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LEUCOCARBO CHALCONOTUS OTAGO SHAG GRACALUS CHALCONOTUS G. R. GRAY, 1845 Diagnosis. A species of Leucocarbo most closely related to L. stewarti and L. onslowi but distinguished from these species by the plumage characters and allometries outlined in Table 1. Distribution. Formerly the eastern South, NZ. Leucocarbo chalconotus bones have been recorded from Late Quaternary and archaeological deposits along the entire eastern coastline of South (e.g., Worthy, 1998 a; Smith, 2011). Now restricted to Otago from Lake Ki-Wainono to The Sisters (based on historical museum skins and modern specimens), with modern vagrants north to Banks Peninsula (see Fig. 1). Rare modern beach wrecks on Stewart (Rawlence et al., 2015)., Published as part of Nicolas J. Rawlence, R. Paul Scofield, Hamish G. Spencer, Chris Lalas, Luke J. Easton, Alan J. D. Tennyson, Mark Adams, Eric Pasquet, Cody Fraser, Jonathan M. Waters & Martyn Kennedy, 2016, Genetic and morphological evidence for two species of Leucocarbo shag (Aves, Pelecaniformes, Phalacrocoracidae) from southern South Island of New Zealand, pp. 676-694 in Zoological Journal of the Linnean Society 177 on page 687, DOI: 10.1111/zoj.12376, http://zenodo.org/record/270312, {"references":["Gray GR. 1845. Birds. In: Richardson JE, Gray GR, eds. The Zoology of the Voyage of H. M. S. Erebus and Terror, During the Years 1839 - 43. London: E. W. Janson, 1 - 20.","Worthy TH. 1998 a. A remarkable fossil and archaeological avifauna from Marfells Beach, Lake Grassmere, South Island, New Zealand. Records of the Canterbury Museum 12: 79 - 176.","Smith IWG. 2011. Estimating the magnitude of pre-European Maori marine harvest in two New Zealand study areas. New Zealand Aquatic Environment and Biodiversity Report No. 82.","Rawlence NJ, Kennedy M, Anderson CNK, Till CE, Smith I, Scofield RP, Tennyson AJD, Hamel J, Lalas C, Matisoo-Smith EA, Waters JM. 2015. Geographically contrasting biodiversity reductions in a widespread New Zealand seabird. Molecular Ecology 18: 4605 - 4616."]}

LEUCOCARBO CHALCONOTUS (G. R. GRAY, 1845) OTAGO SHAG. Graucalus auritus Lesson, 1831: G. R. Gray, 1843, in E. Dieffenbach, Travels in N. Z. 2: 201. Not Carbo auritus Lesson, 1831. Gracalus chalconotus G. R. Gray, 1845; in Richardson & J. E. Gray (eds), Zool. Voy. Erebus & Terror: Birds 1 (8): 20, pl. 21 ��� Otago Province, possibly Karitane (Scofield et al., 2012). Graculus glaucus ���Gray���: Reichenbach, 1850, Nov. Syn. Avium; pl. 49, fig. 2553 ��� Otago. Phalacrocorax glaucus Hombron & Jacquinot, 1853, in Dumont d���Urville, Voyage �� Sud, Zoologie 3: 127. Graculus chalconotus (G. R. Gray): G. R. Gray, 1862, Ibis 4: 252. Graculus glaucus (Hombron & Jacquinot, 1853): Bonaparte, 1855, Consp. Gen. Avium 2: 171. Phalacrocorax chalconotus (G. R. Gray): Buller, 1873, Hist. Birds N. Z. (1 st ed.), 334. Buller, 1888, Hist. Birds N. Z. (2 nd ed.) 2: 162. Phalacrocorax carunculatus (Gmelin): Buller, 1888, Hist. Birds N. Z. (2 nd ed.) 2: 155. In part. Phalacrocorax chalconotus (G. R. Gray): Buller, 1888, Hist. Birds N. Z. (2 nd ed.) 2: 162. Phalacrocorax glaucus Hombron & Jacquinot: Buller, 1888, Hist. Birds N. Z. (2 nd ed.) 2: 163. Phalacrocorax huttoni Buller, 1888: Hist. Birds N. Z. (2 nd ed.) 2: 174 ��� ocean beach near Dunedin, most likely Ocean Beach at Saint Clair, Dunedin (this study). Phalacrocorax chalconotus (G. R. Gray): Ogilvie- Grant, 1898, Cat. Birds Brit. Mus. 26: 369. In part. Hypoleucus campbelli stewarti (Ogilvie-Grant): Mathews & Iredale, 1913, Ibis 1 (10 th Ser.): 413. In part. Hypoleucus chalconotus (G. R. Gray): Mathews & Iredale, 1913, Ibis 1 (10 th Ser.): 413. In part. Hypoleucus