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<mods:title id="C5B8E2A8C698B8247F7C66FC3919860D">Genetic and morphological evidence for two species of Leucocarbo shag (Aves, Pelecaniformes, Phalacrocoracidae) from southern South Island of New Zealand</mods:title>
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<mods:namePart id="12BE549F6FB5C531B3DE2EF77A51C1C2">Nicolas J. Rawlence</mods:namePart>
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<mods:namePart id="FAAFED0AB3B72FB647314D2BEF9BBC27">R. Paul Scofield</mods:namePart>
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<mods:namePart id="170F2C6C8EFB3B380BC2FD8725287CA9">Hamish G. Spencer</mods:namePart>
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<mods:namePart id="53645F6447B55E4221C919E7602A6EA3">Chris Lalas</mods:namePart>
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<mods:namePart id="7FB4989C45FF3AA6F785811FAD845BC2">Luke J. Easton</mods:namePart>
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<mods:namePart id="A7FA8BBBB8E0DEBE5E596F138D743F66">Alan J. D. Tennyson</mods:namePart>
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<mods:namePart id="B0A5C9578857CFAE34DF45C6BC397303">Mark Adams</mods:namePart>
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<mods:namePart id="B810DBE614665CA92609476C97BFB385">Eric Pasquet</mods:namePart>
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<mods:namePart id="0A035FD14F936C520F5128C4A2234F8A">Jonathan M. Waters</mods:namePart>
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<mods:namePart id="E6251A0404186AF838E4A679DE3524AA">Martyn Kennedy</mods:namePart>
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<treatment id="039387CFFF94D842683FFF1CFF41681C" ID-DOI="http://doi.org/10.5281/zenodo.5685567" ID-GBIF-Taxon="121268891" ID-Zenodo-Dep="5685567" LSID="urn:lsid:plazi:treatment:039387CFFF94D842683FFF1CFF41681C" httpUri="http://treatment.plazi.org/id/039387CFFF94D842683FFF1CFF41681C" lastPageId="13" lastPageNumber="689" pageId="11" pageNumber="687">
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<taxonomicName id="4C3A4D5AFF94D844683FFF1CFBA36FC7" box="[807,1076,214,235]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="11" pageNumber="692" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF94D844683FFF1CFBA36FC7" bold="true" box="[807,1076,214,235]" italics="true" pageId="11" pageNumber="687">LEUCOCARBO STEWARTI</emphasis>
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FOVEAUX SHAG
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<taxonomicName id="4C3A4D5AFF94D8446820FF38FAEC6E26" authority="Ogilvie-Grant, 1898" authorityName="Ogilvie-Grant" authorityYear="1898" box="[824,1403,240,267]" class="Aves" family="Phalacrocoracidae" genus="Phalacrocorax" kingdom="Animalia" order="Suliformes" pageId="11" pageNumber="693" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF94D8446820FF38FBF86E25" bold="true" box="[824,1135,242,266]" italics="true" pageId="11" pageNumber="687">PHALACROCORAX STEWARTI</emphasis>
<bibRefCitation id="EFAB4B28FF94D8446F60FF3AFAEC6E26" author="Ogilvie-Grant WR." box="[1144,1403,240,267]" journalOrPublisher="London: Trustees of the British Museum [Natural History]" pageId="11" pageNumber="687" part="26" refString="Ogilvie-Grant WR. 1898. Catalogue of Birds of the British Museum, vol. 26. London: Trustees of the British Museum [Natural History]." title="Catalogue of Birds of the British Museum" type="journal volume" year="1898">OGILVIE-GRANT, 1898</bibRefCitation>
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<emphasis id="B94EEACBFF94D844683FFEE8FC0A6E1B" bold="true" box="[807,925,290,311]" italics="true" pageId="11" pageNumber="687">Diagnosis.</emphasis>
A species of 
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<emphasis id="B94EEACBFF94D8446F7BFEE8FB746E14" bold="true" box="[1123,1251,290,312]" italics="true" pageId="11" pageNumber="687">Leucocarbo</emphasis>
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most closely related to 
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<emphasis id="B94EEACBFF94D84468BDFE8BFBDC6E7A" bold="true" box="[933,1099,320,342]" italics="true" pageId="11" pageNumber="687">L. chalconotus</emphasis>
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and 
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<emphasis id="B94EEACBFF94D8446F97FE8BFA926E7A" bold="true" box="[1167,1285,320,342]" italics="true" pageId="11" pageNumber="687">L. onslowi</emphasis>
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but distinguished from these species by the plumage characters and allometries outlined in 
<tableCitation id="C6B80362FF94D8446F82FEB4FB666EBF" box="[1178,1265,382,404]" captionStart="Table 1" captionStartId="12.[125,191,471,490]" captionTargetBox="[125,1811,557,1626]" captionText="Table 1. Species diagnoses for the Otago shag Leucocarbo chalconotus (G. R. Gray, 1845), Foveaux shag Leucocarbo stewarti (Ogilvie-Grant, 1898), and Chatham Island shag Leucocarbo onslowi (Forbes, 1893)" httpUri="http://table.plazi.org/id/DF456651FF93D8436B65FE1DFD306D2B" pageId="11" pageNumber="687" tableUuid="DF456651FF93D8436B65FE1DFD306D2B">Table 1</tableCitation>
.
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<emphasis id="B94EEACBFF94D8446827FE56FC5D6E9E" bold="true" box="[831,970,412,434]" italics="true" pageId="11" pageNumber="687">Distribution</emphasis>
. Restricted to Foveaux Strait and Stewart (based on historical museum skins and modern specimens). 
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<emphasis id="B94EEACBFF94D8446F4DFE13FAA86EC3" bold="true" box="[1109,1343,473,495]" italics="true" pageId="11" pageNumber="687">Leucocarbo stewarti</emphasis>
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bones have been recorded from Late Quaternary and archaeological deposits from this region (e.g., 
<bibRefCitation id="EFAB4B28FF94D844683FFDFFFC4B6D67" author="Worthy TH." box="[807,988,565,587]" journalOrPublisher="Polynesian middens or natural sites? Records of the Canterbury Museum" pageId="11" pageNumber="687" pagination="49 - 82" part="12" refString="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." title="Fossil avifaunas from Old Neck and Native Island, Stewart Island" type="journal article" year="1998" yearSuffix="b">Worthy, 1998b</bibRefCitation>
). Rare archaeological and modern beach wrecks in Otago (
<bibRefCitation id="EFAB4B28FF94D8446F6CFD9EFA1E6D45" author="Rawlence NJ &amp; Till CE &amp; Scofield RP &amp; Tennyson AJD &amp; Collins CJ &amp; Lalas C &amp; Loh G &amp; Matisoo-Smith E &amp; Waters JM &amp; Spencer HG &amp; Kennedy M." box="[1140,1417,595,618]" journalOrPublisher="PLoS ONE" pageId="11" pageNumber="687" pagination="90769" part="9" refString="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." title="Strong phylogeographic structure in a sedentary seabird, the Stewart Island Shag (Leucocarbo chalconotus)" type="journal article" year="2014">
Rawlence 
<emphasis id="B94EEACBFF94D8446FE0FD9EFABC6D45" bold="true" box="[1272,1323,595,617]" italics="true" pageId="11" pageNumber="687">et al</emphasis>
., 2014
</bibRefCitation>
, 
<bibRefCitation id="EFAB4B28FF94D844683FFDB8FCF66DA4" author="Rawlence NJ &amp; Kennedy M &amp; Anderson CNK &amp; Till CE &amp; Smith I &amp; Scofield RP &amp; Tennyson AJD &amp; Hamel J &amp; Lalas C &amp; Matisoo-Smith EA &amp; Waters JM." box="[807,865,626,648]" journalOrPublisher="Molecular Ecology" pageId="11" pageNumber="687" pagination="4605 - 4616" part="18" refString="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." title="Geographically contrasting biodiversity reductions in a widespread New Zealand seabird" type="journal article" year="2015">2015</bibRefCitation>
).
