A new fossil species of Procellaria (Aves: Procellariiformes) from the Pliocene of New Zealand

Abstract

We describe a new Procellaria petrel species from the late Pliocene of Taranaki, New Zealand. The new species is most similar morphologically to the White-Chinned Petrel (P. aequinoctialis), Spectacled Petrel (P. conspicillata) and the Westland Petrel (P. westlandica). Compared with those taxa, the new species has a deeper and shorter premaxilla, longer coracoid and shorter wings, while its legs are a similar size. Today, New Zealand is the centre of global diversity of the genus, with four breeding species. This is the first fossil species of Procellaria to be described from New Zealand, attesting to a reasonably long history of this genus in the region.

Keywords.
Procellaria altirostris sp. nov.; Petrel; Piacenzian; Procellariidae; Taranaki

INTRODUCTION

New Zealand is a global centre of diversity for seabirds today, however this contrasts with its sparse pre-Pleistocene fossil record - while penguins are well represented, other seabirds are not (Gill et al., 2010) - so the history of seabirds in the region is poorly understood. For instance, Procellariiformes are presently the most diverse group of seabirds in New Zealand (Gill et al., 2010), but only three fossil pre-Pleistocene taxa are described: a Pliocene albatross (Mayr & Tennyson, 2020) and a shearwater (Tennyson & Mannering, 2018), both from Taranaki, and a Miocene diving petrel from St Bathans (Worthy et al., 2007).

The Pliocene outcrops in the Taranaki region continue to produce a rich and globally significant fauna of marine vertebrates, including cetaceans, seals, fishes and other birds, the latest being a new species of penguin (McKee, 1994; Thomas et al., 2020). Here we describe the latest discovery from Taranaki: a new procellariiform species.

MATERIAL AND METHODS

The new fossil was identified as a bird from the Procellaria genus and thus compared with all living members of this genus: White-Chinned Petrel (P. aequinoctialis Linnaeus, 1758), Spectacled Petrel (P. conspicillata (Gould, 1844)), Westland Petrel (P. westlandica Falla, 1946), Grey Petrel (P. cinerea (Gmelin, 1789)), and Black Petrel (P. parkinsoni G.R. Gray, 1862).

We measured 155 skeletons of Procellaria spp. deposited in the collections of the Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand (NMNZ) and the Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZUSP). In order to compare the measurements of the different species with the fossil, we only measured beaks from skulls where the ramphotecal plates were removed. Thus, the total sample size for individual measurements varies. For the principal component analysis (see below), we only used individuals that had all measurements, thus the PCA only includes a single P. conspicillata specimen. The primary comparative specimens used are listed in ^{Appendix 1}.

We took the following skeletal measurements (also see Fig. 1): skull minimum interorbital width; premaxilla maximum length from nasal hinge to tip, depth at the middle of the apertura nasi ossea, width at the middle of the apertura nasi ossea; coracoid maximum omal-sternal length; humerus maximum length, maximum proximal dorso-ventral width, maximum distal dorso-ventral width, mid-shaft dorso-ventral width; ulna maximum length; radius maximum length; femur maximum length; tibiotarsus maximum length including cnemial crest, maximum length excluding cnemial crest; tarsometatarsus maximum length. Osteological nomenclature follows Baumel & Witmer (1993). All measurements were taken using vernier digital calipers (precison 0.01 mm) or a ruler (precision 0.1 mm) when skeletal elements were larger than 150 mm.

Figure 1
Plate illustrating how each bone element was measured.

Analyses were carried out in R version 3.6.1 (R Core Team, 2017). Firstly, measurements were compared using descriptive statistics (mean, standard error and measurement range). Then, we carried out a principal component analysis (PCA) on the z-standardized measurements (mean centred at 0 and standard deviation at 1) that allowed us to detect whether the fossil would cluster together with one of the other species.

RESULTS

Systematic Palaeontology

Order Procellariiformes Fürbringer, 1888

Family Procellariidae Leach, 1820

Procellaria Linnaeus, 1758

Procellaria altirostris sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:093F56E5-D07E-44C3-8E75-059C486CB222

Etymology: From the unusually deep premaxilla in comparison to other members of the Procellaria genus.

Holotype: NMNZ S.46691; partial skeleton embedded, but largely exposed, in a concretion. Elements preserved: partial cranium (crushed) with premaxilla, furcula, both coracoids, right scapula, right humerus, both ulnae, right radius, synsacrum, left femur, both tibiotarsi, both tarsometatarsi but the left is missing both ends, one pedal phalange, and other fragments. The fossil is from a fully grown individual based on bone ossification (Fig. 2).

Figure 2
Pliocene fossil petrel Procellaria altirostris sp. nov., holotype NMNZ S.46691. Photo: Jean-Claude Stahl (Te Papa). (1) cranium; (2) premaxilla; (3) furcula; (4) right scapula; (5) left coracoid; (6) right coracoid; (7a) proximal right humerus; (7b) distal right humerus; (8) left ulna; (9a) proximal right ulna; (9b) distal right ulna; (10) right radius; (11) synsacrum; (12a) proximal left femur; (12b) distal left femur; (13a) proximal left tibiotarsus; (13b) distal left tibiotarsus; (14) right tibiotarsus; (15) left tarsometatarsus; (16) right tarsometatarsus; (17) pedal phalange.

Type Locality, Type Horizon and Age: Ohawe Beach, south Taranaki, North Island, New Zealand (New Zealand Fossil Record Electronic Database Number Q21/F0175, Fig. 3). The fossil was found inside a marine mudstone concretion loose on the beach in 2015, and prepared by John Buchanan-Brown in the same year. The concretion is presumed to have eroded from an adjacent cliff in the Tangahoe Formation. This Formation is 3.0-3.4 Ma (local Waipipian stage, late Pliocene, Piacenzian; McKee, 1994; Naish et al., 2005; Thomas et al., 2020). Such concretions are thought to have formed on the ocean floor at a depth of up to 50 m or more (Tennyson & Mannering, 2018).

Figure 3
General location in Taranaki where the fossil was found.

Generic attribution:Procellaria altirostris sp. nov. is referred to Procellaria because it is a large petrel falling in the size range of Procellaria species and outside the size range of every other recent genus in Procellariiformes: all recent Diomedeidae are larger and all Hydrobatidae and Pelecanoididae are smaller. Within the family Procellaridae, birds in the Procellaria genus are the largest, with the exception of Macronectes Richmond, 1905, which is similar in size to some albatross taxa. Based on the narrow and deep beak, the strongly hooked premaxilla and the prominent pila supranasalis, the genus Procellaria can be easily distinguished from almost all other genera within the Procellaridae; the exceptions being Pterodroma Bonaparte, 1856 and Pseudobulweria Mathews, 1936. Generally, Pterodroma taxa have a narrower and shorter beak than Procellaria taxa; moreover, as in all Procellaria, the fossil has a longer humerus than ulna, while Pterodroma taxa have longer ulnae than humeri (Tennyson & Mannering, 2018). The apertura nasi ossea of Pseudobulweria is smaller than in Procellaria taxa; moreover, the proximal end of the humerus of Pseudobulweria taxa lack the tubercle on the cranial surface of the ventral margin in the bicipital area (Tennyson & Mannering, 2018). The shape of the apertura nasi ossea and this humeral tubercle of the fossil are features found in Procellaria taxa (Tennyson & Mannering, 2018).

Differential diagnosis:Procellaria altirostris sp. nov. is identified as a member of the Procellariiformes by the beak having a sharply hooked tip, lateral furrows rostral of the nostrils (nasolabial grooves) and marked supraorbital fossae for nasal glands (fossa glandulae nasalis), i.e., salt glands.

Procellaria altirostris sp. nov. is most similar to the White-Chinned Petrel (P. aequinoctialis), Spectacled Petrel (P. conspicillata) and Westland Petrel (P. westlandica). Compared with these taxa, the new species has a deeper and shorter premaxilla, a larger coracoid and shorter wings, while its legs are a similar size. It is slightly larger than the Grey Petrel (P. cinerea) and notably larger than the Black Petrel (P. parkinsoni). The Grey Petrel also differs in having a more slender beak (width and depth relative to length) than all other members of the genus. Procellaria species show minor sexual dimorphism in size, with males averaging a little larger (Marchant & Higgins, 1990; Warham, 1990). However, regardless of the sex of the fossil, it still falls outside the range of variation seen in other species (Table 1, Fig. 2, Fig. 4, ^{Appendix 2}).

Table 1
Measurements (in mm) of skeletal elements of the holotype of P. altirostris sp. nov., compared with measurements of other Procellaria species. For measurement details, see Methods.