chalconotus (G. R. Gray): Mathews, 1927, Syst. Avium Australasianarum 1: 227. In part. Phalacrocorax chalconotus (G. R. Gray): Oliver, 1930, N. Z. Birds (1 st ed.): 192. In part. Phalacrocorax huttoni Buller: Oliver, 1930, N. Z. Birds (1 st ed.): 190. In part. Hypoleucus chalconotus (G. R. Gray): Mathews, 1931, List Birds Australasia: 146. In part. Phalacrocorax (Leucocarbo) huttoni Buller: Falla, 1932, Rec. Auck. Inst. Mus. 1: 147. In part. Phalacrocorax (Leucocarbo) chalconotus (G. R. Gray): Falla, 1932, Rec. Auck. Inst. Mus. 1: 148. In part. Hypoleucus chalconotus (G. R. Gray): Mathews, 1936, Suppl. Birds Norfolk & Lord Howe Islands: 142. In part. Hypoleucus huttoni (Buller): Mathews, 1936, Suppl. Birds Norfolk & Lord Howe Islands: 141. In part. Phalacrocorax (Leucocarbo) carunculatus chalconotus (G. R. Gray): Checklist Committee Ornithological Society of New Zealand, 1953, Checklist N. Z. Birds: 29. In part. Leucocarbo carunculatus chalconotus (G. R. Gray): Checklist Committee OSNZ, 1970, Annot. Checklist Birds N. Z. (2 nd ed.): 32. In part. Leucocarbo chalconotus (G. R. Gray): van Tets, 1976, Proc. 16 th Int. Ornith. Cong.: 122. In part. Phalacrocorax carunculatus chalconotus (G. R. Gray): Dorst & Mougin, 1979, in Peters, Check-list Birds World 1 (2 nd ed.): 176. In part. Euleucocarbo chalconotus (G. R. Gray): Siegel-Causey, 1988, Condor 90: 892. In part. Leucocarbo chalconotus (G. R. Gray): Checklist Committee OSNZ, 1990, Checklist Birds N. Z. (3 rd ed.): 83. In part. Phalacrocorax chalconotus (G. R. Gray): O���Brien, 1990, in Marchant & Higgins, Handbook Aust. N. Z. Antarct. Birds: 876. In part. Leucocarbo (carunculatus) chalconotus (G. R. Gray): Johnsgard, 1993, Cormorants, Darters and Pelicans World: 271. In part. Leucocarbo chalconotus (G. R. Gray): Checklist Committee OSNZ, 2010, Checklist Birds N. Z. (4 th ed.): 147. In part. Leucocarbo chalconotus ���Otago���: Rawlence et al., 2014, PLoS ONE 9: e 90769: 1., Published as part of Nicolas J. Rawlence, R. Paul Scofield, Hamish G. Spencer, Chris Lalas, Luke J. Easton, Alan J. D. Tennyson, Mark Adams, Eric Pasquet, Cody Fraser, Jonathan M. Waters & Martyn Kennedy, 2016, Genetic and morphological evidence for two species of Leucocarbo shag (Aves, Pelecaniformes, Phalacrocoracidae) from southern South Island of New Zealand, pp. 676-694 in Zoological Journal of the Linnean Society 177 on page 693, DOI: 10.1111/zoj.12376, http://zenodo.org/record/270312

LEUCOCARBO STEWARTI FOVEAUX SHAG PHALACROCORAX STEWARTI OGILVIE-GRANT, 1898 Diagnosis. A species of Leucocarbo most closely related to L. chalconotus and L. onslowi but distinguished from these species by the plumage characters and allometries outlined in Table 1. Distribution. Restricted to Foveaux Strait and Stewart (based on historical museum skins and modern specimens). Leucocarbo stewarti bones have been recorded from Late Quaternary and archaeological deposits from this region (e.g., Worthy, 1998 b). Rare archaeological and modern beach wrecks in Otago (Rawlence et al., 2014, 2015). The type locality of Gracalus chalconotus G. R. Gray, 1845 is currently considered to be Otago Province. However, the Otago Province included Southland and Foveaux Strait until 1861. We consider that the likely type locality is in fact Karitane, where the type specimen was likely collected by Percy Earl in 1843 (Scofield et al., 2012). Although Earl spent the majority of his time at Karitane, Earl only travelled as far south as the Clutha River, which is still within the range of the Otago lineage, but may have had Maori��� collect for him elsewhere (Scofield et al., 2012). The type locality of Phalacrocorax stewarti Ogilvie-Grant, 1898 is Bluff (a town in Southland), where specimens were collected by Baron A. von Hugel on 13 February 1875 (Ogilvie-Grant, 1898 contra Stewart, Gill et al., 2010). Ogilvie-Grant (1898) designated three syntypes (NHMUK 1880.5. 