© The Author, 2010. Journal compilation © Australian Museum, Sydney, 2010 Records of the Australian Museum (2010) Vol. 62: 115-122. ISSN 0067-1975 doi: 10.3853/j.0067-1975.62.2010.1535 Regional Comparisons of the Thickness of Moa Eggshell Fragments (Aves: Dinornithiformes) B J. Gill Auckland War Memorial Museum, Private Bag 92018, Auckland, New Zealand bgill @ aucklandmuseum.com Abstract. Samples of moa eggshell fragments from eight sites throughout New Zealand were measured to investigate the usefulness of the graphed distribution of eggshell-thicknesses in reflecting the moa fauna of the site. Assuming larger moa species laid eggs with thicker eggshells, a frequency histogram of eggshell thicknesses for each site seems to mirror what is known (from bones) of the incidence and relative abundance of large and small moa species at the site. This is particularly so for North Island sites which had lower moa diversity than South Island sites. At North Cape and Tokerau Beach abundant thin eggshell (mode at 0.90-0.94 mm) was probably produced by Euryapteryx “curtus-gravis” and Pachyornis geranoides, and rarer thick eggshell (mostly 1.2-1.7 mm) by Dinornis novaezealandiae. At both Puketitiri and Castle Point there were broadly unimodal distributions of thin to medium-thickness eggshell, with thick eggshell almost absent. At Puketitiri the eggshell is assumed to be mainly from Anomalopteryx didiformis, and the slightly smaller P. geranoides, and averages thicker with a broader range than at Castle Point where the assumed identity of the eggshell lies with two small species (P. geranoides and Eu. “curtus-gravis"). At the four South Island sites the correlation to species is less clear. The modal thicknesses at Wairau Bar, Oamaru, Chatto Creek and Shag River are all in the range 1.15-1.44 mm and probably largely attributable to Eu. “curtus-gravis” which has a large form in the South Island and dominates the bones at all four sites. However, several other moas could have contributed the thinnest and thickest eggshells in most of the South Island samples. Archaeological sites had similar large ranges of eggshell-thickness to natural sites, suggesting that Maoris collected moa eggs from all available species and not just the largest ones. The study demonstrates the usefulness of eggshell-thickness histograms at particular sites as an adjunct to, or surrogate for, information on the relative abundance of moa bones, especially for North Island sites. Gill, B.J., 2010. Regional comparisons of the thickness of moa eggshell fragments (Aves: Dinornithiformes). In Proceedings of the VII International Meeting of the Society of Avian Paleontology and Evolution, ed. W.E. Boles and T.H. Worthy. Records of the Australian Museum 62(1): 115-122. Fossil bones and eggshell fragments are the main evidence Zealand. However, there has been little research on the nature of the former presence in New Zealand of moas (Dinomithi- and characteristics of moa eggshell fragments, and reports of formes), the extinct large ratite birds currently thought to archaeological and paleontological excavations typically and number 10 species (nomenclature after Worthy & Holdaway, unhelpfully record “moa eggshell”, without further analysis. 2002; Bunce et al., 2003; Worthy, 2005). Moa eggshell One aim of my recent work on moa eggs and eggshell (Gill, fragments are common, often in large quantities, in various 2000, 2006, 2007) has been to seek a better understanding archaeological and Holocene fossil sites throughout New of unassociated, broken moa eggshell. 