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<mods:title id="A5B3E88329C30C2B0A290A1F58F39E27">Cryptic subterranean diversity: regional phylogeography of the sand termite Psammotermes allocerus Silvestri, 1908 in the wider Namib region</mods:title>
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<mods:namePart id="32E85C6580643E8F2B579E7ACFA12594">Gunter, Felicitas</mods:namePart>
<mods:affiliation id="0F0C6767D76AB8E8425099B3219DA728">Biodiversity, Evolution and Ecology of Plants % BEE), Institute of Plant Science and Microbiology, University of Hamburg, Ohnhorststrasse 18, 22609 Hamburg, Germany</mods:affiliation>
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<mods:namePart id="C9141D981EC77B0B5A807D30750EDCFE">Oldeland, Jens</mods:namePart>
<mods:affiliation id="273DB94A9662B4EEDCD7594C68F9F1E4">Biodiversity, Evolution and Ecology of Plants % BEE), Institute of Plant Science and Microbiology, University of Hamburg, Ohnhorststrasse 18, 22609 Hamburg, Germany</mods:affiliation>
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<mods:namePart id="6FB7BA99FEEAFD3235AC8134BD7E567A">Picker, Mike D.</mods:namePart>
<mods:affiliation id="F7B52246A88B042DC3E08B0F202A9E5A">Department of Biological Sciences, University of Cape Town, Cape Town, Rondebosch 7701, South Africa</mods:affiliation>
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<mods:namePart id="311B27C0E1F15FCE6586B115C209FE75">Jürgens, Norbert</mods:namePart>
<mods:affiliation id="9A7106A40A7B3E23CBECF5302EAA2AA0">Biodiversity, Evolution and Ecology of Plants % BEE), Institute of Plant Science and Microbiology, University of Hamburg, Ohnhorststrasse 18, 22609 Hamburg, Germany</mods:affiliation>
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<treatment id="B721F3218F27F11EFCD7FB5CFB76FA6B" ID-DOI="http://doi.org/10.5281/zenodo.12801601" ID-Zenodo-Dep="12801601" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B" httpUri="http://treatment.plazi.org/id/B721F3218F27F11EFCD7FB5CFB76FA6B" lastPageId="3" lastPageNumber="142" pageId="2" pageNumber="141">
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<heading id="647FF55B8F27F11FFCD7FB5CFC17FB5D" bold="true" box="[816,919,1217,1247]" fontSize="11" level="1" pageId="2" pageNumber="141" reason="1">
<taxonomicName id="F88839B48F27F11FFCD7FB5CFC17FB5D" ID-CoL="4ND6T" authority="Silvestri, 1908" authorityName="Silvestri" authorityYear="1908" box="[816,919,1217,1247]" class="Insecta" family="Rhinotermitidae" genus="Psammotermes" kingdom="Animalia" order="Blattodea" pageId="2" pageNumber="141" phylum="Arthropoda" rank="species" species="allocerus">
<emphasis id="0DFC9E258F27F11FFCD7FB5CFC17FB5D" bold="true" box="[816,919,1217,1247]" pageId="2" pageNumber="141">Results</emphasis>
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<emphasis id="0DFC9E258F27F11FFCD7FA98FBA0FAA2" bold="true" box="[816,1056,1285,1312]" pageId="2" pageNumber="141">Phylogeny analyses</emphasis>
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<paragraph id="3F3742378F27F11EFCD7FAD4FE4EFEF8" blockId="2.[816,1453,1353,1911]" lastBlockId="3.[135,773,153,1911]" lastPageId="3" lastPageNumber="142" pageId="2" pageNumber="141">
The mitochondrial sequences were visually examined according to the presence of NUMTs %nuclear mitochondrial sequences). Characteristics such as stop codons, frameshift mutations and insertions-deletions were not detected. Finally, the COI marker %362 bp) contained 137 constant, 70 uninformative, and 155 parsimony-informative positions. For the second marker, COII %448 bp), 259 positions were constant, 89 were uninformative, and 124 were parsimony-informative. The concatenated alignment %810 bp) resulted in 507 constant positions. A low number of 92 positions were uninformative, and 211 were parsimony-informative. All mitochondrial COI and COII sequences were concatenated in one alignment, and the resulting Bayesian Inference %BI) analysis shows a strongly supported differentiation of the
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65
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<emphasis id="0DFC9E258F27F11FFC35F880FBD7F8B6" box="[978,1111,1820,1844]" italics="true" pageId="2" pageNumber="141">P. allocerus</emphasis>
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samples from the outgroup %1 posterior probabilities %pp)). We identified seven genetic groups which may represent putative species clusters according to the high posterior probability values of the nodes %pp&gt; 0.95), higher values of p-distances between these clusters as within the clusters as well haplotype analyses according to Roy et al. %2006) and Hausberger et al. %2011). The seven strongly supported clades %0.99 pp,