3.1, 1880.5. 3.2, and 1880.5.3.6), all pied morphs. Warren (1966) only segregated and listed the syntype 1880.5. 3.6 for inclusion in the NHMUK type collection, but this action does not affect the status of the remaining unselected syntypes. NHMUK 1880.5. 3.1 and 1880.5. 3.2 are currently labelled as Phalacrocorax campbelli huttoni (reflecting a previous taxonomic treatment) and we did not attempt to obtain DNA from them. As all three syntypes were collected from the same locality at the same time, and 1880.5. 3.6 clusters with individuals from Foveaux Strait (Figs. 4, 6, Appendix 1), we refer 1880.5. 3.1 and 1880.5. 3.2 to L. stewarti. PLUMAGE EVOLUTION AND PHYLOGEOGRAPHICAL CONSIDERATIONS Phylogenetic relationships amongst the Otago, Foveaux, and Chatham shags suggest a geologically recent colonization of the Chatham Islands from a mainland South source population, consistent with numerous phylogeographical studies from the region (Goldberg, Trewick & Paterson, 2008; Mitchell et al., 2014; Wood et al., 2014). All New Zealand /sub-Antarctic Leucocarbo shags have subtle differences in plumage patterns (e.g. scapulars, prenuptial head crest; Marchant & Higgins, 1990). However, the Otago and Foveaux shags are the only Leucocarbo species with adult dimorphic plumage patterns ��� pied (black and white) and bronze (wholly dark) (Figs 1, 2). Given our phylogeny (Fig. 4), we can hypothesize that the common ancestor of L. chalconotus, L. onslowi, and L. stewarti evolved dimorphic plumage. Divergence between L. chalconotus and L. stewarti is probably a result of isolation in refugia during the Pleistocene glacial cycles, when the majority of offshore islands and rock stacks in Foveaux Strait and around Stewart favoured by L. stewarti were landlocked because of lower sea levels. The Chatham Islands were colonized by progenitors of L. chalconotus, which evolved into L. onslowi. Colonization was either by pied L. chalconotus individuals or a mixture of pied and bronze individuals, with subsequent genetic drift resulting in the loss of the bronze morph. Given the lack of well-supported structure within the Otago shag we cannot definitively test evolutionary scenarios for the bronze morph, but it seems much simpler to assume that the bronze morph evolved once and was subsequently lost in L. onslowi than that it evolved independently twice (in the Otago shag and in the Foveaux shag). CONSERVATION IMPLICATIONS Prior to our splitting of the Stewart shag into two distinct taxa, the International Union for Conservation of Nature Red List categorized this species as Vulnerable with population estimates varying between 1600 and 1800 pairs to fewer than 5000 birds and threats including fisheries interactions and nest disturbance (Birdlife International, 2012). Given their revised status, Otago and Foveaux shags should be managed separately, with further population-genetic research to determine levels of genetic variation at faster-evolving nuclear loci and potential inbreeding (e.g. Calderon et al., 2014). The Foveaux shag has been characterized by population and range stability, in contrast to the Otago shag that has undergone a pronounced population bottleneck and range contraction (Rawlence et al., 2015). Although the Otago shag has recolonized part of its former distribution since the European era, its population size continues to decline (Lalas & Perriman, 2009)., Published as part of Nicolas J. Rawlence, R. Paul Scofield, Hamish G. Spencer, Chris