116 Records of the Australian Museum (2010) Vol. 62 Materials and methods Sites. I examined samples of moa eggshell fragments at eight sites or geographic areas chosen because they had yielded large samples held in museum collections, and the sites were spread latitudinally throughout New Zealand (Fig. 1). Five are natural sites, known (or assumed) to be of Holocene age, i.e. no older than 10,000 years before present. Three archaeological sites, all in the South Island, are assumed to date from within 100-200 years of first human settlement of New Zealand (which was ca. 1250 A.D.; Anderson, 1991; Wilmshurst et al., 2008). The sites were as follows (see Table 3 for latitudes and sample sizes; see Appendix for museum registration numbers): • North Cape, Northland (natural sites) • Tokerau Beach, Northland (natural sites; same sample analysed in Gill, 2000) • Puketitiri, Hawke’s Bay (natural sites) • Castle Point region, Wairarapa Coast (natural sites) • Wairau Bar, Marlborough (archaeological site) • Oamaru, North Otago (archaeological site) • Chatto Creek, Central Otago (natural sites) • Shag River mouth, Otago (archaeological site) Samples. Eggshell fragments from the selected sites were examined closely and any with significant signs of surface-wear were rejected. For each acceptable fragment (n = 3565), one measurement of eggshell thickness was made to the nearest 0.01 mm (with vernier callipers or a screw micrometer) at an undamaged point on the edge of the fragment. Thicknesses were plotted as histograms with £ 0.05 mm intervals to give a very finely-divided spread of thickness classes (0.50-0.54,0.55-0.59,0.60-0.64 etc.). All Figure 1. Locality map of New Zealand showing eight sites (four fragments were white (or shades close to white) except for from each of North and South Islands) for which samples of moa six of the 79 fragments in OM AV7477, which were green. eggshell fragments were measured in this study. Archaeological Green eggshell belongs to Megalapteryx didinus (see Gill, sites are marked with an asterisk (*). 2007), but the identity of white eggshell is open to conjecture. Moa faunas. The moa faunas at each site are known from the identity and relative abundance of bones. The taxonomy and nomenclature of moas is still under review. Bunce et al. It is well known that moa eggshell fragments vary in (2003) showed that instead of three species of giant moas thickness, and differences between “thick” eggshell (ca. Dinornis throughout New Zealand there is just one North 1.3-1.7 mm) and “thin” eggshell (ca. 0.5-1.0 mm) were Island species (D. novaezealandiae) and one South Island noted earlier (e.g., Archey, 1931). Thicker eggshell fragments species (D. robustus). This has simplified the moa fauna at presumably derive from larger eggs that were produced by most sites. Worthy (2005) showed that the stout-legged moa, the larger species of moas. Support for this comes from a previously called Euryapteryx geranoides, should now be general positive correlation between the lengths of whole called Eu. gravis, and that Mappin’s moa Pachyomis mappini moa eggs and their eggshell thicknesses (Gill, 2007: fig. 5). is now P. geranoides. The moa fauna in a given region is a subset of the known It is also likely that Eu. gravis (North and South Islands) species, and it follows that the thicknesses of the moa and Eu. curtus (North Island only) are conspecific, with eggshell fragments at a fossil site should reflect the sizes, size-differences in the bone samples reflecting various and thus identities, of the resident species of moa. The combinations of temporal, geographic and sexual variation relative abundance of eggshell of different thicknesses at a (Tennyson & Martinson, 2006: 36, 146; T.H. Worthy, pers. site might also reflect the relative abundance of the species comm. 2008). For example, Eu. gravis is recorded from that produced them. Tokerau Beach but such bones probably belonged to the The aim of this study was to analyze the thickness of moa biggest of the larger (female) individuals of Euryapteryx eggshell fragments at a range of sites and look for agreement at that site. Worthy (1987) showed that histograms of between the thickness profiles and the diversity and relative bone-lengths of Euryapteryx from Tokerau Beach were abundance of the moa species at the sites as determined all bimodal, and he considered the different size-classes from bones. It was a test of the notion that a histogram of to reflect sexual dimorphism. The Holocene bones