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<treatment id="372087BCFFF29121FCEB3DE914557E44" ID-DOI="http://doi.org/10.5281/zenodo.7195081" ID-GBIF-Taxon="202610623" ID-Zenodo-Dep="7195081" LSID="urn:lsid:plazi:treatment:372087BCFFF29121FCEB3DE914557E44" httpUri="http://treatment.plazi.org/id/372087BCFFF29121FCEB3DE914557E44" lastPageId="11" lastPageNumber="934" pageId="9" pageNumber="933">
<subSubSection id="F7936521FFF29123FCEB3DE9131279B5" box="[855,1379,1746,1769]" pageId="9" pageNumber="933" type="nomenclature">
<paragraph id="BF3636AAFFF29123FCEB3DE9131279B5" blockId="9.[855,1379,1746,1769]" box="[855,1379,1746,1769]" pageId="9" pageNumber="933">
<heading id="E47E81C6FFF29123FCEB3DE9131279B5" box="[855,1379,1746,1769]" centered="true" fontSize="9" level="2" pageId="9" pageNumber="933" reason="2">
<taxonomicName id="78894D29FFF29123FCEB3DE9126679B5" ID-CoL="7Q5FB" authorityName="Enghoff" authorityYear="2017" box="[855,1047,1746,1769]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="9" pageNumber="933" phylum="Arthropoda" rank="genus">
<emphasis id="8DFDEAB8FFF29123FCEB3DE9126679B5" box="[855,1047,1746,1769]" italics="true" pageId="9" pageNumber="933">TROPOSTREPTUS</emphasis>
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PHYLOGENY AND EVOLUTION
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Twenty-one of our included mitochondrial genomes belong to
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<emphasis id="8DFDEAB8FFF29123FC1B3C2212237872" box="[935,1106,1817,1838]" italics="true" pageId="9" pageNumber="933">Tropostreptus</emphasis>
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, allowing for a thorough investigation of the evolution of this genus in the Eastern Arc. We observe a clear genetic structure in
<taxonomicName id="78894D29FFF19120FF1F3BFD17337F87" authorityName="Enghoff" authorityYear="2017" box="[163,322,198,219]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="genus">
<emphasis id="8DFDEAB8FFF19120FF1F3BFD17337F87" box="[163,322,198,219]" italics="true" pageId="10" pageNumber="934">Tropostreptus</emphasis>
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, with distinct lineages (both inter- and intraspecific) being defined by the mountain blocks (
<figureCitation id="27B22A2FFFF19120FF103A3916857E44" box="[172,244,258,280]" captionStart="Figure 4" captionStartId="8.[163,241,1164,1186]" captionTargetBox="[165,1447,197,1122]" captionTargetId="figure-260@8.[163,1447,195,1125]" captionTargetPageId="8" captionText="Figure 4. Maximum likelihood-based phylogeny, with 100 bootstrap replicates and using all the 26 mitochondrial genomes generated in this study. The dataset was supplemented with Thyropygus sp. and Abacion magnum as outgroups, with sequences derived from GenBank. GenBank accession numbers are given in parentheses. Colours represent Tropostreptus sample origins. The upper right inset shows the topology of the Tropostreptus hamatus lineage, enlarged to clarify the branchingorder.Onlysupportvalues&lt;100 areshown.*Thyropygus sp.(redfont)isverylikely to beaspecies misidentification; for more information, see Discussion text." figureDoi="http://doi.org/10.5281/zenodo.7184596" httpUri="https://zenodo.org/record/7184596/files/figure.png" pageId="10" pageNumber="934">Fig. 4</figureCitation>
). This is consistent with previous genetic results of Eastern Arc gene pools (e.g. cat snakes:
<bibRefCitation id="DB184B5BFFF19120FF1F3A7B17167E09" author="Gravlund P" box="[163,359,320,342]" pageId="10" pageNumber="934" pagination="46 - 56" refId="ref10104" refString="Gravlund P. 2002. Molecular phylogeny of Tornier's cat snake (Crotaphopeltis tornieri), endemic to East African mountain forests: biogeography, vicariance events and problematic species boundaries. Journal of Zoological Systematics and Evolutionary Research 40: 46 - 56." type="journal article" year="2002">Gravlund, 2002</bibRefCitation>