Lalas, Luke J. Easton, Alan J. D. Tennyson, Mark Adams, Eric Pasquet, Cody Fraser, Jonathan M. Waters & Martyn Kennedy, 2016, Genetic and morphological evidence for two species of Leucocarbo shag (Aves, Pelecaniformes, Phalacrocoracidae) from southern South Island of New Zealand, pp. 676-694 in Zoological Journal of the Linnean Society 177 on pages 687-689, DOI: 10.1111/zoj.12376, http://zenodo.org/record/270312, {"references":["Ogilvie-Grant WR. 1898. Catalogue of Birds of the British Museum, vol. 26. London: Trustees of the British Museum [Natural History].","Worthy TH. 1998 b. Fossil avifaunas from Old Neck and Native Island, Stewart Island - Polynesian middens or natural sites? Records of the Canterbury Museum 12: 49 - 82.","Rawlence NJ, Till CE, Scofield RP, Tennyson AJD, Collins CJ, Lalas C, Loh G, Matisoo-Smith E, Waters JM, Spencer HG, Kennedy M. 2014. Strong phylogeographic structure in a sedentary seabird, the Stewart Island Shag (Leucocarbo chalconotus). PLoS ONE 9: e 90769.","Rawlence NJ, Kennedy M, Anderson CNK, Till CE, Smith I, Scofield RP, Tennyson AJD, Hamel J, Lalas C, Matisoo-Smith EA, Waters JM. 2015. Geographically contrasting biodiversity reductions in a widespread New Zealand seabird. Molecular Ecology 18: 4605 - 4616.","Gray GR. 1845. Birds. In: Richardson JE, Gray GR, eds. The Zoology of the Voyage of H. M. S. Erebus and Terror, During the Years 1839 - 43. London: E. W. Janson, 1 - 20.","Scofield RP, Cooper JH, Turvey ST. 2012. A naturalist of the very first order? Percy William Earl (1811 - 1846) in New Zealand. Records of the Canterbury Museum 26: 1 - 20.","Gill BJ, Bell BD, Chambers GK, Medway DG, Palma RL, Scofield RP, Tennyson AJD, Worthy TH. 2010. Checklist of the Birds in New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency Antarctica, 4 th edn. Wellington: Te Papa Press in association with the Ornithological Society of New Zealand.","Warren RLM. 1966. Type-specimens of Birds in the British Museum (Natural History), Vol. 1. Non-passerines. London: Trustees of the British Museum (Natural History).","Goldberg J, Trewick SA, Paterson AM. 2008. Evolution of New Zealand's terrestrial fauna: a review of molecular evidence. Philosophical Transactions of the Royal Society Biological Series 363: 3319 - 3334.","Mitchell KJ, Wood JR, Scofield RP, Llamas B, Cooper A. 2014. Ancient mitochondrial genome reveals unsuspected taxonomic affinity of the extinct Chatham duck (Pachyanas chathamica) and resolves divergence times for New Zealand and sub-Antarctic brown teals. Molecular Phylogenetics and Evolution 70: 420 - 428.","Clements JF, Schulenberg TS, IIiff NJ, Robertson D, Fredericks TA, Sullivan BL, Wood CL. 2014. The eBird / Clements checklist of birds of the world: Version 6.9. Available at: http: // www. birds. cornell. edu / clementscheck list / download /.","Marchant S, Higgins PJ. 1990. Handbook of Australian, New Zealand and Antarctic birds (Volume 1, Part B pelican to ducks). Melbourne: Oxford University Press.","Birdlife International. 2012. Phalacrocorax chalconotus. The IUCN red list of threatened species (Version 2014.3). www. iucnredlist. org.","Calderon L, Quintana F, Cabanne GS, Lougheed SC, Tubaro PL. 2014. Phylogeography and genetic structure of two Patagonian shag species (Aves: Phalacrocoracidae). Molecular Phylogenetics and Evolution 72: 42 - 53.","Lalas C, Perriman L. 2009. Nest counts of Stewart Island shags / mapua (Leucocarbo chalconotus) in Otago. Department of Conservation Research and Development Series 314: 1 - 30."]}