of thicknesses taken from a large sample of eggshell fragments at Euryapteryx vary clinally in size, tending larger towards the a site gives a profile reflective of moa composition at the site. south (Worthy, 1987). The two nominal species are referred Gill: Moa eggshell thickness 117 Table 1. Presumed relative abundance of moa taxa (based on bones) at North Island sites where eggshell thickness was assessed. ••• predominant, •• present, • rare, — absent. Euryapteryx P. geranoides A. didiformis D. novaezealandiae North Cape ••• •• — • Tokerau Beach ••• • — • Puketitiri — •• ••• • Castle Point ••• ••• •• •• Table 2. Presumed relative abundance of moa taxa (based on bones) at South Island sites where eggshell thickness was assessed. ••• predominant, •• present, • rare, — absent. Euryapteryx P. elephantopus A. didiformis E. crassus M. didinus D. robustus Wairau Bar ••• • • • • • • Oamaru ••• • • ? • • ? ? Chatto Creek • •• • • — • • • • • • Shag River • •• • — • • — • • to here together as Eu. “curtus-gravis”. Tables 1 and 2, and Castle Point. Pachyornis geranoides and Eu. “curtus- the following notes, summarize the Holocene moa faunas gravis” were co-dominant in the area with lesser numbers at the sites in this study. of A. didiformis and D. novaezealandiae (T.H. Worthy, pers. comm. 2008). The moa fauna was therefore very like that North Cape. The most abundant moas were Euryapteryx at North Cape. “curtus-gravis ” and Pachyornis geranoides, while Dinornis novaezealandiae was rarer (Atkinson & Millener, 1991). Wairau Bar. Euryapteryx “curtus-gravis” predominated Evidence from bones is that P. geranoides was nearly as with lesser numbers of Emeus crassus and rare examples common as Em. “curtus-gravis”. of A. didiformis, Megalapteryx didinus, Pachyornis elephantopus and Dinornis robustus (Scofield et al., 2003). Tokerau Beach. About 95% of skeletons in the area are Eu. “curtus-gravis” (Worthy & Holdaway, 2002: 184). Rarer Oamaru. Euryapteryx “curtus-gravis” dominated and P. species at the site (evidence discussed by Gill, 2000) were elephantopus was much less common (Worthy & Holdaway, P. geranoides and D. novaezealandiae. 2002: 181). Emeus crassus was present (Trotter, 1970). Puketitiri. Anomalopteryx didiformis was the main moa Chatto Creek. In the Central Otago area Eu. “curtus- found at caves and rock-shelters of inland Hawke’s Bay, gravis ” was the dominant moa, with lesser numbers of P. closely followed by P. geranoides, with D. novaezealandiae elephantopus, Em. crassus, M. didinus and D. robustus rarer (Worthy & Holdaway, 2000). (Worthy, 1998). Shag River. Euryapteryx “curtus-gravis ” was the dominant moa with lesser numbers of Em. crassus and D. robustus (Anderson et al., 1996). Results Table 3 summarizes thickness for the samples of eggshell fragments at the eight sites in this study, and for whole moa eggs, and shows that minimum, mean and maximum thicknesses tend to be higher for South Island than for North Island samples. Fig. 2 shows the relationship between mean thickness and latitude for the eight broken eggshell samples (open circles). The eggshell thicknesses of whole eggs (Fig. 2; closed circles) show a very similar regression to the eggshell fragments, though data for the North Island are few. Figures 3-10 show the histograms of moa eggshell thickness for each of the eight sites. North Cape (Fig. 3) and Tokerau Beach (Fig. 4) show a similar pattern—a broadly Figure 2. Regressions of mean eggshell thickness on latitude for bimodal distribution with a spread of numerous thin eggshell the eight geographic samples in this study (Table 3) and for the 20 fragments (mode at 0.90-0.94 mm) and a second spread of whole moa eggs for which measurements of eggshell thickness are rarer thicker fragments (mostly 1.2-1.7 mm thick). The thin possible (Gill, 2007). For eggshell fragments (open circles), the eggshell was presumably produced by Euryapteryx “curtus- four North Island data-points are at bottom left and the four South Island ones at middle right. The regression lines