</paragraph>
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LOCALITIES
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The 
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locality of 
<taxonomicName id="4C3A4D5AFF94D8446F14FD26FCCA6C0D" authority="G. R. Gray, 1845" authorityName="G. R. Gray" authorityYear="1845" class="Aves" family="Phalacrocoracidae" genus="Gracalus" kingdom="Animalia" order="Suliformes" pageId="11" pageNumber="687" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF94D8446F14FD26FA956C2E" bold="true" box="[1036,1282,748,770]" italics="true" pageId="11" pageNumber="687">Gracalus chalconotus</emphasis>
<bibRefCitation id="EFAB4B28FF94D8446E12FD27FCCA6C0D" author="Gray GR." editor="Richardson JE" journalOrPublisher="London: E. W. Janson, 1 - 20" pageId="11" pageNumber="687" pagination="1839 - 43" refString="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." title="Birds" type="book chapter" volumeTitle="The Zoology of the Voyage of H" year="1845">G. R. Gray, 1845</bibRefCitation>
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is currently considered to be Otago Province. However, the Otago Province included Southland and Foveaux Strait until 1861. We consider that the likely 
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locality is in fact Karitane, where the 
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specimen was likely collected by Percy Earl in 1843 (
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Scofield 
<emphasis id="B94EEACBFF94D84468CAFC6EFB926C95" bold="true" box="[978,1029,931,953]" italics="true" pageId="11" pageNumber="687">et al</emphasis>
., 2012
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). 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 (
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Scofield 
<emphasis id="B94EEACBFF94D8446E54FBD4FAE86B18" bold="true" box="[1356,1407,1054,1076]" italics="true" pageId="11" pageNumber="687">et al</emphasis>
., 2012
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). The 
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locality of 
<taxonomicName id="4C3A4D5AFF94D8446F67FBF6FB9A6B5C" authority="Ogilvie-Grant, 1898" authorityName="Ogilvie-Grant" authorityYear="1898" class="Aves" family="Phalacrocoracidae" genus="Phalacrocorax" kingdom="Animalia" order="Suliformes" pageId="11" pageNumber="687" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF94D8446F67FBF6FA1A6B7E" bold="true" box="[1151,1421,1084,1106]" italics="true" pageId="11" pageNumber="687">Phalacrocorax stewarti</emphasis>
<bibRefCitation id="EFAB4B28FF94D844683FFB91FB9A6B5C" author="Ogilvie-Grant WR." box="[807,1037,1115,1137]" journalOrPublisher="London: Trustees of the British Museum [Natural History]" pageId="11" pageNumber="687" part="26" refString="Ogilvie-Grant WR. 1898. Catalogue of Birds of the British Museum, vol. 26. London: Trustees of the British Museum [Natural History]." title="Catalogue of Birds of the British Museum" type="journal volume" year="1898">Ogilvie-Grant, 1898</bibRefCitation>
</taxonomicName>
is Bluff (a town in Southland), where specimens were collected by Baron A. von Hugel on 
<date id="FF841019FF94D844688FFB53FBF46B82" box="[919,1123,1176,1198]" pageId="11" pageNumber="687" value="1875-02-13">13 February 1875</date>
(
<bibRefCitation id="EFAB4B28FF94D8446F6BFB52FAC16B82" author="Ogilvie-Grant WR." box="[1139,1366,1176,1198]" journalOrPublisher="London: Trustees of the British Museum [Natural History]" pageId="11" pageNumber="687" part="26" refString="Ogilvie-Grant WR. 1898. Catalogue of Birds of the British Museum, vol. 26. London: Trustees of the British Museum [Natural History]." title="Catalogue of Birds of the British Museum" type="journal volume" year="1898">Ogilvie-Grant, 1898</bibRefCitation>
<emphasis id="B94EEACBFF94D8446E47FB53FCDE6BE0" bold="true" italics="true" pageId="11" pageNumber="687">contra</emphasis>
Stewart, 
<bibRefCitation id="EFAB4B28FF94D8446F00FB7DFB406BE1" author="Gill BJ &amp; Bell BD &amp; Chambers GK &amp; Medway DG &amp; Palma RL &amp; Scofield RP &amp; Tennyson AJD &amp; Worthy TH." box="[1048,1239,1206,1229]" journalOrPublisher="Wellington: Te Papa Press in association with the Ornithological Society of New Zealand" pageId="11" pageNumber="687" refString="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." title="Checklist of the Birds in New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency Antarctica, 4 th edn" type="book" year="2010">
Gill 
<emphasis id="B94EEACBFF94D8446F57FB7DFB156BE0" bold="true" box="[1103,1154,1206,1228]" italics="true" pageId="11" pageNumber="687">et al</emphasis>
., 2010
</bibRefCitation>
). 
<bibRefCitation id="EFAB4B28FF94D8446FF7FB7DFCFA6BC0" author="Ogilvie-Grant WR." journalOrPublisher="London: Trustees of the British Museum [Natural History]" pageId="11" pageNumber="687" part="26" refString="Ogilvie-Grant WR. 1898. Catalogue of Birds of the British Museum, vol. 26. London: Trustees of the British Museum [Natural History]." title="Catalogue of Birds of the British Museum" type="journal volume" year="1898">Ogilvie-Grant (1898)</bibRefCitation>
designated three 
<typeStatus id="5481887BFF94D8446F8DFB1CFB6E6BC7" box="[1173,1273,1238,1259]" pageId="11" pageNumber="687" type="syntype">syntypes</typeStatus>
(
<collectionCode id="ED2BAE1CFF94D8446E3DFB1CFA1A6BC7" box="[1317,1421,1238,1259]" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" pageId="11" pageNumber="687">NHMUK</collectionCode>
1880.5.3.1, 1880.5.3.2, and 1880.5.3.6), all pied morphs. 
<bibRefCitation id="EFAB4B28FF94D844688AFAD9FBA96A04" author="Warren RLM." box="[914,1086,1298,1320]" journalOrPublisher="London: Trustees of the British Museum (Natural History)" pageId="11" pageNumber="687" refString="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)." title="Type-specimens of Birds in the British Museum (Natural History), Vol. 1. Non-passerines" type="book" year="1966">Warren (1966)</bibRefCitation>
only segregated and listed the 
<typeStatus id="5481887BFF94D844684EFAF8FC386A6B" box="[854,943,1330,1351]" pageId="11" pageNumber="687" type="syntype">syntype</typeStatus>
1880.5.3.6 for inclusion in the 
<collectionCode id="ED2BAE1CFF94D8446E3EFAF8FA1A6A6B" box="[1318,1421,1330,1351]" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" pageId="11" pageNumber="687">NHMUK</collectionCode>
<typeStatus id="5481887BFF94D844683FFA9AFCCF6A49" box="[807,856,1360,1381]" pageId="11" pageNumber="687">type</typeStatus>
collection, but this action does not affect the status of the remaining unselected 
<typeStatus id="5481887BFF94D8446FB7FAA5FA816AA8" box="[1199,1302,1391,1412]" pageId="11" pageNumber="687" type="syntype">syntypes</typeStatus>
. 
<collectionCode id="ED2BAE1CFF94D8446E3EFAA5FA1A6AA8" box="[1318,1421,1391,1412]" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34665" name="Natural History Museum, London" pageId="11" pageNumber="687">NHMUK</collectionCode>
1880.5.3.1 and 1880.5.3.2 are currently labelled as 
<taxonomicName id="4C3A4D5AFF94D844683FFA61FB336AED" box="[807,1188,1451,1473]" class="Aves" family="Phalacrocoracidae" genus="Phalacrocorax" kingdom="Animalia" order="Suliformes" pageId="11" pageNumber="687" phylum="Chordata" rank="subSpecies" species="campbelli" subSpecies="huttoni">
<emphasis id="B94EEACBFF94D844683FFA61FB336AED" bold="true" box="[807,1188,1451,1473]" italics="true" pageId="11" pageNumber="687">Phalacrocorax campbelli huttoni</emphasis>
</taxonomicName>
(reflecting a previous taxonomic treatment) and we did not attempt to obtain 
<collectionCode id="ED2BAE1CFF94D8446862FA23FC246AD2" box="[890,947,1513,1534]" country="Australia" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15536" name="Department of Natural Resources, Environment, The Arts and Sport" pageId="11" pageNumber="687">DNA</collectionCode>
from them. As all three 
<typeStatus id="5481887BFF94D8446FFFFA23FADC6AD2" box="[1255,1355,1513,1534]" pageId="11" pageNumber="687" type="syntype">syntypes</typeStatus>
were collected from the same locality at the same time, and 1880.5.3.6 clusters with individuals from Foveaux Strait (
<figureCitation id="13012A5CFF94D84468EDF98EFBD26975" box="[1013,1093,1604,1626]" captionStart="Figure 4" captionStartId="7.[144,223,1026,1045]" captionTargetBox="[151,1413,198,993]" captionTargetId="figure@7.[151,1413,198,993]" captionTargetPageId="7" captionText="Figure 4. The Bayesian phylogram of Leucocarbo shags from an analysis with a single individual of each haplotype (from BEAST v. 1.7.4). Apart from the two outgroup taxa, the labels show the species (Leucocarbo chalconotus, Leucocarbo onslowi, and Leucocarbo stewarti) with the haplotype in parentheses (for haplotype designation see Table S 5). The" httpUri="https://zenodo.org/record/270316/files/figure.png" pageId="11" pageNumber="687">Figs. 4</figureCitation>
, 
<figureCitation id="13012A5CFF94D8446F43F98EFBFC6976" box="[1115,1131,1604,1626]" captionStart="Figure 6" captionStartId="9.[144,223,1559,1578]" captionTargetBox="[476,1094,948,1525]" captionTargetId="figure@9.[476,1094,948,1526]" captionTargetPageId="9" captionText="Figure 6. Principal component analysis (PCA) of pooled measurements from museum skins for 21 Otago shags (blue squares), 34 Foveaux shags (red triangles), and seven Chatham Island shags (green diamonds). Type specimens for Otago shag (Gracalus chalconotus, Graculus glaucus, and Phalacrocorax huttoni) and Foveaux shag (Phalacrocorax stewarti) are shown with labelled black circles. PC 1 loadings&gt; 0.5 / &lt;- 0.5, PC 2 loading Phal 0.82." httpUri="https://zenodo.org/record/270318/files/figure.png" pageId="11" pageNumber="687">6</figureCitation>
, Appendix 1), we refer 1880.5.3.1 and 1880.5.3.2 to 
<emphasis id="B94EEACBFF94D8446F77F9A9FB656954" bold="true" box="[1135,1266,1635,1656]" italics="true" pageId="11" pageNumber="687">
<taxonomicName id="4C3A4D5AFF94D8446F77F9A9FB7A6954" box="[1135,1261,1635,1656]" class="Aves" family="Phalacrocoracidae" genus="Phalacrocorax" kingdom="Animalia" order="Suliformes" pageId="11" pageNumber="687" phylum="Chordata" rank="species" species="stewarti">L. stewarti</taxonomicName>
.