<figureCitation id="A7B35EB28F26F11EFF6FFEDDFF4BFEDA" box="[136,203,320,344]" captionStart="Fig" captionStartId="4.[136,165,156,177]" captionTargetBox="[557,1447,158,1812]" captionTargetId="figure-86@4.[553,1451,154,1817]" captionTargetPageId="4" captionText="Fig. 1 Phylogeny of 65 P. allocerus collections inferred by the Bayesian analysis of COI and COII markers. Support values are given in posterior probability. Clades are coloured according to the genetic group. Dark blue: Succulent Karoo; Light Blue: Southern Namib; Dark green: East Gariep; Light Green: Southwestern Kalahari; Yellow: Nama; Ochre: Western Kalahari Basin; Red: Northern Namib. Abbreviations of study sites are shown in Fig. 2" figureDoi="http://doi.org/10.5281/zenodo.12764917" httpUri="https://zenodo.org/record/12764917/files/figure.png" pageId="3" pageNumber="142">Fig. 1</figureCitation>
), roughly fit a differentiation from south to north within the study area %
<figureCitation id="A7B35EB28F26F11EFE6FFEFCFE3EFEF8" box="[392,446,353,378]" captionStart="Fig" captionStartId="5.[136,165,1077,1098]" captionTargetId="figure-67@5.[314,1274,154,1030]" captionTargetPageId="5" captionText="Fig.2 Distribution map of all Psammotermes allocerus collections and their assignment to the genetic groups found in the phylogenetic analyses %colours). Circles: Collections used for the phylogeny. Black lines: Country borders. Northern Namib: %AH) Aba Huab rivier, %GV) Giribesvlakte, %HD) Hartmann Dunes, %HO) Hoada, %IO) Iona, %MF) Marienfluss, %PR) Purros, %ROE) Rössing mountain, %SO) Sorris Sorris, %TM) Tomakas; Western Kalahari Basin: %AHV) Alt-Hartebeestvlei, %AM) Alex Muranda, %GU) Gariganus, %KM) Katima Mulilo, %RI) Rimini, %RS) Rundu South, %RV) Ravenna, %SA) Samehaling, %SK) Swartkop, %WF2) Warmfontein; Nama: %BY) Barby, %FRC) Fish river Canyon, %GO) Goageb, %SH) Seeheim, %WV) Witvley; Southwestern" figureDoi="http://doi.org/10.5281/zenodo.12801593" httpUri="https://zenodo.org/record/12801593/files/figure.png" pageId="3" pageNumber="142">Fig. 2</figureCitation>
).
</paragraph>
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The most basal clade within
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<emphasis id="0DFC9E258F26F11EFE05FE19FD07FE19" box="[482,647,388,411]" italics="true" pageId="3" pageNumber="142">Psammotermes</emphasis>
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%dark blue,
<figureCitation id="A7B35EB28F26F11EFF6EFE39FF48FE3E" box="[137,200,420,444]" captionStart="Fig" captionStartId="4.[136,165,156,177]" captionTargetBox="[557,1447,158,1812]" captionTargetId="figure-86@4.[553,1451,154,1817]" captionTargetPageId="4" captionText="Fig. 1 Phylogeny of 65 P. allocerus collections inferred by the Bayesian analysis of COI and COII markers. Support values are given in posterior probability. Clades are coloured according to the genetic group. Dark blue: Succulent Karoo; Light Blue: Southern Namib; Dark green: East Gariep; Light Green: Southwestern Kalahari; Yellow: Nama; Ochre: Western Kalahari Basin; Red: Northern Namib. Abbreviations of study sites are shown in Fig. 2" figureDoi="http://doi.org/10.5281/zenodo.12764917" httpUri="https://zenodo.org/record/12764917/files/figure.png" pageId="3" pageNumber="142">Fig. 1</figureCitation>
) was the Succulent Karoo clade, as all seven samples are located within the Succulent Karoo biome in the north-western part of the winter rainfall region of
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%
<figureCitation id="A7B35EB28F26F11EFF0CFD95FEA2FDA2" box="[235,290,520,544]" captionStart="Fig" captionStartId="5.[136,165,1077,1098]" captionTargetId="figure-67@5.[314,1274,154,1030]" captionTargetPageId="5" captionText="Fig.2 Distribution map of all Psammotermes allocerus collections and their assignment to the genetic groups found in the phylogenetic analyses %colours). Circles: Collections used for the phylogeny. Black lines: Country borders. Northern Namib: %AH) Aba Huab rivier, %GV) Giribesvlakte, %HD) Hartmann Dunes, %HO) Hoada, %IO) Iona, %MF) Marienfluss, %PR) Purros, %ROE) Rössing mountain, %SO) Sorris Sorris, %TM) Tomakas; Western Kalahari Basin: %AHV) Alt-Hartebeestvlei, %AM) Alex Muranda, %GU) Gariganus, %KM) Katima Mulilo, %RI) Rimini, %RS) Rundu South, %RV) Ravenna, %SA) Samehaling, %SK) Swartkop, %WF2) Warmfontein; Nama: %BY) Barby, %FRC) Fish river Canyon, %GO) Goageb, %SH) Seeheim, %WV) Witvley; Southwestern" figureDoi="http://doi.org/10.5281/zenodo.12801593" httpUri="https://zenodo.org/record/12801593/files/figure.png" pageId="3" pageNumber="142">Fig. 2</figureCitation>
). It included the collection of the
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near
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%LUE,
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).