and correlation gravis ” with a lesser contribution by Pachyornis geranoides. coefficients are: y = 0.028x + 0.027, r = 0.75 (fragments; lower Both are small moas, and the former is particularly small in line); y = 0.034x-0.17, r = 0.44 (whole eggs; upper line). the north (Worthy, 1987). The latter species would have had a 118 Records of the Australian Museum (2010) Vol. 62 ) % ( y c n e u q e r F ocoomomomom looiootoomomoiooiooio «5iO<0(£)NNffl(B©0) oooooooooo 18 16 Tokerau Beach 14 ) % ( 12 y c 10 n e 8 u q e 6 r F 4 2 n n^nrHI hnnn 0 J3. oO tOn oi— OCM IPTi) oCO CLTOJ 0'l1 0 0 0ifl (0D I0D 0h h- COO tCoO OOOOOOOOOO^— T— T— T— T— T— T— T— T— T— T— T— T— is 16 Puketitiri 14 ) % ( 12 y c 10 n ue 8 q e 6 r F 4 2 a 0 l~l r~i l~~l OiotiOntOQltODOr--tOr'-OcotOeoOctnOo OCO rO- TUO OM (OM OO OO 0^¥^*0i0o1io0o0(1oi0^0-r1--0o0o*o0o oooooooooo ) % ( y c n e u q e r F OIOOLOOIOOIOOLD uDOLOoiooiooLOOLnoiooiooio ddifi dtO dCO dS dN d03 d0O d(T) dCJ) Di-T-NNfOco^'tinifiiDiOKsmco Thickness (mm) Figures 3-6. Histograms of moa eggshell thicknesses for samples of (Fig. 3) 612 fragments from North Cape, Northland; (Fig. 4) 1042 fragments from Tokerau Beach, Northland; (Fig. 5) 273 fragments from Puketitiri, Hawke’s Bay; and (Fig. 6) 431 fragments from Castle Point, Wairarapa. Gill: Moa eggshell thickness 119 ) % ( y c n e u q e r F ) % ( y c n e u q e r F O O O O O O O O O O T“ T- T“ T“ T~ T” T“ T—' T—¥ T-' 1— T-' T- T~ T~ T~ ) % ( y c n e u q e r F LnoinoLooiDoiootooLDomoLno LO O LO ¥ mo looto^i^coojaiffioqrrpjcyjcinii; ^ lo m (O ID N OOOOOOOOOt~t— T-'r-T-T-i—¥i—''—¥ ) % ( y c n e u q e r F Thickness (mm) Figures 7-10. Histograms of moa eggshell thicknesses for samples of (Fig. 7) 340 fragments from the Wairau Bar archaeological site, Marlborough; (Fig. 8) 595 fragments from the Oamaru archaeological site; (Fig. 9) 147 fragments from Chatto Creek, Central Otago; and (Fig. 10) 125 fragments from the Shag River archaeological site, Otago. 120 Records of the Australian Museum (2010) Vol. 62 Table 3. Latitude (°S) and descriptive statistics for eggshell thickness (mm) for samples of moa eggshell fragments from eight sites around New Zealand (Fig. 1). In brackets after range is the arithmetic extent of the range (mm), i.e. maximum minus minimum. Maximum and minimum eggshell thicknesses (mm) for the North and South Islands (overall) are shown for the samples of eggshell fragments in this study, and for the whole eggs for which eggshell thicknesses can be measured (Gill, 2007). locality mean n s.d. range latitude North Island North Cape 1.08 612 0.287 0.54-1.71 (1.17) 34.4 Tokerau Beach 0.96 1042 0.210 0.56-1.69(1.13) 34.9 Puketitiri 1.08 273 0.129 0.65-1.41 (0.76) 39.3 Castle Point 0.97 431 0.109 0.70-1.48 (0.78) 41.0 fragments (overall) 0.54-1.71 whole eggs 0.90-1.50 South Island Wairau Bar 1.31 340 0.188 0.66-1.82(1.16) 41.5 Oamaru 1.21 595 0.156 0.77-1.73 (0.96) 45.1 Chatto Creek 1.37 147 0.161 0.89-1.65 (0.76) 45.1 Shag River 1.35 125 0.167 0.83-1.69 (0.86) 45.5 fragments (overall) 0.66-1.82 whole eggs 1.02-1.89 minor contribution at Tokerau Beach and a greater contribution A few fragments from Chatto Creek (4%) were green, i.e. at North Cape, according to the relative abundance of bones attributable to M. didinus, and all were 0.89-1.13 mm thick, at these sites, but the eggshells of the two species seem placing them at the very left-hand tail of the histogram (Fig. inseparable in the histograms. The thick eggshell at the 9). White eggshell in that thickness range was also present, two northern sites (Figs. 3, 4) was presumably produced by which at Chatto Creek could be Megalapteryx eggshell Dinornis novaezealandiae, the only large moa present. that has faded to white (see Gill, 2007), or attributable to Puketitiri (Fig. 5) and Castle Point (Fig. 6) show E. crassus. broadly unimodal thickness histograms of thin to medium¬ North Cape and Tokerau Beach have larger ranges of thickness eggshell, with thick eggshell almost absent. At eggshell thicknesses (1.17 mm, 1.13 mm; Table 3) than all Puketitiri