; chameleons:
<bibRefCitation id="DB184B5BFFF19120FDA13A7B15727E09" author="Tolley KA &amp; Tilbury CR &amp; Measey GJ &amp; Menegon M &amp; Branch WR &amp; Matthee CA" box="[541,771,320,342]" pageId="10" pageNumber="934" pagination="1748 - 1760" refId="ref11779" refString="Tolley KA, Tilbury CR, Measey GJ, Menegon M, Branch WR, Matthee CA. 2011. Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hotspot. Journal of Biogeography 3 8: 1748 - 1760." type="journal article" year="2011">
Tolley
<emphasis id="8DFDEAB8FFF19120FDC93A7A14C47E09" box="[629,693,320,342]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2011
</bibRefCitation>
; African violets:
<bibRefCitation id="DB184B5BFFF19120FEDC3A6514247E28" author="Dimitrov D &amp; Nogues-Bravo D &amp; Scharff N" box="[352,597,350,372]" pageId="10" pageNumber="934" pagination="48908" refId="ref9610" refString="Dimitrov D, Nogues-Bravo D, Scharff N. 2012. Why do tropical mountains support exceptionally high biodiversity? The Eastern Arc mountains and the drivers of Saintpaulia diversity. PLoS One 7: e 48908." type="journal article" year="2012">
Dimitrov
<emphasis id="8DFDEAB8FFF19120FE6F3A64147E7E2F" box="[467,527,350,372]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2012
</bibRefCitation>
), where forestadapted species inhabit the montane forests and are absent from the adjacent savannah lowlands. Today, the mountains capture the oceanic winds from the Indian Ocean, which maintains sufficient humidity for dense rain forest to grow, resulting in the forest ‘sky islands’ (
<bibRefCitation id="DB184B5BFFF19120FEB7392C17DB7D70" author="Lovett JC" box="[267,426,535,556]" pageId="10" pageNumber="934" pagination="23 - 29" refId="ref10593" refString="Lovett JC. 1993 a. Climatic history and forest distribution in eastern Africa. In: JC Lovett, SK Wasser, eds. Biogeography and ecology of the rain forests of Eastern Africa. Cambridge: Cambridge University Press, 23 - 29." type="book chapter" year="1993">Lovett, 1993a</bibRefCitation>
, b;
<bibRefCitation id="DB184B5BFFF19120FE6F392C14C67D70" author="Burgess N &amp; Butynski TM &amp; Cordeiro NJ &amp; Doggart NH &amp; Fjeldsa J &amp; Howell KM &amp; Kilahama FB &amp; Loader SP &amp; Lovett JC &amp; Mbilinyi B &amp; Menegon M" box="[467,695,534,556]" pageId="10" pageNumber="934" pagination="209 - 231" refId="ref9320" refString="Burgess N, Butynski TM, Cordeiro NJ, Doggart NH, Fjeldsa J, Howell KM, Kilahama FB, Loader SP, Lovett JC, Mbilinyi B, Menegon M. 2007. The biological importance of the Eastern Arc Mountains of Tanzania and Kenya. Biological Conservation 134: 209 - 231." type="journal article" year="2007">
Burgess
<emphasis id="8DFDEAB8FFF19120FD84392C14037D77" box="[568,626,534,556]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2007
</bibRefCitation>
). Until 30 Mya, the Eastern Arc region is thought to have been covered by rain forest (
<bibRefCitation id="DB184B5BFFF19120FE51396F14E17D35" author="Rodgers WA" box="[493,656,596,618]" pageId="10" pageNumber="934" pagination="7 - 18" refId="ref11353" refString="Rodgers WA. 1998. An introduction to the conservation of the Eastern Arc Mountains. Journal of East African Natural History 87: 7 - 18." type="journal article" year="1998">Rodgers, 1998</bibRefCitation>
;
<bibRefCitation id="DB184B5BFFF19120FD21396F175F7DD4" author="Couvreur TLP &amp; Chatrou LW &amp; Sosef MSM &amp; Richardson JE" pageId="10" pageNumber="934" pagination="54" refId="ref9512" refString="Couvreur TLP, Chatrou LW, Sosef MSM, Richardson JE. 2008. Molecular phylogenetics reveal multiple tertiary vicariance origins of the African rain forest trees. BMC Biology 6: 54." type="journal article" year="2008">
Couvreur
<emphasis id="8DFDEAB8FFF19120FF1F394816937DDB" box="[163,226,626,648]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2008