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next clearly distinguishable genetic group was the
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clade, with the majority of the samples located at the eastern margin of
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%
<figureCitation id="A7B35EB28F26F11EFE92FD10FE2BFD24" box="[373,427,653,678]" captionStart="Fig" captionStartId="5.[136,165,1077,1098]" captionTargetId="figure-67@5.[314,1274,154,1030]" captionTargetPageId="5" captionText="Fig.2 Distribution map of all Psammotermes allocerus collections and their assignment to the genetic groups found in the phylogenetic analyses %colours). Circles: Collections used for the phylogeny. Black lines: Country borders. Northern Namib: %AH) Aba Huab rivier, %GV) Giribesvlakte, %HD) Hartmann Dunes, %HO) Hoada, %IO) Iona, %MF) Marienfluss, %PR) Purros, %ROE) Rössing mountain, %SO) Sorris Sorris, %TM) Tomakas; Western Kalahari Basin: %AHV) Alt-Hartebeestvlei, %AM) Alex Muranda, %GU) Gariganus, %KM) Katima Mulilo, %RI) Rimini, %RS) Rundu South, %RV) Ravenna, %SA) Samehaling, %SK) Swartkop, %WF2) Warmfontein; Nama: %BY) Barby, %FRC) Fish river Canyon, %GO) Goageb, %SH) Seeheim, %WV) Witvley; Southwestern" figureDoi="http://doi.org/10.5281/zenodo.12801593" httpUri="https://zenodo.org/record/12801593/files/figure.png" pageId="3" pageNumber="142">Fig. 2</figureCitation>
).
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localities are very well supported %&gt; 0.99 pp), and only two are sisters to one other %RO and DV).
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next higher positioned clade was split into two larger subclades at an equal level of differentiation, one occurring in south-eastern
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, adjacent to
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.
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other clade includes samples from northern
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and
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.
<location id="3A5714EC8F26F11EFE92FCC8FE1FFCEF" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE92FCC8FE1FFCEF" box="[373,415,853,877]" country="South Africa" county="Luderitz" municipality="The" name="The" pageId="3" pageNumber="142" stateProvince="//Karas">The</location>
well-supported subclade %
<figureCitation id="A7B35EB28F26F11EFD23FCC8FD7AFCEC" box="[708,762,853,878]" captionStart="Fig" captionStartId="4.[136,165,156,177]" captionTargetBox="[557,1447,158,1812]" captionTargetId="figure-86@4.[553,1451,154,1817]" captionTargetPageId="4" captionText="Fig. 1 Phylogeny of 65 P. allocerus collections inferred by the Bayesian analysis of COI and COII markers. Support values are given in posterior probability. Clades are coloured according to the genetic group. Dark blue: Succulent Karoo; Light Blue: Southern Namib; Dark green: East Gariep; Light Green: Southwestern Kalahari; Yellow: Nama; Ochre: Western Kalahari Basin; Red: Northern Namib. Abbreviations of study sites are shown in Fig. 2" figureDoi="http://doi.org/10.5281/zenodo.12764917" httpUri="https://zenodo.org/record/12764917/files/figure.png" pageId="3" pageNumber="142">Fig. 1</figureCitation>
) includes collections from south-eastern
<collectingCountry id="479F02A78F26F11EFDADFCEAFD28FC0D" box="[586,680,887,911]" name="Namibia" pageId="3" pageNumber="142">Namibia</collectingCountry>
and the
<location id="3A5714EC8F26F11EFF6DFC04FEE8FC32" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFF6DFC04FEE8FC32" box="[138,360,920,944]" country="South Africa" county="Luderitz" municipality="The" name="eastern Richtersveld" pageId="3" pageNumber="142" stateProvince="//Karas">eastern Richtersveld</location>
%
<collectingCountry id="479F02A78F26F11EFE66FC05FD8CFC32" box="[385,524,920,944]" name="South Africa" pageId="3" pageNumber="142">South Africa</collectingCountry>
).
<location id="3A5714EC8F26F11EFDF8FC05FD54FC32" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFDF8FC05FD54FC32" box="[543,724,920,944]" country="South Africa" county="Luderitz" municipality="The" name="Phylogenetically" pageId="3" pageNumber="142" stateProvince="//Karas">Phylogenetically</location>
it is subdivided into three genetic groups:
<location id="3A5714EC8F26F11EFDF9FC27FD29FC53" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFDF9FC27FD29FC53" box="[542,681,953,978]" country="South Africa" county="Luderitz" municipality="The" name="East Gariep'" pageId="3" pageNumber="142" stateProvince="//Karas">East Gariep</location>
,
<location id="3A5714EC8F26F11EFD5FFC24FECBFC71" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFD5FFC24FECBFC71" country="South Africa" county="Luderitz" municipality="The" name="Southwestern Kalahari'" pageId="3" pageNumber="142" stateProvince="//Karas">Southwestern Kalahari</location>
and
<location id="3A5714EC8F26F11EFE61FC46FE54FC71" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE61FC46FE54FC71" box="[390,468,987,1011]" country="South Africa" county="Luderitz" municipality="The" name="Nama'" pageId="3" pageNumber="142" stateProvince="//Karas">Nama</location>
.