Measurement	NMNZ S.46691
skull
min. intraorbital width	15	14.24 ± 0.09 (50)	14.4 ± 0.12 (35)	13.57 ± 0.22 (7)	13.76 ± 0.10 (37)	11.4 ± 0.15 (16)
premaxilla
max. length	55.56	62.22 ± 0.34 (32)	59.62 ± 0.36 (29)	57.22 (1)	57.15 ± 0.26 (37)	49.48 ± 0.57 (15)
depth	14.52	11.96 ± 0.08 (34)	12.12 ± 0.11 (34)	11.95 (1)	9.95 ± 0.07 (38)	9.81 ± 0.13 (17)
width	13.5	13.76 ± 0.13 (34)	14.7 ± 0.17 (34)	14.23 (1)	12.42 ± 0.13 (38)	11.98 ± 0.21 (17)
coracoid
max. length	41.99	37.93 ± 0.20 (27)	39.12 ± 0.15 (33)	38.92 ± 0.86 (2)	36.15 ± 0.15 (35)	33.35 ± 0.28 (17)
humerus
max. length	143.22	150.71 ± 0.62 (29)	149.65 ± 0.48 (37)	151.22 ± 1.59 (4)	136.76 ± 0.44 (37)	130.06 ± 0.74 (17)
proximal width	32.26	32.34 ± 0.16 (29)	32.37 ± 0.15 (37)	31.77 ± 0.36 (4)	30.54 ± 0.15 (37)	28.19 ± 0.17 (17)
distal width	17.68	17.49 ± 0.07 (29)	17.12 ± 0.08 (36)	17.77 ± 0.22 (4)	16.4 ± 0.05 (37)	15.04 ± 0.10 (18)
shaft width	7.44	7.58 ± 0.05 (28)	7.44 ± 0.05 (37)	7.38 ± 0.07 (4)	7.41 ± 0.05 (37)	6.52 ± 0.06 (18)
ulna
max. length	138.53	151.13 ± 0.50 (29)	149.91 ± 0.50 (37)	149.84 ± 0.67 (4)	135.09 ± 0.47 (36)	131.29 ± 0.84 (18)
radius
max. length	132.95	147.57 ± 0.53 (29)	146.33 ± 0.52 (36)	146.08 ± 0.37 (4)	131.51 ± 0.45 (36)	128.19 ± 0.81 (18)
femur
max. length	50.05	50.64 ± 0.30 (28)	50.9 ± 0.22 (31)	50.52 ± 0.67 (4)	48.62 ± 0.24 (34)	42.28 ± 0.48 (16)
tibiotarsus
max. length incl. cnemial crest	116.56	114.01 ± 0.52 (27)	111.86 ± 0.40 (31)	114.48 ± 0.65 (4)	109.94 ± 0.35 (32)	94.89 ± 0.70 (14)
max. length excl. cnemial crest	99.53	99.5 ± 0.40 (30)	96.94 ± 0.33 (35)	99.43 ± 0.69 (4)	95.54 ± 0.33 (35)	82.36 ± 0.67 (14)
tarsometatarsus
max. length	64.82	65.99 ± 0.31 (30)	63.65 ± 0.26 (34)	65.33 ± 0.16 (4)	61.66 ± 0.22 (35)	54.53 ± 0.45 (15)

Figure 4
Principal component analysis for the skeletal measurements of the three Procellaria species that are most similar in morphology to the new species: P. aequinoctialis, P. conspicillata and P. westlandica. (A) Biplot of the first and second principal component scores. (B) Projection of the principal component vectors (loadings) of the tested parameters onto the PC1-PC2 biplot. Colours represent the different species: P. aequinoctialis = green; P. conspicillata = blue; P. westlandica = yellow; P. altirostris sp. nov. = red.

Measurements: see Table 1, Fig. 1.

Distribution: The holotype is the only known specimen of Procellaria altirostris sp. nov. All members of the genus Procellaria travel thousands of kilometres at sea when foraging and/or migrating (Brooke, 2004), so we expect that Procellaria altirostris sp. nov. had similar behaviours and would have ranged widely in the seas around proto - New Zealand and probably been a long-distance migrant like other members of this genus (Brooke, 2004).

DISCUSSION

The discovery of a well-preserved specimen of a Procellaria petrel from the Pliocene adds important new knowledge about the history of this genus. Only two previous studies have identified this genus in Pliocene deposits and those were disarticulated remains: a single partial humerus from the early Pliocene of South Africa (Olson, 1985a) and three bones from the early Pliocene of North Carolina, USA (Olson & Rasmussen, 2001). No older fossils of the genus are known (Olson, 1985b). Otherwise only much younger (Holocene) fossils have been documented (e.g.,Tennyson, 2020; Worthy & Jouventin, 1999).

This lack of a good fossil record for the genus has impaired more detailed analyses of its evolution. Our description of a Pliocene Procellaria provides further evidence to support the phylogeny proposed by Penhallurick & Wink (2004), which suggested that crown-group Procellaria taxa were already present in the Miocene.

While we can only speculate on the functional significance of the minor skeletal differences between P. altirostris sp. nov. and other members of the genus, the fossil species probably had similar feeding strategies to living Procellaria. For example, surface feeding and shallow diving, with a diet consisting mainly of fishes, cephalopods and crustaceans (Brooke, 2004). When compared with other Procellaria, the relatively short wings, particularly the short ulna compared with the humerus length, suggest that it may have been less of a glider (Kuroda, 1955), perhaps using diving more often as a feeding strategy.

Finally, while New Zealand today is the worldwide centre of diversity for species of procellariiforms, including Procellaria (Dickinson & Remsen, 2013), the new Pliocene fossil is the only evidence of Procellaria petrels being present in the country prior to the Late Pleistocene (Gill et al., 2010) and demonstrates a longer history for the genus in the region.

ACKNOWLEDGMENTS

We thank the collector, John Buchanan-Brown, for enabling the accession of this specimen into the NMNZ collection, and for preparing it; Luis Fabio Silveira and Rafael Nascimento (MZUSP) who hosted BMT at the MZUSP and assisted in locating specimens and retrieving their data; and Jean-Claude Stahl (NMNZ) for photographing NMNZ specimens. BMT was supported by a Rubicon fellowship of the Netherlands Organization for Scientific Research (NWO; Rubicon #019.181EN.005).

REFERENCES

• Baumel, J.J. & Witmer, L.M. 1993. Osteologia. In: Baumel, J.J.; King, A.S.; Breazile, J.E.; Evans, H.E. & van den Berge, J.C. (Eds.). Handbook of avian anatomy: nomina anatomica avium. Cambridge, Nuttall Ornithological Club. p. 45-132.
• Brooke, M. 2004. Albatrosses and petrels across the world. Oxford, Oxford University Press.
• Dickinson, E.C. & Remsen, J.V. 2013. The Howard & Moore complete checklist of the birds of the World. 4.ed. London, Edition. A. & C. Black.
• Gill, B.J.; Bell, B.D.; Chambers, G.K.; Medway, D.G.; Palma, R.L.; Scofield, R.P.; Tennyson, A.J.D. & Worthy, T.H. 2010. Checklist of the birds of New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency, Antarctica. Wellington, Te Papa Press.
• Kuroda, N. 1955. On the osteology of the gadfly-petrels, Pterodroma. Annotationes Zoologicae Japonenses, 28: 171-177.
• Marchant, S. & Higgins, P.J. 1990. Handbook of Australian, New Zealand and Antarctic birds. Vol. 1: ratites to ducks. Melbourne, Oxford University Press.
• Mayr, G. & Tennyson, A.J.D. 2020. A small, narrow-beaked albatross from the Pliocene of New Zealand demonstrates a higher past diversity in the feeding ecology of the Diomedeidae. Ibis, 162(3): 723-734.
• McKee, J. 1994. Geology and vertebrate palaentology of the Tangahoe Formation, South Taranaki Coast, New Zealand. Geological Society of New Zealand Miscellaneous Publications B, 80: 63-91.
• Naish, T.R.; Wehland, F.; Wilson, G.S.; Browne, G.H.; Cook, R.A.; Morgans, H.E.G.; Rosenberg, M.; King, P.R.; Smale, D.; Nelson, C.S.; Kamp, P.J.J. & Ricketts, B. 2005. An integrated sequence stratigraphic, palaeoenvironmental, and chronostratigraphic analysis of the Tangahoe Formation, southern Taranaki coast, with implications for mid-Pliocene (c. 3.4-3.0 Ma) glacio-eustatic sea-level changes. Journal of the Royal Society of New Zealand, 35: 151-196.
• Olson, S.L. 1985a. An early Pliocene marine avifauna from Duinefontein, Cape Province, South Africa. Annals of the South African Museum, 95: 147-164.
• Olson, S.L. 1985b. The fossil record of birds. Avian Biology, 8: 79-238.
• Olson, S.L. & Rasmussen, P.C. 2001. Miocene and Pliocene birds from the Lee Creek Mine, North Carolina. In: Ray, C.R. & Bohaska, D.J. (Eds.). Geology and paleontology of the Lee Creek Mine, North Carolina, III. Washington, D.C., Smithsonian Institution Press. p. 233-365.
• Penhallurick, J. & Wink, M. 2004. Analysis of the taxonomy and nomenclature of the Procellariiformes based on complete nucleotide sequences of the mitochondrial cytochrome b gene. Emu, 104: 125-147.
• Tennyson, A.J.D. 2020. Holocene bird bones found at the subantarctic Auckland Islands. Notornis, 67: 269-294.
• Tennyson, A.J.D. & Mannering, A.A. 2018. A new species of Pliocene shearwater (Aves: Procellariidae) from New Zealand. Tuhinga, 29: 1-19.
• The R Project for Statistical Computing (R Core Team). 2017. R: A language and environment for statistical computing. Vienna, R Foundation for Statistical Computing.
• Thomas, D.B.; Tennyson, A.J.D.; Scofield, R.P.; Heath, T.A.; Pett, W. & Ksepka, D.T. 2020. Ancient crested penguin constrains timing of recruitment into seabird hotspot. Proceedings of the Royal Society B, 287: 20201497.
• Warham, J. 1990. The petrels: their ecology and breeding systems. London, A. & C. Black.
• Worthy, T.H. & Jouventin, P. 1999. The fossil avifauna of Amsterdam Island, Indian Ocean. Washington, D.C., Smithsonian Contributions to Paleontology, 89: 39-65.
• Worthy, T.H.; Tennyson, A.J.D.; Jones, C.; McNamara, J.A. & Douglas, B.J. 2007. Miocene waterfowl and other birds from central Otago, New Zealand. Journal of Systematic Palaeontology, 5: 1-39.