</bibRefCitation>
), and an uplifting of the Eastern Arc Mountains is believed to have occurred within the last 7 Myr (although this is debated), changing the whole topography of East Africa (
<bibRefCitation id="DB184B5BFFF19120FE6B39F5140F7DB8" author="Griffiths CJ" box="[471,638,718,740]" pageId="10" pageNumber="934" pagination="9 - 22" refId="ref10149" refString="Griffiths CJ. 1993. The geological evolution of East Africa. In: JC Lovett, SK Wasser, eds. Biogeography and Ecology of the Rain Forests of Eastern Africa, Cambridge: Cambridge University Press, 9 - 22." type="book chapter" year="1993">Griffiths, 1993</bibRefCitation>
;
<bibRefCitation id="DB184B5BFFF19120FD3739F515747DB8" author="Ring U" box="[651,773,718,740]" pageId="10" pageNumber="934" pagination="132 - 146" refId="ref11291" refString="Ring U. 2014. The East African rift system. Austrian Journal of Earth Sciences 107: 132 - 146." type="journal article" year="2014">Ring, 2014</bibRefCitation>
;
<bibRefCitation id="DB184B5BFFF19120FF1F39D5172C7C5F" author="Macgregor D" box="[163,349,749,771]" pageId="10" pageNumber="934" pagination="232 - 252" refId="ref10682" refString="Macgregor D. 2015. History of the development of the East African Rift System: a series of interpreted maps through time. Journal of African Earth Sciences 101: 232 - 252." type="journal article" year="2015">Macgregor, 2015</bibRefCitation>
). Climatic and geological fluctuations through time have thus repeatedly affected the forest cover and, presumably, resulted in a multitude of vicariance events when species were isolated in patchy forest remnants (
<bibRefCitation id="DB184B5BFFF19120FED4385314727C21" author="Lovett JC" box="[360,515,872,893]" pageId="10" pageNumber="934" pagination="23 - 29" refId="ref10593" refString="Lovett JC. 1993 a. Climatic history and forest distribution in eastern Africa. In: JC Lovett, SK Wasser, eds. Biogeography and ecology of the rain forests of Eastern Africa. Cambridge: Cambridge University Press, 23 - 29." type="book chapter" year="1993">Lovett, 1993a</bibRefCitation>
;
<bibRefCitation id="DB184B5BFFF19120FDAC385315747C22" author="Sepulchre P &amp; Ramstein G &amp; Fluteau F &amp; Schuster M &amp; Tiercelin J-J &amp; Brunet M" box="[528,773,872,894]" pageId="10" pageNumber="934" pagination="1419 - 1423" refId="ref11473" refString="Sepulchre P, Ramstein G, Fluteau F, Schuster M, Tiercelin J-J, Brunet M. 2006. Tectonic uplift and Eastern Africa aridification. Science 313: 1419 - 1423." type="journal article" year="2006">
Sepulchre
<emphasis id="8DFDEAB8FFF19120FD35385214B07C21" box="[649,705,872,894]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2006
</bibRefCitation>
;
<bibRefCitation id="DB184B5BFFF19120FF1F38BC17E37CC0" author="Couvreur TLP &amp; Chatrou LW &amp; Sosef MSM &amp; Richardson JE" box="[163,402,902,924]" pageId="10" pageNumber="934" pagination="54" refId="ref9512" refString="Couvreur TLP, Chatrou LW, Sosef MSM, Richardson JE. 2008. Molecular phylogenetics reveal multiple tertiary vicariance origins of the African rain forest trees. BMC Biology 6: 54." type="journal article" year="2008">
Couvreur
<emphasis id="8DFDEAB8FFF19120FEAA38BC173F7CC7" box="[278,334,902,924]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2008
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). For these reasons, the splitting order we observe in the
<taxonomicName id="78894D29FFF19120FE10389D14397CE7" authorityName="Enghoff" authorityYear="2017" box="[428,584,934,955]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="genus">
<emphasis id="8DFDEAB8FFF19120FE10389D14397CE7" box="[428,584,934,955]" italics="true" pageId="10" pageNumber="934">Tropostreptus</emphasis>
</taxonomicName>
phylogeny might well reflect forest fragmentation in ancient times.