<location id="3A5714EC8F26F11EFE38FC46FD89FC71" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE38FC46FD89FC71" box="[479,521,987,1011]" country="South Africa" county="Luderitz" municipality="The" name="The" pageId="3" pageNumber="142" stateProvince="//Karas">The</location>
<location id="3A5714EC8F26F11EFDF2FC46FD23FC71" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFDF2FC46FD23FC71" box="[533,675,987,1011]" country="South Africa" county="Luderitz" municipality="The" name="East Gariep'" pageId="3" pageNumber="142" stateProvince="//Karas">East Gariep</location>
group is represented by 13 sites, of which six are located in the
<location id="3A5714EC8F26F11EFD25FC61FEE7FBB7" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFD25FC61FEE7FBB7" country="South Africa" county="Luderitz" municipality="The" name="northeastern Richtersveld" pageId="3" pageNumber="142" stateProvince="//Karas">northeastern Richtersveld</location>
and the adjacent larger
<location id="3A5714EC8F26F11EFD97FB83FF4AFBD5" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFD97FB83FF4AFBD5" country="South Africa" county="Luderitz" municipality="The" name="Orange River valley" pageId="3" pageNumber="142" stateProvince="//Karas">Orange River valley</location>
in
<collectingCountry id="479F02A78F26F11EFF0BFBA2FEC8FBD5" box="[236,328,1087,1111]" name="Namibia" pageId="3" pageNumber="142">Namibia</collectingCountry>
%
<figureCitation id="A7B35EB28F26F11EFE87FBA2FE15FBD5" box="[352,405,1087,1111]" captionStart="Fig" captionStartId="5.[136,165,1077,1098]" captionTargetId="figure-67@5.[314,1274,154,1030]" captionTargetPageId="5" captionText="Fig.2 Distribution map of all Psammotermes allocerus collections and their assignment to the genetic groups found in the phylogenetic analyses %colours). Circles: Collections used for the phylogeny. Black lines: Country borders. Northern Namib: %AH) Aba Huab rivier, %GV) Giribesvlakte, %HD) Hartmann Dunes, %HO) Hoada, %IO) Iona, %MF) Marienfluss, %PR) Purros, %ROE) Rössing mountain, %SO) Sorris Sorris, %TM) Tomakas; Western Kalahari Basin: %AHV) Alt-Hartebeestvlei, %AM) Alex Muranda, %GU) Gariganus, %KM) Katima Mulilo, %RI) Rimini, %RS) Rundu South, %RV) Ravenna, %SA) Samehaling, %SK) Swartkop, %WF2) Warmfontein; Nama: %BY) Barby, %FRC) Fish river Canyon, %GO) Goageb, %SH) Seeheim, %WV) Witvley; Southwestern" figureDoi="http://doi.org/10.5281/zenodo.12801593" httpUri="https://zenodo.org/record/12801593/files/figure.png" pageId="3" pageNumber="142">Fig. 2</figureCitation>
).
<location id="3A5714EC8F26F11EFE4BFBA2FE56FBD5" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE4BFBA2FE56FBD5" box="[428,470,1087,1111]" country="South Africa" county="Luderitz" municipality="The" name="The" pageId="3" pageNumber="142" stateProvince="//Karas">The</location>
weakly supported %0.52 pp) combined groups
<location id="3A5714EC8F26F11EFEA8FBFDFE1BFBFA" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFEA8FBFDFE1BFBFA" box="[335,411,1120,1144]" country="South Africa" county="Luderitz" municipality="The" name="Nama'" pageId="3" pageNumber="142" stateProvince="//Karas">Nama</location>
and
<location id="3A5714EC8F26F11EFE32FBFDFD5AFBFA" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE32FBFDFD5AFBFA" box="[469,730,1120,1144]" country="South Africa" county="Luderitz" municipality="The" name="Southwestern Kalahari'" pageId="3" pageNumber="142" stateProvince="//Karas">Southwestern Kalahari</location>
are sisters to the
<location id="3A5714EC8F26F11EFEC7FB1FFE2FFB1B" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFEC7FB1FFE2FFB1B" box="[288,431,1153,1178]" country="South Africa" county="Luderitz" municipality="The" name="East Gariep'" pageId="3" pageNumber="142" stateProvince="//Karas">East Gariep</location>
group.
<location id="3A5714EC8F26F11EFDE1FB1CFDBBFB1B" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFDE1FB1CFDBBFB1B" box="[518,571,1153,1177]" country="South Africa" county="Luderitz" municipality="The" name="Both" pageId="3" pageNumber="142" stateProvince="//Karas">Both</location>
individual groups %
<location id="3A5714EC8F26F11EFF7CFB3EFE07FB39" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFF7CFB3EFE07FB39" box="[155,391,1187,1211]" country="South Africa" county="Luderitz" municipality="The" name="Southwestern Kalahari" pageId="3" pageNumber="142" stateProvince="//Karas">Southwestern Kalahari</location>
and
<location id="3A5714EC8F26F11EFE5DFB3EFE7BFB39" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE5DFB3EFE7BFB39" box="[442,507,1187,1211]" country="South Africa" county="Luderitz" municipality="The" name="Nama" pageId="3" pageNumber="142" stateProvince="//Karas">Nama</location>
) are very well supported %1 pp).
<location id="3A5714EC8F26F11EFF30FB59FE81FB5E" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFF30FB59FE81FB5E" box="[215,257,1220,1244]" country="South Africa" county="Luderitz" municipality="The" name="The" pageId="3" pageNumber="142" stateProvince="//Karas">The</location>
<location id="3A5714EC8F26F11EFEEBFB59FD91FB5E" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFEEBFB59FD91FB5E" box="[268,529,1220,1244]" country="South Africa" county="Luderitz" municipality="The" name="Southwestern Kalahari'" pageId="3" pageNumber="142" stateProvince="//Karas">Southwestern Kalahari</location>
clade comprises eight collection sites situated in dunes and sand valleys east of the
<collectingRegion id="FD4C8CD58F26F11EFF55FA9AFF71FA9C" box="[178,241,1287,1310]" country="Namibia" name="//Karas" pageId="3" pageNumber="142">Karas</collectingRegion>
mountains.