Leucocarbo Bonaparte, 1857 [type species (by subsequent designation, Ogilvie-Grant, 1898) Carbo bougainvillii Lesson, 1837], Published as part of Nicolas J. Rawlence, R. Paul Scofield, Hamish G. Spencer, Chris Lalas, Luke J. Easton, Alan J. D. Tennyson, Mark Adams, Eric Pasquet, Cody Fraser, Jonathan M. Waters & Martyn Kennedy, 2016, Genetic and morphological evidence for two species of Leucocarbo shag (Aves, Pelecaniformes, Phalacrocoracidae) from southern South Island of New Zealand, pp. 676-694 in Zoological Journal of the Linnean Society 177 on page 687, DOI: 10.1111/zoj.12376, http://zenodo.org/record/270312, {"references":["Ogilvie-Grant WR. 1898. Catalogue of Birds of the British Museum, vol. 26. London: Trustees of the British Museum [Natural History]."]}

LEUCOCARBO STEWARTI (OGILVIE-GRANT, 1898) FOVEAUX SHAG. Phalacrocorax colensoi Buller, 1888: Hist. Birds N. Z. (2 nd ed.): 161 ��� Bluff, Southland. In part. Phalacrocorax stewarti Ogilvie-Grant, 1898: Cat. Birds Brit. Mus. 26: 385 ��� Bluff, South (contra Stewart, Gill et al., 2010). Hypoleucus campbelli stewarti (Ogilvie-Grant): Mathews & Iredale, 1913, Ibis 1 (10 th Ser.): 413. In part. Hypoleucus chalconotus (G. R. Gray): Mathews & Iredale, 1913, Ibis 1 (10 th Ser.): 413. In part. Hypoleucus campbelli stewarti (Ogilvie-Grant): Mathews, 1927, Syst. Avium Australasianarum 1: 227. In part. Hypoleucus chalconotus (G. R. Gray): Mathews, 1927, Syst. Avium Australasianarum 1: 227. In part. Phalacrocorax huttoni Buller: Oliver, 1930, N.Z. Birds (1 st ed.): 190. In part. Phalacrocorax chalconotus (G. R. Gray): Oliver, 1930, N.Z. Birds (1 st ed.): 192. Hypoleucus campbelli stewarti (Ogilvie-Grant): Mathews, 1931, List Birds Australasia: 146. Hypoleucus chalconotus (G. R. Gray): Mathews, 1931, List Birds Australasia: 146. In part. Phalacrocorax (Leucocarbo) huttoni Buller: Falla, 1932, Rec. Auck. Inst. Mus. 1: 147. In part. Phalacrocorax (Leucocarbo) chalconotus (G. R. Gray): Falla, 1932, Rec. Auck. Inst. Mus. 1: 148. In part. Hypoleucus huttoni (Buller): Mathews, 1936, Suppl. Birds Norfolk & Lord Howe Islands: 141. In part. Hypoleucus chalconotus (G. R. Gray): Mathews, 1936, Suppl. Birds Norfolk & Lord Howe Islands: 142. In part. Phalacrocorax (Leucocarbo) carunculatus chalconotus (G. R. Gray): Checklist Committee OSNZ, 1953, Checklist N.Z. Birds: 29. In part. Leucocarbo carunculatus chalconotus (G. R. Gray): Checklist Committee OSNZ, 1970, Annot. Checklist Birds N.Z.: 32. In part. Leucocarbo chalconotus (G. R. Gray); van Tets, 1976, Proc. 16 th Int. Ornith. Cong.: 122. In part. Phalacrocorax carunculatus chalconotus (G. R. Gray): Dorst & Mougin, 1979, in Peters, Check-list Birds World 1 (2 nd ed.): 176. In part. Euleucocarbo chalconotus (G. R. Gray): Siegel-Causey, 1988, Condor 90: 892. In part. Leucocarbo chalconotus (G. R. Gray): Checklist Committee OSNZ, 1990, Checklist Birds N. Z. (3 rd ed.): 83. In part. Phalacrocorax chalconotus (G. R. Gray): O���Brien, 1990, in Marchant & Higgins, Handbook Aust. N. Z. Antarct. Birds: 876. In part. Leucocarbo (carunculatus) chalconotus (G. R. Gray): Johnsgard, 1993, Cormorants, Darters and Pelicans World: 271. In part. Leucocarbo chalconotus (G. R. Gray): Checklist Committee OSNZ, 2010, Checklist Birds N. Z. (4 th ed.): 147. In part. Leucocarbo chalconotus ���Foveaux Strait���: Rawlence et al., 2014, PLoS ONE 9: e 90769: 1., Published as part of Nicolas J. Rawlence, R. Paul Scofield, Hamish G. Spencer, Chris Lalas, Luke J. Easton, Alan J. D. Tennyson, Mark Adams, Eric Pasquet, Cody Fraser, Jonathan M. Waters & Martyn Kennedy, 2016, Genetic and morphological evidence for two species of Leucocarbo shag (Aves, Pelecaniformes, Phalacrocoracidae) from southern South Island of New Zealand, pp. 676-694 in Zoological Journal of the Linnean Society 177 on pages 693-694, DOI: 10.1111/zoj.12376, http://zenodo.org/record/270312