</paragraph>
<paragraph id="BF3636AAFFF19120FF0738D817C97ABE" blockId="10.[161,780,197,1906]" pageId="10" pageNumber="934">
We observe a general trend, whereby northern lineages appear to split off first. The earliest split in
<taxonomicName id="78894D29FFF19120FF1F3F1A17377B6A" authorityName="Enghoff" authorityYear="2017" box="[163,326,1057,1078]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="genus">
<emphasis id="8DFDEAB8FFF19120FF1F3F1A17377B6A" box="[163,326,1057,1078]" italics="true" pageId="10" pageNumber="934">Tropostreptus</emphasis>
</taxonomicName>
separates the
<taxonomicName id="78894D29FFF19120FE463F1A157A7B6A" baseAuthorityName="Attems" baseAuthorityYear="1950" box="[506,779,1057,1078]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="austerus">
<emphasis id="8DFDEAB8FFF19120FE463F1A157A7B6A" box="[506,779,1057,1078]" italics="true" pageId="10" pageNumber="934">Tropostreptus austerus</emphasis>
</taxonomicName>
+
<taxonomicName id="78894D29FFF19120FF073F0417B37B08" box="[187,450,1087,1108]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="severus">
<emphasis id="8DFDEAB8FFF19120FF073F0417B37B08" box="[187,450,1087,1108]" italics="true" pageId="10" pageNumber="934">Tropostreptus severus</emphasis>
</taxonomicName>
lineage from the rest, and the second split separates these two species, today occupying Nguru and Usambara Mountains in the north. A similar intraspecific pattern is evident in more recent splits in
<taxonomicName id="78894D29FFF19120FE083F8114A27B92" box="[436,723,1209,1231]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="hamatus">
<emphasis id="8DFDEAB8FFF19120FE083F8114A27B92" box="[436,723,1209,1231]" italics="true" pageId="10" pageNumber="934">Tropostreptus hamatus</emphasis>
</taxonomicName>
and
<taxonomicName id="78894D29FFF19120FF1F3FE214717BB1" box="[163,512,1240,1262]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="sigmatospinus">
<emphasis id="8DFDEAB8FFF19120FF1F3FE214717BB1" box="[163,512,1240,1262]" italics="true" pageId="10" pageNumber="934">Tropostreptus sigmatospinus</emphasis>
</taxonomicName>
(
<figureCitation id="27B22A2FFFF19120FDAF3FE314297BB1" box="[531,600,1240,1262]" captionStart="Figure 4" captionStartId="8.[163,241,1164,1186]" captionTargetBox="[165,1447,197,1122]" captionTargetId="figure-260@8.[163,1447,195,1125]" captionTargetPageId="8" captionText="Figure 4. Maximum likelihood-based phylogeny, with 100 bootstrap replicates and using all the 26 mitochondrial genomes generated in this study. The dataset was supplemented with Thyropygus sp. and Abacion magnum as outgroups, with sequences derived from GenBank. GenBank accession numbers are given in parentheses. Colours represent Tropostreptus sample origins. The upper right inset shows the topology of the Tropostreptus hamatus lineage, enlarged to clarify the branchingorder.Onlysupportvalues&lt;100 areshown.*Thyropygus sp.(redfont)isverylikely to beaspecies misidentification; for more information, see Discussion text." figureDoi="http://doi.org/10.5281/zenodo.7184596" httpUri="https://zenodo.org/record/7184596/files/figure.png" pageId="10" pageNumber="934">Fig. 4</figureCitation>
), suggesting a repeated pattern of vicariance events occurring first in the north. A separation of species between northern and southern mountains has also been observed in several other Eastern Arc taxa, including amphibians (