<location id="3A5714EC8F26F11EFE9FFA9AFE22FA9D" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE9FFA9AFE22FA9D" box="[376,418,1287,1311]" country="South Africa" county="Luderitz" municipality="The" name="The" pageId="3" pageNumber="142" stateProvince="//Karas">The</location>
<location id="3A5714EC8F26F11EFE4AFA9AFE79FA9D" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE4AFA9AFE79FA9D" box="[429,505,1287,1311]" country="South Africa" county="Luderitz" municipality="The" name="Nama'" pageId="3" pageNumber="142" stateProvince="//Karas">Nama</location>
group %yellow) includes six study sites in the
<location id="3A5714EC8F26F11EFE94FAB5FDC8FAC2" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE94FAB5FDC8FAC2" box="[371,584,1320,1344]" country="South Africa" county="Luderitz" municipality="The" name="southern Namaland" pageId="3" pageNumber="142" stateProvince="//Karas">southern Namaland</location>
of the
<location id="3A5714EC8F26F11EFD7EFAB5FC84FAC2" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFD7EFAB5FC84FAC2" box="[665,772,1320,1344]" country="South Africa" county="Luderitz" municipality="The" name="Namibian" pageId="3" pageNumber="142" stateProvince="//Karas">Namibian</location>
<collectingRegion id="FD4C8CD58F26F11EFF6EFAD7FE9DFAE3" box="[137,285,1353,1377]" country="Namibia" name="//Karas" pageId="3" pageNumber="142">Karas Region</collectingRegion>
.
<location id="3A5714EC8F26F11EFEC0FAD4FED1FAE3" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFEC0FAD4FED1FAE3" box="[295,337,1353,1377]" country="South Africa" county="Luderitz" municipality="The" name="The" pageId="3" pageNumber="142" stateProvince="//Karas">The</location>
second larger subclade is well supported %0.92 pp) and included the groups
<location id="3A5714EC8F26F11EFE1FFAF6FC83FA01" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE1FFAF6FC83FA01" box="[504,771,1387,1411]" country="South Africa" county="Luderitz" municipality="The" name="Western Kalahari Basin'" pageId="3" pageNumber="142" stateProvince="//Karas">Western Kalahari Basin</location>
and
<location id="3A5714EC8F26F11EFF5AFA11FE05FA26" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFF5AFA11FE05FA26" box="[189,389,1420,1444]" country="South Africa" county="Luderitz" municipality="The" name="Northern Namib'" pageId="3" pageNumber="142" stateProvince="//Karas">Northern Namib</location>
, both very well supported %1 pp).
<location id="3A5714EC8F26F11EFF6DFA30FF34FA47" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFF6DFA30FF34FA47" box="[138,180,1453,1477]" country="South Africa" county="Luderitz" municipality="The" name="The" pageId="3" pageNumber="142" stateProvince="//Karas">The</location>
<location id="3A5714EC8F26F11EFF59FA33FE4AFA47" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFF59FA33FE4AFA47" box="[190,458,1453,1478]" country="South Africa" county="Luderitz" municipality="The" name="Western Kalahari Basin'" pageId="3" pageNumber="142" stateProvince="//Karas">Western Kalahari Basin</location>
comprises 11 collection sites ordered in a polytomy.
<location id="3A5714EC8F26F11EFE6AFA52FE3EFA65" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE6AFA52FE3EFA65" box="[397,446,1487,1511]" country="South Africa" county="Luderitz" municipality="The" name="This" pageId="3" pageNumber="142" stateProvince="//Karas">This</location>
group includes sites that are located from south-eastern
<collectingCountry id="479F02A78F26F11EFE4FFA6DFD83F98A" box="[424,515,1520,1544]" name="Namibia" pageId="3" pageNumber="142">Namibia</collectingCountry>
%SK3, SA) to the
<location id="3A5714EC8F26F11EFD59FA6CFF38F9AB" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFD59FA6CFF38F9AB" country="South Africa" county="Luderitz" municipality="The" name="Waterberg" pageId="3" pageNumber="142" stateProvince="//Karas">Waterberg</location>
%RI and RV) and the north-easternmost site
<location id="3A5714EC8F26F11EFD55F98CFF54F9C9" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFD55F98CFF54F9C9" country="South Africa" county="Luderitz" municipality="The" name="Katima Mulilo" pageId="3" pageNumber="142" stateProvince="//Karas">Katima Mulilo</location>
%KM).
<location id="3A5714EC8F26F11EFECEF9AEFED3F9C9" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFECEF9AEFED3F9C9" box="[297,339,1587,1611]" country="South Africa" county="Luderitz" municipality="The" name="The" pageId="3" pageNumber="142" stateProvince="//Karas">The</location>
<location id="3A5714EC8F26F11EFE87F9AEFDA4F9C9" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFE87F9AEFDA4F9C9" box="[352,548,1587,1611]" country="South Africa" county="Luderitz" municipality="The" name="Northern Namib'" pageId="3" pageNumber="142" stateProvince="//Karas">Northern Namib</location>
group encompassed all remaining study sites, located from the
<location id="3A5714EC8F26F11EFDA2F9C9FC83F9EE" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFDA2F9C9FC83F9EE" box="[581,771,1620,1644]" country="South Africa" county="Luderitz" municipality="The" name="Rossing mountain" pageId="3" pageNumber="142" stateProvince="//Karas">Rössing mountain</location>
%ROE) to the
<location id="3A5714EC8F26F11EFEFFF9EBFE62F90F" LSID="urn:lsid:plazi:treatment:B721F3218F27F11EFCD7FB5CFB76FA6B:3A5714EC8F26F11EFEFFF9EBFE62F90F" box="[280,482,1653,1677]" country="South Africa" county="Luderitz" municipality="The" name="Iona National Park" pageId="3" pageNumber="142" stateProvince="//Karas">Iona National Park</location>
in
<collectingCountry id="479F02A78F26F11EFDE3F9E8FDD3F90F" box="[516,595,1653,1677]" name="Angola" pageId="3" pageNumber="142">Angola</collectingCountry>
%IO).