• Published with the financial support of the "Programa de Apoio às Publicações Científicas Periódicas da USP"

APPENDIX 1

List of the comparative specimens measured. All specimens are either part of the Museum of New Zealand Te Papa Tongarewa collection (NMNZ, including specimens with’S’and ‘OR’prefixes) or the collection of the Museu de Zoologia da Universidade de São Paulo (MZUSP).

Museum Registration Taxa Skull premaxilla premaxilla premaxilla coracoid humerus humerus humerus humerus ulna radius femur tibiotarsus tibiotarsus tarsometatarsus min. intraorbital width max. length depth width max. length max. length Proximal width Distal width Shaft width max. length max. length max. length max. length incl. cnemial crest max. Length excl. cnemial crest max. length Te Papa S.046691 altirostris 15 55,56 14,52 13,5 41,99 143,22 32,26 17,68 7,44 138,53 132,95 50,05 116,56 99,53 64,82 MZUSP MZUSP 87937 aequinoctialis 14,6 40,07 152,1 33,43 17,2 7,87 149,62 145,2 51,35 115,48 101,4 66,26 MZUSP MZUSP 113761 aequinoctialis 14,01 MZUSP MZUSP 113915 aequinoctialis 14,42 MZUSP MZUSP 113917 aequinoctialis 14,56 MZUSP MZUSP 113918 aequinoctialis 14,98 MZUSP MZUSP 113919 aequinoctialis 14,11 MZUSP MZUSP 113920 aequinoctialis 12,7 MZUSP MZUSP 113921 aequinoctialis 14,05 MZUSP MZUSP 113922 aequinoctialis 15,22 MZUSP MZUSP 113923 aequinoctialis 13,75 MZUSP MZUSP 113926 aequinoctialis 14,65 MZUSP MZUSP 113929 aequinoctialis 14,38 MZUSP MZUSP 113930 aequinoctialis 14,26 MZUSP MZUSP 113932 aequinoctialis 13,18 MZUSP MZUSP 113933 aequinoctialis 13,37 MZUSP MZUSP 114258 aequinoctialis 13,83 12,18 13,61 38,26 150,14 31,98 17,91 7,91 152,06 149,83 51,15 115,39 100,7 66,19 Te Papa OR.21098 aequinoctialis 14,41 63,8 11,68 13,95 Te Papa OR.21096 aequinoctialis 13,77 62,67 12,05 14,18 Te Papa OR.21097 aequinoctialis 14,54 60,76 12,67 14,27 Te Papa OR.28949 aequinoctialis 12,87 10,69 12,66 Te Papa OR.18904 aequinoctialis 13,86 60,68 11,43 13,47 37,26 149,13 31,53 17,2 7,01 149,15 145,43 50,99 101,03 65,01 Te Papa OR.21095 aequinoctialis 13,8 38,2 153,5 32,52 17,49 7,52 151,9 148,98 51,21 113,08 99,02 66,11 Te Papa OR.21841 aequinoctialis 14,57 60,62 12,2 13,32 36,5 155 31,56 17,27 7,23 153,5 151,14 117,92 102,98 69 Te Papa OR.26612 aequinoctialis 14,08 64,03 12,49 13,1 Te Papa OR.608 aequinoctialis 14,62 63,15 12,71 13,67 39,05 151,5 33,93 17,96 150,43 146,37 98,21 64,08 Te Papa OR.542-S aequinoctialis 14,38 60,81 11,52 13,56 48,61 107,29 94,45 63,34 Te Papa OR.543-S aequinoctialis 13,09 60,76 11,7 13,9 Te Papa OR.16054 aequinoctialis 14,91 66,21 12,66 14,22 158 33,83 18,36 7,89 156 152,8 53,78 115,7 101,94 68,03 Te Papa OR.25171 aequinoctialis 15,39 63,5 12,1 13,58 38,46 151,3 32,19 17,82 7,6 153 149,56 50,01 100,6 65,11 Te Papa OR.29968 aequinoctialis 12,81 59,71 11,94 14,21 146,66 31,71 16,8 7,2 148,86 145,86 49,83 111,82 97,16 65,52 Te Papa OR.19809 aequinoctialis 15,15 61,94 11,66 13,5 37,35 152,1 30,93 16,96 7,68 152,13 148,44 50,02 115,92 100,51 68,03 Te Papa OR.22411 aequinoctialis 15,2 59,52 11,7 13,28 35,61 142,58 31,57 16,76 7,66 145,94 142,56 47,04 111,05 96,83 62,87 Te Papa OR.22077 aequinoctialis 14,48 62,26 11,61 12,53 37,47 152,41 31,94 17,67 7,72 154 150,95 50,7 115,47 100,37 68,11 Te Papa OR.21108 aequinoctialis 14,03 64,77 12,09 14,09 38,05 147,39 31,47 17,25 7,61 148,2 145,28 49,58 114,94 100,25 66,27 Te Papa OR.22078 aequinoctialis 14,43 61,55 12,05 14,01 39,39 151,32 32,5 17,75 7,88 152,51 148,35 52,68 117,25 102,34 67,65 Te Papa OR.21863 aequinoctialis 14,7 62,69 12,34 13,82 38,05 151,58 32,56 18,1 7,7 150,15 146,22 49,99 114,3 99,68 66,58 Te Papa OR.15931 aequinoctialis 13,41 59,6 11,44 14,46 37,22 147,14 31,71 17,17 7,3 147,69 144,3 49,74 110,22 97,42 63,2 Te Papa OR.25169 aequinoctialis 14,56 64,76 12,53 15,49 37,36 150,68 32,45 17,79 7,78 151,31 147,02 51,68 114,09 98,85 65,73 Te Papa OR.536-S aequinoctialis 14,29 62,09 12,39 13,48 37,62 149,68 31,47 17,39 7,32 152,17 148,37 50,04 112,03 98,37 66,17 Te Papa OR.535-S aequinoctialis 13,98 59,92 11,47 13,11 36,51 146,51 31,42 17,31 7,4 148,69 145,48 50,76 111,53 97,42 65,38 Te Papa OR.17184 aequinoctialis 14,61 59,95 11,95 14,63 36,34 144,99 31,59 16,93 7,44 145,73 142,51 47,58 108,82 95 63,19 Te Papa OR.17185 aequinoctialis 14,42 58,89 11,18 12,34 38,33 148,59 31,81 17,54 7,12 150,93 146,67 49,81 111,41 96,97 64,84 Te Papa OR.25170 aequinoctialis 14,15 63,6 11,79 13,18 38,42 154 34,35 17,38 7,78 155,5 150,66 51,74 118,88 102,55 68,54 Te Papa OR.23463 aequinoctialis 14,92 62,7 12,44 15,15 38,85 148,67 33,6 17,22 7,97 148,59 144,87 51,19 116 102,14 65,88 Te Papa OR.22506 aequinoctialis 14,28 62,8 12,87 15,38 37,77 151,62 32,73 17,72 8,17 150,26 146,05 52,18 114,53 100 65,52 Te Papa OR.25172 aequinoctialis 13,99 61,97 12 13,48 37,1 150,74 32,86 17,64 7,42 152,03 148,72 50 114,58 98,76 65,35 Te Papa OR.23529 aequinoctialis 14,53 63,18 11,93 14,08 39,39 149,92 32,32 17,62 7,5 150,46 146,57 50,14 114,38 99,31 66,7 Te Papa OR.30424 aequinoctialis 14,99 64,94 11,74 12,87 39,43 156 32,49 17,56 7,35 156 154 54,47 115,32 99,86 68,89 Te Papa OR.25633 aequinoctialis 14,26 65,92 11,53 13,77 38,14 154 33,58 17,93 7,6 154,5 150,86 51,5 116,67 101,91 66,88 Te Papa OR.25634 aequinoctialis 14,53 61,43 11,95 13,61 37,99 153,32 31,72 17,46 7,55 151,36 146,57 50,18 114,14 98,92 65,24 Te Papa OR.12035 cinerea 36,6 141,64 30,56 16,54 7,37 140,21 136,44 Te Papa OR.16486 cinerea 13,23 55,13 8,81 12,94 34,48 