<bibRefCitation id="DB184B5BFFF19120FF173E4A17917ADA" author="Blackburn DC &amp; Measey GJ" box="[171,480,1393,1415]" pageId="10" pageNumber="934" pagination="1904 - 1915" refId="ref9107" refString="Blackburn DC, Measey GJ. 2009. Dispersal to or from an African biodiversity hotspot? Molecular Ecology 18: 1904 - 1915." type="journal article" year="2009">Blackburn &amp; Measey, 2009</bibRefCitation>
), gastropods (
<bibRefCitation id="DB184B5BFFF19120FD383E4A176F7AF9" author="Tattersfield P &amp; Seddon MB &amp; Meena C" pageId="10" pageNumber="934" pagination="119 - 138" refId="ref11630" refString="Tattersfield P, Seddon MB, Meena C. 1998. Ecology and conservation of the land-snails of the Eastern Arc Mountains. Journal of East African Natural History 87: 119 - 138." type="journal article" year="1998">
Tattersfield
<emphasis id="8DFDEAB8FFF19120FF1F3EAA16AA7AF9" box="[163,219,1424,1446]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 1998
</bibRefCitation>
) and reptiles (
<bibRefCitation id="DB184B5BFFF19120FE793EAB14077AF9" author="Gravlund P" box="[453,630,1424,1446]" pageId="10" pageNumber="934" pagination="46 - 56" refId="ref10104" refString="Gravlund P. 2002. Molecular phylogeny of Tornier's cat snake (Crotaphopeltis tornieri), endemic to East African mountain forests: biogeography, vicariance events and problematic species boundaries. Journal of Zoological Systematics and Evolutionary Research 40: 46 - 56." type="journal article" year="2002">Gravlund, 2002</bibRefCitation>
;
<bibRefCitation id="DB184B5BFFF19120FD3D3EAB16AA7A98" author="Tolley KA &amp; Tilbury CR &amp; Measey GJ &amp; Menegon M &amp; Branch WR &amp; Matthee CA" pageId="10" pageNumber="934" pagination="1748 - 1760" refId="ref11779" refString="Tolley KA, Tilbury CR, Measey GJ, Menegon M, Branch WR, Matthee CA. 2011. Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hotspot. Journal of Biogeography 3 8: 1748 - 1760." type="journal article" year="2011">
Tolley
<emphasis id="8DFDEAB8FFF19120FD713EAA15747AF9" box="[717,773,1424,1446]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2011
</bibRefCitation>
), but also in well-dispersing taxa, such as birds (
<bibRefCitation id="DB184B5BFFF19120FF173EF617D87ABE" author="Fjeldsa J &amp; Bowie RCK" box="[171,425,1485,1507]" pageId="10" pageNumber="934" pagination="235 - 247" refId="ref10041" refString="Fjeldsa J, Bowie RCK. 2008. New perspectives on the origin and diversification of Africa's forest avifauna. African Journal of Ecology 46: 235 - 247." type="journal article" year="2008">Fjeldså &amp; Bowie, 2008</bibRefCitation>
).
</paragraph>
<paragraph id="BF3636AAFFF19120FF073ED7159E7E09" blockId="10.[161,780,197,1906]" lastBlockId="10.[824,1444,197,1906]" pageId="10" pageNumber="934">
This indicates a forest retraction southwards during dry periods, resulting in vicariance events, followed by forest expansion and thus northward recolonization of species during periods with higher humidity. Northward migration is also observed in other Eastern Arc species, such as chameleons (
<bibRefCitation id="DB184B5BFFF19120FD3D3DBE16AA79E5" author="Tolley KA &amp; Tilbury CR &amp; Measey GJ &amp; Menegon M &amp; Branch WR &amp; Matthee CA" pageId="10" pageNumber="934" pagination="1748 - 1760" refId="ref11779" refString="Tolley KA, Tilbury CR, Measey GJ, Menegon M, Branch WR, Matthee CA. 2011. Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hotspot. Journal of Biogeography 3 8: 1748 - 1760." type="journal article" year="2011">
Tolley
