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<front>
<journal-meta>
<journal-id journal-id-type="nlm-ta">PLoS ONE</journal-id>
<journal-id journal-id-type="publisher-id">plos</journal-id>
<journal-id journal-id-type="pmc">plosone</journal-id>
<journal-title-group>
<journal-title>PLOS ONE</journal-title>
</journal-title-group>
<issn pub-type="epub">1932-6203</issn>
<publisher>
<publisher-name>Public Library of Science</publisher-name>
<publisher-loc>San Francisco, CA USA</publisher-loc>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.1371/journal.pone.0247873</article-id>
<article-id pub-id-type="publisher-id">PONE-D-20-38105</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Research Article</subject>
</subj-group>
<subj-group subj-group-type="Discipline-v3">
<subject>Biology and life sciences</subject><subj-group><subject>Organisms</subject><subj-group><subject>Eukaryota</subject><subj-group><subject>Animals</subject><subj-group><subject>Vertebrates</subject><subj-group><subject>Amniotes</subject><subj-group><subject>Birds</subject></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>Biology and life sciences</subject><subj-group><subject>Zoology</subject><subj-group><subject>Animals</subject><subj-group><subject>Vertebrates</subject><subj-group><subject>Amniotes</subject><subj-group><subject>Birds</subject></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>People and places</subject><subj-group><subject>Geographical locations</subject><subj-group><subject>Oceania</subject><subj-group><subject>New Zealand</subject></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>Earth sciences</subject><subj-group><subject>Seasons</subject></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>Biology and life sciences</subject><subj-group><subject>Psychology</subject><subj-group><subject>Behavior</subject><subj-group><subject>Habits</subject><subj-group><subject>Nesting habits</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>Social sciences</subject><subj-group><subject>Psychology</subject><subj-group><subject>Behavior</subject><subj-group><subject>Habits</subject><subj-group><subject>Nesting habits</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>Biology and life sciences</subject><subj-group><subject>Evolutionary biology</subject><subj-group><subject>Evolutionary processes</subject><subj-group><subject>Speciation</subject><subj-group><subject>Cryptic speciation</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>Biology and life sciences</subject><subj-group><subject>Psychology</subject><subj-group><subject>Behavior</subject><subj-group><subject>Animal behavior</subject><subj-group><subject>Animal sexual behavior</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>Social sciences</subject><subj-group><subject>Psychology</subject><subj-group><subject>Behavior</subject><subj-group><subject>Animal behavior</subject><subj-group><subject>Animal sexual behavior</subject></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>Biology and life sciences</subject><subj-group><subject>Zoology</subject><subj-group><subject>Animal behavior</subject><subj-group><subject>Animal sexual behavior</subject></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>Biology and life sciences</subject><subj-group><subject>Organisms</subject><subj-group><subject>Eukaryota</subject><subj-group><subject>Animals</subject><subj-group><subject>Vertebrates</subject><subj-group><subject>Amniotes</subject><subj-group><subject>Birds</subject><subj-group><subject>Passerines</subject></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>Biology and life sciences</subject><subj-group><subject>Zoology</subject><subj-group><subject>Animals</subject><subj-group><subject>Vertebrates</subject><subj-group><subject>Amniotes</subject><subj-group><subject>Birds</subject><subj-group><subject>Passerines</subject></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group></subj-group><subj-group subj-group-type="Discipline-v3">
<subject>Ecology and environmental sciences</subject><subj-group><subject>Conservation science</subject></subj-group></subj-group></article-categories>
<title-group>
<article-title>Evaluation of counting methods for monitoring populations of a cryptic alpine passerine, the rock wren (Passeriformes, Acanthisittidae, <italic>Xenicus gilviventris</italic>)</article-title>
<alt-title alt-title-type="running-head">Monitoring techniques for alpine passerines</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes" xlink:type="simple">
<contrib-id authenticated="true" contrib-id-type="orcid">https://orcid.org/0000-0001-9391-380X</contrib-id>
<name name-style="western">
<surname>Monks</surname>
<given-names>Joanne M.</given-names>
</name>
<role content-type="https://casrai.org/credit/">Conceptualization</role>
<role content-type="https://casrai.org/credit/">Data curation</role>
<role content-type="https://casrai.org/credit/">Formal analysis</role>
<role content-type="https://casrai.org/credit/">Funding acquisition</role>
<role content-type="https://casrai.org/credit/">Investigation</role>
<role content-type="https://casrai.org/credit/">Methodology</role>
<role content-type="https://casrai.org/credit/">Writing – original draft</role>
<role content-type="https://casrai.org/credit/">Writing – review &amp; editing</role>
<xref ref-type="aff" rid="aff001"><sup>1</sup></xref>
<xref ref-type="corresp" rid="cor001">*</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>O’Donnell</surname>
<given-names>Colin F. J.</given-names>
</name>
<role content-type="https://casrai.org/credit/">Conceptualization</role>
<role content-type="https://casrai.org/credit/">Data curation</role>
<role content-type="https://casrai.org/credit/">Formal analysis</role>
<role content-type="https://casrai.org/credit/">Funding acquisition</role>
<role content-type="https://casrai.org/credit/">Investigation</role>
<role content-type="https://casrai.org/credit/">Methodology</role>
<role content-type="https://casrai.org/credit/">Writing – original draft</role>
<xref ref-type="aff" rid="aff002"><sup>2</sup></xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple">
<contrib-id authenticated="true" contrib-id-type="orcid">https://orcid.org/0000-0002-4183-0719</contrib-id>
<name name-style="western">
<surname>Greene</surname>
<given-names>Terry C.</given-names>
</name>
<role content-type="https://casrai.org/credit/">Conceptualization</role>
<role content-type="https://casrai.org/credit/">Formal analysis</role>
<role content-type="https://casrai.org/credit/">Methodology</role>
<role content-type="https://casrai.org/credit/">Writing – review &amp; editing</role>
<xref ref-type="aff" rid="aff002"><sup>2</sup></xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Weston</surname>
<given-names>Kerry A.</given-names>
</name>
<role content-type="https://casrai.org/credit/">Conceptualization</role>
<role content-type="https://casrai.org/credit/">Data curation</role>
<role content-type="https://casrai.org/credit/">Formal analysis</role>
<role content-type="https://casrai.org/credit/">Investigation</role>
<role content-type="https://casrai.org/credit/">Project administration</role>
<role content-type="https://casrai.org/credit/">Writing – original draft</role>
<xref ref-type="aff" rid="aff002"><sup>2</sup></xref>
</contrib>
</contrib-group>
<aff id="aff001"><label>1</label> <addr-line>Biodiversity Group, Department of Conservation, Dunedin, New Zealand</addr-line></aff>
<aff id="aff002"><label>2</label> <addr-line>Biodiversity Group, Department of Conservation, Christchurch, New Zealand</addr-line></aff>
<contrib-group>
<contrib contrib-type="editor" xlink:type="simple">
<name name-style="western">
<surname>Yue</surname>
<given-names>Bi-Song</given-names>
</name>
<role>Editor</role>
<xref ref-type="aff" rid="edit1"/>
</contrib>
</contrib-group>
<aff id="edit1"><addr-line>Sichuan University, CHINA</addr-line></aff>
<author-notes>
<fn fn-type="conflict" id="coi001">
<p>All authors are employees of the government department that funded this work.</p>
</fn>
<corresp id="cor001">* E-mail: <email xlink:type="simple">jmonks@doc.govt.nz</email></corresp>
</author-notes>
<pub-date pub-type="epub">
<day>4</day>
<month>3</month>
<year>2021</year>
</pub-date>
<pub-date pub-type="collection">
<year>2021</year>
</pub-date>
<volume>16</volume>
<issue>3</issue>
<elocation-id>e0247873</elocation-id>
<history>
<date date-type="received">
<day>3</day>
<month>12</month>
<year>2020</year>
</date>
<date date-type="accepted">
<day>16</day>
<month>2</month>
<year>2021</year>
</date>
</history>
<permissions>
<copyright-year>2021</copyright-year>
<copyright-holder>Monks et al</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
<license-p>This is an open access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">Creative Commons Attribution License</ext-link>, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
</license>
</permissions>
<self-uri content-type="pdf" xlink:href="info:doi/10.1371/journal.pone.0247873"/>
<abstract>
<p>Developing and validating methods to determine trends in populations of threatened species is essential for evaluating the effectiveness of conservation interventions. For cryptic species inhabiting remote environments, this can be particularly challenging. Rock wrens, <italic>Xenicus gilviventris</italic>, are small passerines endemic to the alpine zone of southern New Zealand. They are highly vulnerable to predation by introduced mammalian predators. Establishing a robust, cost-effective monitoring tool to evaluate population trends in rock wrens is a priority for conservation of both the species and, more broadly, as part of a suite of indicators for evaluating effectiveness of management in New Zealand’s alpine ecosystems. We assessed the relative accuracy and precision of three population estimation techniques (mark-resight, distance sampling and simple counts on line transects) for two populations of rock wrens in the Southern Alps over six breeding seasons (2012–2018). The performance of these population estimators was compared to known rock wren population size derived from simultaneous territory mapping. Indices of abundance derived from counts on transects were correlated with territory mapping at both study areas, and performed better than either mark-resight methods or distance sampling. Simple counts on standardised line transects are a highly cost-effective method of monitoring birds because they do not require banding a population. As such, we recommend that line transect counts using the design outlined in this paper be adopted as a standard method for long-term monitoring of rock wren populations. Although species-specific testing is required to validate use of low-cost population indices, our results may have utility for the monitoring of other cryptic passerines in relatively open habitats.</p>
</abstract>
<funding-group>
<award-group id="award001">
<funding-source>
<institution>New Zealand Department of Conservation</institution>
</funding-source>
</award-group>
<funding-statement>The New Zealand Department of Conservation (<ext-link ext-link-type="uri" xlink:href="https://www.doc.govt.nz/" xlink:type="simple">https://www.doc.govt.nz/</ext-link>) funded the work presented in this paper. The funder did not play any role in any aspect of the research or manuscript preparation, but all work was done by employees of the government department that funded the work.</funding-statement>
</funding-group>
<counts>
<fig-count count="3"/>
<table-count count="5"/>
<page-count count="18"/>
</counts>
<custom-meta-group>
<custom-meta id="data-availability">
<meta-name>Data Availability</meta-name>
<meta-value>All relevant data are within the paper and its <xref ref-type="sec" rid="sec027">Supporting Information</xref> files.</meta-value>
</custom-meta>
</custom-meta-group>
</article-meta>
</front>
<body>
<sec id="sec001" sec-type="intro">
<title>Introduction</title>
<p>Developing monitoring methods for threatened species that are difficult to detect (i.e. ‘cryptic’) can be problematic, particularly in challenging environments. However, information on population trends is one of the metrics used in assessing species status (e.g. by the International Union for the Conservation of Nature) and is essential for evaluating the effectiveness of conservation interventions. Therefore, development and validation of an appropriate method to estimate abundance and derived population trends, that is achievable within budget constraints, is vital for all threatened species prioritised for management [<xref ref-type="bibr" rid="pone.0247873.ref001">1</xref>–<xref ref-type="bibr" rid="pone.0247873.ref004">4</xref>].</p>
<p>Absolute measures of population abundance and density are often extremely difficult and costly to obtain. There has been considerable recent debate over the assumptions and application of indices [<xref ref-type="bibr" rid="pone.0247873.ref005">5</xref>] as well as development of estimation methods that explicitly address concerns regarding variable detectability [<xref ref-type="bibr" rid="pone.0247873.ref006">6</xref>–<xref ref-type="bibr" rid="pone.0247873.ref008">8</xref>]. However, newer methods often involve restrictive assumptions, complex field designs and analyses and, therefore, high costs [<xref ref-type="bibr" rid="pone.0247873.ref009">9</xref>]. Cost-effective and robust methods that can be used to confidently detect population trends are essential for conservation managers [<xref ref-type="bibr" rid="pone.0247873.ref003">3</xref>,<xref ref-type="bibr" rid="pone.0247873.ref010">10</xref>].</p>
<p>Alpine ecosystems in New Zealand are under increasing pressure from the interacting effects of climate change, invasive browsers and predators [<xref ref-type="bibr" rid="pone.0247873.ref011">11</xref>]. Yet, information about alpine ecology is lacking. Despite encompassing 11% of the land mass of New Zealand [<xref ref-type="bibr" rid="pone.0247873.ref012">12</xref>], a dearth of biodiversity monitoring in the alpine zone means that no alpine taxa are included in national scale trend reporting [<xref ref-type="bibr" rid="pone.0247873.ref013">13</xref>]. The development of robust and logistically feasible monitoring methods for species above the timberline has been identified as an urgent requirement [<xref ref-type="bibr" rid="pone.0247873.ref011">11</xref>,<xref ref-type="bibr" rid="pone.0247873.ref012">12</xref>,<xref ref-type="bibr" rid="pone.0247873.ref014">14</xref>] so that a suite of alpine indicators can be used to measure trends, including response to management, in this nationally significant ecosystem.</p>
<p>Rock wrens (<italic>Xenicus gilviventris</italic>; also known as pīwauwau, mātuitui, and tuke) are small (14–20 g) passerines endemic to the alpine zone of New Zealand’s South Island. They are poor fliers, and difficult to detect given their small size and the challenging high-altitude environment they occupy amongst alpine scrub, boulder fields and rocky bluffs [<xref ref-type="bibr" rid="pone.0247873.ref015">15</xref>]. Further, rock wrens have become increasingly rare, largely due to unsustainable predation by invasive mammalian predators [<xref ref-type="bibr" rid="pone.0247873.ref012">12</xref>,<xref ref-type="bibr" rid="pone.0247873.ref016">16</xref>]. Human-induced climate change is also predicted to place further pressure on rock wren populations occupying geographically disjunct mountain ranges [<xref ref-type="bibr" rid="pone.0247873.ref017">17</xref>,<xref ref-type="bibr" rid="pone.0247873.ref018">18</xref>]. The IUCN conservation status of Endangered reflects the ongoing threats to the species [<xref ref-type="bibr" rid="pone.0247873.ref018">18</xref>]. Recent genetic work also identified two distinct lineages within the population [<xref ref-type="bibr" rid="pone.0247873.ref017">17</xref>], the southern lineage is currently classified as Nationally Endangered, and the northern lineage as Nationally Critical [<xref ref-type="bibr" rid="pone.0247873.ref019">19</xref>]. Establishing a robust, cost-effective monitoring tool to evaluate long-term population trends in rock wrens is a priority for conservation of the species [<xref ref-type="bibr" rid="pone.0247873.ref018">18</xref>]. We aimed to develop a cost-effective method of monitoring rock wrens that would enable monitoring of population trends over time and ultimately assess the effectiveness of conservation interventions (landscape-scale predator control and translocations) with confidence.</p>
</sec>
<sec id="sec002" sec-type="materials|methods">
<title>Methods</title>
<p>We reviewed potential monitoring techniques for rock wrens based on established bird counting techniques used for similar rare and cryptic taxa (<xref ref-type="table" rid="pone.0247873.t001">Table 1</xref>). Based on this review, we decided to proceed with a comparison of three methods (mark-resight, distance sampling and simple counts on line transects) against territory mapping, the latter of which we considered the ‘gold standard’ [<xref ref-type="bibr" rid="pone.0247873.ref002">2</xref>,<xref ref-type="bibr" rid="pone.0247873.ref020">20</xref>]. Territory mapping enables estimation of population size in the breeding season when complete censuses are not possible [<xref ref-type="bibr" rid="pone.0247873.ref020">20</xref>,<xref ref-type="bibr" rid="pone.0247873.ref021">21</xref>]. Mark-resight and distance sampling involve incomplete counts that estimate absolute density and account for variable detection probabilities. Counts on line transects are indices that estimate relative abundance, but do not adjust for detection probabilities. We decided against trialling site occupancy as a method because detectability can vary with rock wren behaviour in relation to the nesting season and because it is difficult to compare with other methods (<xref ref-type="table" rid="pone.0247873.t001">Table 1</xref>).</p>
<table-wrap id="pone.0247873.t001" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0247873.t001</object-id>
<label>Table 1</label> <caption><title>Potential monitoring techniques considered for rock wrens based on techniques used for similar rare and cryptic taxa.</title></caption>
<alternatives>
<graphic id="pone.0247873.t001g" mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0247873.t001" xlink:type="simple"/>
<table>
<colgroup>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
</colgroup>
<thead>
<tr>
<th align="left" style="border-top:thick;border-bottom:thick">Method<xref ref-type="table-fn" rid="t001fn001"><sup>a</sup></xref></th>
<th align="left" style="border-top:thick;border-bottom:thick">Main assumptions</th>
<th align="left" style="border-top:thick;border-bottom:thick">Advantages for rock wrens</th>
<th align="left" style="border-top:thick;border-bottom:thick">Disadvantages and biases for rock wrens</th>
<th align="left" style="border-top:thick;border-bottom:thick">Cost</th>
<th align="left" style="border-top:thick;border-bottom:thick">References</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left">Territory mapping</td>
<td align="left">Observer is good at finding and identifying birds.<break/>Records are plotted accurately.<break/>There is a reasonable chance of detecting every territory that overlaps with the defined sampling area<break/>Biases are standardised.</td>
<td align="left">A thorough method for determining territories that would give a good indication of decline of rock wrens over time particularly in small populations.<break/>Do not need to mark individual rock wren (though this would give more accurate results).<break/>Method could be used to assess breeding success.</td>
<td align="left">May only capture estimates of breeding population and not account for other individuals.<break/>Territories of rock wrens overlapping with the sampling area may be missed where birds primarily occupy bluff habitat and rarely use accessible habitat.<break/>Can be intensive and time consuming requires many repeat visits.<break/>Subject to variation in time, weather, observer abilities and so need to standardise these biases.</td>
<td align="left">Medium to high.<break/>Repeat surveys can be time and resource consuming.</td>
<td align="left">[<xref ref-type="bibr" rid="pone.0247873.ref020">20</xref>,<xref ref-type="bibr" rid="pone.0247873.ref022">22</xref>]</td>
</tr>
<tr>
<td align="left">Mark-resight</td>
<td align="left">Birds have same probability of been caught.<break/>Population is closed for the survey.<break/>Marks are permanent.</td>
<td align="left">Highly precise result if assumptions are met.<break/>Analysis of data is straightforward with NOREMARK.<break/>Good for estimating other useful information such as survival, population trends and recruitment which is required for rock wrens.<break/>Closed population assumption should hold true for rock wrens during short sampling sessions.</td>
<td align="left">May be difficult to obtain high level of re-sightings.<break/>Requires 40% of birds to be individually marked, which is time and resource consuming for rock wrens.<break/>Requires constant up-keep of colour banding if monitoring is to be long term.</td>
<td align="left">High.<break/>Need to maintain a banded population.<break/>Banding birds is time and resource consuming.</td>
<td align="left">[<xref ref-type="bibr" rid="pone.0247873.ref020">20</xref>,<xref ref-type="bibr" rid="pone.0247873.ref023">23</xref>]</td>
</tr>
<tr>
<td align="left">Site occupancy</td>
<td align="left">Sites remain occupied or unoccupied for duration of survey.<break/>Species are available for detection for duration of survey.</td>
<td align="left">Only need to detect species once by sight, sound or other cues in each site.<break/>Cost effective and efficient method for covering large areas making it ideal for range and distribution.</td>
<td align="left">Need to survey sites a number of times to improve accuracy of probability function.<break/>Cannot estimate population size from method.<break/>Rock wrens may not always be available for detection inside site (e.g. they are may travel up cliff faces).</td>
<td align="left">Medium.<break/>Repeat surveys can be time and resource consuming.</td>
<td align="left">[<xref ref-type="bibr" rid="pone.0247873.ref024">24</xref>,<xref ref-type="bibr" rid="pone.0247873.ref025">25</xref>]</td>
</tr>
<tr>
<td align="left">Distance sampling (from point counts or line transects)</td>
<td align="left">All birds on the line or point are detected.<break/>Birds do not move towards or away from the observer.<break/>Distance to observations are measured accurately.</td>
<td align="left">Good method for robust, unbiased abundance estimation.<break/>Reduces the incomplete detectability resulting from simple counts.<break/>Estimates of population can be compared across time and habitat.<break/>No need to count all birds in area.</td>
<td align="left">Violation of assumptions can lead to large errors.<break/>Minimum number of detections required. 60 for line and 80 for point transects. This may be difficult to obtain for rock wrens.<break/>Often only hear rock wren so may create inaccuracies in distance measurement.</td>
<td align="left">Low to med.<break/>Requires no capturing or marking of birds.<break/>May need some observer training.</td>
<td align="left" style="background-color:#FFFFFF">[<xref ref-type="bibr" rid="pone.0247873.ref006">6</xref>,<xref ref-type="bibr" rid="pone.0247873.ref026">26</xref>]</td>
</tr>
<tr>
<td align="left" style="border-bottom:thick">Simple counts (from point counts or line transects)</td>
<td align="left" style="border-bottom:thick">Sample points distribution random, systematic or stratified.<break/>Detection probabilities remain constant.<break/>Relationship between index and true abundance is linear.<break/>Must calibrate if using to estimate density.<break/>Birds not double counted.</td>
<td align="left" style="border-bottom:thick">Cost effective and efficient method for covering large areas of alpine habitat.<break/>Method may be more suited to alpine habitat than other more intensive methods as actual distance to birds or identification of individual birds are not required.</td>
<td align="left" style="border-bottom:thick">Indices only, and the relationship with true density of rock wrens would not be known, unless calibrated with known population size and the relationship doesn’t change over time.<break/>Unadjusted for detectability, which will change with season, habitat and observer (though can be minimised through careful design and analysis).<break/>May not reflect subtle changes in populations.</td>
<td align="left" style="border-bottom:thick">Low. Set up cost is minimal.</td>
<td align="left" style="border-bottom:thick">[<xref ref-type="bibr" rid="pone.0247873.ref020">20</xref>,<xref ref-type="bibr" rid="pone.0247873.ref027">27</xref>]</td>
</tr>
</tbody>
</table>
</alternatives>
<table-wrap-foot>
<fn id="t001fn001"><p><sup>a</sup>See <xref ref-type="sec" rid="sec002">Methods</xref> text and references included within this table for definitions of techniques.</p></fn>
<fn id="t001fn002"><p>5MBC = 5-minute bird counts.</p></fn>
</table-wrap-foot>
</table-wrap>
<sec id="sec003">
<title>Study areas</title>
<p>Rock wrens were monitored during the austral spring, summer and autumn seasons (October-May) in two alpine study areas: the Homer-Gertrude cirque in the head of the Hollyford Valley, Fiordland and Haast Range, South Westland in the Southern Alps of New Zealand [<xref ref-type="bibr" rid="pone.0247873.ref012">12</xref>]. The Homer-Gertrude cirque (44° 45’ S, 168° 0’ E), is a vertical sided U-shaped valley in the Darran Mountains at the head of the Hollyford Valley. Rock wren territories were located primarily in extensive boulder fields and talus slopes interspersed with subalpine scrub and patchy <italic>Chionochloa</italic> grasslands 700–1100 m a.s.l. The Haast Range study area, between the Waiatoto and Arawhata Rivers, was centred on Lake Greaney (44° 5’ S, 168° 47’E). The study area was dominated by <italic>Chionochloa</italic> grasslands interspersed with scrub-covered cliff systems and the occasional boulder patch and talus slope, 1000–1400 m a.s.l.</p>
</sec>
<sec id="sec004">
<title>Count methods</title>
<p>We tested four counting methods in each of six breeding seasons between 2012 and 2018, and three sampling periods within each of the first four breeding seasons. Sampling periods were defined as: (1) nesting, when birds were nest building, incubating or feeding chicks on the nest (October-December), (2) fledging, once nestlings had left the nest (January-February) and (3) post-fledging (March onwards). Data from all four methods were collected during the fledging period annually during this period; however, data were not collected annually for all methods in the nesting and post-fledging periods due to logistical constraints and issues with detectability of rock wrens (see <xref ref-type="sec" rid="sec018">Results</xref> for details).</p>
<sec id="sec005">
<title>Ethical statement</title>
<p>Rock wrens were banded by qualified banders under New Zealand’s National Bird Banding Certification process. Because all other animal data collected in this study were strictly observational (i.e. no other animal handling occurred), an Animal Ethics permit was not required. Rock wrens are a protected species in New Zealand and the work occurred in two of the country’s National Parks. Management of both the species and the Public Conservation Land in which they reside rests with our employer, Te Papa Atawhai—Department of Conservation. As such, a Wildlife Act permit was not required for the research.</p>
</sec>
<sec id="sec006">
<title>Territory mapping</title>
<p>Territory mapping was undertaken as a benchmark against which to compare population size with other count methods [<xref ref-type="bibr" rid="pone.0247873.ref008">8</xref>,<xref ref-type="bibr" rid="pone.0247873.ref020">20</xref>]. Throughout the study, rock wrens were uniquely colour banded. As many birds as possible were banded, with most banding conducted at the beginning of each breeding season, but with further birds marked opportunistically throughout (including fledglings as they became available). Sightings of known individuals, and their unbanded mates or family members, were collected throughout the nesting and fledging periods so that territories could be mapped. The locations of all birds were recorded with GPS (Garmin GPSMAP 64s) by field teams, generally of 2–4 people, whenever birds were encountered, and records entered into a sightings database. Locations of birds were also recorded on maps in the field for visualisation and cross-checking. Field teams searched the study areas for birds and nests on a weekly basis through the breeding season. They spent time following birds, identifying nesting sites and recording nesting phenology every few days, to get an idea of the core areas and limits of their ranges by recording multiple GPS locations. These maps distinguished between adult males and females, which have subtly different plumages [<xref ref-type="bibr" rid="pone.0247873.ref028">28</xref>] and fledglings.</p>
</sec>
<sec id="sec007">
<title>Mark-resight</title>
<p>Analysis of the number of marked animals seen within a population over multiple resighting surveys allows abundance to be estimated using the ratio of marked to unmarked individuals, if &gt;40% of individuals in a population are marked [<xref ref-type="bibr" rid="pone.0247873.ref029">29</xref>–<xref ref-type="bibr" rid="pone.0247873.ref031">31</xref>]. Resightings of colour-banded and unmarked birds throughout the study areas were recorded during the three sampling periods (nesting, fledging and post-fledging) each breeding season. Survey routes covered all rock wren territories mapped in the study areas to ensure equal probability of detection for all individuals and included the line transect routes used for other counts (see below). The observers walked the routes slowly four times during each sampling session in fine weather. If birds were detected, they were followed until it could be confirmed if they were banded or not.</p>
</sec>
<sec id="sec008">
<title>Simple counts on transects</title>
<p>Simple counts were conducted along multiple 250 m line transects located randomly within suitable rock wren habitat in each study area. We defined suitable habitat as boulder fields, talus slopes, cliff systems, subalpine scrub based on mapping in ArcGIS Version 9 using SPOT satellite imagery. The number of transects sampled were representative of rock wren habitat in each study area: 14 transects at Homer-Gertrude cirque and 27 at Haast Range. Each transect was selected sequentially from a paired list of randomised start points and bearings, with a minimum of 250 m between transects. The observer walked slowly along the transect, recording all rock wrens seen or heard. Each transect took ca. 20 minutes to walk (mean = 22.5 minutes/transect). To maximise detectability of rock wrens, each transect was counted four times in fine weather (no rain, thick fog or strong winds), generally on consecutive days and each time by a different observer, where possible. Surveys took 3–8 days, depending on interruptions during adverse weather.</p>
</sec>
<sec id="sec009">
<title>Distance sampling</title>
<p>Distance sampling, either from transects or points, is a widely used method for estimating abundance by modelling the detectability of an animal as its distance from the observer increases [<xref ref-type="bibr" rid="pone.0247873.ref003">3</xref>,<xref ref-type="bibr" rid="pone.0247873.ref009">9</xref>,<xref ref-type="bibr" rid="pone.0247873.ref029">29</xref>]. Rock wren counts from line transects described above were also used in calculating population estimates using distance sampling. In addition, perpendicular distance from the birds’ location when first detected to the transect line was estimated visually or using Bushnell Yardage Pro 500 rangefinders to the nearest metre.</p>
</sec>
<sec id="sec010">
<title>Co-variates</title>
<p>We recorded weather and environmental variables during each transect count. Variables included were: (1) temperature: cold = 0−5°C, cool = &gt;5−11°C, mild = &gt;11−16°C, warm = &gt;16−22°C, hot = &gt;22°C; (2) visibility at ground level in metres, averaged over the count; (3) overhead sunshine in minutes; (4) precipitation: 0 = none, 1 = misty, 2 = drizzle, 3 = light, 4 = moderate, 5 = heavy; and (5) wind: 0 = leaves still or moving without noise, 1 = leaves rustling, 2 = leaves and branchlets in constant motion, 3 = branches or trees swaying. These covariates were tested as predictors of rock wren detections on line transects (see below).</p>
</sec>
</sec>
<sec id="sec011">
<title>Analysis</title>
<sec id="sec012">
<title>Territory mapping</title>
<p>Field maps from each field trip were checked against data entered in our resighting and nesting databases and combined to create single territory maps for the nesting and fledging periods each breeding season. Numbers of adults and fledglings and the territories they occupied were summed to estimate population size for each sampling period in each study area.</p>
</sec>
<sec id="sec013">
<title>Mark-resight</title>
<p>The number of banded birds available for resighting was calculated from the territory maps and sightings database for each season, adjusted for additional birds banded between surveys and any deaths detected while monitoring nests. Data were analysed using the mark–resight modelling program NOREMARK for closed populations [<xref ref-type="bibr" rid="pone.0247873.ref030">30</xref>,<xref ref-type="bibr" rid="pone.0247873.ref031">31</xref>]. We used Bowden’s estimator to compute mark–resight abundance estimates [<xref ref-type="bibr" rid="pone.0247873.ref030">30</xref>]. Bowden’s estimator assumes that the probability of capture and resight is the same for all animals, but relaxes assumptions that the population is closed; allowing temporary movement out of the study area (in this case over inaccessible cliff edges), variation in resighting probabilities, sampling with replacement, and not requiring all animals to be correctly identified during each sampling session [<xref ref-type="bibr" rid="pone.0247873.ref009">9</xref>,<xref ref-type="bibr" rid="pone.0247873.ref032">32</xref>]. Although other models often appear to have greater precision, they can be overly precise and perform poorly when estimating confidence intervals compared with the Bowden’s estimator [<xref ref-type="bibr" rid="pone.0247873.ref009">9</xref>,<xref ref-type="bibr" rid="pone.0247873.ref033">33</xref>].</p>
</sec>
<sec id="sec014">
<title>Simple counts on transects</title>
<p>Despite constraining transect counts to relatively fine weather (no rain, thick fog or strong winds), we tested the effects of climatic variables on rock wren detections to evaluate the need for adjusting indices to account for any such effects. To do this we used a zero-inflated generalised mixed-effects model in RStudio (version 1.1.423) for a reduced dataset for which all covariate data were available (see <xref ref-type="sec" rid="sec018">Results</xref>). We included the following in the models: (1) rock wrens seen or heard on transects as the response variable, (2) temperature, sun (% direct sunshine during transect count), visibility, precipitation level and wind as fixed factors, and (3) transect nested within study area as the random effect. For comparison with other methods, rock wren detections on line transects were summarised as detections per transect per season for each year (<xref ref-type="table" rid="pone.0247873.t002">Table 2</xref>).</p>
<table-wrap id="pone.0247873.t002" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0247873.t002</object-id>
<label>Table 2</label> <caption><title>A comparison of rock wren, <italic>Xenicus gilviventris</italic>, estimates and indices of abundance from four monitoring methods at two study areas in the Southern Alps of New Zealand, 2012–2018.</title></caption>
<alternatives>
<graphic id="pone.0247873.t002g" mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0247873.t002" xlink:type="simple"/>
<table>
<colgroup>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
</colgroup>
<thead>
<tr>
<th align="center">Study area</th>
<th align="center">Year</th>
<th align="center">Sampling period</th>
<th align="center">Territory map</th>
<th align="center">Mark-resight (± 95% CI)</th>
<th align="center">Distance sampling (± 95% CI)</th>
<th align="center">Simple counts (mean ± SE)</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left">Haast</td>
<td align="center">2012</td>
<td align="left">Nesting</td>
<td align="center">76</td>
<td align="right">81 (64–102)</td>
<td align="right">102 (79–133)</td>
<td align="right">0.54 (± 0.09)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2013</td>
<td align="left"/>
<td align="center">61</td>
<td align="right">80 (60–107)</td>
<td align="right">13 (9–18)</td>
<td align="right">0.28 (± 0.06)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2014</td>
<td align="left"/>
<td align="center">58</td>
<td align="right">74 (55–100)</td>
<td align="right">9 (6–14)</td>
<td align="right">0.21 (± 0.05)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2015</td>
<td align="left"/>
<td align="center">43</td>
<td align="right">59 (36–100)</td>
<td align="right">35 (23–53)</td>
<td align="right">0.22 (± 0.07)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2013</td>
<td align="left">Fledging</td>
<td align="center">108</td>
<td align="right">90 (71–113)</td>
<td align="right">113 (91–139)</td>
<td align="right">0.61 (± 0.08)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2014</td>
<td align="left"/>
<td align="center">87</td>
<td align="right">98 (68–142)</td>
<td align="right">14 (10–19)</td>
<td align="right">0.25 (± 0.06)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2015</td>
<td align="left"/>
<td align="center">84</td>
<td align="right">116 (71–191)</td>
<td align="right">8 (5–12)</td>
<td align="right">0.29 (± 0.06)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2016</td>
<td align="left"/>
<td align="center">67</td>
<td align="right">226 (108–474)</td>
<td align="right">52 (35–79)</td>
<td align="right">0.31 (± 0.06)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2017</td>
<td align="left"/>
<td align="center">88</td>
<td align="right">226 (123–418)</td>
<td align="right">39 (29–51)</td>
<td align="right">0.69 (± 0.12)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2018</td>
<td align="left"/>
<td align="center">253</td>
<td align="right">285 (244–333)</td>
<td align="right">44 (36–54)</td>
<td align="right">0.99 (± 0.12)</td>
</tr>
<tr>
<td align="left">Homer/</td>
<td align="center">2012</td>
<td align="left">Nesting</td>
<td align="center">56</td>
<td align="right">66 (53–82)</td>
<td align="right">38 (16–69)</td>
<td align="right">0.56 (± 0.19)</td>
</tr>
<tr>
<td align="left">Gertrude</td>
<td align="center">2013</td>
<td align="left"/>
<td align="center">67</td>
<td align="right">94 (64–138)</td>
<td align="right">42 (17–75)</td>
<td align="right">0.39 (± 0.11)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2014</td>
<td align="left"/>
<td align="center">36</td>
<td align="right">35 (26–47)</td>
<td align="right">68 (25–116)</td>
<td align="right">0.73 (± 0.15)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2015</td>
<td align="left"/>
<td align="center">69</td>
<td align="right">66 (52–83)</td>
<td align="right">82 (46–126)</td>
<td align="right">0.90 (± 0.14)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2013</td>
<td align="left">Fledging</td>
<td align="center">34</td>
<td align="right">35 (30–41)</td>
<td align="right">40 (19–64)</td>
<td align="right">0.45 (± 0.09)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2014</td>
<td align="left"/>
<td align="center">83</td>
<td align="right">93 (62–142)</td>
<td align="right">61 (32–94)</td>
<td align="right">0.53 (± 0.14)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2015</td>
<td align="left"/>
<td align="center">76</td>
<td align="right">87 (64–119)</td>
<td align="right">55 (30–85)</td>
<td align="right">0.69 (± 0.14)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2016</td>
<td align="left"/>
<td align="center">135</td>
<td align="right">209 (136–332)</td>
<td align="right">212 (128–311)</td>
<td align="right">2.38 (± 0.29)</td>
</tr>
<tr>
<td align="right"/>
<td align="center">2017</td>
<td align="left"/>
<td align="center">92</td>
<td align="right">168 (101–282)</td>
<td align="right">28 (18–39)</td>
<td align="right">1.20 (± 0.18)</td>
</tr>
<tr>
<td align="left"/>
<td align="center">2018</td>
<td align="left"/>
<td align="center">129</td>
<td align="right">120 (98–147)</td>
<td align="right">24 (14–39)</td>
<td align="right">1.11 (± 0.23)</td>
</tr>
</tbody>
</table>
</alternatives>
<table-wrap-foot>
<fn id="t002fn001"><p>Note that a major predation event at The Homer-Gertrude cirque in spring 2012 that resulted in failure of 100% of nests monitored and mortality of several adult females on nests [<xref ref-type="bibr" rid="pone.0247873.ref012">12</xref>] coincided with the monitoring comparison; as such these data should be treated with caution. The post-fledging sampling period was omitted due to low detection rates during this time (see <xref ref-type="sec" rid="sec018">Results</xref>).</p></fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec id="sec015">
<title>Distance sampling</title>
<p>Data obtained by distance sampling were analysed using the program Distance 6.2 [<xref ref-type="bibr" rid="pone.0247873.ref034">34</xref>]. Observed differences in the general topography, vegetation composition and structure between the two study areas meant that detection probabilities for rock wrens at each study area were likely to differ. Data were therefore analysed independently for each study area [<xref ref-type="bibr" rid="pone.0247873.ref026">26</xref>]. As distances to birds were recorded to the nearest metre, distances were left ungrouped rather than being aggregated into distance classes.</p>
<p>To increase sample size and estimate precision, data from all surveys at each study area were pooled and global detection functions were calculated for each study area [<xref ref-type="bibr" rid="pone.0247873.ref026">26</xref>]. Using these global detection functions, data were post-stratified by survey and histograms of perpendicular distance measurements constructed. A selection of robust models and appropriate expansion functions recommended by [<xref ref-type="bibr" rid="pone.0247873.ref026">26</xref>] were then fitted. Model fit was assessed using Akaike’s Information Criterion, Goodness of Fit and Q-Q plots and associated statistics, and the most parsimonious model was selected [<xref ref-type="bibr" rid="pone.0247873.ref006">6</xref>,<xref ref-type="bibr" rid="pone.0247873.ref026">26</xref>,<xref ref-type="bibr" rid="pone.0247873.ref035">35</xref>].</p>
<p>For the Homer-Gertrude study area, good model fit was achieved using half-normal and hazard rate models, and in the Haast Range, uniform and hazard rate models with varying numbers of adjustment terms in both areas (<xref ref-type="table" rid="pone.0247873.t003">Table 3</xref>). The largest five percent of distance measurements were truncated to improve estimate precision [<xref ref-type="bibr" rid="pone.0247873.ref026">26</xref>].</p>
<table-wrap id="pone.0247873.t003" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0247873.t003</object-id>
<label>Table 3</label> <caption><title>Models evaluated to produce population estimates from distance sampling.</title></caption>
<alternatives>
<graphic id="pone.0247873.t003g" mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0247873.t003" xlink:type="simple"/>
<table>
<colgroup>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
</colgroup>
<thead>
<tr>
<th align="center">Location</th>
<th align="center">Model (key +adjustment)<xref ref-type="table-fn" rid="t003fn002"><sup>1</sup></xref></th>
<th align="center">No. parameters</th>
<th align="center">ΔAIC<xref ref-type="table-fn" rid="t003fn003"><sup>2</sup></xref></th>
<th align="center"><inline-formula id="pone.0247873.e001"><alternatives><graphic id="pone.0247873.e001g" mimetype="image" position="anchor" xlink:href="info:doi/10.1371/journal.pone.0247873.e001" xlink:type="simple"/><mml:math display="inline" id="M1"><mml:mover accent="true"><mml:mi mathvariant="bold">D</mml:mi><mml:mo>⌢</mml:mo></mml:mover></mml:math></alternatives></inline-formula>ensity (ha<sup>-1</sup>)</th>
<th align="center">95% Confidence Interval</th>
<th align="center">%CV<xref ref-type="table-fn" rid="t003fn004"><sup>3</sup></xref></th>
</tr>
</thead>
<tbody>
<tr>
<td align="left">Haast</td>
<td align="left">Uniform + simple poly. (Poisson)<xref ref-type="table-fn" rid="t003fn001">*</xref></td>
<td align="center">2</td>
<td align="center">0</td>
<td align="center">0.080</td>
<td align="center">0.072–0.090</td>
<td align="center">5.7</td>
</tr>
<tr>
<td align="left"/>
<td align="left">Uniform + cos.</td>
<td align="center">1</td>
<td align="center">0.909</td>
<td align="center">0.080</td>
<td align="center">0.069–0.093</td>
<td align="center">7.4</td>
</tr>
<tr>
<td align="left"/>
<td align="left">Hazard rate + simple poly.</td>
<td align="center">2</td>
<td align="center">1.248</td>
<td align="center">0.077</td>
<td align="center">0.065–0.091</td>
<td align="center">8.5</td>
</tr>
<tr>
<td align="left"/>
<td align="left">Hazard rate + cos.</td>
<td align="center">2</td>
<td align="center">1.248</td>
<td align="center">0.077</td>
<td align="center">0.065–0.091</td>
<td align="center">8.5</td>
</tr>
<tr>
<td align="left"/>
<td align="left">Half normal + hermite poly.</td>
<td align="center">1</td>
<td align="center">4.091</td>
<td align="center">0.082</td>
<td align="center">0.070–0.096</td>
<td align="center">7.9</td>
</tr>
<tr>
<td align="left"/>
<td align="left">Half normal +cos.</td>
<td align="center">1</td>
<td align="center">4.091</td>
<td align="center">0.082</td>
<td align="center">0.070–0.096</td>
<td align="center">7.9</td>
</tr>
<tr>
<td align="left">Homer-Gertrude</td>
<td align="left">Half normal +cos. (Poisson)<xref ref-type="table-fn" rid="t003fn001">*</xref></td>
<td align="center">3</td>
<td align="center">0</td>
<td align="center">0.304</td>
<td align="center">0.256–0.360</td>
<td align="center">8.7</td>
</tr>
<tr>
<td align="left"/>
<td align="left">Hazard rate + simple poly.</td>
<td align="center">2</td>
<td align="center">1.787</td>
<td align="center">0.291</td>
<td align="center">0.238–0.354</td>
<td align="center">10.1</td>
</tr>
<tr>
<td align="left"/>
<td align="left">Hazard rate + cos.</td>
<td align="center">2</td>
<td align="center">1.787</td>
<td align="center">0.291</td>
<td align="center">0.238–0.354</td>
<td align="center">10.1</td>
</tr>
<tr>
<td align="left"/>
<td align="left">Uniform + cos.</td>
<td align="center">1</td>
<td align="center">3.726</td>
<td align="center">0.251</td>
<td align="center">0.225–0.281</td>
<td align="center">5.7</td>
</tr>
<tr>
<td align="left"/>
<td align="left">Half normal + hermite poly.</td>
<td align="center">1</td>
<td align="center">4.962</td>
<td align="center">0.247</td>
<td align="center">0.219–0.278</td>
<td align="center">6.0</td>
</tr>
<tr>
<td align="left"/>
<td align="left">Uniform + simple poly.</td>
<td align="center">2</td>
<td align="center">6.120</td>
<td align="center">0.248</td>
<td align="center">0.220–0.280</td>
<td align="center">6.1</td>
</tr>
</tbody>
</table>
</alternatives>
<table-wrap-foot>
<fn id="t003fn001"><p>*Global model with lowest ΔAIC selected to compute post-stratified seasonal estimates in <xref ref-type="table" rid="pone.0247873.t002">Table 2</xref>.</p></fn>
<fn id="t003fn002"><p><sup>1</sup>Model consisting of a key function and an adjustment term.</p></fn>
<fn id="t003fn003"><p><sup>2</sup>AIC values rescaled as simple differences between models.</p></fn>
<fn id="t003fn004"><p><sup>3</sup>% Coefficient of Variation.</p></fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec id="sec016">
<title>Comparison of methods</title>
<p>We used linear models in RStudio to evaluate the relationship between estimates from territory maps and the other estimates/indices. We initially constructed global linear models to evaluate the influence of study area (Haast or Homer-Gertrude) and sampling period (nesting or fledging) in addition to each method (mark-resight, distance sampling, simple counts; separately) on estimates from territory maps. We then used linear models to explore correlations between territory maps and the other methods during the fledging period (for which most data exist; <xref ref-type="table" rid="pone.0247873.t002">Table 2</xref>) for each study area separately.</p>
</sec>
<sec id="sec017">
<title>Estimating costs of methods</title>
<p>By far the greatest cost involved in monitoring rock wrens was the wages paid to field staff. As such, we recorded the number of days and number of people required to undertake each technique as the basis for a cost comparison between methods. We assumed an hourly pay rate of NZD50 as the basis for calculations. We separated costs into: (1) costs associated with setting up the monitoring technique in year 1, (2) annual maintenance costs, if applicable to the method, and (3) annual monitoring costs.</p>
</sec>
</sec>
</sec>
<sec id="sec018" sec-type="results">
<title>Results</title>
<p>We collected rock wren sightings data to create territory maps and compare these with population estimates from mark-resight and distance sampling data, and with indices from simple counts, across six years, 2012–2018 (see <xref ref-type="supplementary-material" rid="pone.0247873.s001">S1 Table</xref> for raw data). Robust data were collected in all six years during the fledging period (January–February) and in four of six years during the nesting period (October–December). However, detection rates in post-fledging period (March onwards) were too low to create meaningful estimates from territory maps (our ‘gold standard’), and so we excluded this period from analyses. Sample sizes presented below reflect only data collected in the nesting and fledging periods.</p>
<sec id="sec019">
<title>Territory mapping</title>
<p>Four hundred and eighty birds were colour banded during the study (Homer-Gertrude = 197; Haast = 283). Although not all birds in the study areas were marked, the proportion of banded birds was sufficient to map territories and distinguish unbanded birds and breeding pairs, based on behaviours of marked and adjacent pairs, locations of nests, and behaviours at nests. Numbers of rock wrens counted each season varied between 34 and 135 adult birds at Homer-Gertrude and 43 and 253 at Haast (<xref ref-type="table" rid="pone.0247873.t002">Table 2</xref>). The numbers of breeding rock wrens in the Haast study area declined over the first four breeding seasons (<xref ref-type="fig" rid="pone.0247873.g001">Fig 1A</xref>). However, following instigation of alpine predator trapping targeted at stoats (<italic>Mustela erminea</italic>) part way into the 2015/16 breeding season, productivity increased considerably, contributing to an increase in the study population over the following two seasons [<xref ref-type="bibr" rid="pone.0247873.ref012">12</xref>]. In contrast, the number of breeding birds at The Homer-Gertrude cirque increased steadily, though with some variability (<xref ref-type="fig" rid="pone.0247873.g001">Fig 1B</xref>), through the monitoring period during which time nests were protected using a similar predator trapping programme [<xref ref-type="bibr" rid="pone.0247873.ref012">12</xref>].</p>
<fig id="pone.0247873.g001" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0247873.g001</object-id>
<label>Fig 1</label>
<caption>
<title/>
<p><bold>Numbers of adult and fledgling rock wrens, <italic>Xenicus gilviventris</italic>, detected through territory mapping during the fledging (January-February) period 2012–2018 at: (a) the Haast Range, South Westland, and (b) the Homer-Gertrude cirque, Fiordland</bold>.</p>
</caption>
<graphic mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0247873.g001" xlink:type="simple"/>
</fig>
</sec>
<sec id="sec020">
<title>Mark-resight</title>
<p>We undertook 10 mark-resight surveys during nesting and fledging in each study area across six breeding seasons, with equal survey effort in each study area. The proportion of each population banded (available for resighting) in each sampling period averaged 54.5 ± 17.3% SD (ra = 35.8–79.4%) at Homer-Gertrude and 50.9 ± 8.5% (ra = 41.1–65.8%) at Haast. Of these banded birds, on average, 50.1 ± 8.5% SD (Homer-Gertrude) and 49.0 ± 6.85% (Haast) were resighted on mark-resight surveys during nesting, with slightly lower proportions resighted during fledging (44.1 ± 14.8% and 30.6 ± 18.3% respectively).</p>
<p>Accuracy of recording band combinations was difficult to measure, but the rates of known partial or incorrect band combinations recorded give confidence that the majority of band combinations were recorded correctly. On average 3.6 ± 7.6% SD of sightings at Homer-Gertrude and 1.4 ± 3.4% of sighting at Haast were partial band combinations and incorrect band combinations were recorded on five (5.3%) of the mark-resight surveys (both study areas combined).</p>
<p>Population estimates derived from mark-resight data using Bowden’s estimator ranged from 59 to 285 at the Haast study area and from 35 to 209 at the Homer-Gertrude cirque (<xref ref-type="table" rid="pone.0247873.t002">Table 2</xref>). Precision of these estimates were highly variable in both study areas, with 95% confidence intervals being largest when population estimates were highest (<xref ref-type="table" rid="pone.0247873.t002">Table 2</xref>).</p>
</sec>
<sec id="sec021">
<title>Distance sampling</title>
<p>Field observations showed that rock wren behaviours almost certainly violated assumptions of distance sampling (<xref ref-type="table" rid="pone.0247873.t001">Table 1</xref>), particularly: (1) birds moving towards or away from the observer, and (2) all birds on the line are detected (i.e. birds obscured beneath large boulders on the line). The number of distance measurements to individual rock wrens for any given survey were highly variable, ranging from 14–120 birds at Homer-Gertrude and 18–110 birds for Haast Range. This corresponded to an encounter rate on transects of 1.0–9.2 birds detected/km surveyed at Homer-Gertrude and 0.2–2.4 birds/km at Haast. Pooling detections for each study area (assuming detectability at each study area remains the same over time), applying a global detection function, and post-stratifying by survey period provided a partial solution to the lack of data for some surveys. However, extremely low sample sizes in some surveys compromised the precision of abundance estimates [<xref ref-type="bibr" rid="pone.0247873.ref026">26</xref>] (<xref ref-type="table" rid="pone.0247873.t002">Table 2</xref>).</p>
<p>Abundance estimates and corresponding 95% confidence intervals (only bootstrapped for the Homer-Gertrude study area because estimates for Haast failed to converge) ranged from 8 to 113 for Haast and from 24 to 212 in Homer-Gertrude (<xref ref-type="table" rid="pone.0247873.t002">Table 2</xref> and <xref ref-type="fig" rid="pone.0247873.g002">Fig 2</xref>). Confidence Intervals for some survey periods were extremely wide for both study areas, particularly for surveys where modelled estimates of abundance were large (<xref ref-type="fig" rid="pone.0247873.g002">Fig 2</xref>).</p>
<fig id="pone.0247873.g002" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0247873.g002</object-id>
<label>Fig 2</label>
<caption>
<title/>
<p><bold>Rock wren, <italic>Xenicus gilviventris</italic>, estimates and indices of abundance from four methods over six summer fledging periods at two alpine study areas ((a) Haast and (b) Homer-Gertrude) in the Southern Alps of New Zealand.</bold> Symbols are as follows: (1) territory map = black circles; (2) mark resight = open circles; (3) distance sampling = black triangles; (4) simple counts = black diamonds. The left axis shows the territory map figures and estimates from mark resight and distance sampling; the right axis shows the index of abundance from line transects. See <xref ref-type="table" rid="pone.0247873.t002">Table 2</xref> for variance estimates.</p>
</caption>
<graphic mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0247873.g002" xlink:type="simple"/>
</fig>
</sec>
<sec id="sec022">
<title>Simple counts on transects</title>
<p>Between 2012 and 2018 we recorded 968 rock wrens (i.e. birds seen or heard) during 1647 transect counts conducted in the nesting and fledging periods. Indices of abundance (i.e. number of birds detected per transect during a sampling period) ranged from 0.21 to 0.99 birds per transect at Haast and 0.39 to 2.38 birds per transect at Homer-Gertrude (<xref ref-type="table" rid="pone.0247873.t002">Table 2</xref>).</p>
<p>The full range of weather variables was measured on 922 transect counts; these counts were used to evaluate relationships between rock wren counts and weather variables (air temperature, sunshine, visibility, precipitation and wind). Within the sampling constraints imposed (counts were conducted during fine weather; that is, no rain, thick fog or strong wind), we detected no significant correlations between rock wren counts and weather recorded (P &gt; 0.05 in all cases). As such, we did not need to adjust the indices of abundance obtained from line transect sampling for weather variables prior to comparison with estimates derived from territory mapping.</p>
</sec>
<sec id="sec023">
<title>Comparison of techniques</title>
<p>Estimates of rock wren populations based on territory mapping during the fledging period (January-February) were positively correlated with indices of abundance from simple counts on line transects at both study areas during the same period (Haast: t<sub>1</sub> = 3.041, P = 0.038; Homer-Gertrude: t<sub>1</sub> = 2.555, P = 0.063; <xref ref-type="table" rid="pone.0247873.t004">Table 4</xref> and <xref ref-type="fig" rid="pone.0247873.g003">Fig 3</xref>). These territory map estimates were also positively correlated with estimates derived from mark-resight surveys in the Homer-Gertrude cirque during the fledging period (t<sub>1</sub> = 2.922, P = 0.043), but not at the Haast study area (t<sub>1</sub> = 1.351, P = 0.248). However, rock wren population estimates from distance sampling were not correlated with estimates from territory maps in either study area (P &gt; 0.1 for both study areas; <xref ref-type="table" rid="pone.0247873.t004">Table 4</xref> and <xref ref-type="fig" rid="pone.0247873.g003">Fig 3</xref>).</p>
<fig id="pone.0247873.g003" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0247873.g003</object-id>
<label>Fig 3</label>
<caption>
<title/>
<p><bold>Comparison of three methods (mark-resight, distance sampling and line transects) against territory mapping for rock wrens, <italic>Xenicus gilviventris</italic>, at the (a) Haast and (b) Homer-Gertrude alpine study areas in the South Island of New Zealand.</bold> Filled circles = summer; open circles = spring; dashed regression line added for significant linear relationships between methods (see <xref ref-type="sec" rid="sec018">Results</xref>).</p>
</caption>
<graphic mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0247873.g003" xlink:type="simple"/>
</fig>
<table-wrap id="pone.0247873.t004" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0247873.t004</object-id>
<label>Table 4</label> <caption><title>Correlation coefficients (r<sup>2</sup> values from linear models) between abundance derived from territory mapping of rock wrens, <italic>Xenicus gilviventris</italic>, and population estimates from both mark-resight and distance sampling and indices of relative abundance from simple counts on line transects.</title></caption>
<alternatives>
<graphic id="pone.0247873.t004g" mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0247873.t004" xlink:type="simple"/>
<table>
<colgroup>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
</colgroup>
<thead>
<tr>
<th align="left"/>
<th align="center" colspan="2">Haast</th>
<th align="center" colspan="2">Homer-Gertrude</th>
</tr>
<tr>
<th align="left"/>
<th align="center">Overall</th>
<th align="center">Fledging period</th>
<th align="center">Overall</th>
<th align="center">Fledging period</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left">Territory ~ mark-resight</td>
<td align="center">0.47</td>
<td align="center">0.31</td>
<td align="center">0.76</td>
<td align="center">0.68</td>
</tr>
<tr>
<td align="left">Territory ~ distance sampling</td>
<td align="center">0.02</td>
<td align="center">0.01</td>
<td align="center">0.20</td>
<td align="center">0.25</td>
</tr>
<tr>
<td align="left">Territory ~ simple counts</td>
<td align="center">0.72</td>
<td align="center">0.70</td>
<td align="center">0.58</td>
<td align="center">0.49</td>
</tr>
</tbody>
</table>
</alternatives>
<table-wrap-foot>
<fn id="t004fn001"><p>Data from the nesting period are not presented separately due to insufficient data points (n = 4 per study area), but are included within the overall metrics.</p></fn>
</table-wrap-foot>
</table-wrap>
<sec id="sec024">
<title>Estimating costs of methods</title>
<p>Set-up costs for the methods requiring banding rock wrens (territory mapping and mark-resight) were 25 times higher than methods for which birds were not banded (distance sampling and simple counts; <xref ref-type="table" rid="pone.0247873.t005">Table 5</xref>). Annual costs (maintenance and monitoring costs combined) were greatest for territory mapping at NZD 22,000 p.a., intermediate for mark-resight at NZD 13,200 p.a. and lowest for distance sampling and simple counts, both of which costs NZD 3,200 p.a. (<xref ref-type="table" rid="pone.0247873.t005">Table 5</xref>). As such, annual costs of territory mapping were 6.9 times higher than annual costs both distance sampling and simple counts.</p>
<table-wrap id="pone.0247873.t005" position="float">
<object-id pub-id-type="doi">10.1371/journal.pone.0247873.t005</object-id>
<label>Table 5</label> <caption><title>Cost comparison for four monitoring methods for rock wrens, <italic>Xenicus gilviventris</italic>.</title></caption>
<alternatives>
<graphic id="pone.0247873.t005g" mimetype="image" position="float" xlink:href="info:doi/10.1371/journal.pone.0247873.t005" xlink:type="simple"/>
<table>
<colgroup>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
<col align="left" valign="middle"/>
</colgroup>
<thead>
<tr>
<th align="left">Technique</th>
<th align="center" colspan="3">Set-up</th>
<th align="center" colspan="3">Annual maintenance</th>
<th align="center" colspan="2">Annual monitoring</th>
</tr>
<tr>
<th align="left"/>
<th align="center">Tasks</th>
<th align="center">Time</th>
<th align="center">Cost</th>
<th align="center">Tasks</th>
<th align="center">Time</th>
<th align="center">Cost</th>
<th align="center">Time</th>
<th align="center">Cost</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left">Territory mapping</td>
<td align="left">Scope study area Catch initial sample of birds</td>
<td align="left">10 days x 5 people</td>
<td align="left">$20,000</td>
<td align="left">Keeping a sample of birds tagged</td>
<td align="left">5 days x 5 people</td>
<td align="left">$10,000</td>
<td align="left">15 days x 2 people</td>
<td align="left">$12,000</td>
</tr>
<tr>
<td align="left">Mark-resight</td>
<td align="left">Scope study area Catch initial sample of birds</td>
<td align="left">10 days x 5 people</td>
<td align="left">$20,000</td>
<td align="left">Keeping a sample of birds tagged</td>
<td align="left">5 days x 5 people</td>
<td align="left">$10,000</td>
<td align="left">4 days x 2 people</td>
<td align="left">$3,200</td>
</tr>
<tr>
<td align="left">Distance sampling</td>
<td align="left">Plan &amp; mark transects</td>
<td align="left">2 person days</td>
<td align="left">$800</td>
<td align="left">None</td>
<td align="left"/>
<td align="left">$0</td>
<td align="left">4 days x 2 people</td>
<td align="left">$3,200</td>
</tr>
<tr>
<td align="left">Simple counts</td>
<td align="left">Plan &amp; mark transects</td>
<td align="left">2 person days</td>
<td align="left">$800</td>
<td align="left">None</td>
<td align="left"/>
<td align="left">$0</td>
<td align="left">4 days x 2 people</td>
<td align="left">$3,200</td>
</tr>
</tbody>
</table>
</alternatives>
<table-wrap-foot>
<fn id="t005fn001"><p>All costs are presented in New Zealand Dollars (NZD).</p></fn>
</table-wrap-foot>
</table-wrap>
</sec>
</sec>
</sec>
<sec id="sec025" sec-type="conclusions">
<title>Discussion</title>
<p>The strong, positive correlations between estimates from territory maps (the ‘gold standard’ technique) and indices of relative abundance derived from simple counts on line transects suggest that these counts can be used as a low-cost, reliable, technique to monitor trends in rock wren populations over time. Banding birds (required for territory mapping and mark-resight) is time-intensive and dependent on the availability of skilled personnel to coincide with fair weather conditions, which can be problematic in the alpine zone of an oceanic island like New Zealand. We estimate that annual costs of territory mapping are 6.9 times higher, and annual costs of mark-resight are 4.1 times higher than those of simple counts and distance sampling. Because simple counts don’t require banding of rock wrens, they provide a much more cost-effective monitoring tool that we have now validated against a benchmark. We do, however, recommend that results obtained through this method be interpreted with appropriate caution given that index methods lack incorporation of detection probabilities, which may vary across habitats, densities and time [<xref ref-type="bibr" rid="pone.0247873.ref036">36</xref>,<xref ref-type="bibr" rid="pone.0247873.ref037">37</xref>].</p>
<p>Although territory mapping is often considered the ‘gold standard’ for monitoring bird species [<xref ref-type="bibr" rid="pone.0247873.ref020">20</xref>], this technique still does not represent a true census. Because birds are highly cryptic, and frequently occupy inaccessible cliff habitats, it is difficult to accurately measure immigration, emigration and mortality during a season. Crypsis in rock wrens increases as the season wears on, with the proportion of birds detected in surveys in the post-fledging period being markedly lower than in both the nesting and fledging periods due to fledglings becoming independent and dispersing throughout the landscape [<xref ref-type="bibr" rid="pone.0247873.ref038">38</xref>], as has been previously reported for house wrens, <italic>Troglodytes aedon</italic>, in Ohio, USA [<xref ref-type="bibr" rid="pone.0247873.ref039">39</xref>]. Nevertheless, we felt population estimates derived from territory mapping in the nesting season were accurate, and changes in population sizes were what was expected given our monitoring of high predation rates, particularly by invasive stoats, prior to introduction of population scale trapping and documented recovery afterwards at both study areas [<xref ref-type="bibr" rid="pone.0247873.ref012">12</xref>]. We were unable to obtain sufficient data for all methods trialled to undertake a statistical comparison of methods during the post-fledging period. However, based on the limited data we were able to collect during this period, we suggest that results from all methods are more variable and less reliable in the post-fledging period than earlier in the breeding season. The increased crypsis later in the season suggests that monitoring of trends in rock wrens should be timed consistently each year and occur prior to dispersal of fledglings.</p>
<p>Accuracy and precision of population estimates derived using Bowden’s mark-resight estimator were highly variable. Incomplete identification of marked individuals is potentially a major source of bias in mark-resight abundance estimators [<xref ref-type="bibr" rid="pone.0247873.ref040">40</xref>] applicable to rock wrens because full colour band combinations are not always seen when rock wrens are only glimpsed briefly. Population estimates generated from mark-resight data from visual surveys were only strongly correlated with estimates from territory mapping at one of our two study areas. Rock wrens were highly cryptic, with a low proportion detected on any one survey. They often fed underground in the extensive boulder fields and dense subalpine scrub that characterises their habitat. Further, mark-resight surveys in the alpine zone were very labour intensive. In addition to the considerable effort involved in banding a meaningful proportion of the population, the thorough surveys required to sample all territories in the population were very extensive. For example, the route surveyed in the Haast Range was c. 35 km long to sample c. 30 rock wren territories. Thus, increasing the effort to complete more than four surveys to further increase the proportion of marked birds detected and the accuracy of the technique would use considerable additional resources with no guarantee of achieving this objective.</p>
<p>Distance sampling has recently been used successfully to estimate densities of several forest birds and to evaluate their long-term responses to conservation management [<xref ref-type="bibr" rid="pone.0247873.ref003">3</xref>,<xref ref-type="bibr" rid="pone.0247873.ref041">41</xref>]. However, population estimates for rock wrens derived from distance sampling were not correlated with population estimates produced from territory maps. At least two of the three key assumptions of distance sampling (<xref ref-type="table" rid="pone.0247873.t001">Table 1</xref>) are regularly violated in rock wren monitoring. Firstly, the assumption of 100% detectability on the zero line (i.e. the transect) was not achieved where large boulders with sub-terranean space were present on the transect line and rock wrens frequently used that space to forage or take refuge, without making many calls. This regularly occurred when attempts were made to catch rock wrens for banding or when rock wrens were nesting beneath boulders. Secondly, the assumption that birds do not approach or avoid observers was violated in open habitat types where the birds frequently flew away from observers on approach. The poor performance of distance sampling as a technique for producing population estimates in rock wrens is similar to that seen in bellbirds, <italic>Anthornis melanura</italic>, which were also variable in conspicuousness and moved away from the line transect when approached by an observer [<xref ref-type="bibr" rid="pone.0247873.ref041">41</xref>]. However, it is somewhat surprising that distance sampling performed so poorly in comparison to indices of relative abundance which were derived from the same line transects. One potential explanation is that the estimated distances were so inaccurate that including them in the population estimation process introduced more error than it removed. Initial attempts to use range finders to measure distance were often thwarted by misty conditions in the alpine zone and we resorted to visual estimation of distance in most cases. Further, it is possible that systematic bias was induced by estimated distances being correlated with habitat.</p>
<p>More promisingly, indices of abundance generated from the simple counts on line transects were strongly correlated with territory map estimates at both of our alpine study sites over a six-year period. Repeated sampling on line transects has shown similar promise in open fen mire habitat for aquatic warblers, <italic>Acrocephalus paludicola</italic>, in Central Europe [<xref ref-type="bibr" rid="pone.0247873.ref004">4</xref>] and in forest for endemic passerines, <italic>Mohoua ochrecephala</italic>, in New Zealand [<xref ref-type="bibr" rid="pone.0247873.ref042">42</xref>]. In the latter study, monitoring of 14 <italic>Mohoua ochrecephala</italic> populations on line transects at 12 sites over up to 11 years revealed one population extinction and a further five populations in decline [<xref ref-type="bibr" rid="pone.0247873.ref042">42</xref>]. This led to an understanding that a species previously thought to be secure was endangered and in need of immediate conservation intervention [<xref ref-type="bibr" rid="pone.0247873.ref042">42</xref>]. When applying our findings to recommending a new standard monitoring technique for rock wrens we acknowledge that the relationship between the index and real density may not remain the same over time. Lower correlation coefficients at the Homer-Gertrude study area can be partially attributed to the index overestimating the population in the fledging period of 2016 when the population was at its highest during the sampling period (2012–2018). This may hint at a non-linear relationship between detectability and density whereby birds are disproportionately active and vocal as density increases. A non-linear relationship between detectability and density was also observed for South Island robins, <italic>Petroica australis</italic>; however, in the case of robins, detectability increased at lower population density due to an increase in calling by males when females are scarce in the population [<xref ref-type="bibr" rid="pone.0247873.ref009">9</xref>]. Therefore, while we recommend that indices of abundance on line transects be adopted as a standard low-cost technique for monitoring trends in rock wren populations, we suggest that care be taken in interpreting results derived from indices, for which detection probabilities are not accounted [<xref ref-type="bibr" rid="pone.0247873.ref036">36</xref>,<xref ref-type="bibr" rid="pone.0247873.ref037">37</xref>]. Further targeted testing of the line transect method at even higher densities should be undertaken if populations increase beyond levels observed during this study. Promisingly, this seems likely for populations where effective alpine predator control in the form of population-scale trapping [<xref ref-type="bibr" rid="pone.0247873.ref012">12</xref>] or landscape-scale toxin application [<xref ref-type="bibr" rid="pone.0247873.ref043">43</xref>] leads to further increases in rock wrens.</p>
</sec>
<sec id="sec026" sec-type="conclusions">
<title>Conclusion</title>
<p>Monitoring population trends in montane and alpine birds is becoming increasingly important to determine potential impacts of climate change and increased anthropogenic disturbance [<xref ref-type="bibr" rid="pone.0247873.ref037">37</xref>,<xref ref-type="bibr" rid="pone.0247873.ref044">44</xref>–<xref ref-type="bibr" rid="pone.0247873.ref046">46</xref>]. In general, calibrating bird monitoring methods has been undertaken infrequently, limiting comparability among monitoring programmes [<xref ref-type="bibr" rid="pone.0247873.ref047">47</xref>,<xref ref-type="bibr" rid="pone.0247873.ref048">48</xref>]. Our comparison of monitoring methods for alpine passerines in New Zealand, and finding that a low-cost index technique is strongly correlated with estimates from territory mapping which is regarded as a ‘gold standard’ in bird monitoring, has applicability to other open habitat bird species as well as highlighting the importance of validating potential monitoring techniques, prior to using them as the basis of monitoring protocols.</p>
</sec>
<sec id="sec027" sec-type="supplementary-material">
<title>Supporting information</title>
<supplementary-material id="pone.0247873.s001" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet" position="float" xlink:href="info:doi/10.1371/journal.pone.0247873.s001" xlink:type="simple">
<label>S1 Table</label>
<caption>
<title>Raw data collected on rock wrens, <italic>Xenicus gilviventris</italic>, using four techniques (territory mapping, mark-resight, distance sampling and simple counts) at two sites in the Southern Alps of New Zealand, 2012–2018.</title>
<p>(XLSX)</p>
</caption>
</supplementary-material>
</sec>
</body>
<back>
<ack>
<p>Thanks to Sue Heath, Bruce Robertson, Megan Willans and James Reardon for input into study design and Richard Earl for assistance with mapping rock wren habitat in the study areas. Thanks also to Kathrin Affeld, Will Batson, Becky Bell, Crystal Brindle, Iris Broekema, Bevan Cameron, Jo Carpenter, Jono Dobbs, Phil Evans, Clare Kilner, Ian Clark, Eric Edwards, Sarah Forder, Ruth Garland, Flo Gaud, Lynette Hartley, Adam Ingram, Athene Irvine, Rebecca Jackson, Franziska Landesberger, Rose Lanman, Jamie McAulay, Bruce McKinlay, Fraser Maddigan, James Maunder, Grant Maslowski, Kathy Morrison, Dan Palmer, Moira Pryde, Emma Richardson, Bruce Robertson, Lucy Rossiter, Sam Rowland, Anne Schlesselmann, Sanjay Thakur, Jo Tilson, Kerry Uren, Antje Wahlberg, Jim Watts, David Webb, Jemma van Beek, Maddie van de Wetering, Jason van de Wetering, Megan Willans, Kailash Willis and Rebecca Wilson for assistance conducting counts in the field and banding rock wrens, Kathrin Affeld for data management, Karina Sidaway for technical editing and the staff of the Haast and Te Anau Department of Conservation offices for considerable assistance with logistics.</p>
</ack>
<ref-list>
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<named-content content-type="letter-date">4 Jan 2021</named-content>
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<p>PONE-D-20-38105</p>
<p>Evaluation of counting methods for monitoring populations of a cryptic alpine passerine, the rock wren (<italic>Xenicus gilviventris</italic>)</p>
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<p>Reviewer #1: This is a well-done study that evaluates different monitoring methods and their appropriateness for a particular species of wren in New Zealand. The authors have done an excellent job conveying what they did and its importance for species conservation. I have a few remaining questions and suggestions.</p>
<p>- I (not an ornithologist) struggled to understand why the numbers in the simple count method were given as abundance indices, and not converted to a number that would be directly comparable to the territory mapping. Would doing so have altered the statistical results in a meaningful way?</p>
<p>- Line 84: I’d like a lot more detail about this literature review, particularly since it is called a lit review. Are these methods that are usually used in alpine bird species? Was anything in particular discarded, and if so, why?</p>
<p>- In table 1, please add a column with a short description of each method. This information is in the body of the text but not the first time these terms are used.</p>
<p>- Why have the methods been ordered this way in the table? As a reader, it seems to make more logical sense that territory mapping would be described first or last (as the standard to which all other methods are compared)</p>
<p>The line numbers disappeared after Table 1.</p>
<p>- In the Distance Sampling methods paragraph (page 10), please explain what conventional methods are in a short sentence or two.</p>
<p>- Page 11 – NOREMARK is a different spelling than in Table 1. I think this spelling is correct and the table is wrong, but please check.</p>
<p>- In the results, page 14, you mention that territory mapping is only reliable through the fledgling period. There is a bit of discussion of this issue on page 19, but I’d like more – is this expected to be true of all of these methods? Might one method be better for surveys in the post-fledgling period?</p>
<p>- Fig 4 – I don’t’ see an explanation of panel a vs b (they are clearly the two study sites but it’s not clear which is which).</p>
<p>Reviewer #2: Please include in the title the Order and the Family of the bird species.</p>
<p>This study departs from a relevant perspective and searching for cost-effective and cost-less methods is valid if your output is equally valid, replicable, and renders strong data to be used in long-term monitoring and conservation. Comparison between different methods is also relevant, and this study is particularly important in bringing some light on this neglected subject. One of the authors' concerns is about the cost of each method x quality of the results, and this statement appears along with the manuscript. However, this topic is not explored with the details requirable in the discussion, and according to the authors, in a situation where the results must be "achievable within budget constraints" I hope to see this topic explored with the necessary details. As the authors have the raw data &amp; costs to obtain these data in each methodology I strongly suggest that they can cover this topic more accurately. I agree that banding birds is the most expensive method and potentially harmful specially for delicate and threatened species. Although I agree that pointing counts is an effective tool, I believe the authors must provide stronger evidence in the discussion (maybe the last paragraph should be subjected to a more lengthy explanation).</p>
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<article-title>Author response to Decision Letter 0</article-title>
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<p>
<named-content content-type="author-response-date">14 Feb 2021</named-content>
</p>
<p>Bi-Song Yue</p>
<p>Academic Editor</p>
<p>PLOS ONE</p>
<p>Dear Dr Bi-Song Yue</p>
<p>Thank you very much for the constructive feedback on our manuscript. We have thoroughly addressed all points raised during review. Our detailed responses to reviewers’ comments are in italics below.</p>
<p>We hope that our revised manuscript is now suitable for publication in PLOS One and look forward to hearing from you again.</p>
<p>With best wishes,</p>
<p>Jo Monks (on behalf of all authors)</p>
<p>Journal requirements:</p>
<p>1. Please ensure that your manuscript meets PLOS ONE's style requirements, including those for file naming. </p>
<p>&gt; Done. </p>
<p>2. We note that Figure 1 in your submission contain map images which may be copyrighted. We require you to either (1) present written permission from the copyright holder to publish these figures specifically under the CC BY 4.0 license, or (2) remove the figures from your submission.</p>
<p>&gt; On reflection, Figure 1 (a map showing location of the two study sites) is not essential to this manuscript, so we have decided to remove it rather than seeking copyright permission. Further, we now refer to another paper which includes a map showing site locations. </p>
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<p>&gt; Done.</p>
<p>Reviewer #1: </p>
<p>This is a well-done study that evaluates different monitoring methods and their appropriateness for a particular species of wren in New Zealand. The authors have done an excellent job conveying what they did and its importance for species conservation. I have a few remaining questions and suggestions.</p>
<p>- I (not an ornithologist) struggled to understand why the numbers in the simple count method were given as abundance indices, and not converted to a number that would be directly comparable to the territory mapping. Would doing so have altered the statistical results in a meaningful way?</p>
<p>&gt; Simple counts are presented as abundance indices because the number of transects surveyed at each site differed according to availability of suitable habitat for rock wrens and, due to weather and logistical constraints, it was not always possible to survey all transects an equal number of times per sampling period. Furthermore, attempting to present simple counts as a number to be directly compared with estimates from territory mapping would be misleading, because they do not sample the full area and because they do not account for detectability.</p>
<p>- Line 84: I’d like a lot more detail about this literature review, particularly since it is called a lit review. Are these methods that are usually used in alpine bird species? Was anything in particular discarded, and if so, why?</p>
<p>&gt; Reviewer 1 makes a good point that we didn’t conduct a formal literature review in creating this table, rather researched the applicability of established bird counting techniques to rock wrens and this is what we present in Table 1. We have removed reference to the term ‘literature review’.</p>
<p>&gt; Based on our understanding from the literature, there is no ‘usual’ method for alpine bird species. Some of these techniques are applied elsewhere in the world in alpine, or more usually open, habitats, but we didn’t find strong evidence of consistency. This was part of our motivation for the present study, which we hope will contribute to the literature on methods for alpine birds.</p>
<p>&gt; Site occupancy was discarded because detectability can vary with rock wren behaviour in relation to the nesting season and because it is difficult to compare with other methods. See lines 93 to 95 of the Methods for this explanation.</p>
<p>- In table 1, please add a column with a short description of each method. This information is in the body of the text but not the first time these terms are used.</p>
<p>&gt; We attempted to do this, but it is impossible to describe these techniques briefly in a way amenable to inclusion in a table; the detail is important! We have instead referred the reader to the Methods text and references included within the table in the footnotes of the table for these descriptions.</p>
<p>- Why have the methods been ordered this way in the table? As a reader, it seems to make more logical sense that territory mapping would be described first or last (as the standard to which all other methods are compared)</p>
<p>&gt; This is a good point. We have reordered the table to begin with territory mapping (as the gold standard), followed by the other techniques in decreasing order of cost.</p>
<p>The line numbers disappeared after Table 1.</p>
<p>&gt; Good point! Now fixed.</p>
<p>- In the Distance Sampling methods paragraph (page 10), please explain what conventional methods are in a short sentence or two.</p>
<p>&gt; Done. We’ve added a description of distance sampling and rewritten the paragraph for clarity.</p>
<p>- Page 11 – NOREMARK is a different spelling than in Table 1. I think this spelling is correct and the table is wrong, but please check.</p>
<p>&gt; Thanks – we’ve corrected the spelling error in Table 1.</p>
<p>- In the results, page 14, you mention that territory mapping is only reliable through the fledgling period. There is a bit of discussion of this issue on page 19, but I’d like more – is this expected to be true of all of these methods? Might one method be better for surveys in the post-fledgling period?</p>
<p>&gt; We have added the following two sentences to the paragraph Reviewer 1 refers to in order to address this point: </p>
<p>“We were unable to obtain sufficient data for all methods trialled to undertake a statistical comparison of methods during the post-fledging period. However, based on the limited data we were able to collect during this period, we suggest that results from all methods are more variable and less reliable in the post-fledging period than earlier in the breeding season.”</p>
<p>- Fig 4 – I don’t’ see an explanation of panel a vs b (they are clearly the two study sites but it’s not clear which is which).</p>
<p>&gt; Good point! We’ve added this information into the figure title.</p>
<p>Reviewer #2: </p>
<p>Please include in the title the Order and the Family of the bird species.</p>
<p>&gt; Done.</p>
<p>This study departs from a relevant perspective and searching for cost-effective and cost-less methods is valid if your output is equally valid, replicable, and renders strong data to be used in long-term monitoring and conservation. Comparison between different methods is also relevant, and this study is particularly important in bringing some light on this neglected subject. One of the authors' concerns is about the cost of each method x quality of the results, and this statement appears along with the manuscript. However, this topic is not explored with the details requirable in the discussion, and according to the authors, in a situation where the results must be "achievable within budget constraints" I hope to see this topic explored with the necessary details. As the authors have the raw data &amp; costs to obtain these data in each methodology I strongly suggest that they can cover this topic more accurately. I agree that banding birds is the most expensive method and potentially harmful specially for delicate and threatened species.</p>
<p>&gt; This is a very good point. In response we collated information on set-up, annual maintenance and annual monitoring costs for all four field methods. This information is now presented in a new table (Table 5), described in the methods and results section, and referred to in the discussion.</p>
<p>Although I agree that pointing counts is an effective tool, I believe the authors must provide stronger evidence in the discussion (maybe the last paragraph should be subjected to a more lengthy explanation).</p>
<p>&gt; We have expanded this paragraph of the discussion to strengthen the evidence for utility of repeated counts on line transects as a suitable technique for trend monitoring in rock wrens and potentially other passerines in open habitat. In doing so, we made it explicit that in our study indices of abundance from line transects were strongly correlated with estimates from territory mapping at both study sites over a 6-year period. We also added a case study in which the technique was successfully applied to a forest-dwelling passerine and resulted in strong management recommendations.</p>
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<surname>Yue</surname>
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<p>
<named-content content-type="letter-date">16 Feb 2021</named-content>
</p>
<p>Evaluation of counting methods for monitoring populations of a cryptic alpine passerine, the rock wren (Passeriformes, Acanthisittidae, Xenicus gilviventris)</p>
<p>PONE-D-20-38105R1</p>
<p>Dear Dr. Monks,</p>
<p>We’re pleased to inform you that your manuscript has been judged scientifically suitable for publication and will be formally accepted for publication once it meets all outstanding technical requirements.</p>
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<p>Academic Editor</p>
<p>PLOS ONE</p>
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<given-names>Bi-Song</given-names>
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<p>
<named-content content-type="letter-date">23 Feb 2021</named-content>
</p>
<p>PONE-D-20-38105R1 </p>
<p>Evaluation of counting methods for monitoring populations of a cryptic alpine passerine, the rock wren (Passeriformes, Acanthisittidae, <italic>Xenicus gilviventris</italic>) </p>
<p>Dear Dr. Monks:</p>
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