the eggshell is assumed to be mainly from the other sites except Wairau Bar (1.16 mm; Table 3). However, small species Anomalopteryx didiformis and the even the thickness range at Wairau is exaggerated by one outlying smaller P. geranoides. At Castle Point the assumption is thin fragment in the 0.65-0.69 mm range (Fig. 7) which that the eggshell is a mixed sample attributable largely to P. could conceivably belong to a large bird other than a moa. geranoides and Eu. “curtus-gravis”, as at North Cape and Tokerau Beach. Indeed, the frequency distribution at Castle Discussion Point, with a mode at about 1.0 mm, is very like that of thin eggshell at the two northern-most sites. Anomalopteryx The latitudinal trend in moa eggshell thickness, with thicker didiformis probably had thicker eggshell than P. geranoides, eggshells in the south (Fig. 2), has two probable explanations. being larger, and this may be reflected in the thicker eggshell One underlying factor will be the presence of certain large overall at Puketitiri than at Castle Point. Some of the thickest moa species in the South Island (most notably Pachyornis eggshell at Puketitiri, and the few pieces in the 1.40-1.49 elephantopus) that are absent in the North Island. However, mm range at Castle Point, may belong to Dinornis. Bergmann’s Rule may also be involved. For New Zealand The four South Island samples are all broadly unimodal birds, this predicts that where a species varies geographically and towards the thicker end of the spectrum. Very thin in size, individuals in cooler (e.g., southern) areas will be eggshell is rare. The modal thicknesses at Wairau Bar (Fig. larger. Bergmann’s Rule is thought to have operated in 7), Chatto Creek (Fig. 9) and Shag River (Fig. 10) are all at Holocene populations of various moas, e.g., Euryapteryx 1.35-1.44 mm. Oamaru(Fig. 8) has a lower mode (1.15-1.19 (Worthy, 1987, 1992) and Dinornis (Worthy et al., 2005), mm) but greater modal thickness than for the two North whose populations are present in both islands. If the birds of Island unimodal samples (Puketitiri and Castle Point). Eu. these species varied clinally in body-size, then so too may “curtus-gravis” dominated the bones at all four South Island the size of their eggs and the thickness of their eggshells. sites, and much of the eggshell presumably belongs to this In general, the histograms of moa eggshell thickness seem moa, which is large-statured in the South Island. to mirror closely what is known of the diversity and relative However, the South Island situation is compounded by abundance of moas at the sites. For example, the bimodally a greater diversity of moas at each site (usually 4-6 spp.; distributed thick and thin eggshell at the two northern-most Table 2) than in the North Island (3-4 spp.; Table 1). Emeus sites (Figs 3, 4) seem to fit with the indication from bones crassus, Anomalopteryx didiformis and Megalapteryx of numerous small moas (.Euryapteryx “curtus-gravis ” and didinus could have contributed the thinnest eggshells in most Pachyornis geranoides) and rarer large moas (Dinornis of the South Island samples, and Pachyornis elephantopus novaezealandiae). The extreme spread of eggshell thicknesses and Dinornis robustus were present at most sites to contribute at each of the two northernmost sites must reflect the presence the thickest eggshell fragments. there of moas of extreme sizes—the smallest known members Gill: Moa eggshell thickness 121 of the Euryapteryx “curtus-gravis ” complex and a Dinornis. didinus. Three whole green eggs, two from Chatto Creek The situation is now much simpler and clearer than when three (Gill, 2006: Eggs 22 and 23) and one from Mt Aspiring species of Dinornis and two of Eury apteryx were thought to National Park (Egg 29), had similarly thin eggshell (1.1-1.2 occur at Tokerau Beach (Gill, 2000). mm thick; Gill, 2007). A whole egg from the Shag River site We can infer from the histograms that both Euryapteryx has an eggshell-thickness of 1.73-1.89 mm (Gill, 2007: Egg “curtus-gravis” and