<emphasis id="8DFDEAB8FFF19120FD713DBD157479C6" box="[717,773,1669,1691]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2011
</bibRefCitation>
;
<bibRefCitation id="DB184B5BFFF19120FF553D9F179679E5" author="Ceccarelli FS &amp; Menegon M &amp; Tolley KA &amp; Tilbury CR &amp; Gower DJ &amp; Laserna MH &amp; Kasahun R &amp; Prieto A &amp; Hagmann R &amp; Loader SP" box="[233,487,1700,1722]" pageId="10" pageNumber="934" pagination="125 - 136" refId="ref9424" refString="Ceccarelli FS, Menegon M, Tolley KA, Tilbury CR, Gower DJ, Laserna MH, Kasahun R, Rodriguez- Prieto A, Hagmann R, Loader SP. 2014. Evolutionary relationships, species delimitation and biogeography of Eastern Afromontane horned chameleons (Chamaeleonidae: Trioceros). Molecular Phylogenetics and Evolution 80: 125 - 136." type="journal article" year="2014">
Ceccarelli
<emphasis id="8DFDEAB8FFF19120FEDA3D9E17D079E5" box="[358,417,1700,1722]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2014
</bibRefCitation>
). A recent cycle of forest expansion/retraction can explain why
<taxonomicName id="78894D29FFF19120FDD53DF8177E79AA" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="hamatus">
<emphasis id="8DFDEAB8FFF19120FDD53DF8177E79AA" italics="true" pageId="10" pageNumber="934">Tropostreptus hamatus</emphasis>
</taxonomicName>
and
<taxonomicName id="78894D29FFF19120FEE93DD914B279AB" box="[341,707,1762,1783]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="sigmatospinus">
<emphasis id="8DFDEAB8FFF19120FEE93DD914B279AB" box="[341,707,1762,1783]" italics="true" pageId="10" pageNumber="934">Tropostreptus sigmatospinus</emphasis>
</taxonomicName>
exist across several of the mountains without having evolved into distinct species yet. Other events have isolated
<taxonomicName id="78894D29FFF19120FEB93C05178F780E" box="[261,510,1853,1875]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="kipunji">
<emphasis id="8DFDEAB8FFF19120FEB93C05178F780E" box="[261,510,1853,1875]" italics="true" pageId="10" pageNumber="934">Tropostreptus kipunji</emphasis>
</taxonomicName>
in the forest on Mount Rungwe, the most south-westerly occurring species in the Eastern Arc region, in addition to
<taxonomicName id="78894D29FFF19120FABD3BFD15947FA5" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="sigmatospinus">
<emphasis id="8DFDEAB8FFF19120FABD3BFD15947FA5" italics="true" pageId="10" pageNumber="934">Tropostreptus sigmatospinus</emphasis>
</taxonomicName>
in
<collectingCountry id="C79E763AFFF19120FBB13BDF12077FA6" box="[1037,1142,228,250]" name="Tanzania" pageId="10" pageNumber="934">Zanzibar</collectingCountry>
and, potentially, also the Rondo Plateau, from where
<taxonomicName id="78894D29FFF19120FB303A38135E7E44" authorityName="Enghoff" authorityYear="2017" box="[1164,1327,259,280]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="genus">
<emphasis id="8DFDEAB8FFF19120FB303A38135E7E44" box="[1164,1327,259,280]" italics="true" pageId="10" pageNumber="934">Tropostreptus</emphasis>
</taxonomicName>
has been observed but for which molecular data are still lacking (
<bibRefCitation id="DB184B5BFFF19120FCFF3A7B15917E09" author="Enghoff H" box="[835,992,320,342]" pageId="10" pageNumber="934" pagination="501 - 530" refId="ref9734" refString="Enghoff H. 2017. A new East African genus of spirostreptid millipedes (Diplopoda, Spirostreptida, Spirostreptidae), with notes on their fungal ectoparasite Rickia gigas. Zootaxa 4273: 501 - 530." type="journal article" year="2017">Enghoff, 2017</bibRefCitation>
).