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C3206552FF94D842685FF965FD0E6B5D" lastPageId="13" lastPageNumber="689" pageId="11" pageNumber="687" type="discussion">
<paragraph id="8B8536D9FF94D844685FF965FAFA69E8" blockId="11.[839,1389,1711,1762]" box="[839,1389,1711,1732]" pageId="11" pageNumber="687">
<heading id="D0CD81B5FF94D844685FF965FAFA69E8" allCaps="true" box="[839,1389,1711,1732]" centered="true" fontSize="8" level="6" pageId="11" pageNumber="687" reason="5">PLUMAGE EVOLUTION AND PHYLOGEOGRAPHICAL</heading>
</paragraph>
<paragraph id="8B8536D9FF94D84468E3F91BFB2F69CE" blockId="11.[839,1389,1711,1762]" box="[1019,1208,1745,1762]" pageId="11" pageNumber="687">
<heading id="D0CD81B5FF94D84468E3F91BFB2F69CE" allCaps="true" box="[1019,1208,1745,1762]" centered="true" fontSize="7" level="6" pageId="11" pageNumber="687" reason="4">CONSIDERATIONS</heading>
</paragraph>
<paragraph id="8B8536D9FF94D842683FF931FD0E6B5D" blockId="11.[807,1421,1787,1901]" lastBlockId="13.[144,759,199,1137]" lastPageId="13" lastPageNumber="689" pageId="11" pageNumber="687">
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 (
<bibRefCitation id="EFAB4B28FF92D8426A6AFF2FFF5E6E35" author="Goldberg J &amp; Trewick SA &amp; Paterson AM." journalOrPublisher="Philosophical Transactions of the Royal Society Biological Series 363" pageId="13" pageNumber="689" pagination="3319 - 3334" refString="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." title="Evolution of New Zealand's terrestrial fauna: a review of molecular evidence" type="book chapter" year="2008">Goldberg, Trewick &amp; Paterson, 2008</bibRefCitation>
; 
<bibRefCitation id="EFAB4B28FF92D8426BC2FECEFE5C6E35" author="Mitchell KJ &amp; Wood JR &amp; Scofield RP &amp; Llamas B &amp; Cooper A." box="[218,459,259,282]" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="13" pageNumber="689" pagination="420 - 428" part="70" refString="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." title="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" type="journal article" year="2014">
Mitchell 
<emphasis id="B94EEACBFF92D8426A5DFECEFEE96E35" bold="true" box="[325,382,259,281]" italics="true" pageId="13" pageNumber="689">et al.</emphasis>
, 2014
</bibRefCitation>
; 
<bibRefCitation id="EFAB4B28FF92D8426AF2FECEFD2A6E36" author="Clements JF &amp; Schulenberg TS &amp; IIiff NJ &amp; Robertson D &amp; Fredericks TA &amp; Sullivan BL &amp; Wood CL." box="[490,701,259,282]" journalOrPublisher="Available at" pageId="13" pageNumber="689" publicationUrl="http://www.birds.cornell.edu/clementscheck" refString="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 /." title="The eBird / Clements checklist of birds of the world: Version 6.9" type="book" year="2014">
Wood 
<emphasis id="B94EEACBFF92D842692DFECEFDF86E35" bold="true" box="[565,623,259,281]" italics="true" pageId="13" pageNumber="689">et al.</emphasis>
, 2014
</bibRefCitation>
). All 
<collectingCountry id="F32D7649FF92D8426B88FEE9FEBA6E14" box="[144,301,290,312]" name="New Zealand" pageId="13" pageNumber="689">New Zealand</collectingCountry>
/sub-Antarctic 
<taxonomicName id="4C3A4D5AFF92D8426AF8FEE8FDF76E14" box="[480,608,290,312]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="genus">
<emphasis id="B94EEACBFF92D8426AF8FEE8FDF76E14" bold="true" box="[480,608,290,312]" italics="true" pageId="13" pageNumber="689">Leucocarbo</emphasis>
</taxonomicName>
shags have subtle differences in plumage patterns (e.g. scapulars, prenuptial head crest; 
<bibRefCitation id="EFAB4B28FF92D8426AE9FE95FF5D6EB8" author="Marchant S &amp; Higgins PJ." journalOrPublisher="New Zealand and Antarctic birds" pageId="13" pageNumber="689" part="1" refString="Marchant S, Higgins PJ. 1990. Handbook of Australian, New Zealand and Antarctic birds (Volume 1, Part B pelican to ducks). Melbourne: Oxford University Press." title="Handbook of Australian" type="journal volume" year="1990">Marchant &amp; Higgins, 1990</bibRefCitation>
). However, the Otago and Foveaux shags are the only 
<taxonomicName id="4C3A4D5AFF92D8426A1DFE56FE126E9E" box="[261,389,412,434]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="genus">
<emphasis id="B94EEACBFF92D8426A1DFE56FE126E9E" bold="true" box="[261,389,412,434]" italics="true" pageId="13" pageNumber="689">Leucocarbo</emphasis>
</taxonomicName>
species with adult dimorphic plumage patterns – pied (black and white) and bronze (wholly dark) (
<figureCitation id="13012A5CFF92D8426A94FE10FE436EC3" box="[396,468,474,496]" captionStart="Figure 1" captionStartId="2.[166,245,996,1015]" captionTargetBox="[393,1217,198,963]" captionTargetId="figure@2.[393,1218,198,963]" captionTargetPageId="2" captionText="Figure 1. Distributional and morphological data for Otago, Foveaux, and Chatham Island shags. A, map of New Zeal- and showing the location of the Chatham Islands and Otago / Foveaux Strait study sites. Blue circles represent the prehistoric distribution of Otago shag outside the study area. B, distribution of Chatham Island shag breeding colonies and roosting sites (green circles). The Chatham Island shag exhibits pied plumage only (white pie chart) with pronounced bright orange caruncles in breeding plumage (orange pie chart). C, distribution of Otago (blue circles) and Foveaux (red circles) shag breeding colonies and roosting sites. Otago shag populations have 20 – 30 % pied morphs (pie charts; black: dark-bronze; white: pied) and 50: 50 % small bright orange caruncles: dark orange scattered papillae in prenuptial breeding plumage (pie charts; yellow: small bright orange caruncles; grey: dark orange scattered papillae) compared with 50 – 60 % pied morph and dark orange scattered papillae in prenuptial breeding plumage in the Foveaux shag. In (C), multiple breeding colonies and roosting sites are represented at the following locations (from north to south): Otago Peninsula including Long Beach, Aramoana, Otago Harbour, Taiaroa Head, Boulder Beach, Papanui Beach, Allans Beach, Wharekakahu Island, and Gull Rocks; Seal Rocks and Ruapuke Island; Easy Harbour and Shag Rock. In (C), the Leucocarbo shag species illustrating plumage characters are: pied morph (Otago shag), bronze morph (Foveaux shag), small caruncles (Otago shag), and scattered papillae (Foveaux shag). Figure adapted from Rawlence et al. (2014)." httpUri="https://zenodo.org/record/270313/files/figure.png" pageId="13" pageNumber="689">Figs 1</figureCitation>
, 
<figureCitation id="13012A5CFF92D8426AFBFE10FE646EDC" box="[483,499,474,496]" captionStart="Figure 2" captionStartId="3.[144,223,564,583]" captionTargetBox="[151,1413,198,531]" captionTargetId="figure@3.[151,1413,198,531]" captionTargetPageId="3" captionText="Figure 2. Carunculation and gular pouch colour in pied morphs of Chatham Island, Otago, and Foveaux shags. The Chatham Island shag (left) has pronounced bright orange facial caruncles and gular pouch; the Otago shag (centre) has equal frequencies of small bright orange facial caruncles (shown) and dark orange scattered papillae (not shown), and bright to dark orange (shown) or purple (not shown) gular pouch; the Foveaux shag (right) has scattered dark orange papillae (shown) and bright to dark orange (shown) gular pouch. Photos (not to scale) courtesy of New Zealand Birds Online (John Kendrick, left and Brian Bell, right) and Hiltrun Ratz (centre)." httpUri="https://zenodo.org/record/270314/files/figure.png" pageId="13" pageNumber="689">2</figureCitation>
). Given our phylogeny (
<figureCitation id="13012A5CFF92D8426B80FE32FF4B6D22" box="[152,220,504,526]" captionStart="Figure 4" captionStartId="7.[144,223,1026,1045]" captionTargetBox="[151,1413,198,993]" captionTargetId="figure@7.[151,1413,198,993]" captionTargetPageId="7" captionText="Figure 4. The Bayesian phylogram of Leucocarbo shags from an analysis with a single individual of each haplotype (from BEAST v. 1.7.4). Apart from the two outgroup taxa, the labels show the species (Leucocarbo chalconotus, Leucocarbo onslowi, and Leucocarbo stewarti) with the haplotype in parentheses (for haplotype designation see Table S 5). The" httpUri="https://zenodo.org/record/270316/files/figure.png" pageId="13" pageNumber="689">Fig. 4</figureCitation>
), we can hypothesize that the common ancestor of 
<taxonomicName id="4C3A4D5AFF92D8426BF3FDDDFE066D00" box="[235,401,534,556]" class="Aves" family="Phalacrocoracidae" genus="Gracalus" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF92D8426BF3FDDDFE066D00" bold="true" box="[235,401,534,556]" italics="true" pageId="13" pageNumber="689">L. chalconotus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D5AFF92D8426AB3FDDDFDB56D00" box="[427,546,534,556]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="species" species="onslowi">
<emphasis id="B94EEACBFF92D8426AB3FDDDFDB56D00" bold="true" box="[427,546,534,556]" italics="true" pageId="13" pageNumber="689">L. onslowi</emphasis>
</taxonomicName>
, and 
<taxonomicName id="4C3A4D5AFF92D8426962FDDDFD616D00" box="[634,758,535,556]" class="Aves" family="Phalacrocoracidae" genus="Phalacrocorax" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF92D8426962FDDDFD616D00" bold="true" box="[634,758,535,556]" italics="true" pageId="13" pageNumber="689">L. stewarti</emphasis>
</taxonomicName>
evolved dimorphic plumage. Divergence between 
<taxonomicName id="4C3A4D5AFF92D8426B88FD9EFEA16D45" box="[144,310,595,617]" class="Aves" family="Phalacrocoracidae" genus="Gracalus" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF92D8426B88FD9EFEA16D45" bold="true" box="[144,310,595,617]" italics="true" pageId="13" pageNumber="689">L. chalconotus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D5AFF92D8426A6AFD9EFE786D45" box="[370,495,596,617]" class="Aves" family="Phalacrocoracidae" genus="Phalacrocorax" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF92D8426A6AFD9EFE786D45" bold="true" box="[370,495,596,617]" italics="true" pageId="13" pageNumber="689">L. stewarti</emphasis>
</taxonomicName>
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 
<taxonomicName id="4C3A4D5AFF92D8426A83FD04FD806DCF" box="[411,535,718,739]" class="Aves" family="Phalacrocoracidae" genus="Phalacrocorax" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF92D8426A83FD04FD806DCF" bold="true" box="[411,535,718,739]" italics="true" pageId="13" pageNumber="689">L. stewarti</emphasis>
</taxonomicName>
were landlocked because of lower sea levels. The Chatham Islands were colonized by progenitors of 
<taxonomicName id="4C3A4D5AFF92D8426951FCC6FD786C0D" box="[585,751,779,801]" class="Aves" family="Phalacrocoracidae" genus="Gracalus" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF92D8426951FCC6FD786C0D" bold="true" box="[585,751,779,801]" italics="true" pageId="13" pageNumber="689">L. chalconotus</emphasis>
</taxonomicName>
, which evolved into 
<taxonomicName id="4C3A4D5AFF92D8426A8CFCE0FD9C6C13" box="[404,523,809,831]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="species" species="onslowi">
<emphasis id="B94EEACBFF92D8426A8CFCE0FD9C6C13" bold="true" box="[404,523,809,831]" italics="true" pageId="13" pageNumber="689">L. onslowi</emphasis>
</taxonomicName>
. Colonization was either by pied 
<taxonomicName id="4C3A4D5AFF92D8426A5DFC83FE7B6C72" box="[325,492,840,862]" class="Aves" family="Phalacrocoracidae" genus="Gracalus" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF92D8426A5DFC83FE7B6C72" bold="true" box="[325,492,840,862]" italics="true" pageId="13" pageNumber="689">L. chalconotus</emphasis>
</taxonomicName>
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 
<taxonomicName id="4C3A4D5AFF92D8426B88FBF7FE916B7E" box="[144,262,1084,1106]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="13" pageNumber="689" phylum="Chordata" rank="species" species="onslowi">
<emphasis id="B94EEACBFF92D8426B88FBF7FE916B7E" bold="true" box="[144,262,1084,1106]" italics="true" pageId="13" pageNumber="689">L. onslowi</emphasis>
</taxonomicName>
than that it evolved independently twice (in the Otago shag and in the Foveaux shag).