Pachyornis geranoides had eggshell 27), the thickest moa eggshell ever recorded, and thought roughly 0.5-1.2 mm thick at North Cape (Fig. 3) and 0.6-1.1 to belong to P. elephantopus. This extreme thickness was mm at Tokerau Beach (Fig. 4). A whole egg from Tokerau not represented in the broken eggshell sample from that Beach thought to belong to Eu. “curtus-gravis ” (Gill, 2006: site (Fig. 10); the sample of 125 fragments was clearly not Egg 2) has eggshell about 0.9 mm thick (Archey, 1931), large enough. However, the Shag River sample has a big which places it at the mode in both thickness histograms peak at 1.60-1.64 mm, more so than for any other sample. and in the centre of the spread of thin eggshell fragments. These thickest fragments in the Shag River sample may be The same histograms suggest that the eggshell of Dinornis attributable to P. elephantopus. novaezealandiae was approximately 1.1-1.7 mm thick The archaeological eggshell samples at Wairau Bar, but there is no information from whole eggs on eggshell Oamaru and Shag River are presumably biased towards the thickness in this species. The largest North Island egg moa eggs that local Maoris collected and discarded. Yet the (Gill, 2006: Egg 5, Waitomo) is probably Dinornis but the range of eggshell thicknesses at these midden sites is very eggshell fragments are assembled in a way that prevents their large, and seems no different from the range of thicknesses thickness being measured. The thickest eggshell measured at natural sites (Table 3). This suggests that moa eggs of all from North Island whole eggs is only 1.5 mm (Table 3) and available sizes (i.e. from all available species) were collected was attributed to Anomalopteryx (Gill, 2006: Egg 8). for human use, not just the largest ones. We can predict from Fig. 5 (Puketitiri) that P. geranoides This study suggests a usefulness in histograms of moa had eggshell from about 0.7 mm thick to an unknown eggshell thicknesses generated for specific paleontological maximum, and that A. didiformis at the same site had eggshell and archaeological sites. Moa eggshell thickness profiles from an unknown minimum to 1.4 mm thick. No whole eggs seem broadly to reflect the moa faunas at the sites, and may of P. geranoides are known but four whole eggs from inland be a helpful adjunct to, or surrogate for, information on the North Island sites and most likely attributable to A. didiformis relative abundance of moa bones at sites. This is especially have data on eggshell thickness (Gill, 2006, 2007: Eggs 7, so in the North Island where moas are usually less diverse 8, 13 and 14). The attribution of the eggs to Anomalopteryx at a site than in the South Island. is based on that small species dominating at the inland sites where the eggs were found and on the eggs being small (i.e. 152-175 mm long). Thirty-five eggshell thickness measurements from these four eggs had a range of 1.1-1.5 Acknowledgments. I thank the following curatorial staff for mm. This supports the notion that at least the thicker eggshell access to moa eggshell fragments in their collections: Neville at Puketitiri belonged to A. didiformis. By elimination, much Hudson (Geology Department, Auckland University); Heather of the thinner eggshell at Puketitiri, especially in the range Sadler and Ian Smith (UO); Paul Scofield (CMC); Daniel McKnight 0.7-1.0 mm, must belong to P. geranoides. At Castle Point, and Kaaren Mitcalfe (HBM); Alan Tennyson (NMNZ); and Ilka where the small and similarly-sized P. geranoides and Eu. Sohle and Abigail Blair (OM). The New Zealand Lottery Grants “curtus-gravis” dominated (on bone evidence), most of the Board financially supported field trips to Northland dune-sites in eggshell was indeed in that range (Fig. 6). the 1990s, and Fred Brook provided transport and companionship Given that the large South Island form of Eu. “curtus- on some trips. Trevor Worthy and two referees (Alan Tennyson and gravis” dominated the bones at all four South Island sites, another) gave helpful comments