</paragraph>
<paragraph id="BF3636AAFFF19120FCEF3A65123C7A50" blockId="10.[824,1444,197,1906]" pageId="10" pageNumber="934">
Regarding the timing of the species splits (
<figureCitation id="27B22A2FFFF19120FAC93A6515387ECF" captionStart="Figure 5" captionStartId="9.[146,228,881,903]" captionTargetBox="[148,1425,198,836]" captionTargetId="figure-497@9.[145,1428,195,842]" captionTargetPageId="9" captionText="Figure 5. Bayesian phylogeny, with species divergence age estimates reconstructed with BEAST using all the 26 mitochondrial genomes generated in this study. The dataset was supplemented with Thyropygus sp. and Abacion magnum as outgroups, derived from GenBank. GenBank accession numbers are provided in parentheses. Blue bars indicate the 95% highest probability density intervals for node ages. Age estimation for lineage divergence was based on a general arthropod mitochondrial DNA substitution rate and should be considered with caution. *Thyropygus sp. (red font) is very likely to be a misidentification; for more information, see the Discussion." figureDoi="http://doi.org/10.5281/zenodo.7184598" httpUri="https://zenodo.org/record/7184598/files/figure.png" pageId="10" pageNumber="934">Fig. 5</figureCitation>
), several major events might have played a role. Around 30 Mya the Antarctic ice sheet started to form (
<bibRefCitation id="DB184B5BFFF19120FCFF3A8012427E8C" author="Couvreur TLP &amp; Chatrou LW &amp; Sosef MSM &amp; Richardson JE" box="[835,1075,442,464]" pageId="10" pageNumber="934" pagination="54" refId="ref9512" refString="Couvreur TLP, Chatrou LW, Sosef MSM, Richardson JE. 2008. Molecular phylogenetics reveal multiple tertiary vicariance origins of the African rain forest trees. BMC Biology 6: 54." type="journal article" year="2008">
Couvreur
<emphasis id="8DFDEAB8FFF19120FC0A3A80159E7E93" box="[950,1007,442,464]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2008
</bibRefCitation>
), along with rifting that started to occur in northern East Africa (
<bibRefCitation id="DB184B5BFFF19120FB0B3AE213407EB2" author="Ring U" box="[1207,1329,473,495]" pageId="10" pageNumber="934" pagination="132 - 146" refId="ref11291" refString="Ring U. 2014. The East African rift system. Austrian Journal of Earth Sciences 107: 132 - 146." type="journal article" year="2014">Ring, 2014</bibRefCitation>
), possibly initiating the fragmentation of the pan-African forest. Through millions of years, the rifting would continue southwards (
<bibRefCitation id="DB184B5BFFF19120FC69390E12257D16" author="Ring U" box="[981,1108,565,587]" pageId="10" pageNumber="934" pagination="132 - 146" refId="ref11291" refString="Ring U. 2014. The East African rift system. Austrian Journal of Earth Sciences 107: 132 - 146." type="journal article" year="2014">Ring, 2014</bibRefCitation>
), affecting the topology and possibly related to the forest fragmentation responsible for the divergence of
<taxonomicName id="78894D29FFF19120FBF03948131B7DD4" baseAuthorityName="Attems" baseAuthorityYear="1950" box="[1100,1386,627,648]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="austerus">
<emphasis id="8DFDEAB8FFF19120FBF03948131B7DD4" box="[1100,1386,627,648]" italics="true" pageId="10" pageNumber="934">Tropostreptus austerus</emphasis>
</taxonomicName>
and
<taxonomicName id="78894D29FFF19120FC8739A9125F7DFB" box="[827,1070,658,679]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="severus">
<emphasis id="8DFDEAB8FFF19120FC8739A9125F7DFB" box="[827,1070,658,679]" italics="true" pageId="10" pageNumber="934">Tropostreptus severus</emphasis>
</taxonomicName>
observed ~22 Mya. The observed divergence of
<taxonomicName id="78894D29FFF19120FC59398A128C7D99" box="[997,1277,688,710]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="hamatus">
<emphasis id="8DFDEAB8FFF19120FC59398A128C7D99" box="[997,1277,688,710]" italics="true" pageId="10" pageNumber="934">Tropostreptus hamatus</emphasis>
</taxonomicName>
and the split between
<taxonomicName id="78894D29FFF19120FC1439F412CC7DB8" baseAuthorityName="Attems" baseAuthorityYear="1950" box="[936,1213,719,740]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="austerus">
<emphasis id="8DFDEAB8FFF19120FC1439F412CC7DB8" box="[936,1213,719,740]" italics="true" pageId="10" pageNumber="934">Tropostreptus austerus</emphasis>
</taxonomicName>
and
<taxonomicName id="78894D29FFF19120FB4239F415FC7C5F" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="severus">