134,22 28,95 16,01 7,3 49,25 108,2 93,99 61,36 Te Papa OR.18136 cinerea 14,22 58,04 10,14 12 35,4 135,7 31,13 16,47 8,05 132,54 128,53 48,15 109,8 94,95 Te Papa OR.21862 cinerea 13,19 57,27 10,34 11,39 37,62 138,89 32,27 16,28 7,19 133,9 131,13 50,84 96,33 61,55 Te Papa OR.607-S cinerea 12,7 55,45 10,25 11,94 134,78 29,77 16 7,19 131,41 128,77 Te Papa OR.791-S cinerea 9,94 12,83 96,42 60,53 Te Papa OR.853-S cinerea 13,49 59,16 9,8 11,99 61,73 Te Papa OR.602-S cinerea 36,86 138,46 31,49 16,5 8,04 135,18 131,99 Te Papa OR.24981 cinerea 14,48 57,55 10,21 12,03 137,23 30,6 16,17 7,14 132,81 130,32 48,36 109,26 95,33 63,35 Te Papa OR.26232 cinerea 12,98 56,32 9,88 12,61 36,58 138,93 31,63 16,76 7,41 133,86 131,2 47,75 96,29 61,01 Te Papa OR.25168 cinerea 14,14 56,83 10,18 12,6 Te Papa OR.24975 cinerea 13,7 56,8 10,22 11,7 35,35 137,7 28,8 16,05 7,14 134,45 131,58 49,41 110,8 96,1 60,76 Te Papa OR.24662 cinerea 14,05 61,52 10,62 13,85 35,25 135,9 28,63 16,4 7,45 134,13 130,5 47,85 109,95 96,02 60,21 Te Papa OR.24972 cinerea 13,75 54,72 8,97 12,65 37,25 139,74 30,65 16,1 7,63 137,57 133,07 48,55 110,95 99,59 61,67 Te Papa OR.24664 cinerea 14,83 55,98 9,7 12,1 35,75 140,82 29,95 16,44 7,87 137,27 133,79 48,67 111,76 97,21 63,28 Te Papa OR.24663 cinerea 13,79 59,26 10,75 13,62 37,3 139,86 30,7 16,55 7,54 137,46 134,42 50,62 111,43 96,36 63,18 Te Papa OR.24974 cinerea 13,58 55,95 9,84 13,94 35,11 135,21 30,17 16,45 7,11 134,14 130,41 48,89 110,48 95,63 62,57 Te Papa OR.17183 cinerea 12,15 55,48 9,74 11,44 35,43 134,07 29,6 15,84 7,63 132,26 128,72 48,21 108 93,83 61,94 Te Papa OR.12475 cinerea 13,11 55,74 9,32 12,2 35,74 134,44 30,38 15,96 7,06 135,29 132,04 44,67 110,63 95,02 61 Te Papa OR.12474 cinerea 13,39 56,6 10,45 13,23 36,16 135,72 29,18 16,28 7,16 134,04 129,98 48,88 110,9 96,6 63,34 Te Papa OR.12248 cinerea 13,45 56,87 9,27 11,88 36,7 137,09 29,9 16,29 6,85 134,03 130,71 47,81 110,64 94,79 61,41 Te Papa OR.24089 cinerea 13,77 59,36 9,9 11,28 36,91 139,56 32,3 16,69 7,81 139,53 136,56 48,94 112,29 97,4 63,55 Te Papa OR.24153 cinerea 13,8 60,13 10,73 13,65 37,2 138,15 30,9 17,14 7,3 133,67 129,81 49,2 110,8 95,35 61,8 Te Papa OR.24215 cinerea 14,62 57,32 10 13,26 35,46 137,88 30,78 16,73 7,69 138,5 134,65 49,99 111,71 97,53 62,92 Te Papa OR.19305 cinerea 14,61 59,56 10,24 12,18 37,95 140,73 31,9 16,64 7,14 141,29 137,26 51,28 113,64 99,29 63,43 Te Papa OR.792-S cinerea 13,33 55,78 10,24 11,36 35,69 136,54 29,57 16,14 7,37 135,1 132,04 48,43 111,32 97,1 62,08 Te Papa OR.918-S cinerea 12,8 58,4 10,1 12,83 36,48 137,29 31,27 16,71 7 136,44 133 51,64 113,41 98,79 62,76 Te Papa OR.24977 cinerea 14,24 58,55 10,01 13,14 37,64 138,88 31,05 16,09 7,51 139,43 135,77 49,6 109,91 95,87 60,74 Te Papa OR.24976 cinerea 14,39 56,56 9,8 11,28 35,21 139,69 31,4 16,91 7,31 137,11 133,88 49,61 110,64 95,76 62,42 Te Papa OR.24980 cinerea 13,15 56,97 10,05 13,38 35,4 135,48 30,26 16,27 7,42 133,55 129,79 48,71 108,9 93,91 61,28 Te Papa OR.24978 cinerea 13,87 58,1 10,02 12,65 36,2 136,83 30,22 16,26 7,13 134,48 131,8 49,79 111,94 97,08 63,28 Te Papa OR.24982 cinerea 14,32 57,1 9,73 12,9 35,37 135,4 30,67 16,31 7,42 133,3 130,09 46,38 107,9 93,74 59,59 Te Papa OR.24511 cinerea 13,85 56,97 10,54 11,69 36,58 134,78 30,48 16,7 7,31 133,29 130,33 47,19 108,05 93,94 60,81 Te Papa OR.24614 cinerea 14,06 57,05 10,25 13,59 36,54 136,64 30,81 16,4 7,48 136,24 132,02 49,21 111,57 96,19 61,82 Te Papa OR.24512 cinerea 13,97 57,69 9,88 12,5 36,58 138,64 30,24 16,65 7,54 133,93 130,63 48,34 108,64 94,08 61,91 Te Papa OR.24510 cinerea 13,57 57,35 9,92 11,95 36,21 134,36 30,9 16,13 7,91 138,78 128,4 47,51 106,42 92,59 60,8 Te Papa OR.24659 cinerea 13,77 55,4 9,02 11,6 34,46 132,82 30,02 16,25 7,03 133,54 129,87 47,45 106,76 93,06 60,9 Te Papa OR.24432 cinerea 13,81 53,99 10,06 11,47 35,88 132,71 30,66 16,57 7,5 132,87 130,45 47,8 108,75 95,04 61,3 Te Papa OR.24983 cinerea 14,5 56,49 9,94 13,15 36 134,29 30,84 16,68 7,63 135,1 131,12 46,75 107,06 92,21 61,01 Te Papa OR.28850 cinerea 14,4 57,16 9,4 11,04 35,87 128,98 31,53 16,34 7,57 126,71 123,18 47,41 105,41 90,4 56,88 MZUSP MZUSP 113526 conspicillata 12,93 38,06 148,04 32,39 17,63 7,16 148,29 145,34 50,01 113,83 98,75 65,11 MZUSP MZUSP 113543 conspicillata 13,73 152,62 31,35 17,49 7,43 150,21 146,67 49,9 115,37 100,17 65,09 MZUSP MZUSP 113688 conspicillata 13,54 149,22 30,96 17,54 7,48 149,4 145,54 49,65 112,96 97,86 65,31 MZUSP MZUSP 113924 conspicillata 13,11 MZUSP MZUSP 113927 conspicillata 13,35 MZUSP MZUSP 113934 conspicillata 13,56 MZUSP MZUSP 99377 conspicillata 14,74 57,22 11,95 14,23 39,79 155 32,39 18,42 7,45 151,46 146,79 52,54 115,75 100,94 65,8 Te Papa OR.18907 parkinsoni 10,41 49,02 9,43 11,4 31,4 124,61 27,32 14,86 6,28 126,4 123,77 40,01 92,84 79,71 53,37 Te Papa OR.18906 parkinsoni 11,75 48,71 9,69 12,18 32,17 128,93 27,61 14,82 6,6 128,48 125,76 41,04 94,61 82,23 54,66 Te Papa OR.17258 parkinsoni 11,55 8,93 10,59 33,42 132,37 28,3 14,53 6,12 133,52 130,21 44,61 95,46 83,55 53,96 Te Papa OR.793-S parkinsoni 10,83 53,18 10,03 11,86 15,52 7,03 136,73 133,23 57,6 Te Papa OR.25953a parkinsoni 10,68 53,98 10,37 11,2 33,17 129,72 28,19 14,68 6,11 130,88 128,07 42,9 Te Papa OR.19283 parkinsoni 11,08 46,69 9,14 11,8 32,18 129,47 27,7 14,39 6,44 131,05 128,28 41,83 94,79 82,81 55,51 Te Papa OR.18969 parkinsoni 11,32 46,71 9,25 10,73 32,29 127,58 27,17 14,67 6,45 126,19 123,09 41,96 92 80,3 54,23 Te Papa OR.27276a parkinsoni 10,29 13,07 33,73 131,52 28,09 15,05 6,49 130,8 128,6 43,5 96,18 83,65 54,83 Te Papa OR.24247 parkinsoni 11,02 46,78 10,25 11,98 32,96 129,1 28,59 14,58 6,41 127,66 125,19 Te Papa OR.27119a parkinsoni 34,75 135,82 29,56 15,2 6,55 138,32 134,62 43,78 Te Papa OR.29637 parkinsoni 12,33 49,56 10,36 11,74 34,67 130,58 29,08 15,9 6,88 131,64 127,73 43,18 95,58 83,09 54,22 Te Papa OR.18905 parkinsoni 12,5 49,47 9,25 10,82 33,33 128,67 29,19 14,81 6,68 129,48 126,58 41,19 96,64 83,62 54,22 Te Papa OR.19282 parkinsoni 11,79 48,63 10,92 12,59 32,1 127,3 27,91 15,3 6,78 130,04 127,26 37,73 90,96 78,16 52,12 Te Papa OR.18968 parkinsoni 11,67 47,84 9,25 13,17 32,28 125,09 27,09 15,14 6,96 127,37 123,74 40,57 91,3 78,7 50,94 Te Papa OR.19298 parkinsoni 11,74 50,16 9,88 11,8 34,76 130,41 28,45 14,96 6,41 130,57 126,88 42,02 93,57 81,01 54,42 Te Papa OR.19781 parkinsoni 11,01 52,08 9,71 12,4 34,29 134,66 28,06 15,23 6,3 135,51 131,65 44,05 97,76 85,24 55,25 Te Papa OR.24236 parkinsoni 10,92 49,72 10,04 12,98 34,57 131,55 27,95 15,2 6,69 132,88 129,15 42,76 96,32 83,87 54,9 Te Papa OR.289617 parkinsoni 11,79 49,62 10,06 13,38 34,96 133,72 28,91 15,8 6,21 135,76 133,66 45,39 100,45 87,06 57,68 Te Papa OR.21422 westlandica 14,69 60,88 12,87 14,24 40,4 149,98 32,54 17,18 7,49 148,43 144,33 63,92 Te Papa OR.22084 westlandica 14,63 58,26 11,82 14,72 38,95 144,04 30,86 16,33 7,14 143,99 141,29 Te Papa OR.21423 westlandica 13,93 12,15 15,61 39,45 149,9 34,12 7,31 149,06 144,37 51,89 98,08 64,38 Te Papa OR.21424 westlandica 15,9 59,24 11,73 14,61 38,91 149,5 17,2 7,4 150,2 146,57 51,52 99,6 63,46 Te Papa OR.21106 westlandica 14,35 58,56 12,29 15,05 39,89 151,68 33,1 17,21 8,27 152,82 96,35 Te Papa OR.21094 westlandica 38,82 155 32,91 17,1 7,09 150,94 147,81 51,46 97,62 64,66 Te Papa OR.24272 westlandica 15,26 12,64 17,03 40,63 150,21 32,9 17,46 7,94 148,28 145,25 50,2 112,52 97,43 64,09 Te Papa OR.11494 westlandica 14,44 Te Papa OR.838-S westlandica 38,61 150,94 33,3 17,3 7,73 148,63 145,69 111,4 96,6 62,5 Te Papa OR.15900 westlandica 13,84 12,24 15,73 39,91 146,4 32,22 17,02 7,38 146,7 142,16 50,38 109,22 93,83 60,87 Te Papa OR.599-S westlandica 38,6 32,61 50,56 Te Papa OR.11511 westlandica 12,59 56,32 10,94 14,32 37,95 144,98 32 16,52 6,98 147,46 143,7 50,93 111,04 96,07 63,1 Te Papa OR.22968 westlandica 151,72 31,87 16,96 7,52 158 155 114,99 99,37 66,13 Te Papa OR.22933 westlandica 14,4 57,49 10,92 13,51 148,52 32,04 16,97 6,81 151,83 147,81 49,38 112,81 98,38 63,15 Te Papa OR.22085 westlandica 13,27 11,34 13,32 38,53 150,38 31,87 16,9 7 151,13 146,9 51 112,29 95,52 63,54 Te Papa OR.23766 westlandica 14,32 59,27 12,38 11,49 149,35 32,5 17,43 7,54 153,5 151,19 50,02 109,16 93,94 62,31 Te Papa OR.28672 westlandica 14,39 62,2 12,1 13,98 40,19 151,29 31,48 17,3 7,07 151,27 147,12 51,32 112,42 97,88 64,85 Te Papa OR.23155 westlandica 14,24 12,65 13,73 150,31 32,62 16,75 7,39 151,74 146,7 111,73 97,53 Te Papa OR.23151 westlandica 13,45 57,41 11,1 14,39 142,7 31,15 15,85 7,2 144,87 140,85 108,15 92,64 59,82 Te Papa OR.19299 westlandica 14,97 61,78 13,04 14,98 38,66 152 31,95 17,27 7,14 150 148 51,88 114,52 99,74 64,34 Te Papa OR.21091 westlandica 13,6 60,13 11,4 15,34 38,9 148,14 31,37 16,74 7,49 150,47 145,83 50,36 109,88 95,91 63,46 Te Papa OR.19281 westlandica 14,95 61,08 12,84 15,78 38,92 146,9 34,5 17,36 8,04 145,8 142,33 51,52 110,85 95,91 63,19 Te Papa OR.19311 westlandica 14,96 60,86 13,38 14,69 39,92 152,82 33,45 17,78 7,42 150,84 147,45 52,58 116,27 100,37 67,33 Te Papa OR.21499 westlandica 14,97 57,47 11,96 14,41 36,94 149,63 30,47 16,28 7,32 152,85 149,63 51,92 112,35 97,74 65,09 Te Papa OR.21466 westlandica 15,42 62,58 13,17 14,85 38,35 150,21 32,38 17,14 7,58 148,45 145,12 52,8 112,42 97,25 63,9 Te Papa OR.21099 westlandica 14,87 60,68 11,37 15 38,83 146,07 32,74 17,47 7,6 146,33 143,21 49,67 107,42 93,27 62,57 Te Papa OR.21092 westlandica 14,37 57,7 12,07 16,3 37,21 150,95 31,87 17,31 7,65 152,75 148,88 52,78 115,58 100,13 65,62 Te Papa OR.21093 westlandica 14,84 58,53 12,11 14,81 39,3 148,58 31,75 17,43 7,52 148,02 145,12 50,42 111,04 96,17 63 Te Papa OR.27314 westlandica 14,49 60,97 12,49 14,3 39,76 148,35 33,27 17,48 7,48 147,37 144,45 50,24 113,19 97,7 64,07 Te Papa OR.27409 westlandica 14,04 57,54 12,52 14,4 38,85 152,17 31,33 16,88 7,83 152,35 149,18 47,74 109,79 94,75 61,36 Te Papa OR.24271 westlandica 13,92 60,97 11,7 13,51 40,33 157 33,91 17,45 7,53 156 151,83 49,88 114,59 98,76 64,61 Te Papa OR.24244 westlandica 13,72 56,51 11,78 14,94 39,35 148,21 32,4 17,11 7,13 146,21 142,19 49,62 114,13 98,05 63,66 Te Papa OR.13566 westlandica 13,3 63,3 12,06 16,24 38,26 152,45 32,83 17,59 7,65 153,03 150,02 49,9 110,27 96,15 65,62 Te Papa OR.13594 westlandica 14,13 58,89 11,86 14,8 38,97 150,84 32,72 17,13 7,4 151,4 146,62 52,81 113,71 98,02 63,32 Te Papa OR.22668 westlandica 13,79 57,8 11,62 14,92 39,47 149,63 31,57 17,06 7,2 149,77 146,76 50,04 109,5 94,71 62,96 Te Papa OR.23734 westlandica 15,25 61,09 12,25 14,61 39,02 143,61 31,89 16,24 7,37 146,68 143,34 50,11 109,15 94,88 61,38 Te Papa OR.23725 westlandica 14,35 61,73 11,95 14,1 39,4 151,45 33,42 17,43 7,7 152,58 149,2 51,29 111,96 97,75 64,26 Te Papa OR.23121 westlandica 14,86 58,93 12,22 14,66 38,94 149,41 31,41 17,45 7,58 149,45 147,13 50,76 111,12 97,11 63,18 Te Papa OR.23306 westlandica 15,51 60,68 13,21 15,53 40,9 151,91 32,32 18,13 7,36 147,37 144,96 53,01 114,13 97,79 64,23