</paragraph>
</subSubSection>
<caption id="DF456651FF93D8436B65FE1DFD306D2B" ID-Table-UUID="DF456651FF93D8436B65FE1DFD306D2B" httpUri="http://table.plazi.org/id/DF456651FF93D8436B65FE1DFD306D2B" pageId="12" pageNumber="688" targetBox="[125,1811,557,1626]" targetIsTable="true" targetPageId="12">
<paragraph id="8B8536D9FF93D8436B65FE1DFD306D2B" blockId="12.[125,1821,132,519]" pageId="12" pageNumber="688">
<emphasis id="B94EEACBFF93D8436B65FE1DFF4D6EC5" bold="true" box="[125,218,470,490]" pageId="12" pageNumber="688">Table 1.</emphasis>
Species diagnoses for the Otago shag 
<taxonomicName id="4C3A4D5AFF93D8436965FE1CFBD16EC5" authority="(G. R. Gray, 1845)" authorityName="G. R. Gray" authorityYear="1845" baseAuthorityName="G. R. Gray" baseAuthorityYear="1845" box="[637,1094,470,490]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF93D8436965FE1CFCE16EC5" bold="true" box="[637,886,470,489]" italics="true" pageId="12" pageNumber="688">Leucocarbo chalconotus</emphasis>
(G. R. Gray, 1845)
</taxonomicName>
, Foveaux shag 
<taxonomicName id="4C3A4D5AFF93D8436FF7FE1CF9396EC6" authority="Ogilvie-Grant, 1898" authorityName="Ogilvie-Grant" authorityYear="1898" box="[1263,1710,470,490]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF93D8436FF7FE1CFA566EC5" bold="true" box="[1263,1473,470,489]" italics="true" pageId="12" pageNumber="688">Leucocarbo stewarti</emphasis>
(Ogilvie-Grant, 1898)
</taxonomicName>
, and Chatham Island shag 
<taxonomicName id="4C3A4D5AFF93D8436A26FE39FD306D2B" authority="(Forbes, 1893)" authorityName="Forbes" authorityYear="1893" baseAuthorityName="Forbes" baseAuthorityYear="1893" box="[318,679,499,519]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="onslowi">
<emphasis id="B94EEACBFF93D8436A26FE39FD9D6D2A" bold="true" box="[318,522,499,518]" italics="true" pageId="12" pageNumber="688">Leucocarbo onslowi</emphasis>
(Forbes, 1893)
</taxonomicName>
</paragraph>
</caption>
<paragraph id="8B8536D9FF93D8436ACBFDE7F9026976" pageId="12" pageNumber="688">
<table id="F93AC479FF9327B06B65FDE7F8846976" box="[125,1811,557,1626]" gridcols="5" gridrows="36" pageId="12" pageNumber="688">
<tr id="350A349BFF9327B06B65FDE7F8846D6C" box="[125,1811,557,576]" gridrow="0" pageId="12" pageNumber="688" rowspan-0="1" rowspan-2="1" rowspan-3="1" rowspan-4="1">
<th id="76DB5DE7FF9327B06ACBFDE7FC996D6C" box="[467,782,557,576]" gridcol="1" gridrow="0" pageId="12" pageNumber="688">Chatham Island shag</th>
</tr>
<tr id="350A349BFF9327B06B65FD80F8846D72" box="[125,1811,586,606]" gridrow="1" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65FD80FE356D72" box="[125,418,586,606]" gridcol="0" gridrow="1" pageId="12" pageNumber="688">Feature</th>
<td id="76DB5DE7FF9327B06ACBFD80FC996D72" box="[467,782,586,606]" gridcol="1" gridrow="1" pageId="12" pageNumber="688">
(
<taxonomicName id="4C3A4D5AFF93D8436AC2FD81FDD06D71" box="[474,583,586,606]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="onslowi">
<emphasis id="B94EEACBFF93D8436AC2FD81FDD06D71" bold="true" box="[474,583,586,606]" italics="true" pageId="12" pageNumber="688">L. onslowi</emphasis>
</taxonomicName>
)
</td>
<td id="76DB5DE7FF9327B06830FD80FBCC6D72" box="[808,1115,586,606]" gridcol="2" gridrow="1" pageId="12" pageNumber="688">
Otago shag (
<taxonomicName id="4C3A4D5AFF93D84368B6FD81FBD06D71" box="[942,1095,586,606]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF93D84368B6FD81FBD06D71" bold="true" box="[942,1095,586,606]" italics="true" pageId="12" pageNumber="688">L. chalconotus</emphasis>
</taxonomicName>
)
</td>
<td id="76DB5DE7FF9327B06F66FD80FA6E6D72" box="[1150,1529,586,606]" gridcol="3" gridrow="1" pageId="12" pageNumber="688">
Foveaux shag (
<taxonomicName id="4C3A4D5AFF93D8436E06FD81FA186D71" box="[1310,1423,586,606]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF93D8436E06FD81FA186D71" bold="true" box="[1310,1423,586,606]" italics="true" pageId="12" pageNumber="688">L. stewarti</emphasis>
</taxonomicName>
)
</td>
<td id="76DB5DE7FF9327B06D0BFD80F8846D72" box="[1555,1811,586,606]" gridcol="4" gridrow="1" pageId="12" pageNumber="688">References</td>
</tr>
<tr id="350A349BFF9327B06B65FD48F8846DBA" box="[125,1811,642,662]" gridrow="2" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65FD48FE356DBA" box="[125,418,642,662]" gridcol="0" gridrow="2" pageId="12" pageNumber="688">Prehistoric distribution</th>
<td id="76DB5DE7FF9327B06ACBFD48FC996DBA" box="[467,782,642,662]" gridcol="1" gridrow="2" pageId="12" pageNumber="688">Chatham Islands</td>
<td id="76DB5DE7FF9327B06830FD48FBCC6DBA" box="[808,1115,642,662]" gridcol="2" gridrow="2" pageId="12" pageNumber="688">Eastern South Island</td>
<td id="76DB5DE7FF9327B06F66FD48FA6E6DBA" box="[1150,1529,642,662]" gridcol="3" gridrow="2" pageId="12" pageNumber="688">Foveaux Strait</td>
<td id="76DB5DE7FF9327B06D0BFD48F8846DBA" box="[1555,1811,642,662]" gridcol="4" gridrow="2" pageId="12" pageNumber="688">
Rawlence 
<emphasis id="B94EEACBFF93D8436D66FD49F9246DB9" bold="true" box="[1662,1715,642,662]" italics="true" pageId="12" pageNumber="688">et al.</emphasis>
(2015)
</td>
</tr>
<tr id="350A349BFF9327B06B65FD55F8846D9F" box="[125,1811,671,691]" gridrow="3" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65FD55FE356D9F" box="[125,418,671,691]" gridcol="0" gridrow="3" pageId="12" pageNumber="688">Modern distribution</th>
<td id="76DB5DE7FF9327B06ACBFD55FC996D9F" box="[467,782,671,691]" gridcol="1" gridrow="3" pageId="12" pageNumber="688">Chatham Islands</td>
<td id="76DB5DE7FF9327B06830FD55FBCC6D9F" box="[808,1115,671,691]" gridcol="2" gridrow="3" pageId="12" pageNumber="688">Otago</td>
<td id="76DB5DE7FF9327B06F66FD55FA6E6D9F" box="[1150,1529,671,691]" gridcol="3" gridrow="3" pageId="12" pageNumber="688">Foveaux Strait</td>
<td id="76DB5DE7FF9327B06D0BFD55F8846D9F" box="[1555,1811,671,691]" gridcol="4" gridrow="3" pageId="12" pageNumber="688">Lalas (1983), Lalas &amp;</td>
</tr>
<tr id="350A349BFF9327B06B65FD77F8846DFC" box="[125,1811,701,720]" gridrow="4" pageId="12" pageNumber="688" rowspan-0="1" rowspan-1="1" rowspan-2="1" rowspan-3="1">
<td id="76DB5DE7FF9327B06D0BFD77F8846DFC" box="[1555,1811,701,720]" gridcol="4" gridrow="4" pageId="12" pageNumber="688">Perriman (2009),</td>
</tr>
<tr id="350A349BFF9327B06B65FD10F8846DC1" box="[125,1811,730,749]" gridrow="5" pageId="12" pageNumber="688" rowspan-0="1" rowspan-1="1" rowspan-2="1" rowspan-3="1">
<td id="76DB5DE7FF9327B06D0BFD10F8846DC1" box="[1555,1811,730,749]" gridcol="4" gridrow="5" pageId="12" pageNumber="688">
Rawlence 
<emphasis id="B94EEACBFF93D8436D92FD10F9286DC1" bold="true" box="[1674,1727,730,749]" italics="true" pageId="12" pageNumber="688">et al.</emphasis>
(2014)
</td>
</tr>
<tr id="350A349BFF9327B06B65FD3DF8846C27" box="[125,1811,759,779]" gridrow="6" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65FD3DFE356C27" box="[125,418,759,779]" gridcol="0" gridrow="6" pageId="12" pageNumber="688">DNA [fixed mtDNA control</th>
<td id="76DB5DE7FF9327B06ACBFD3DFC996C27" box="[467,782,759,779]" gridcol="1" gridrow="6" pageId="12" pageNumber="688">Monophyletic clade ‘daughter’</td>
<td id="76DB5DE7FF9327B06830FD3DFBCC6C27" box="[808,1115,759,779]" gridcol="2" gridrow="6" pageId="12" pageNumber="688">Monophyletic clade,</td>
<td id="76DB5DE7FF9327B06F66FD3DFA6E6C27" box="[1150,1529,759,779]" gridcol="3" gridrow="6" pageId="12" pageNumber="688">Monophyletic clade basal to</td>
<td id="76DB5DE7FF9327B06D0BFD3DF8846C27" box="[1555,1811,759,779]" gridcol="4" gridrow="6" pageId="12" pageNumber="688">
Rawlence 
<emphasis id="B94EEACBFF93D8436D66FD3DF9246C26" bold="true" box="[1662,1715,759,778]" italics="true" pageId="12" pageNumber="688">et al.</emphasis>
(2014,
</td>
</tr>
<tr id="350A349BFF9327B06B65FCDEF8846C04" box="[125,1811,788,808]" gridrow="7" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65FCDEFE356C04" box="[125,418,788,808]" gridcol="0" gridrow="7" pageId="12" pageNumber="688">
region 1 (
<emphasis id="B94EEACBFF93D8436BF5FCDEFE8D6C0B" bold="true" box="[237,282,788,807]" italics="true" pageId="12" pageNumber="688">CR1</emphasis>
) differences,
</th>
<td id="76DB5DE7FF9327B06ACBFCDEFC996C04" box="[467,782,788,808]" gridcol="1" gridrow="7" pageId="12" pageNumber="688">
to 
<taxonomicName id="4C3A4D5AFF93D8436AE2FCDFFD056C0B" box="[506,658,788,808]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF93D8436AE2FCDFFD056C0B" bold="true" box="[506,658,788,808]" italics="true" pageId="12" pageNumber="688">L. chalconotus</emphasis>
</taxonomicName>
.
</td>
<td id="76DB5DE7FF9327B06830FCDEFBCC6C04" box="[808,1115,788,808]" gridcol="2" gridrow="7" pageId="12" pageNumber="688">distinguished from</td>
<td id="76DB5DE7FF9327B06F66FCDEFA6E6C04" box="[1150,1529,788,808]" gridcol="3" gridrow="7" pageId="12" pageNumber="688">
<taxonomicName id="4C3A4D5AFF93D8436F91FCDFFAB56C0B" box="[1161,1314,788,808]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF93D8436F91FCDFFAB56C0B" bold="true" box="[1161,1314,788,808]" italics="true" pageId="12" pageNumber="688">L. chalconotus</emphasis>
</taxonomicName>
/ 
<emphasis id="B94EEACBFF93D8436E30FCDFFA0C6C0B" bold="true" box="[1320,1435,788,808]" italics="true" pageId="12" pageNumber="688">
<taxonomicName id="4C3A4D5AFF93D8436E30FCDFFA0F6C0B" box="[1320,1432,788,808]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="onslowi">L. onslowi</taxonomicName>
.
</emphasis>
</td>
<td id="76DB5DE7FF9327B06D0BFCDEF8846C04" box="[1555,1811,788,808]" gridcol="4" gridrow="7" pageId="12" pageNumber="688">2015)</td>
</tr>
<tr id="350A349BFF9327B06B65FCFBF8846C69" box="[125,1811,817,837]" gridrow="8" pageId="12" pageNumber="688" rowspan-4="1">
<th id="76DB5DE7FF9327B06B65FCFBFE356C69" box="[125,418,817,837]" gridcol="0" gridrow="8" pageId="12" pageNumber="688">
248 bp; 
<emphasis id="B94EEACBFF93D8436BC5FCFBFEA66C68" bold="true" box="[221,305,817,836]" pageId="12" pageNumber="688">full CR</emphasis>
, 1040 bp.
</th>
<td id="76DB5DE7FF9327B06ACBFCFBFC996C69" box="[467,782,817,837]" gridcol="1" gridrow="8" pageId="12" pageNumber="688">Distinguished from</td>
<td id="76DB5DE7FF9327B06830FCFBFBCC6C69" box="[808,1115,817,837]" gridcol="2" gridrow="8" pageId="12" pageNumber="688">
<taxonomicName id="4C3A4D5AFF93D843682CFCF8FC326C69" box="[820,933,818,837]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF93D843682CFCF8FC326C69" bold="true" box="[820,933,818,837]" italics="true" pageId="12" pageNumber="688">L. stewarti</emphasis>
</taxonomicName>
by C-T (92), G-A
</td>
<td id="76DB5DE7FF9327B06F66FCFBFA6E6C69" box="[1150,1529,817,837]" gridcol="3" gridrow="8" pageId="12" pageNumber="688">
Distinguished from 
<taxonomicName id="4C3A4D5AFF93D8436E43FCF8FA636C68" box="[1371,1524,817,837]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF93D8436E43FCF8FA636C68" bold="true" box="[1371,1524,817,837]" italics="true" pageId="12" pageNumber="688">L. chalconotus</emphasis>
</taxonomicName>
</td>
</tr>
<tr id="350A349BFF9327B06B65FC84F8846C4E" box="[125,1811,846,866]" gridrow="9" pageId="12" pageNumber="688" rowspan-4="1">
<th id="76DB5DE7FF9327B06B65FC84FE356C4E" box="[125,418,846,866]" gridcol="0" gridrow="9" pageId="12" pageNumber="688">Numbers in parentheses</th>
<td id="76DB5DE7FF9327B06ACBFC84FC996C4E" box="[467,782,846,866]" gridcol="1" gridrow="9" pageId="12" pageNumber="688">
<taxonomicName id="4C3A4D5AFF93D8436AC6FC85FDE06C4D" box="[478,631,846,866]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF93D8436AC6FC85FDE06C4D" bold="true" box="[478,631,846,866]" italics="true" pageId="12" pageNumber="688">L. chalconotus</emphasis>
</taxonomicName>
by C-T (40),
</td>
<td id="76DB5DE7FF9327B06830FC84FBCC6C4E" box="[808,1115,846,866]" gridcol="2" gridrow="9" pageId="12" pageNumber="688">(110), C-T (148), T-C (229),</td>
<td id="76DB5DE7FF9327B06F66FC84FA6E6C4E" box="[1150,1529,846,866]" gridcol="3" gridrow="9" pageId="12" pageNumber="688">by T-C (92), A-G (110), T-C (148),</td>
</tr>
<tr id="350A349BFF9327B06B65FCA6F8846C53" box="[125,1811,876,895]" gridrow="10" pageId="12" pageNumber="688" rowspan-4="1">
<th id="76DB5DE7FF9327B06B65FCA6FE356C53" box="[125,418,876,895]" gridcol="0" gridrow="10" pageId="12" pageNumber="688">represent the base pair</th>
<td id="76DB5DE7FF9327B06ACBFCA6FC996C53" box="[467,782,876,895]" gridcol="1" gridrow="10" pageId="12" pageNumber="688">T-C (50), and C-T (105).</td>
<td id="76DB5DE7FF9327B06830FCA6FBCC6C53" box="[808,1115,876,895]" gridcol="2" gridrow="10" pageId="12" pageNumber="688">
<emphasis id="B94EEACBFF93D843682CFCA6FCC86C53" bold="true" box="[820,863,876,895]" pageId="12" pageNumber="688">G-A</emphasis>
(
<emphasis id="B94EEACBFF93D8436876FCA6FC026C53" bold="true" box="[878,917,876,895]" pageId="12" pageNumber="688">493</emphasis>
), and 
<emphasis id="B94EEACBFF93D84368C1FCA6FB956C53" bold="true" box="[985,1026,876,895]" pageId="12" pageNumber="688">T-C</emphasis>
(
<emphasis id="B94EEACBFF93D8436F0AFCA6FBAE6C53" bold="true" box="[1042,1081,876,895]" pageId="12" pageNumber="688">628</emphasis>
).
