian genus, Mancalla. Los Angles County Museum
Contributions in Science 203:1-12.
———. Pliocene avian remains from Baja California. Los Angles County Museum
Contributions to Science 217:1-17.
Hunter, F. M., I. L. Jones, J. C. Williams, and G. V. Byrd. 2002. Breeding biology of the
261
Whiskered Auklet (Aethia pygmaea) at Buldir Island, Alaska. Auk 119:1036-1051.
262
Kaler, R. S. A., L. A. Kenney, and B. K. Sandercock. 2009. Breeding Ecology of Kittlitz's
263
Murrelets at Agattu Island, Aleutian Islands, Alaska. Waterbirds 32:363-373.
Smith-supplement264
265
266
267
21
Konyukhov, N. B., and A. S. Kitaysky. 1995. The asian race of the Marbled Murrelet. Pp. 23-29.
USDA Forest Service General Technical Report PSW-152.
Ksepka, D. T. 2007. Phylogeny, histology, and functional morphology of fossil penguins
(Sphenisciformes). Columbia University.
268
Livezey, B. 1988. Morphometrics of flightlessness in the Alcidae. Auk 105:681-698.
269
Lucas, F. A. 1890. The expedition to Funk Island, with observations upon the history and
270
anatomy of the Great Auk. Report of the United States National Museum for 1887-1888
271
1887-1888:493-529.
272
273
274
Martin, J. W. R., C. A. Walker, R. Bonser, and G. J. Dyke. 2001. A new species of large auk
from the Pliocene of Belgium. Oryctos 3:53-60.
Nelson, S. K., and T. E. Hamer. 1995. Nest Success and the Effects of Predation on Marbled
275
Murrelets. Pp. 89-97. USDA Forest Service General Technical Report PSW-152. 1995.
276
Nettleship, D. N., and T. Birkhead. 1985. The Atlantic Alcidae: the evolution, distribution, and
277
biology of the auks inhabiting the Atlantic Ocean and adjacent water areas. Academic
278
Press, London and Orlando.
279
Olson, S. L. 1977. A Great Auk, Pingunis [sic], from the Pliocene of North Carolina (Aves,
280
Alcidae). Proceedings of the Biological Society of Washington 90:690-697.
281
Owen, R. 1864. Description of the skeleton of the Great Auk, or Garfowl (Alca inpennis, L.).
282
Prange, H. D., J. F. Anderson, and H. Rahn. 1979. Scaling of Skeletal Mass to Body-Mass in
283
284
285
Birds and Mammals. American Naturalist 113:103-122.
Raikow, R. J., L. Bicanovosky, and A. H. Bledsoe. 1988. Forelimb joint mobility and the
evolution of wing-propelled diving birds. Auk 105:446-451.
Smith-supplement286
22
Ricklefs, R. E., and J. M. Starck. 1998. Embryonic growth and development. Pp. 31-58. In R. E.
287
Ricklefs, and J. M. Starck, eds. Avian Growth and Development. Oxford University Press,
288
New York.
289
290
291
Smith, N. A. 2011b. Taxonomic revision and phylogenetic analysis of the flightless Mancallinae
(Aves, Pan-Alcidae). ZooKeys 91:1-116.
Smith, N. A., and J. A. Clarke. 2014. Osteological histology of the Pan-Alcidae (Aves,
292
Charadriiformes): correlates of wing-propelled diving and flightlessness. The Anatomical
293
Record 297:188-199.
294
Storer, R. W. 1952. A comparison of variation, behavior and evolution in the sea bird genera
295
Uria and Cepphus. University of California Publications in Zoology 52:121-222.
296
Székely, T., J. D. Reynolds, and J. Figuerola. 2000. Sexual size dimorphism in shorebirds, gulls,
297
and alcids: the influence of sexual and natural selection. Evolution 54:1404-1413.