<emphasis id="8DFDEAB8FFF19120FB4239F415FC7C5F" italics="true" pageId="10" pageNumber="934">Tropostreptus severus</emphasis>
</taxonomicName>
correspond well to the closing of the Tethys Sea (17 Mya), which would have altered ocean currents and, probably, the climate of the area (
<bibRefCitation id="DB184B5BFFF19120FB17381013E57C1C" author="Couvreur TLP &amp; Chatrou LW &amp; Sosef MSM &amp; Richardson JE" box="[1195,1428,810,832]" pageId="10" pageNumber="934" pagination="54" refId="ref9512" refString="Couvreur TLP, Chatrou LW, Sosef MSM, Richardson JE. 2008. Molecular phylogenetics reveal multiple tertiary vicariance origins of the African rain forest trees. BMC Biology 6: 54." type="journal article" year="2008">
Couvreur
<emphasis id="8DFDEAB8FFF19120FAA1381013237C63" box="[1309,1362,810,832]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2008
</bibRefCitation>
). Likewise, the isolation of the
<taxonomicName id="78894D29FFF19120FB1D387113D27C02" box="[1185,1443,841,863]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="kipunji">
<emphasis id="8DFDEAB8FFF19120FB1D387113D27C02" box="[1185,1443,841,863]" italics="true" pageId="10" pageNumber="934">Tropostreptus kipunji</emphasis>
</taxonomicName>
lineage corresponds with the uplifting of Mount Rungwe from ~8 Mya (
<bibRefCitation id="DB184B5BFFF19120FC6038BD12247CC0" author="Ring U" box="[988,1109,902,924]" pageId="10" pageNumber="934" pagination="132 - 146" refId="ref11291" refString="Ring U. 2014. The East African rift system. Austrian Journal of Earth Sciences 107: 132 - 146." type="journal article" year="2014">Ring, 2014</bibRefCitation>
). Finally, between 5 Mya and today, we observe a radiation in
<taxonomicName id="78894D29FFF19120FB23389D13D37CE6" box="[1183,1442,933,955]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="hamatus">
<emphasis id="8DFDEAB8FFF19120FB23389D13D37CE6" box="[1183,1442,933,955]" italics="true" pageId="10" pageNumber="934">Tropostreptus hamatus</emphasis>
</taxonomicName>
and
<taxonomicName id="78894D29FFF19120FCD038FE12DE7C85" box="[876,1199,964,986]" class="Diplopoda" family="Spirostreptidae" genus="Tropostreptus" kingdom="Animalia" order="Spirostreptida" pageId="10" pageNumber="934" phylum="Arthropoda" rank="species" species="sigmatospinus">
<emphasis id="8DFDEAB8FFF19120FCD038FE12DE7C85" box="[876,1199,964,986]" italics="true" pageId="10" pageNumber="934">Tropostreptus sigmatospinus</emphasis>
</taxonomicName>
(
<figureCitation id="27B22A2FFFF19120FB0238FF128C7C86" box="[1214,1277,964,986]" captionStart="Figure 5" captionStartId="9.[146,228,881,903]" captionTargetBox="[148,1425,198,836]" captionTargetId="figure-497@9.[145,1428,195,842]" captionTargetPageId="9" captionText="Figure 5. Bayesian phylogeny, with species divergence age estimates reconstructed with BEAST using all the 26 mitochondrial genomes generated in this study. The dataset was supplemented with Thyropygus sp. and Abacion magnum as outgroups, derived from GenBank. GenBank accession numbers are provided in parentheses. Blue bars indicate the 95% highest probability density intervals for node ages. Age estimation for lineage divergence was based on a general arthropod mitochondrial DNA substitution rate and should be considered with caution. *Thyropygus sp. (red font) is very likely to be a misidentification; for more information, see the Discussion." figureDoi="http://doi.org/10.5281/zenodo.7184598" httpUri="https://zenodo.org/record/7184598/files/figure.png" pageId="10" pageNumber="934">Fig. 5</figureCitation>
). A reasonable explanation for this is the uplift of the Eastern Arc Mountains, shifting the precipitation from the lowlands to the mountains (
<bibRefCitation id="DB184B5BFFF19120FBA53F1B12CC7B69" author="Lovett JC" box="[1049,1213,1056,1077]" pageId="10" pageNumber="934" pagination="23 - 29" refId="ref10593" refString="Lovett JC. 1993 a. Climatic history and forest distribution in eastern Africa. In: JC Lovett, SK Wasser, eds. Biogeography and ecology of the rain forests of Eastern Africa. Cambridge: Cambridge University Press, 23 - 29." type="book chapter" year="1993">Lovett, 1993a</bibRefCitation>
, b), in combination with the Antarctic ice sheet forming, thus decreasing global humidity (
<bibRefCitation id="DB184B5BFFF19120FBBD3F6612A47B2E" author="Polyak L &amp; Alley RB &amp; Andrews JT &amp; Brigham-Grette J &amp; Cronin TM &amp; Darby DA &amp; Dyke AS &amp; Fitzpatrick JJ &amp; Funder S &amp; