ˇ˛<html xmlns:v="urn:schemas-microsoft-com:vml" xmlns:o="urn:schemas-microsoft-com:office:office" xmlns:w="urn:schemas-microsoft-com:office:word" xmlns:m="http://schemas.microsoft.com/office/2004/12/omml" xmlns:mv="http://macVmlSchemaUri" xmlns="http://www.w3.org/TR/REC-html40"> <head> <meta name=Title content="Split Zimmerius minimus from Z. chrysops"> <meta name=Keywords content=""> <meta http-equiv=Content-Type content="text/html; charset=macintosh"> <meta name=ProgId content=Word.Document> <meta name=Generator content="Microsoft Word 14"> <meta name=Originator content="Microsoft Word 14"> <link rel=File-List href="SACCprop550X_files/filelist.xml"> <title>Split Zimmerius minimus from Z. chrysops</title> <!--[if gte mso 9]><xml> <o:DocumentProperties> <o:Author> </o:Author> <o:LastAuthor>James Remsen</o:LastAuthor> <o:Revision>2</o:Revision> <o:TotalTime>8</o:TotalTime> <o:Created>2015-10-25T01:35:00Z</o:Created> <o:LastSaved>2015-10-25T01:35:00Z</o:LastSaved> <o:Pages>1</o:Pages> <o:Words>980</o:Words> <o:Characters>5207</o:Characters> <o:Company>SACC</o:Company> <o:Lines>92</o:Lines> <o:Paragraphs>15</o:Paragraphs> <o:CharactersWithSpaces>6172</o:CharactersWithSpaces> <o:Version>14.0</o:Version> </o:DocumentProperties> <o:OfficeDocumentSettings> <o:PixelsPerInch>96</o:PixelsPerInch> </o:OfficeDocumentSettings> </xml><![endif]--> <link rel=themeData href="SACCprop550X_files/themedata.xml"> <!--[if gte mso 9]><xml> <w:WordDocument> <w:Zoom>116</w:Zoom> <w:DisplayBackgroundShape/> <w:SpellingState>Clean</w:SpellingState> <w:GrammarState>Clean</w:GrammarState> <w:TrackMoves>false</w:TrackMoves> <w:TrackFormatting/> <w:PunctuationKerning/> <w:ValidateAgainstSchemas/> <w:SaveIfXMLInvalid>false</w:SaveIfXMLInvalid> <w:IgnoreMixedContent>false</w:IgnoreMixedContent> <w:AlwaysShowPlaceholderText>false</w:AlwaysShowPlaceholderText> <w:DoNotPromoteQF/> <w:LidThemeOther>EN-US</w:LidThemeOther> <w:LidThemeAsian>JA</w:LidThemeAsian> <w:LidThemeComplexScript>X-NONE</w:LidThemeComplexScript> <w:Compatibility> <w:BreakWrappedTables/> <w:SnapToGridInCell/> <w:WrapTextWithPunct/> <w:UseAsianBreakRules/> <w:DontGrowAutofit/> <w:SplitPgBreakAndParaMark/> </w:Compatibility> <m:mathPr> <m:mathFont m:val="Cambria Math"/> <m:brkBin m:val="before"/> <m:brkBinSub m:val="&#45;-"/> <m:smallFrac m:val="off"/> <m:dispDef m:val="off"/> <m:lMargin m:val="0"/> <m:rMargin m:val="0"/> <m:defJc m:val="centerGroup"/> <m:wrapRight/> <m:intLim m:val="subSup"/> <m:naryLim m:val="subSup"/> </m:mathPr></w:WordDocument> </xml><![endif]--><!--[if gte mso 9]><xml> <w:LatentStyles DefLockedState="false" DefUnhideWhenUsed="false" DefSemiHidden="false" DefQFormat="false" LatentStyleCount="276"> <w:LsdException Locked="false" Priority="1" Name="Default Paragraph Font"/> <w:LsdException Locked="false" Priority="99" Name="Normal (Web)"/> <w:LsdException Locked="false" Priority="99" Name="No List"/> <w:LsdException Locked="false" Priority="34" QFormat="true" Name="List Paragraph"/> </w:LatentStyles> </xml><![endif]--> <style> <!-- /* Font Definitions */ @font-face {font-family:Arial; 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font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p>&nbsp;</o:p></span></p> <p class=MsoNormal align=center style='text-align:center'><b style='mso-bidi-font-weight: normal'><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family: Arial;mso-bidi-font-family:"Times New Roman"'>Split <i style='mso-bidi-font-style: normal'>Zimmerius minimus</i> from <i style='mso-bidi-font-style:normal'>Z. chrysops</i><o:p></o:p></span></b></p> <p class=MsoNormal><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p>&nbsp;</o:p></span></p> <p class=MsoNormal><u><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'>Background</span></u><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family: "Times New Roman"'>:<span style="mso-spacerun:yes">&nbsp;&nbsp; </span>The confusing genus <i style='mso-bidi-font-style:normal'>Zimmerius</i>, as defined by Traylor (1977), includes about a dozen currently recognized species. Morphologically cryptic, <i style='mso-bidi-font-style:normal'>Zimmerius</i> flycatchers can better be distinguished by vocal and molecular characters (Rheindt et al. 2008). One superspecies that has been particularly puzzling is the <i style='mso-bidi-font-style:normal'>Z. viridiflavus</i> complex, which has variously been treated as a single species or a collection of 2-3 species. In a recent study using mitochondrial and vocal evidence, Rheindt et al. (2008) showed that the taxon <i style='mso-bidi-font-style:normal'>albigularis</i> seems to be distantly related to the complex and must be split. The latter authors also presented data in support of a 2-species treatment for the remaining <i style='mso-bidi-font-style:normal'>Z. viridiflavus</i> complex, but along different boundaries than suggested by most previous ornithologists. The Tumbesian taxon <i style='mso-bidi-font-style:normal'>flavidifrons</i> and an unnamed population from the Peruvian departments of Amazonas and San MartÌn (south of the MaraÒÛn River) were vocally and mitochondrially akin to <i style='mso-bidi-font-style:normal'>Z. viridiflavus</i>, even though their plumage resembles <i style='mso-bidi-font-style:normal'>Z. chrysops</i>. SACC (proposal </span><a href="SACCprop173.html"><span style='font-size:14.0pt;mso-bidi-font-size: 12.0pt;font-family:Arial;mso-bidi-font-family:"Times New Roman"'>173</span></a><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family: "Times New Roman"'>) decided to follow Rheindt et al. s (2008) recommendations of aligning those two entities with <i style='mso-bidi-font-style:normal'>Z. viridiflavus</i>, thereby restricting the range of <i style='mso-bidi-font-style: normal'>Z. chrysops</i> to the eastern Andean slopes north of the River MaraÒÛn. Rheindt et al. s (2008) sampling did not include any specimens from the extensive northern part of the range of <i style='mso-bidi-font-style:normal'>Z. chrysops</i> in Colombia and Venezuela; hence, the taxa <i style='mso-bidi-font-style: normal'>cumanensis</i> and <i style='mso-bidi-font-style:normal'>minimus</i> remained unsampled.<o:p></o:p></span></p> <p class=MsoNormal><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p>&nbsp;</o:p></span></p> <p class=MsoNormal><u><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'>New Evidence</span></u><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family: "Times New Roman"'>:<o:p></o:p></span></p> <p class=MsoNormal><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'>Rheindt et al. (in press) added eight specimens of <i style='mso-bidi-font-style:normal'>Z. chrysops</i> (<i style='mso-bidi-font-style:normal'>sensu</i> Rheindt et al., 2008) to the molecular sampling of Rheindt et al (2008), most from Colombia and Venezuela. Although Rheindt et al. (in press) also analyzed a nuclear intron (Fibrinogen 5), it turned out to contain little phylogenetic information at the taxonomic level of interest, so most of their conclusions were reached on the basis of the mitochondrial (ND2) dataset.<o:p></o:p></span></p> <p class=MsoNormal style='text-indent:.5in'><span style='font-size:14.0pt; mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family:"Times New Roman"'>The two Venezuelan specimens from eastern T·chira formed a strongly supported monophyletic group that is sister to &#8211; but deeply diverged (4.7%) from &#8211; all other specimens of <i style='mso-bidi-font-style:normal'>Z. chrysops</i> (<i style='mso-bidi-font-style:normal'>sensu</i> Rheindt et al 2008). Significantly, the larger, more southerly group also contained specimens from as far north as Antioquia (Colombia), which are geographically quite close to T·chira.<o:p></o:p></span></p> <p class=MsoNormal style='text-indent:.5in'><span style='font-size:14.0pt; mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family:"Times New Roman"'>Considering recordings deposited at Xeno-Canto and the Macaulay Sound Library, Rheindt et al. (in press) then went on to compare informally whether the mitochondrial pattern is supported by the primary call vocalization of these tyrannulets. They found that the typical, well-known  peeew call of <i style='mso-bidi-font-style: normal'>Z. chrysops</i> extends from the MaraÒÛn River in the south to the Colombian-Venezuelan border in the north (including extreme western T·chira), whereas populations on the other side of the T·chira Low (starting from eastern T·chira) towards the east utter a distinct, much shorter, harder  pyupp . Therefore, vocal evidence supports mtDNA in indicating the existence of two species.<o:p></o:p></span></p> <p class=MsoNormal style='text-indent:.5in'><span style='font-size:14.0pt; mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family:"Times New Roman"'>To the best of my knowledge, no taxonomic moniker is available for the split-worthy populations of <i style='mso-bidi-font-style:normal'>Z. chrysops</i> in the Venezuelan Andes. However, two nearby taxa have so far escaped molecular sampling: taxon <i style='mso-bidi-font-style:normal'>minimus</i> (Santa Marta Mts) and taxon <i style='mso-bidi-font-style:normal'>cumanensis</i> (Turimiquire Mtns. in eastern Venezuela). Although <i style='mso-bidi-font-style: normal'>cumanensis</i> remains almost unknown in life and may represent yet another species-level taxon, there are two recordings of <i style='mso-bidi-font-style: normal'>minimus</i> at Macaulay (68092, 68093) that sound identical to the short, hard  pyupp of birds from the Venezuelan Andes. Although the Santa Marta Mtns. are geographically isolated from the Venezuelan Andes, vocal evidence supports their conspecificity with Venezuelan populations. Hence, Rheindt et al. (in press) advocated that <i style='mso-bidi-font-style:normal'>Z. minimus</i> from the Santa Marta Mts and the Venezuelan Andes east of the T·chira Low be split from <i style='mso-bidi-font-style:normal'>Z. chrysops</i>. Although unusual, the geographic checkerboard distribution of <i style='mso-bidi-font-style:normal'>Z. minimus</i> could be explained by <i style='mso-bidi-font-style:normal'>Z. chrysops</i> having expanded northwards and displaced <i style='mso-bidi-font-style:normal'>Z. minimus</i>, with the expansion having been halted by the T·chira Low and the coastal plains that isolate the Santa Marta Massif.<o:p></o:p></span></p> <p class=MsoNormal><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p>&nbsp;</o:p></span></p> <p class=MsoNormal><u><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'>Recommendation</span></u><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family: "Times New Roman"'>:<span style="mso-spacerun:yes">&nbsp; </span>Rheindt et al. s (in press) paper may have certain weaknesses: their molecular conclusions are restricted to mitochondrial evidence, and their vocal comparisons are informal and not statistically backed. Nevertheless, given the congruence of vocal and molecular results, this paper should shift the burden of evidence to those who would like to continue subspecific treatment of <i style='mso-bidi-font-style: normal'>minimus</i> for the following reasons:<o:p></o:p></span></p> <p class=MsoNormal><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p>&nbsp;</o:p></span></p> <p class=MsoListParagraphCxSpFirst style='text-indent:-.25in;mso-list:l1 level1 lfo2'><![if !supportLists]><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family:Arial;mso-fareast-font-family: Arial'><span style='mso-list:Ignore'>(1)</span></span><![endif]><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family: "Times New Roman"'> As shown, the use of single nuclear markers adds little to the mitochondrial evidence. If nuclear DNA were to contribute any information, it would have to come from a great number of loci.<o:p></o:p></span></p> <p class=MsoListParagraphCxSpMiddle style='text-indent:-.25in;mso-list:l1 level1 lfo2'><![if !supportLists]><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family:Arial;mso-fareast-font-family: Arial'><span style='mso-list:Ignore'>(2)</span></span><![endif]><i style='mso-bidi-font-style:normal'><span style='font-size:14.0pt;mso-bidi-font-size: 12.0pt;font-family:Arial;mso-bidi-font-family:"Times New Roman"'> Zimmerius</span></i><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family: "Times New Roman"'> calls are distinct 1-2 syllable vocalizations that can easily be distinguished by the human ear. There is nothing equivocal about them. Given that the mtDNA evidence is congruent with the distinct calls of these birds, it would be hard to argue that these vocal differences are based on human perception bias.<o:p></o:p></span></p> <p class=MsoListParagraphCxSpLast><span style='font-size:14.0pt;mso-bidi-font-size: 12.0pt;font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p>&nbsp;</o:p></span></p> <p class=MsoNormal style='text-indent:.25in'><span style='font-size:14.0pt; mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family:"Times New Roman"'>I recommend recognition of <i style='mso-bidi-font-style:normal'>Z. minimus</i> from the Venezuelan Andes and the Santa Marta Mountains as an independent species  Coopmans s Tyrannulet , honoring the man who provided some of the earliest sound recordings of topotypical <i style='mso-bidi-font-style:normal'>Z. minimus</i>. Based on geographical proximity, the taxon <i style='mso-bidi-font-style: normal'>cumanensis</i> would best be recognized as a subspecies of <i style='mso-bidi-font-style:normal'>Z. minimus</i> (not <i style='mso-bidi-font-style: normal'>Z. chrysops</i>) pending data on its vocalizations and DNA.<o:p></o:p></span></p> <p class=MsoNormal><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p>&nbsp;</o:p></span></p> <p class=MsoNormal><u><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'>Literature</span></u><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family: "Times New Roman"'>:<o:p></o:p></span></p> <p><span style='font-size:14.0pt;mso-bidi-font-size:11.0pt;font-family:Arial; mso-bidi-font-family:"Times New Roman"'>Rheindt F.E., Cuervo A.M., Brumfield R.T. Rampant polyphyly indicates cryptic diversity in a clade of Neotropical flycatchers (Aves: Tyrannidae). Biological Journal of the Linnean Society, in press.</span><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p></o:p></span></p> <p><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family:Arial; mso-bidi-font-family:"Times New Roman"'>Rheindt F.E., Norman J.A., Christidis L. 2008. DNA evidence shows vocalizations to be a better indicator of taxonomic limits than plumage patterns in <i style='mso-bidi-font-style:normal'>Zimmerius</i> tyrant- flycatchers. Mol. Phylogenetics and Evolution 48: 150-156.<o:p></o:p></span></p> <p class=MsoNormal><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p>&nbsp;</o:p></span></p> <p class=MsoNormal align=center style='text-align:center'><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt;font-family:Arial;mso-bidi-font-family: "Times New Roman"'>Frank Rheindt, September 2012<o:p></o:p></span></p> <p class=MsoNormal><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p>&nbsp;</o:p></span></p> <div style='mso-element:para-border-div;border:none;border-bottom:double windowtext 2.25pt; padding:0in 0in 1.0pt 0in'> <p class=MsoNormal align=center style='text-align:center;border:none; mso-border-bottom-alt:double windowtext 2.25pt;padding:0in;mso-padding-alt: 0in 0in 1.0pt 0in'><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial'><o:p>&nbsp;</o:p></span></p> </div> <p class=MsoNormal align=center style='text-align:center'><span style='font-size:14.0pt;mso-bidi-font-size:19.0pt;font-family:Arial;mso-bidi-font-family: "Times New Roman"'><o:p>&nbsp;</o:p></span></p> <p class=MsoNormal><span style='font-size:14.0pt;mso-bidi-font-size:12.0pt; font-family:Arial;mso-bidi-font-family:"Times New Roman"'><o:p>&nbsp;</o:p></span></p> </div> </body> </html>