Museum	Registration	Taxa	Skull	premaxilla	premaxilla	premaxilla	coracoid	humerus	humerus	humerus	humerus	ulna	radius	femur	tibiotarsus	tibiotarsus	tarsometatarsus
			min. intraorbital width	max. length	depth	width	max. length	max. length	Proximal width	Distal width	Shaft width	max. length	max. length	max. length	max. length incl. cnemial crest	max. Length excl. cnemial crest	max. length
Te Papa	S.046691	altirostris	15	55,56	14,52	13,5	41,99	143,22	32,26	17,68	7,44	138,53	132,95	50,05	116,56	99,53	64,82
MZUSP	MZUSP 87937	aequinoctialis	14,6				40,07	152,1	33,43	17,2	7,87	149,62	145,2	51,35	115,48	101,4	66,26
MZUSP	MZUSP 113761	aequinoctialis	14,01
MZUSP	MZUSP 113915	aequinoctialis	14,42
MZUSP	MZUSP 113917	aequinoctialis	14,56
MZUSP	MZUSP 113918	aequinoctialis	14,98
MZUSP	MZUSP 113919	aequinoctialis	14,11
MZUSP	MZUSP 113920	aequinoctialis	12,7
MZUSP	MZUSP 113921	aequinoctialis	14,05
MZUSP	MZUSP 113922	aequinoctialis	15,22
MZUSP	MZUSP 113923	aequinoctialis	13,75
MZUSP	MZUSP 113926	aequinoctialis	14,65
MZUSP	MZUSP 113929	aequinoctialis	14,38
MZUSP	MZUSP 113930	aequinoctialis	14,26
MZUSP	MZUSP 113932	aequinoctialis	13,18
MZUSP	MZUSP 113933	aequinoctialis	13,37
MZUSP	MZUSP 114258	aequinoctialis	13,83		12,18	13,61	38,26	150,14	31,98	17,91	7,91	152,06	149,83	51,15	115,39	100,7	66,19
Te Papa	OR.21098	aequinoctialis	14,41	63,8	11,68	13,95
Te Papa	OR.21096	aequinoctialis	13,77	62,67	12,05	14,18
Te Papa	OR.21097	aequinoctialis	14,54	60,76	12,67	14,27
Te Papa	OR.28949	aequinoctialis	12,87		10,69	12,66
Te Papa	OR.18904	aequinoctialis	13,86	60,68	11,43	13,47	37,26	149,13	31,53	17,2	7,01	149,15	145,43	50,99		101,03	65,01
Te Papa	OR.21095	aequinoctialis	13,8				38,2	153,5	32,52	17,49	7,52	151,9	148,98	51,21	113,08	99,02	66,11
Te Papa	OR.21841	aequinoctialis	14,57	60,62	12,2	13,32	36,5	155	31,56	17,27	7,23	153,5	151,14		117,92	102,98	69
Te Papa	OR.26612	aequinoctialis	14,08	64,03	12,49	13,1
Te Papa	OR.608	aequinoctialis	14,62	63,15	12,71	13,67	39,05	151,5	33,93	17,96		150,43	146,37			98,21	64,08
Te Papa	OR.542-S	aequinoctialis	14,38	60,81	11,52	13,56								48,61	107,29	94,45	63,34
Te Papa	OR.543-S	aequinoctialis	13,09	60,76	11,7	13,9
Te Papa	OR.16054	aequinoctialis	14,91	66,21	12,66	14,22		158	33,83	18,36	7,89	156	152,8	53,78	115,7	101,94	68,03
Te Papa	OR.25171	aequinoctialis	15,39	63,5	12,1	13,58	38,46	151,3	32,19	17,82	7,6	153	149,56	50,01		100,6	65,11
Te Papa	OR.29968	aequinoctialis	12,81	59,71	11,94	14,21		146,66	31,71	16,8	7,2	148,86	145,86	49,83	111,82	97,16	65,52
Te Papa	OR.19809	aequinoctialis	15,15	61,94	11,66	13,5	37,35	152,1	30,93	16,96	7,68	152,13	148,44	50,02	115,92	100,51	68,03
Te Papa	OR.22411	aequinoctialis	15,2	59,52	11,7	13,28	35,61	142,58	31,57	16,76	7,66	145,94	142,56	47,04	111,05	96,83	62,87
Te Papa	OR.22077	aequinoctialis	14,48	62,26	11,61	12,53	37,47	152,41	31,94	17,67	7,72	154	150,95	50,7	115,47	100,37	68,11
Te Papa	OR.21108	aequinoctialis	14,03	64,77	12,09	14,09	38,05	147,39	31,47	17,25	7,61	148,2	145,28	49,58	114,94	100,25	66,27
Te Papa	OR.22078	aequinoctialis	14,43	61,55	12,05	14,01	39,39	151,32	32,5	17,75	7,88	152,51	148,35	52,68	117,25	102,34	67,65
Te Papa	OR.21863	aequinoctialis	14,7	62,69	12,34	13,82	38,05	151,58	32,56	18,1	7,7	150,15	146,22	49,99	114,3	99,68	66,58
Te Papa	OR.15931	aequinoctialis	13,41	59,6	11,44	14,46	37,22	147,14	31,71	17,17	7,3	147,69	144,3	49,74	110,22	97,42	63,2
Te Papa	OR.25169	aequinoctialis	14,56	64,76	12,53	15,49	37,36	150,68	32,45	17,79	7,78	151,31	147,02	51,68	114,09	98,85	65,73
Te Papa	OR.536-S	aequinoctialis	14,29	62,09	12,39	13,48	37,62	149,68	31,47	17,39	7,32	152,17	148,37	50,04	112,03	98,37	66,17
Te Papa	OR.535-S	aequinoctialis	13,98	59,92	11,47	13,11	36,51	146,51	31,42	17,31	7,4	148,69	145,48	50,76	111,53	97,42	65,38
Te Papa	OR.17184	aequinoctialis	14,61	59,95	11,95	14,63	36,34	144,99	31,59	16,93	7,44	145,73	142,51	47,58	108,82	95	63,19
Te Papa	OR.17185	aequinoctialis	14,42	58,89	11,18	12,34	38,33	148,59	31,81	17,54	7,12	150,93	146,67	49,81	111,41	96,97	64,84
Te Papa	OR.25170	aequinoctialis	14,15	63,6	11,79	13,18	38,42	154	34,35	17,38	7,78	155,5	150,66	51,74	118,88	102,55	68,54
Te Papa	OR.23463	aequinoctialis	14,92	62,7	12,44	15,15	38,85	148,67	33,6	17,22	7,97	148,59	144,87	51,19	116	102,14	65,88
Te Papa	OR.22506	aequinoctialis	14,28	62,8	12,87	15,38	37,77	151,62	32,73	17,72	8,17	150,26	146,05	52,18	114,53	100	65,52
Te Papa	OR.25172	aequinoctialis	13,99	61,97	12	13,48	37,1	150,74	32,86	17,64	7,42	152,03	148,72	50	114,58	98,76	65,35
Te Papa	OR.23529	aequinoctialis	14,53	63,18	11,93	14,08	39,39	149,92	32,32	17,62	7,5	150,46	146,57	50,14	114,38	99,31	66,7
Te Papa	OR.30424	aequinoctialis	14,99	64,94	11,74	12,87	39,43	156	32,49	17,56	7,35	156	154	54,47	115,32	99,86	68,89
Te Papa	OR.25633	aequinoctialis	14,26	65,92	11,53	13,77	38,14	154	33,58	17,93	7,6	154,5	150,86	51,5	116,67	101,91	66,88
Te Papa	OR.25634	aequinoctialis	14,53	61,43	11,95	13,61	37,99	153,32	31,72	17,46	7,55	151,36	146,57	50,18	114,14	98,92	65,24
Te Papa	OR.12035	cinerea					36,6	141,64	30,56	16,54	7,37	140,21	136,44
Te Papa	OR.16486	cinerea	13,23	55,13	8,81	12,94	34,48	134,22	28,95	16,01	7,3			49,25	108,2	93,99	61,36
Te Papa	OR.18136	cinerea	14,22	58,04	10,14	12	35,4	135,7	31,13	16,47	8,05	132,54	128,53	48,15	109,8	94,95
Te Papa	OR.21862	cinerea	13,19	57,27	10,34	11,39	37,62	138,89	32,27	16,28	7,19	133,9	131,13	50,84		96,33	61,55
Te Papa	OR.607-S	cinerea	12,7	55,45	10,25	11,94		134,78	29,77	16	7,19	131,41	128,77
Te Papa	OR.791-S	cinerea			9,94	12,83										96,42	60,53
Te Papa	OR.853-S	cinerea	13,49	59,16	9,8	11,99											61,73