</paragraph>
<paragraph id="3F3742378F26F11EFF41F90AFE37F8F5" blockId="3.[135,773,153,1911]" pageId="3" pageNumber="142">
Overall, the mean
<emphasis id="0DFC9E258F26F11EFE9DF905FE07F92D" box="[378,391,1688,1711]" italics="true" pageId="3" pageNumber="142">p</emphasis>
-distances are higher between the seven groups than within each group %
<tableCitation id="720A778C8F26F11EFDCAF925FD00F952" box="[557,640,1720,1744]" captionStart="Table 1" captionStartId="5.[136,185,1534,1555]" captionTargetPageId="5" captionText="Table 1 Estimates of evolutionary divergence for the concatenated alignment %COI and COII) for mean p-distances %±SE) between the groups" httpUri="http://table.plazi.org/id/6BF712BF8F20F118FF6FFA63FA13F991" pageId="3" pageNumber="142" tableUuid="6BF712BF8F20F118FF6FFA63FA13F991">Tables 1</tableCitation>
and
<tableCitation id="720A778C8F26F11EFD52F925FD42F952" box="[693,706,1720,1744]" captionStart="Table2" captionStartId="6.[137,184,156,177]" captionTargetPageId="6" captionText="Table2 Estimates of evolutionary divergence for the concatenated alignment %COI and COII) for mean p-distances %±SE) within the groups" httpUri="http://table.plazi.org/id/6BF712BF8F23F11BFF6EFF01FD57FF49" pageId="3" pageNumber="142" tableUuid="6BF712BF8F23F11BFF6EFF01FD57FF49">2</tableCitation>
). The mean
<emphasis id="0DFC9E258F26F11EFF2BF947FF59F973" box="[204,217,1754,1777]" italics="true" pageId="3" pageNumber="142">p</emphasis>
-distance between the groups ranges from 0.0508 %Nama vs. East Gariep) to 0.1007 %Kalahari Basin vs. Succulent Karoo). The values of the mean
<emphasis id="0DFC9E258F26F11EFD73F880FD21F8B6" box="[660,673,1821,1844]" italics="true" pageId="3" pageNumber="142">p</emphasis>
-distance are lower and ranged from 0.0100 %Succulent Karoo) to 0.0363 %Northern Namib).
</paragraph>
<paragraph id="3F3742378F26F11EFCD7FF07FB9FFF37" blockId="3.[816,1055,154,181]" box="[816,1055,154,181]" pageId="3" pageNumber="142">
<heading id="647FF55B8F26F11EFCD7FF07FB9FFF37" bold="true" box="[816,1055,154,181]" fontSize="11" level="1" pageId="3" pageNumber="142" reason="1">
<emphasis id="0DFC9E258F26F11EFCD7FF07FB9FFF37" bold="true" box="[816,1055,154,181]" pageId="3" pageNumber="142">Haplotype analyses</emphasis>
</heading>
</paragraph>
<paragraph id="3F3742378F26F11EFCD7FF43FBDAFB1E" blockId="3.[815,1453,222,1513]" pageId="3" pageNumber="142">
The median-joining network based on the concatenated alignment comprised 31 observed haplotypes, including 18 unique haplotypes %which includes a single sample) and 12 calculated hypothetical haplotypes %
<figureCitation id="A7B35EB28F26F11EFB52FEDFFB69FED8" box="[1205,1257,322,346]" captionStart="Fig" captionStartId="6.[136,165,999,1020]" captionTargetBox="[497,1443,1001,1904]" captionTargetId="figure-514@6.[491,1451,997,1908]" captionTargetPageId="6" captionText="Fig. 3 Median-joining network of the combined COI and COII P. allocerus sequences. Coloured circles represent the observed haplotypes, and the size is proportional to the num- ber of collections. Black circles represent missing haplotypes. Marks show the number of mutation steps. Haplotype num- bers are gained from DnaSP v6. Dotted lines and colours mark haplotypes according to the genetic group of the phylogeny" figureDoi="http://doi.org/10.5281/zenodo.12764919" httpUri="https://zenodo.org/record/12764919/files/figure.png" pageId="3" pageNumber="142">Fig. 3</figureCitation>
). Eight hypothetical haplotypes are situated in the centre of the median-joining network. The observed haplotypes of the seven genetic groups are linked over this centre. The highest value of unique haplotypes was calculated for the Southern Namib group with six haplotypes. In contrast, the Succulent Karoo comprised two haplotypes calculated for six sequences. The two groups, Western Kalahari Basin and Succulent Karoo, are connected to the centre with a high number of mutation steps %8 and 10,
<figureCitation id="A7B35EB28F26F11EFC60FDF3FC46FD04" box="[903,966,622,646]" captionStart="Fig" captionStartId="6.[136,165,999,1020]" captionTargetBox="[497,1443,1001,1904]" captionTargetId="figure-514@6.[491,1451,997,1908]" captionTargetPageId="6" captionText="Fig. 3 Median-joining network of the combined COI and COII P. allocerus sequences. Coloured circles represent the observed haplotypes, and the size is proportional to the num- ber of collections. Black circles represent missing haplotypes. Marks show the number of mutation steps. Haplotype num- bers are gained from DnaSP v6. Dotted lines and colours mark haplotypes according to the genetic group of the phylogeny" figureDoi="http://doi.org/10.5281/zenodo.12764919" httpUri="https://zenodo.org/record/12764919/files/figure.png" pageId="3" pageNumber="142">Fig. 3</figureCitation>
). The number of haplotypes differed slightly between the median-joining network analysis %