on drafts of the paper. the frequency bars in the histograms suggest that its eggshell was about 1.1-1.5 mm thick at Wairau Bar (Fig. 7), 1.0-1.3 References mm at Oamaru (Fig. 8), 1.3-1.5 mm at Chatto Creek (Fig. 9) and 1.2-1.6 mm at Shag River (Fig. 10), or roughly 1.0-1.6 Anderson, A., 1991. The chronology of colonization in New Zealand. Antiquity 65: 767-795. mm for the South Island in general. Considering whole eggs, Anderson, A., T. Worthy & R. McGovern-Wilson, 1996. Chapter there are seven South Island eggs possibly attributable to 14. Moa remains and taphonomy. Shag River Mouth. In Euryapteryx (Gill, 2006,2007): Eggs 16 and 18-21 from the The Archaeology of an Early Southern Maori Village, ed. A. same Wairau Bar site as the eggshell fragments (Fig. 7) and Anderson, B. Allingham, and I. Smith, pp. 200-213. Canberra: Eggs 11 and 32 from the same Oamaru site as in Fig. 8. These ANH Publications. eggs are not objectively linked to Euryapteryx, but they are Archey, G., 1931. Notes on sub-fossil bird remains. Records of the medium-sized (i.e. 194-215 mm long) and assumed to belong Auckland Institute and Museum 1(2): 113-121. to the medium-sized moa that dominates bones at the sites. Archey, G., 1941. The Moa. A Study of the Dinornithiformes. Fifty-three thickness measurements from these seven eggs had Auckland: Auckland Museum. a range of 1.1-1.7 mm. This close agreement in thickness Atkinson, I.A.E., & PR. Millener, 1991. An ornithological glimpse into New Zealand’s pre-human past. Proceedings of the 20th between the broken eggshell and whole eggshells reinforces International Ornithological Congress 1: 129-192. the conjecture that the bulk of the eggshell fragments at Bunce, M., T.H. Worthy, T. Ford, W. Hoppitt, E. Willerslev, A. Wairau Bar, Oamaru, Chatto Creek and Shag River are from Drummond, & A. Cooper, 2003. Extreme reversed sexual size medium-sized eggs that were probably Euryapteryx. dimorphism in the extinct New Zealand moa Dinornis. Nature Green eggshell fragments at Chatto Creek were thin 425: 172-175. (0.89-1.13 mm) and signal the presence of Megalapteryx http://dx.doi. org/10.1038/nature01871 122 Records of the Australian Museum (2010) Vol. 62 Gill, B.J., 2000. Morphometries of moa eggshell fragments (Aves: Worthy, T.H., 1992. A re-examination of the species Euryapteryx Dinornithiformes) from Late Holocene dune-sands of the geranoides (Owen) including comparisons with other emeiin Karikari Peninsula, New Zealand. Journal of the Royal Society moas (Aves: Dinornithiformes). Journal of the Royal Society of New Zealand 30(2): 131-145. of New Zealand 22(1): 19-40. Gill, B J., 2006. A catalogue of moa eggs (Aves: Dinornithiformes). Worthy, T.H., 1998. Quaternary fossil faunas of Otago, South Island, Records of the Auckland Museum 43: 55-80. New Zealand. Journal of the Royal Society of New Zealand Gill, B.J., 2007. Eggshell characteristics of moa eggs (Aves: 28(3): 421-521. Dinornithiformes). Journal of the Royal Society of New Zealand Worthy, T.H., 2005. Rediscovery of the types of Dinornis curtus 37(4): 139-150. Owen and Palapteryx geranoides Owen, with a new synonymy Scofield, R, T.H. Worthy, & H. Schlumpf, 2003. What birds were (Aves: Dinornithiformes). Tuhinga: Records of the Museum of New Zealand’s first people eating?—Wairau Bar’s avian remains New Zealand Te Papa Tongarewa 16: 33^-3. re-examined. Records of the Canterbury Museum 17: 17-35. Worthy, T.H., M. Bunce, A. Cooper, & R.P. Scofield, 2005. Tennyson, A., & P. Martinson, 2006. Extinct Birds of New Zealand. Dinornis—an insular oddity, a taxonomic conundrum reviewed. Wellington: Te Papa Press. In Proceedings of the International Symposium “Insular Trotter, M.M., 1970. North Otago archaeological sites. Part 2. New Vertebrate Evolution: The Palaeontological Approach ”, ed. J.A. Zealand Archaeological Association Newsletter 13(3): 135-142. AlcoverandP. Bover,pp. 377-390. Mallorca: Societatd’Historia Wilmshurst, J. M., A.J. Anderson, T.F.G. Higham, & T.H. Worthy, Natural de les Balears. 