Te Papa	OR.602-S	cinerea					36,86	138,46	31,49	16,5	8,04	135,18	131,99
Te Papa	OR.24981	cinerea	14,48	57,55	10,21	12,03		137,23	30,6	16,17	7,14	132,81	130,32	48,36	109,26	95,33	63,35
Te Papa	OR.26232	cinerea	12,98	56,32	9,88	12,61	36,58	138,93	31,63	16,76	7,41	133,86	131,2	47,75		96,29	61,01
Te Papa	OR.25168	cinerea	14,14	56,83	10,18	12,6
Te Papa	OR.24975	cinerea	13,7	56,8	10,22	11,7	35,35	137,7	28,8	16,05	7,14	134,45	131,58	49,41	110,8	96,1	60,76
Te Papa	OR.24662	cinerea	14,05	61,52	10,62	13,85	35,25	135,9	28,63	16,4	7,45	134,13	130,5	47,85	109,95	96,02	60,21
Te Papa	OR.24972	cinerea	13,75	54,72	8,97	12,65	37,25	139,74	30,65	16,1	7,63	137,57	133,07	48,55	110,95	99,59	61,67
Te Papa	OR.24664	cinerea	14,83	55,98	9,7	12,1	35,75	140,82	29,95	16,44	7,87	137,27	133,79	48,67	111,76	97,21	63,28
Te Papa	OR.24663	cinerea	13,79	59,26	10,75	13,62	37,3	139,86	30,7	16,55	7,54	137,46	134,42	50,62	111,43	96,36	63,18
Te Papa	OR.24974	cinerea	13,58	55,95	9,84	13,94	35,11	135,21	30,17	16,45	7,11	134,14	130,41	48,89	110,48	95,63	62,57
Te Papa	OR.17183	cinerea	12,15	55,48	9,74	11,44	35,43	134,07	29,6	15,84	7,63	132,26	128,72	48,21	108	93,83	61,94
Te Papa	OR.12475	cinerea	13,11	55,74	9,32	12,2	35,74	134,44	30,38	15,96	7,06	135,29	132,04	44,67	110,63	95,02	61
Te Papa	OR.12474	cinerea	13,39	56,6	10,45	13,23	36,16	135,72	29,18	16,28	7,16	134,04	129,98	48,88	110,9	96,6	63,34
Te Papa	OR.12248	cinerea	13,45	56,87	9,27	11,88	36,7	137,09	29,9	16,29	6,85	134,03	130,71	47,81	110,64	94,79	61,41
Te Papa	OR.24089	cinerea	13,77	59,36	9,9	11,28	36,91	139,56	32,3	16,69	7,81	139,53	136,56	48,94	112,29	97,4	63,55
Te Papa	OR.24153	cinerea	13,8	60,13	10,73	13,65	37,2	138,15	30,9	17,14	7,3	133,67	129,81	49,2	110,8	95,35	61,8
Te Papa	OR.24215	cinerea	14,62	57,32	10	13,26	35,46	137,88	30,78	16,73	7,69	138,5	134,65	49,99	111,71	97,53	62,92
Te Papa	OR.19305	cinerea	14,61	59,56	10,24	12,18	37,95	140,73	31,9	16,64	7,14	141,29	137,26	51,28	113,64	99,29	63,43
Te Papa	OR.792-S	cinerea	13,33	55,78	10,24	11,36	35,69	136,54	29,57	16,14	7,37	135,1	132,04	48,43	111,32	97,1	62,08
Te Papa	OR.918-S	cinerea	12,8	58,4	10,1	12,83	36,48	137,29	31,27	16,71	7	136,44	133	51,64	113,41	98,79	62,76
Te Papa	OR.24977	cinerea	14,24	58,55	10,01	13,14	37,64	138,88	31,05	16,09	7,51	139,43	135,77	49,6	109,91	95,87	60,74
Te Papa	OR.24976	cinerea	14,39	56,56	9,8	11,28	35,21	139,69	31,4	16,91	7,31	137,11	133,88	49,61	110,64	95,76	62,42
Te Papa	OR.24980	cinerea	13,15	56,97	10,05	13,38	35,4	135,48	30,26	16,27	7,42	133,55	129,79	48,71	108,9	93,91	61,28
Te Papa	OR.24978	cinerea	13,87	58,1	10,02	12,65	36,2	136,83	30,22	16,26	7,13	134,48	131,8	49,79	111,94	97,08	63,28
Te Papa	OR.24982	cinerea	14,32	57,1	9,73	12,9	35,37	135,4	30,67	16,31	7,42	133,3	130,09	46,38	107,9	93,74	59,59
Te Papa	OR.24511	cinerea	13,85	56,97	10,54	11,69	36,58	134,78	30,48	16,7	7,31	133,29	130,33	47,19	108,05	93,94	60,81
Te Papa	OR.24614	cinerea	14,06	57,05	10,25	13,59	36,54	136,64	30,81	16,4	7,48	136,24	132,02	49,21	111,57	96,19	61,82
Te Papa	OR.24512	cinerea	13,97	57,69	9,88	12,5	36,58	138,64	30,24	16,65	7,54	133,93	130,63	48,34	108,64	94,08	61,91
Te Papa	OR.24510	cinerea	13,57	57,35	9,92	11,95	36,21	134,36	30,9	16,13	7,91	138,78	128,4	47,51	106,42	92,59	60,8
Te Papa	OR.24659	cinerea	13,77	55,4	9,02	11,6	34,46	132,82	30,02	16,25	7,03	133,54	129,87	47,45	106,76	93,06	60,9
Te Papa	OR.24432	cinerea	13,81	53,99	10,06	11,47	35,88	132,71	30,66	16,57	7,5	132,87	130,45	47,8	108,75	95,04	61,3
Te Papa	OR.24983	cinerea	14,5	56,49	9,94	13,15	36	134,29	30,84	16,68	7,63	135,1	131,12	46,75	107,06	92,21	61,01
Te Papa	OR.28850	cinerea	14,4	57,16	9,4	11,04	35,87	128,98	31,53	16,34	7,57	126,71	123,18	47,41	105,41	90,4	56,88
MZUSP	MZUSP 113526	conspicillata	12,93				38,06	148,04	32,39	17,63	7,16	148,29	145,34	50,01	113,83	98,75	65,11
MZUSP	MZUSP 113543	conspicillata	13,73					152,62	31,35	17,49	7,43	150,21	146,67	49,9	115,37	100,17	65,09
MZUSP	MZUSP 113688	conspicillata	13,54					149,22	30,96	17,54	7,48	149,4	145,54	49,65	112,96	97,86	65,31
MZUSP	MZUSP 113924	conspicillata	13,11
MZUSP	MZUSP 113927	conspicillata	13,35
MZUSP	MZUSP 113934	conspicillata	13,56
MZUSP	MZUSP 99377	conspicillata	14,74	57,22	11,95	14,23	39,79	155	32,39	18,42	7,45	151,46	146,79	52,54	115,75	100,94	65,8
Te Papa	OR.18907	parkinsoni	10,41	49,02	9,43	11,4	31,4	124,61	27,32	14,86	6,28	126,4	123,77	40,01	92,84	79,71	53,37
Te Papa	OR.18906	parkinsoni	11,75	48,71	9,69	12,18	32,17	128,93	27,61	14,82	6,6	128,48	125,76	41,04	94,61	82,23	54,66
Te Papa	OR.17258	parkinsoni	11,55		8,93	10,59	33,42	132,37	28,3	14,53	6,12	133,52	130,21	44,61	95,46	83,55	53,96
Te Papa	OR.793-S	parkinsoni	10,83	53,18	10,03	11,86				15,52	7,03	136,73	133,23				57,6
Te Papa	OR.25953a	parkinsoni	10,68	53,98	10,37	11,2	33,17	129,72	28,19	14,68	6,11	130,88	128,07	42,9
Te Papa	OR.19283	parkinsoni	11,08	46,69	9,14	11,8	32,18	129,47	27,7	14,39	6,44	131,05	128,28	41,83	94,79	82,81	55,51
Te Papa	OR.18969	parkinsoni	11,32	46,71	9,25	10,73	32,29	127,58	27,17	14,67	6,45	126,19	123,09	41,96	92	80,3	54,23
Te Papa	OR.27276a	parkinsoni			10,29	13,07	33,73	131,52	28,09	15,05	6,49	130,8	128,6	43,5	96,18	83,65	54,83
Te Papa	OR.24247	parkinsoni	11,02	46,78	10,25	11,98	32,96	129,1	28,59	14,58	6,41	127,66	125,19
Te Papa	OR.27119a	parkinsoni					34,75	135,82	29,56	15,2	6,55	138,32	134,62	43,78
Te Papa	OR.29637	parkinsoni	12,33	49,56	10,36	11,74	34,67	130,58	29,08	15,9	6,88	131,64	127,73	43,18	95,58	83,09	54,22
Te Papa	OR.18905	parkinsoni	12,5	49,47	9,25	10,82	33,33	128,67	29,19	14,81	6,68	129,48	126,58	41,19	96,64	83,62	54,22