<figureCitation id="A7B35EB28F26F11EFADCFD0DFAF3FD2A" box="[1339,1395,656,680]" captionStart="Fig" captionStartId="6.[136,165,999,1020]" captionTargetBox="[497,1443,1001,1904]" captionTargetId="figure-514@6.[491,1451,997,1908]" captionTargetPageId="6" captionText="Fig. 3 Median-joining network of the combined COI and COII P. allocerus sequences. Coloured circles represent the observed haplotypes, and the size is proportional to the num- ber of collections. Black circles represent missing haplotypes. Marks show the number of mutation steps. Haplotype num- bers are gained from DnaSP v6. Dotted lines and colours mark haplotypes according to the genetic group of the phylogeny" figureDoi="http://doi.org/10.5281/zenodo.12764919" httpUri="https://zenodo.org/record/12764919/files/figure.png" pageId="3" pageNumber="142">Fig. 3</figureCitation>
) and the calculated number of haplotypes gained by DnaSP v6 %
<tableCitation id="720A778C8F26F11EFCA4FD4FFC08FD68" box="[835,904,722,746]" captionStart="Table 3" captionStartId="7.[136,184,156,177]" captionTargetPageId="7" captionText="Table 3 Results of the genetic diversity indices calculated for the con- number of segregating sites %S), nucleotide diversity %π), and haplocatenated alignment of COI and COII markers and seven clades within type diversity %Hd), Tajimas D value %nonsignificant=ns, for p&gt;0.10) P. allocerus: number of haplotypes %NH), number of sequences %NS)," httpUri="http://table.plazi.org/id/6BF712BF8F22F11AFF6FFF01FC84FF64" pageId="3" pageNumber="142" tableUuid="6BF712BF8F22F11AFF6FFF01FC84FF64">Table 3</tableCitation>
). The nucleotide diversity %π) ranged from 0.00416 %Western Kalahari Basin) to 0.01597 %Northern Namib,
<tableCitation id="720A778C8F26F11EFCD6FC88FC01FCAF" box="[817,897,789,813]" captionStart="Table 3" captionStartId="7.[136,184,156,177]" captionTargetPageId="7" captionText="Table 3 Results of the genetic diversity indices calculated for the con- number of segregating sites %S), nucleotide diversity %π), and haplocatenated alignment of COI and COII markers and seven clades within type diversity %Hd), Tajimas D value %nonsignificant=ns, for p&gt;0.10) P. allocerus: number of haplotypes %NH), number of sequences %NS)," httpUri="http://table.plazi.org/id/6BF712BF8F22F11AFF6FFF01FC84FF64" pageId="3" pageNumber="142" tableUuid="6BF712BF8F22F11AFF6FFF01FC84FF64">Table 3</tableCitation>
). The lowest haplotype diversity was calculated for the Succulent Karoo with 0.78571 and the highest for the Southern Namib with 1.0000 %
<tableCitation id="720A778C8F26F11EFB46FCC5FB6CFCF2" box="[1185,1260,856,880]" captionStart="Table 3" captionStartId="7.[136,184,156,177]" captionTargetPageId="7" captionText="Table 3 Results of the genetic diversity indices calculated for the con- number of segregating sites %S), nucleotide diversity %π), and haplocatenated alignment of COI and COII markers and seven clades within type diversity %Hd), Tajimas D value %nonsignificant=ns, for p&gt;0.10) P. allocerus: number of haplotypes %NH), number of sequences %NS)," httpUri="http://table.plazi.org/id/6BF712BF8F22F11AFF6FFF01FC84FF64" pageId="3" pageNumber="142" tableUuid="6BF712BF8F22F11AFF6FFF01FC84FF64">Table 3</tableCitation>
). The number of segregating sites %S) was highest within the Southern Namib group with 22 S and lowest within the Succulent Karoo and Western Kalahari Basin with each six S %
<tableCitation id="720A778C8F26F11EFAC2FC21FAECFC56" box="[1317,1388,956,980]" captionStart="Table 3" captionStartId="7.[136,184,156,177]" captionTargetPageId="7" captionText="Table 3 Results of the genetic diversity indices calculated for the con- number of segregating sites %S), nucleotide diversity %π), and haplocatenated alignment of COI and COII markers and seven clades within type diversity %Hd), Tajimas D value %nonsignificant=ns, for p&gt;0.10) P. allocerus: number of haplotypes %NH), number of sequences %NS)," httpUri="http://table.plazi.org/id/6BF712BF8F22F11AFF6FFF01FC84FF64" pageId="3" pageNumber="142" tableUuid="6BF712BF8F22F11AFF6FFF01FC84FF64">Table 3</tableCitation>
). The genetic groups undergo a different selection pressure. A positive Tajimas D value was calculated for the groups: Northern Namib, Western Kalahari Basin and Southwestern Kalahari and a negative Tajimas D value was estimated for the remaining groups %Nama, Southern Namib, East Gariep, Succulent Karoo,
<tableCitation id="720A778C8F26F11EFC1CFB19FBCAFB1E" box="[1019,1098,1156,1180]" captionStart="Table 3" captionStartId="7.[136,184,156,177]" captionTargetPageId="7" captionText="Table 3 Results of the genetic diversity indices calculated for the con- number of segregating sites %S), nucleotide diversity %π), and haplocatenated alignment of COI and COII markers and seven clades within type diversity %Hd), Tajimas D value %nonsignificant=ns, for p&gt;0.10) P. allocerus: number of haplotypes %NH), number of sequences %NS)," httpUri="http://table.plazi.org/id/6BF712BF8F22F11AFF6FFF01FC84FF64" pageId="3" pageNumber="142" tableUuid="6BF712BF8F22F11AFF6FFF01FC84FF64">Table 3</tableCitation>
).