</td>
<td id="76DB5DE7FF9327B06F66FCA6FA6E6C53" box="[1150,1529,876,895]" gridcol="3" gridrow="10" pageId="12" pageNumber="688">
C-T (229), 
<emphasis id="B94EEACBFF93D8436FE2FCA6FAB26C53" bold="true" box="[1274,1317,876,895]" pageId="12" pageNumber="688">A-G</emphasis>
(
<emphasis id="B94EEACBFF93D8436E2CFCA6FACC6C53" bold="true" box="[1332,1371,876,895]" pageId="12" pageNumber="688">493</emphasis>
), and 
<emphasis id="B94EEACBFF93D8436E87FCA6FA5F6C53" bold="true" box="[1439,1480,876,895]" pageId="12" pageNumber="688">C-T</emphasis>
</td>
</tr>
<tr id="350A349BFF9327B06B65FC43F8846CB0" box="[125,1811,905,924]" gridrow="11" pageId="12" pageNumber="688" rowspan-4="1">
<th id="76DB5DE7FF9327B06B65FC43FE356CB0" box="[125,418,905,924]" gridcol="0" gridrow="11" pageId="12" pageNumber="688">position (e.g. 40) in the</th>
<td id="76DB5DE7FF9327B06ACBFC43FC996CB0" box="[467,782,905,924]" gridcol="1" gridrow="11" pageId="12" pageNumber="688">Distinguished from</td>
<td id="76DB5DE7FF9327B06830FC43FBCC6CB0" box="[808,1115,905,924]" gridcol="2" gridrow="11" pageId="12" pageNumber="688">Distinguished from</td>
<td id="76DB5DE7FF9327B06F66FC43FA6E6CB0" box="[1150,1529,905,924]" gridcol="3" gridrow="11" pageId="12" pageNumber="688">
(
<emphasis id="B94EEACBFF93D8436F89FC43FB2F6CB0" bold="true" box="[1169,1208,905,924]" pageId="12" pageNumber="688">628</emphasis>
). Distinguished from
</td>
</tr>
<tr id="350A349BFF9327B06B65FC6CF8846C96" box="[125,1811,934,954]" gridrow="12" pageId="12" pageNumber="688" rowspan-4="1">
<th id="76DB5DE7FF9327B06B65FC6CFE356C96" box="[125,418,934,954]" gridcol="0" gridrow="12" pageId="12" pageNumber="688">
<emphasis id="B94EEACBFF93D8436B91FC6CFF216C95" bold="true" box="[137,182,934,953]" italics="true" pageId="12" pageNumber="688">CR1</emphasis>
sequence where the
</th>
<td id="76DB5DE7FF9327B06ACBFC6CFC996C96" box="[467,782,934,954]" gridcol="1" gridrow="12" pageId="12" pageNumber="688">
<taxonomicName id="4C3A4D5AFF93D8436AC6FC6DFDC76C95" box="[478,592,934,954]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF93D8436AC6FC6DFDC76C95" bold="true" box="[478,592,934,954]" italics="true" pageId="12" pageNumber="688">L. stewarti</emphasis>
</taxonomicName>
by T-C (24), C-T
</td>
<td id="76DB5DE7FF9327B06830FC6CFBCC6C96" box="[808,1115,934,954]" gridcol="2" gridrow="12" pageId="12" pageNumber="688">
<taxonomicName id="4C3A4D5AFF93D843682CFC6DFC376C95" box="[820,928,934,954]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="onslowi">
<emphasis id="B94EEACBFF93D843682CFC6DFC376C95" bold="true" box="[820,928,934,954]" italics="true" pageId="12" pageNumber="688">L. onslowi</emphasis>
</taxonomicName>
by T-C (40), C-T
</td>
<td id="76DB5DE7FF9327B06F66FC6CFA6E6C96" box="[1150,1529,934,954]" gridcol="3" gridrow="12" pageId="12" pageNumber="688">
<taxonomicName id="4C3A4D5AFF93D8436F91FC6DFB616C95" box="[1161,1270,934,954]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="onslowi">
<emphasis id="B94EEACBFF93D8436F91FC6DFB616C95" bold="true" box="[1161,1270,934,954]" italics="true" pageId="12" pageNumber="688">L. onslowi</emphasis>
</taxonomicName>
by C-T (24), T-C (40),
</td>
</tr>
<tr id="350A349BFF9327B06B65FC0EF8846CFB" box="[125,1811,964,983]" gridrow="13" pageId="12" pageNumber="688" rowspan-4="1">
<th id="76DB5DE7FF9327B06B65FC0EFE356CFB" box="[125,418,964,983]" gridcol="0" gridrow="13" pageId="12" pageNumber="688">fixed difference (e.g. C-T)</th>
<td id="76DB5DE7FF9327B06ACBFC0EFC996CFB" box="[467,782,964,983]" gridcol="1" gridrow="13" pageId="12" pageNumber="688">(40), C-T (92), C-T (105),</td>
<td id="76DB5DE7FF9327B06830FC0EFBCC6CFB" box="[808,1115,964,983]" gridcol="2" gridrow="13" pageId="12" pageNumber="688">(50), and T-C (105)</td>
<td id="76DB5DE7FF9327B06F66FC0EFA6E6CFB" box="[1150,1529,964,983]" gridcol="3" gridrow="13" pageId="12" pageNumber="688">T-C (92), T-C (105), A-G (110), C-T</td>
</tr>
<tr id="350A349BFF9327B06B65FC2BF8846CD8" box="[125,1811,993,1012]" gridrow="14" pageId="12" pageNumber="688" rowspan-2="1" rowspan-4="1">
<th id="76DB5DE7FF9327B06B65FC2BFE356CD8" box="[125,418,993,1012]" gridcol="0" gridrow="14" pageId="12" pageNumber="688">occurs.</th>
<td id="76DB5DE7FF9327B06ACBFC2BFC996CD8" box="[467,782,993,1012]" gridcol="1" gridrow="14" pageId="12" pageNumber="688">G-A (110), T-C (117),</td>
<td id="76DB5DE7FF9327B06F66FC2BFA6E6CD8" box="[1150,1529,993,1012]" gridcol="3" gridrow="14" pageId="12" pageNumber="688">
(117), T-C (148), C-T (229), 
<emphasis id="B94EEACBFF93D8436EB5FC2BFA4F6CD8" bold="true" box="[1453,1496,993,1012]" pageId="12" pageNumber="688">A-G</emphasis>
</td>
</tr>
<tr id="350A349BFF9327B06B65FC34F8846B3D" box="[125,1811,1022,1041]" gridrow="15" pageId="12" pageNumber="688" rowspan-0="1" rowspan-2="1" rowspan-4="1">
<td id="76DB5DE7FF9327B06ACBFC34FC996B3D" box="[467,782,1022,1041]" gridcol="1" gridrow="15" pageId="12" pageNumber="688">
C-T (148), T-C (229), 
<emphasis id="B94EEACBFF93D84369A7FC34FD7D6B3D" bold="true" box="[703,746,1022,1041]" pageId="12" pageNumber="688">G-A</emphasis>
</td>
<td id="76DB5DE7FF9327B06F66FC34FA6E6B3D" box="[1150,1529,1022,1041]" gridcol="3" gridrow="15" pageId="12" pageNumber="688">
(
<emphasis id="B94EEACBFF93D8436F89FC34FB2F6B3D" bold="true" box="[1169,1208,1022,1041]" pageId="12" pageNumber="688">493</emphasis>
), and 
<emphasis id="B94EEACBFF93D8436FE4FC34FAB26B3D" bold="true" box="[1276,1317,1022,1041]" pageId="12" pageNumber="688">C-T</emphasis>
(
<emphasis id="B94EEACBFF93D8436E2CFC34FACC6B3D" bold="true" box="[1332,1371,1022,1041]" pageId="12" pageNumber="688">628</emphasis>
).
</td>
</tr>
<tr id="350A349BFF9327B06B65FBD1F8846B03" box="[125,1811,1051,1071]" gridrow="16" pageId="12" pageNumber="688" rowspan-0="1" rowspan-2="1" rowspan-3="1" rowspan-4="1">
<td id="76DB5DE7FF9327B06ACBFBD1FC996B03" box="[467,782,1051,1071]" gridcol="1" gridrow="16" pageId="12" pageNumber="688">
(
<emphasis id="B94EEACBFF93D8436AFEFBD1FD9A6B02" bold="true" box="[486,525,1051,1070]" pageId="12" pageNumber="688">493</emphasis>
), and 
<emphasis id="B94EEACBFF93D8436949FBD1FDED6B02" bold="true" box="[593,634,1051,1070]" pageId="12" pageNumber="688">T-C</emphasis>
(
<emphasis id="B94EEACBFF93D8436991FBD1FD276B02" bold="true" box="[649,688,1051,1070]" pageId="12" pageNumber="688">628</emphasis>
).
</td>
</tr>
<tr id="350A349BFF9327B06B65FBF3F8846B60" box="[125,1811,1081,1100]" gridrow="17" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65FBF3FE356B60" box="[125,418,1081,1100]" gridcol="0" gridrow="17" pageId="12" pageNumber="688">Plumage morphs</th>
<td id="76DB5DE7FF9327B06ACBFBF3FC996B60" box="[467,782,1081,1100]" gridcol="1" gridrow="17" pageId="12" pageNumber="688">Pied (black and white)</td>
<td id="76DB5DE7FF9327B06830FBF3FBCC6B60" box="[808,1115,1081,1100]" gridcol="2" gridrow="17" pageId="12" pageNumber="688">Pied (black and white),</td>
<td id="76DB5DE7FF9327B06F66FBF3FA6E6B60" box="[1150,1529,1081,1100]" gridcol="3" gridrow="17" pageId="12" pageNumber="688">Pied (black and white), bronze</td>
<td id="76DB5DE7FF9327B06D0BFBF3F8846B60" box="[1555,1811,1081,1100]" gridcol="4" gridrow="17" pageId="12" pageNumber="688">Lalas (1983); Marchant</td>
</tr>
<tr id="350A349BFF9327B06B65FB9CF8846B45" box="[125,1811,1110,1129]" gridrow="18" pageId="12" pageNumber="688" rowspan-0="1" rowspan-1="1">
<td id="76DB5DE7FF9327B06830FB9CFBCC6B45" box="[808,1115,1110,1129]" gridcol="2" gridrow="18" pageId="12" pageNumber="688">bronze (wholly dark)</td>
<td id="76DB5DE7FF9327B06F66FB9CFA6E6B45" box="[1150,1529,1110,1129]" gridcol="3" gridrow="18" pageId="12" pageNumber="688">(wholly dark)</td>
<td id="76DB5DE7FF9327B06D0BFB9CF8846B45" box="[1555,1811,1110,1129]" gridcol="4" gridrow="18" pageId="12" pageNumber="688">&amp; Higgins (1990);</td>
</tr>
<tr id="350A349BFF9327B06B65FBB9F8846BAA" box="[125,1811,1139,1158]" gridrow="19" pageId="12" pageNumber="688" rowspan-0="1" rowspan-1="1" rowspan-2="1" rowspan-3="1">
<td id="76DB5DE7FF9327B06D0BFBB9F8846BAA" box="[1555,1811,1139,1158]" gridcol="4" gridrow="19" pageId="12" pageNumber="688">
Rawlence 
<emphasis id="B94EEACBFF93D8436D92FBB9F9286BAA" bold="true" box="[1674,1727,1139,1158]" italics="true" pageId="12" pageNumber="688">et al.</emphasis>
(2014)
</td>
</tr>
<tr id="350A349BFF9327B06B65FB5AF8846B88" box="[125,1811,1168,1188]" gridrow="20" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65FB5AFE356B88" box="[125,418,1168,1188]" gridcol="0" gridrow="20" pageId="12" pageNumber="688">Plumage morph</th>
<td id="76DB5DE7FF9327B06ACBFB5AFC996B88" box="[467,782,1168,1188]" gridcol="1" gridrow="20" pageId="12" pageNumber="688">100% pied</td>
<td id="76DB5DE7FF9327B06830FB5AFBCC6B88" box="[808,1115,1168,1188]" gridcol="2" gridrow="20" pageId="12" pageNumber="688">~20–30% pied</td>
<td id="76DB5DE7FF9327B06F66FB5AFA6E6B88" box="[1150,1529,1168,1188]" gridcol="3" gridrow="20" pageId="12" pageNumber="688">~50–60% pied</td>
<td id="76DB5DE7FF9327B06D0BFB5AF8846B88" box="[1555,1811,1168,1188]" gridcol="4" gridrow="20" pageId="12" pageNumber="688">Lalas (1983); Rawlence</td>
</tr>
<tr id="350A349BFF9327B06B65FB67F8846BED" box="[125,1811,1197,1217]" gridrow="21" pageId="12" pageNumber="688" rowspan-1="1" rowspan-2="1" rowspan-3="1">
<th id="76DB5DE7FF9327B06B65FB67FE356BED" box="[125,418,1197,1217]" gridcol="0" gridrow="21" pageId="12" pageNumber="688">percentages</th>
<td id="76DB5DE7FF9327B06D0BFB67F8846BED" box="[1555,1811,1197,1217]" gridcol="4" gridrow="21" pageId="12" pageNumber="688">
<emphasis id="B94EEACBFF93D8436D06FB64F9C46BEC" bold="true" box="[1566,1619,1197,1217]" italics="true" pageId="12" pageNumber="688">et al.</emphasis>
(2014)
</td>
</tr>
<tr id="350A349BFF9327B06B65FB01F8846BF2" box="[125,1811,1227,1246]" gridrow="22" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65FB01FE356BF2" box="[125,418,1227,1246]" gridcol="0" gridrow="22" pageId="12" pageNumber="688">Carunculation (breeding</th>
<td id="76DB5DE7FF9327B06ACBFB01FC996BF2" box="[467,782,1227,1246]" gridcol="1" gridrow="22" pageId="12" pageNumber="688">Pronounced bright orange</td>
<td id="76DB5DE7FF9327B06830FB01FBCC6BF2" box="[808,1115,1227,1246]" gridcol="2" gridrow="22" pageId="12" pageNumber="688">Equal frequencies of small,</td>
<td id="76DB5DE7FF9327B06F66FB01FA6E6BF2" box="[1150,1529,1227,1246]" gridcol="3" gridrow="22" pageId="12" pageNumber="688">Scattered dark orange papillae</td>
<td id="76DB5DE7FF9327B06D0BFB01F8846BF2" box="[1555,1811,1227,1246]" gridcol="4" gridrow="22" pageId="12" pageNumber="688">Lalas (1983); Rawlence</td>
</tr>
<tr id="350A349BFF9327B06B65FB22F8846BD7" box="[125,1811,1256,1275]" gridrow="23" pageId="12" pageNumber="688" rowspan-3="1">
<th id="76DB5DE7FF9327B06B65FB22FE356BD7" box="[125,418,1256,1275]" gridcol="0" gridrow="23" pageId="12" pageNumber="688">plumage)</th>
<td id="76DB5DE7FF9327B06ACBFB22FC996BD7" box="[467,782,1256,1275]" gridcol="1" gridrow="23" pageId="12" pageNumber="688">facial caruncles</td>
<td id="76DB5DE7FF9327B06830FB22FBCC6BD7" box="[808,1115,1256,1275]" gridcol="2" gridrow="23" pageId="12" pageNumber="688">bright orange facial</td>
<td id="76DB5DE7FF9327B06D0BFB22F8846BD7" box="[1555,1811,1256,1275]" gridcol="4" gridrow="23" pageId="12" pageNumber="688">
<emphasis id="B94EEACBFF93D8436D06FB22F9C46BD7" bold="true" box="[1566,1619,1256,1275]" italics="true" pageId="12" pageNumber="688">et al.</emphasis>
(2014)
</td>
</tr>
<tr id="350A349BFF9327B06B65FACFF8846A35" box="[125,1811,1285,1305]" gridrow="24" pageId="12" pageNumber="688" rowspan-0="1" rowspan-1="1" rowspan-3="1" rowspan-4="1">
<td id="76DB5DE7FF9327B06830FACFFBCC6A35" box="[808,1115,1285,1305]" gridcol="2" gridrow="24" pageId="12" pageNumber="688">caruncles and dark orange</td>
</tr>
<tr id="350A349BFF9327B06B65FAE8F8846A19" box="[125,1811,1314,1333]" gridrow="25" pageId="12" pageNumber="688" rowspan-0="1" rowspan-1="1" rowspan-3="1" rowspan-4="1">
<td id="76DB5DE7FF9327B06830FAE8FBCC6A19" box="[808,1115,1314,1333]" gridcol="2" gridrow="25" pageId="12" pageNumber="688">scattered papillae</td>
</tr>
<tr id="350A349BFF9327B06B65FA8AF8846A7F" box="[125,1811,1344,1363]" gridrow="26" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65FA8AFE356A7F" box="[125,418,1344,1363]" gridcol="0" gridrow="26" pageId="12" pageNumber="688">Gular pouch (breeding</th>
<td id="76DB5DE7FF9327B06ACBFA8AFC996A7F" box="[467,782,1344,1363]" gridcol="1" gridrow="26" pageId="12" pageNumber="688">Bright orange</td>
<td id="76DB5DE7FF9327B06830FA8AFBCC6A7F" box="[808,1115,1344,1363]" gridcol="2" gridrow="26" pageId="12" pageNumber="688">Bright to dark orange or</td>
<td id="76DB5DE7FF9327B06F66FA8AFA6E6A7F" box="[1150,1529,1344,1363]" gridcol="3" gridrow="26" pageId="12" pageNumber="688">Bright to dark orange</td>
<td id="76DB5DE7FF9327B06D0BFA8AF8846A7F" box="[1555,1811,1344,1363]" gridcol="4" gridrow="26" pageId="12" pageNumber="688">Lalas (1983); Rawlence</td>
</tr>
<tr id="350A349BFF9327B06B65FA96F8846A5C" box="[125,1811,1372,1392]" gridrow="27" pageId="12" pageNumber="688" rowspan-1="1" rowspan-3="1">
<th id="76DB5DE7FF9327B06B65FA96FE356A5C" box="[125,418,1372,1392]" gridcol="0" gridrow="27" pageId="12" pageNumber="688">plumage)</th>
<td id="76DB5DE7FF9327B06830FA96FBCC6A5C" box="[808,1115,1372,1392]" gridcol="2" gridrow="27" pageId="12" pageNumber="688">purple</td>
<td id="76DB5DE7FF9327B06D0BFA96F8846A5C" box="[1555,1811,1372,1392]" gridcol="4" gridrow="27" pageId="12" pageNumber="688">
<emphasis id="B94EEACBFF93D8436D06FA97F9C46A43" bold="true" box="[1566,1619,1372,1392]" italics="true" pageId="12" pageNumber="688">et al.</emphasis>
(2014)
</td>
</tr>
<tr id="350A349BFF9327B06B65FAB0F8846AA2" box="[125,1811,1402,1422]" gridrow="28" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65FAB0FE356AA2" box="[125,418,1402,1422]" gridcol="0" gridrow="28" pageId="12" pageNumber="688">Morphometics/allometries</th>
<td id="76DB5DE7FF9327B06ACBFAB0FC996AA2" box="[467,782,1402,1422]" gridcol="1" gridrow="28" pageId="12" pageNumber="688">Smaller on average than both</td>
<td id="76DB5DE7FF9327B06830FAB0FBCC6AA2" box="[808,1115,1402,1422]" gridcol="2" gridrow="28" pageId="12" pageNumber="688">Larger on average than both</td>
<td id="76DB5DE7FF9327B06F66FAB0FA6E6AA2" box="[1150,1529,1402,1422]" gridcol="3" gridrow="28" pageId="12" pageNumber="688">Smaller on average than</td>
<td id="76DB5DE7FF9327B06D0BFAB0F8846AA2" box="[1555,1811,1402,1422]" gridcol="4" gridrow="28" pageId="12" pageNumber="688">Lalas (1983); Rawlence</td>
</tr>
<tr id="350A349BFF9327B06B65FA5DF8846A87" box="[125,1811,1431,1451]" gridrow="29" pageId="12" pageNumber="688" rowspan-0="1">
<td id="76DB5DE7FF9327B06ACBFA5DFC996A87" box="[467,782,1431,1451]" gridcol="1" gridrow="29" pageId="12" pageNumber="688">
<taxonomicName id="4C3A4D5AFF93D8436AC6FA52FDE06A86" box="[478,631,1431,1451]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF93D8436AC6FA52FDE06A86" bold="true" box="[478,631,1431,1451]" italics="true" pageId="12" pageNumber="688">L. chalconotus</emphasis>
</taxonomicName>
and
</td>
<td id="76DB5DE7FF9327B06830FA5DFBCC6A87" box="[808,1115,1431,1451]" gridcol="2" gridrow="29" pageId="12" pageNumber="688">
<taxonomicName id="4C3A4D5AFF93D843682CFA52FC326A86" box="[820,933,1431,1451]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF93D843682CFA52FC326A86" bold="true" box="[820,933,1431,1451]" italics="true" pageId="12" pageNumber="688">L. stewarti</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D5AFF93D84368C4FA52FBDE6A86" box="[988,1097,1431,1451]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="onslowi">
<emphasis id="B94EEACBFF93D84368C4FA52FBDE6A86" bold="true" box="[988,1097,1431,1451]" italics="true" pageId="12" pageNumber="688">L. onslowi</emphasis>
</taxonomicName>
</td>
<td id="76DB5DE7FF9327B06F66FA5DFA6E6A87" box="[1150,1529,1431,1451]" gridcol="3" gridrow="29" pageId="12" pageNumber="688">
<taxonomicName id="4C3A4D5AFF93D8436F91FA52FAB56A86" box="[1161,1314,1431,1451]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="chalconotus">
<emphasis id="B94EEACBFF93D8436F91FA52FAB56A86" bold="true" box="[1161,1314,1431,1451]" italics="true" pageId="12" pageNumber="688">L. chalconotus</emphasis>
</taxonomicName>
, and similar in size
</td>
<td id="76DB5DE7FF9327B06D0BFA5DF8846A87" box="[1555,1811,1431,1451]" gridcol="4" gridrow="29" pageId="12" pageNumber="688">
<emphasis id="B94EEACBFF93D8436D06FA52F9C46A86" bold="true" box="[1566,1619,1431,1451]" italics="true" pageId="12" pageNumber="688">et al.</emphasis>
(2014); this study
</td>
</tr>
<tr id="350A349BFF9327B06B65FA7EF8846AE4" box="[125,1811,1460,1480]" gridrow="30" pageId="12" pageNumber="688" rowspan-0="1" rowspan-2="1" rowspan-4="1">
<td id="76DB5DE7FF9327B06ACBFA7EFC996AE4" box="[467,782,1460,1480]" gridcol="1" gridrow="30" pageId="12" pageNumber="688">
<taxonomicName id="4C3A4D5AFF93D8436AC6FA7FFDC76AEB" box="[478,592,1460,1480]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="stewarti">
<emphasis id="B94EEACBFF93D8436AC6FA7FFDC76AEB" bold="true" box="[478,592,1460,1480]" italics="true" pageId="12" pageNumber="688">L. stewarti</emphasis>
</taxonomicName>
</td>
<td id="76DB5DE7FF9327B06F66FA7EFA6E6AE4" box="[1150,1529,1460,1480]" gridcol="3" gridrow="30" pageId="12" pageNumber="688">
to 
<taxonomicName id="4C3A4D5AFF93D8436FBDFA7FFA866AEB" box="[1189,1297,1460,1480]" class="Aves" family="Phalacrocoracidae" genus="Leucocarbo" kingdom="Animalia" order="Suliformes" pageId="12" pageNumber="688" phylum="Chordata" rank="species" species="onslowi">
<emphasis id="B94EEACBFF93D8436FBDFA7FFA866AEB" bold="true" box="[1189,1297,1460,1480]" italics="true" pageId="12" pageNumber="688">L. onslowi</emphasis>
</taxonomicName>
</td>
</tr>
<tr id="350A349BFF9327B06B65FA18F8846AC9" box="[125,1811,1490,1509]" gridrow="31" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65FA18FE356AC9" box="[125,418,1490,1509]" gridcol="0" gridrow="31" pageId="12" pageNumber="688">Osteology</th>
<td id="76DB5DE7FF9327B06ACBFA18FC996AC9" box="[467,782,1490,1509]" gridcol="1" gridrow="31" pageId="12" pageNumber="688">The foramen vasculare</td>
<td id="76DB5DE7FF9327B06830FA18FBCC6AC9" box="[808,1115,1490,1509]" gridcol="2" gridrow="31" pageId="12" pageNumber="688">The foramen vasculare</td>
<td id="76DB5DE7FF9327B06F66FA18FA6E6AC9" box="[1150,1529,1490,1509]" gridcol="3" gridrow="31" pageId="12" pageNumber="688">The foramen vasculare distale on</td>
<td id="76DB5DE7FF9327B06D0BFA18F8846AC9" box="[1555,1811,1490,1509]" gridcol="4" gridrow="31" pageId="12" pageNumber="688">Worthy (2011); this</td>
</tr>
<tr id="350A349BFF9327B06B65FA25F884692E" box="[125,1811,1519,1538]" gridrow="32" pageId="12" pageNumber="688" rowspan-0="1">
<td id="76DB5DE7FF9327B06ACBFA25FC99692E" box="[467,782,1519,1538]" gridcol="1" gridrow="32" pageId="12" pageNumber="688">distale on the distal</td>
<td id="76DB5DE7FF9327B06830FA25FBCC692E" box="[808,1115,1519,1538]" gridcol="2" gridrow="32" pageId="12" pageNumber="688">distale on the distal</td>
<td id="76DB5DE7FF9327B06F66FA25FA6E692E" box="[1150,1529,1519,1538]" gridcol="3" gridrow="32" pageId="12" pageNumber="688">the distal tarsometatarsus is small</td>
<td id="76DB5DE7FF9327B06D0BFA25F884692E" box="[1555,1811,1519,1538]" gridcol="4" gridrow="32" pageId="12" pageNumber="688">study</td>
</tr>
<tr id="350A349BFF9327B06B65F9C6F884690C" box="[125,1811,1548,1568]" gridrow="33" pageId="12" pageNumber="688" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<td id="76DB5DE7FF9327B06ACBF9C6FC99690C" box="[467,782,1548,1568]" gridcol="1" gridrow="33" pageId="12" pageNumber="688">tarsometatarsus is small</td>
<td id="76DB5DE7FF9327B06830F9C6FBCC690C" box="[808,1115,1548,1568]" gridcol="2" gridrow="33" pageId="12" pageNumber="688">tarsometatarsus varies in</td>
</tr>
<tr id="350A349BFF9327B06B65F9E3F8846910" box="[125,1811,1577,1596]" gridrow="34" pageId="12" pageNumber="688" rowspan-0="1" rowspan-1="1" rowspan-3="1" rowspan-4="1">
<td id="76DB5DE7FF9327B06830F9E3FBCC6910" box="[808,1115,1577,1596]" gridcol="2" gridrow="34" pageId="12" pageNumber="688">size</td>
</tr>
<tr id="350A349BFF9327B06B65F98DF8846976" box="[125,1811,1607,1626]" gridrow="35" pageId="12" pageNumber="688">
<th id="76DB5DE7FF9327B06B65F98DFE356976" box="[125,418,1607,1626]" gridcol="0" gridrow="35" pageId="12" pageNumber="688">Onset of breeding season</th>
<td id="76DB5DE7FF9327B06ACBF98DFC996976" box="[467,782,1607,1626]" gridcol="1" gridrow="35" pageId="12" pageNumber="688">September–February</td>
<td id="76DB5DE7FF9327B06830F98DFBCC6976" box="[808,1115,1607,1626]" gridcol="2" gridrow="35" pageId="12" pageNumber="688">May–September</td>
<td id="76DB5DE7FF9327B06F66F98DFA6E6976" box="[1150,1529,1607,1626]" gridcol="3" gridrow="35" pageId="12" pageNumber="688">September onwards</td>
<td id="76DB5DE7FF9327B06D0BF98DF8846976" box="[1555,1811,1607,1626]" gridcol="4" gridrow="35" pageId="12" pageNumber="688">Lalas (1983)</td>
</tr>
</table>
</paragraph>
<subSubSection id="C3206552FF92D8426A02FB62FF41681C" pageId="13" pageNumber="689" type="conservation">
<paragraph id="8B8536D9FF92D8426A02FB62FDFC6B90" blockId="13.[282,619,1192,1213]" box="[282,619,1192,1213]" pageId="13" pageNumber="689">
<heading id="D0CD81B5FF92D8426A02FB62FDFC6B90" allCaps="true" box="[282,619,1192,1213]" centered="true" fontSize="8" level="6" pageId="13" pageNumber="689" reason="3">CONSERVATION IMPLICATIONS</heading>
</paragraph>
<paragraph id="8B8536D9FF92D8426B88FB1FFF41681C" blockId="13.[144,759,1237,1840]" pageId="13" pageNumber="689">
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 
<date id="FF841019FF92D8426BE5FA9AFE256A49" box="[253,434,1360,1382]" pageId="13" pageNumber="689" value="1600" valueMax="1800">1600 and 1800</date>
pairs to fewer than 5000 birds and threats including fisheries interactions and nest disturbance (
<bibRefCitation id="EFAB4B28FF92D8426AB8FA47FD7C6A8F" author="Birdlife International" box="[416,747,1421,1443]" journalOrPublisher="The IUCN red list of threatened species (Version 2014.3). www. iucnredlist. org" pageId="13" pageNumber="689" refString="Birdlife International. 2012. Phalacrocorax chalconotus. The IUCN red list of threatened species (Version 2014.3). www. iucnredlist. org." title="Phalacrocorax chalconotus" type="book" year="2012">Birdlife International, 2012</bibRefCitation>
). 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. 
<bibRefCitation id="EFAB4B28FF92D8426A86F9ECFD3D6910" author="Calderon L &amp; Quintana F &amp; Cabanne GS &amp; Lougheed SC &amp; Tubaro PL." box="[414,682,1573,1596]" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="13" pageNumber="689" pagination="42 - 53" part="72" refString="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." title="Phylogeography and genetic structure of two Patagonian shag species (Aves: Phalacrocoracidae)" type="journal article" year="2014">
Calderon 
<emphasis id="B94EEACBFF92D8426903F9ECFDC36917" bold="true" box="[539,596,1573,1595]" italics="true" pageId="13" pageNumber="689">et al.</emphasis>
, 2014
</bibRefCitation>
). 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 (
<bibRefCitation id="EFAB4B28FF92D8426AF2F96AFD7C699A" author="Rawlence NJ &amp; Kennedy M &amp; Anderson CNK &amp; Till CE &amp; Smith I &amp; Scofield RP &amp; Tennyson AJD &amp; Hamel J &amp; Lalas C &amp; Matisoo-Smith EA &amp; Waters JM." box="[490,747,1695,1718]" journalOrPublisher="Molecular Ecology" pageId="13" pageNumber="689" pagination="4605 - 4616" part="18" refString="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." title="Geographically contrasting biodiversity reductions in a widespread New Zealand seabird" type="journal article" year="2015">
Rawlence 
<emphasis id="B94EEACBFF92D842697CF96AFD0A6999" bold="true" box="[612,669,1695,1717]" italics="true" pageId="13" pageNumber="689">et al.</emphasis>
, 2015
</bibRefCitation>
). Although the Otago shag has recolonized part of its former distribution since the European era, its population size continues to decline (
<bibRefCitation id="EFAB4B28FF92D842690FF931FF5D681C" author="Lalas C &amp; Perriman L." journalOrPublisher="Department of Conservation Research and Development Series 314" pageId="13" pageNumber="689" pagination="1 - 30" refString="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." title="Nest counts of Stewart Island shags / mapua (Leucocarbo chalconotus) in Otago" type="book chapter" year="2009">Lalas &amp; Perriman, 2009</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>