2008. Dating the late prehistoric dispersal of Polynesians to New Worthy, T.H., & R.N. Holdaway, 2000. Terrestrial fossil vertebrate Zealand using the commensal Pacific rat. Proceedings of the faunas from inland Hawke’s Bay, North Island, New Zealand. National Academy of Sciences of the United States of America Part 1. Records of the Canterbury Museum 14: 89-154. 105(22): 7676-7680. Worthy, T.H., & R.N. Holdaway, 2002. The Lost World of the Moa. http://dx.doi.org/10.1073/pnas.0801507105 Prehistoric Life of New Zealand. Bloomington, Indiana: Indiana Worthy, T.H., 1987. Sexual dimorphism and temporal variation in University Press. the North Island moa species Euryapteryx curtus (Owen) and Pachyornis mappini Archey. National Museum of New Zealand Records 3(6): 59-70. Appendix Museum registration numbers of the samples from which moa eggshell fragments were measured for thickness (number of fragments measured shown in brackets). Where the number measured was not the entire sample, fragments were chosen at random. Museum codes: UO, Anthropology Department, University of Otago, Dunedin; AIM, Auckland Museum, Auckland; CMC, Canterbury Museum, Christchurch; HBM, Hawke’s Bay Museum, Napier; NMNZ, Museum of New Zealand Te Papa Tongarewa, Wellington; OM, Otago Museum, Dunedin. North Cape (24 samples, 612 fragments). Samples collected 1976-1999 by F.J. Brook, N. Douglas, B.J. Gill, R. Renwick and H. Seelye.—Tom Bowling Bay (22 samples, 591 fragments). AIM LB6774 (1), LB6775 (7), LB7930 (81), LB7931 (39), LB7932 (27), LB7933 (16), LB7934 (16), LB7935 (31), LB7937 (30), LB8487 (23), LB8488 (4), LB8510 (65), LB8511 (4), LB8862 (5), LB8863 (29), LB9223 (8), LB9224 (1), LB9226 (27), LB9227 (37), LB9228 (34), LB9230 (39), LB9231 (61).—Waikuku Beach (2 samples, 21 fragments) AIM LB7936 (18), LB8825 (3). Tokerau Beach (97 samples, 1,042 fragments). See Gill (2000: Appendix 1). Puketitiri (5 samples, 273 fragments). Samples collected 1950s to 1961 by W.H. Hartree and J.C. Yaldwyn. In separate labelled containers at HBM; not registered or numbered. “Bush Face No. 1” (140), “Hukanui No. 7b 1959-1960” (25), “Hukanui No. 7b May 1960” (40), Hukanui No. 7b (cigarette tin) (60), Hukanui No. 7b (another cigarette tin) (8). Castle Point (11 samples, 431 fragments). Samples collected 1934-2000 by E. Barton, T. Cairns, I. Cameron, I. Dandermam, E. Smith and A.J.D. Tennyson.—Coast between Mataikona and Whakataki NMNZ S23166 (52), S23167 (15), S36733 (3), S37890 (20), S37910 (13), S37913 (92), S37923 (23), S38403 (97), S38441 (56), S38460 (10), S40676 (50). Wairau Bar (23 samples, 340 fragments). Archaeological excavations by J.R. Eyles, R. Duff, R. Perano and Canterbury Museum Archaeological Society 1942-1959. CMC AY19947 (4), AV19948 (50), AV19951 (20), AV19960 (25), AV19966 (27), AV19968 (7), AVI9971 (22), AV19974 (10), AV19978 (13), AV19979 (12), AV19981 (16), AV19986 (6), AV19987 (35), AV19990 (6), AV20038 (8), AV21128 (27), AV21129 (10), AV2U37 (5), AV21167 (28), AV25620 (1), AV25882 (1), AV29513 (4), AV29528 (3). Oamaru (23 samples, 595 fragments). Archaeological excavations by W.B.D. Mantell ca. 1852. The Mantell collection at AIM (LB4014) is a large assemblage of eggshell fragments believed to have been collected from a Maori midden site at Awamoa, near Oamaru, North Otago (Archey, 1941: 74). Mantell grouped the fragments on some basis. Within each group, numerous fragments are joined in an attempt to reconstruct sections of the original eggs, but thickness varies greatly within most of these groupings so they cannot represent separate individual eggs. Boxes A-D (20), Box E (1), Boxes F-G (2), Box H (56), Box I (23), Boxes J-K (20), Box L (17), Boxes M-N (23), Box O (21), Box P (17), Box Q (20), Box R (30), Box S (27), Box 4014/1 (56), Box 4014/2 (43), Box 4014/3 (92), Box 4014/4 (22). Chatto Creek (2 samples, 147 fragments). Samples collected ca. 1954 (collector not recorded). OM AV7376 (68), AV7477 (79). Shag River (2 samples, 125 fragments). Archaeological excavations led by A. Anderson 1988-1989. UO SM/B (23), SM/C (102).