</paragraph>
<paragraph id="3F3742378F26F11EFCA8FB38FB76FA6B" blockId="3.[815,1453,222,1513]" pageId="3" pageNumber="142">
Two additional features distinguish the Succulent Karoo group from the other groups. The tapetum of the tunnels within the nest system of
<taxonomicName id="F88839B48F26F11EFBA1FB75FB46FA82" authorityName="% Uys" authorityYear="2002" box="[1094,1222,1256,1280]" class="Insecta" family="Rhinotermitidae" genus="Psammotermes" kingdom="Animalia" order="Blattodea" pageId="3" pageNumber="142" phylum="Arthropoda" rank="species" species="allocerus">
<emphasis id="0DFC9E258F26F11EFBA1FB75FB46FA82" box="[1094,1222,1256,1280]" italics="true" pageId="3" pageNumber="142">P. allocerus</emphasis>
</taxonomicName>
colonies in the Succulent Karoo is whitish. In all other groups, it is blackish %
<figureCitation id="A7B35EB28F26F11EFCA4FAB7FCF7FAC0" box="[835,887,1322,1346]" captionStart="Fig" captionStartId="7.[136,165,1319,1340]" captionTargetBox="[491,1450,1318,1908]" captionTargetId="figure-645@7.[491,1451,1317,1908]" captionTargetPageId="7" captionText="Fig.4 Differences in the tape- tum colour of three P. allocerus colonies and the royal pair. A Whitish tapetum of a nest from the Springklipplain from the Succulent Karoo group %South Africa, 26 September 2016). B First image of the king and queen of P. allocerus from Yellow Dune %Succulent Karoo, South Africa, 08 March 2015). C Blackish tapetum of a nest from Dieprivier %Southern Namib, Namibia, 05 April 2017). D Blackish tapetum and chambers filled with foraged grass from Iona %Northern Namib, Angola, 26 September 2016). Images taken by Norbert Jürgens, Felicitas Gunter" figureDoi="http://doi.org/10.5281/zenodo.12764921" httpUri="https://zenodo.org/record/12764921/files/figure.png" pageId="3" pageNumber="142">Fig. 4</figureCitation>
). Only in the Succulent Karoo nests was it possible to locate the royal chambers with paired kings and queens at a depth of
<quantity id="F870EFD28F26F11EFC58FAF0FBC0FA06" box="[959,1088,1389,1413]" metricMagnitude="-1" metricUnit="m" metricValue="2.25" metricValueMax="3.0" metricValueMin="1.5" pageId="3" pageNumber="142" unit="cm" value="22.5" valueMax="30.0" valueMin="15.0">15 to 30 cm</quantity>
below the soil surface %
<figureCitation id="A7B35EB28F26F11EFAADFAF0FAFEFA07" box="[1354,1406,1389,1413]" captionStart="Fig" captionStartId="7.[136,165,1319,1340]" captionTargetBox="[491,1450,1318,1908]" captionTargetId="figure-645@7.[491,1451,1317,1908]" captionTargetPageId="7" captionText="Fig.4 Differences in the tape- tum colour of three P. allocerus colonies and the royal pair. A Whitish tapetum of a nest from the Springklipplain from the Succulent Karoo group %South Africa, 26 September 2016). B First image of the king and queen of P. allocerus from Yellow Dune %Succulent Karoo, South Africa, 08 March 2015). C Blackish tapetum of a nest from Dieprivier %Southern Namib, Namibia, 05 April 2017). D Blackish tapetum and chambers filled with foraged grass from Iona %Northern Namib, Angola, 26 September 2016). Images taken by Norbert Jürgens, Felicitas Gunter" figureDoi="http://doi.org/10.5281/zenodo.12764921" httpUri="https://zenodo.org/record/12764921/files/figure.png" pageId="3" pageNumber="142">Fig. 4</figureCitation>
). In all other groups, one of the authors %NJ) never locate royal chambers or kings or queens in the first
<quantity id="F870EFD28F26F11EFB2DFA2DFA97FA45" box="[1226,1303,1456,1480]" metricMagnitude="0" metricUnit="m" metricValue="1.0" pageId="3" pageNumber="142" unit="cm" value="100.0">100 cm</quantity>
below the soil surface, despite some 100 nest excavations.
</paragraph>
</subSubSection>
</treatment>
</document>