Te Papa	OR.19282	parkinsoni	11,79	48,63	10,92	12,59	32,1	127,3	27,91	15,3	6,78	130,04	127,26	37,73	90,96	78,16	52,12
Te Papa	OR.18968	parkinsoni	11,67	47,84	9,25	13,17	32,28	125,09	27,09	15,14	6,96	127,37	123,74	40,57	91,3	78,7	50,94
Te Papa	OR.19298	parkinsoni	11,74	50,16	9,88	11,8	34,76	130,41	28,45	14,96	6,41	130,57	126,88	42,02	93,57	81,01	54,42
Te Papa	OR.19781	parkinsoni	11,01	52,08	9,71	12,4	34,29	134,66	28,06	15,23	6,3	135,51	131,65	44,05	97,76	85,24	55,25
Te Papa	OR.24236	parkinsoni	10,92	49,72	10,04	12,98	34,57	131,55	27,95	15,2	6,69	132,88	129,15	42,76	96,32	83,87	54,9
Te Papa	OR.289617	parkinsoni	11,79	49,62	10,06	13,38	34,96	133,72	28,91	15,8	6,21	135,76	133,66	45,39	100,45	87,06	57,68
Te Papa	OR.21422	westlandica	14,69	60,88	12,87	14,24	40,4	149,98	32,54	17,18	7,49	148,43	144,33				63,92
Te Papa	OR.22084	westlandica	14,63	58,26	11,82	14,72	38,95	144,04	30,86	16,33	7,14	143,99	141,29
Te Papa	OR.21423	westlandica	13,93		12,15	15,61	39,45	149,9	34,12		7,31	149,06	144,37	51,89		98,08	64,38
Te Papa	OR.21424	westlandica	15,9	59,24	11,73	14,61	38,91	149,5		17,2	7,4	150,2	146,57	51,52		99,6	63,46
Te Papa	OR.21106	westlandica	14,35	58,56	12,29	15,05	39,89	151,68	33,1	17,21	8,27	152,82				96,35
Te Papa	OR.21094	westlandica					38,82	155	32,91	17,1	7,09	150,94	147,81	51,46		97,62	64,66
Te Papa	OR.24272	westlandica	15,26		12,64	17,03	40,63	150,21	32,9	17,46	7,94	148,28	145,25	50,2	112,52	97,43	64,09
Te Papa	OR.11494	westlandica	14,44
Te Papa	OR.838-S	westlandica					38,61	150,94	33,3	17,3	7,73	148,63	145,69		111,4	96,6	62,5
Te Papa	OR.15900	westlandica	13,84		12,24	15,73	39,91	146,4	32,22	17,02	7,38	146,7	142,16	50,38	109,22	93,83	60,87
Te Papa	OR.599-S	westlandica					38,6		32,61					50,56
Te Papa	OR.11511	westlandica	12,59	56,32	10,94	14,32	37,95	144,98	32	16,52	6,98	147,46	143,7	50,93	111,04	96,07	63,1
Te Papa	OR.22968	westlandica						151,72	31,87	16,96	7,52	158	155		114,99	99,37	66,13
Te Papa	OR.22933	westlandica	14,4	57,49	10,92	13,51		148,52	32,04	16,97	6,81	151,83	147,81	49,38	112,81	98,38	63,15
Te Papa	OR.22085	westlandica	13,27		11,34	13,32	38,53	150,38	31,87	16,9	7	151,13	146,9	51	112,29	95,52	63,54
Te Papa	OR.23766	westlandica	14,32	59,27	12,38	11,49		149,35	32,5	17,43	7,54	153,5	151,19	50,02	109,16	93,94	62,31
Te Papa	OR.28672	westlandica	14,39	62,2	12,1	13,98	40,19	151,29	31,48	17,3	7,07	151,27	147,12	51,32	112,42	97,88	64,85
Te Papa	OR.23155	westlandica	14,24		12,65	13,73		150,31	32,62	16,75	7,39	151,74	146,7		111,73	97,53
Te Papa	OR.23151	westlandica	13,45	57,41	11,1	14,39		142,7	31,15	15,85	7,2	144,87	140,85		108,15	92,64	59,82
Te Papa	OR.19299	westlandica	14,97	61,78	13,04	14,98	38,66	152	31,95	17,27	7,14	150	148	51,88	114,52	99,74	64,34
Te Papa	OR.21091	westlandica	13,6	60,13	11,4	15,34	38,9	148,14	31,37	16,74	7,49	150,47	145,83	50,36	109,88	95,91	63,46
Te Papa	OR.19281	westlandica	14,95	61,08	12,84	15,78	38,92	146,9	34,5	17,36	8,04	145,8	142,33	51,52	110,85	95,91	63,19
Te Papa	OR.19311	westlandica	14,96	60,86	13,38	14,69	39,92	152,82	33,45	17,78	7,42	150,84	147,45	52,58	116,27	100,37	67,33
Te Papa	OR.21499	westlandica	14,97	57,47	11,96	14,41	36,94	149,63	30,47	16,28	7,32	152,85	149,63	51,92	112,35	97,74	65,09
Te Papa	OR.21466	westlandica	15,42	62,58	13,17	14,85	38,35	150,21	32,38	17,14	7,58	148,45	145,12	52,8	112,42	97,25	63,9
Te Papa	OR.21099	westlandica	14,87	60,68	11,37	15	38,83	146,07	32,74	17,47	7,6	146,33	143,21	49,67	107,42	93,27	62,57
Te Papa	OR.21092	westlandica	14,37	57,7	12,07	16,3	37,21	150,95	31,87	17,31	7,65	152,75	148,88	52,78	115,58	100,13	65,62
Te Papa	OR.21093	westlandica	14,84	58,53	12,11	14,81	39,3	148,58	31,75	17,43	7,52	148,02	145,12	50,42	111,04	96,17	63
Te Papa	OR.27314	westlandica	14,49	60,97	12,49	14,3	39,76	148,35	33,27	17,48	7,48	147,37	144,45	50,24	113,19	97,7	64,07
Te Papa	OR.27409	westlandica	14,04	57,54	12,52	14,4	38,85	152,17	31,33	16,88	7,83	152,35	149,18	47,74	109,79	94,75	61,36
Te Papa	OR.24271	westlandica	13,92	60,97	11,7	13,51	40,33	157	33,91	17,45	7,53	156	151,83	49,88	114,59	98,76	64,61
Te Papa	OR.24244	westlandica	13,72	56,51	11,78	14,94	39,35	148,21	32,4	17,11	7,13	146,21	142,19	49,62	114,13	98,05	63,66
Te Papa	OR.13566	westlandica	13,3	63,3	12,06	16,24	38,26	152,45	32,83	17,59	7,65	153,03	150,02	49,9	110,27	96,15	65,62
Te Papa	OR.13594	westlandica	14,13	58,89	11,86	14,8	38,97	150,84	32,72	17,13	7,4	151,4	146,62	52,81	113,71	98,02	63,32
Te Papa	OR.22668	westlandica	13,79	57,8	11,62	14,92	39,47	149,63	31,57	17,06	7,2	149,77	146,76	50,04	109,5	94,71	62,96
Te Papa	OR.23734	westlandica	15,25	61,09	12,25	14,61	39,02	143,61	31,89	16,24	7,37	146,68	143,34	50,11	109,15	94,88	61,38
Te Papa	OR.23725	westlandica	14,35	61,73	11,95	14,1	39,4	151,45	33,42	17,43	7,7	152,58	149,2	51,29	111,96	97,75	64,26
Te Papa	OR.23121	westlandica	14,86	58,93	12,22	14,66	38,94	149,41	31,41	17,45	7,58	149,45	147,13	50,76	111,12	97,11	63,18
Te Papa	OR.23306	westlandica	15,51	60,68	13,21	15,53	40,9	151,91	32,32	18,13	7,36	147,37	144,96	53,01	114,13	97,79	64,23

APPENDIX 2

Means (solid symbols) and standard errors (error bars) of each of the 15 skeletal measurements used for the fossil description for each Procellaria species. Open symbols represent the individual measurements. Colours represent different species: yellow = P. westlandica; blue = P. conspicillata; green = P. aequinoctialis; red = P. altirostris sp. nov.; orange = P. cinerea; pink = P. parkinsoni.

Edited by

Edited by: Luís Fábio Silveira

Publication Dates

• Publication in this collection
  12 Feb 2021
• Date of issue
  2021

History

• Received
  06 Oct 2020
• Accepted
  13 Nov 2020
• Published
  29 Jan 2021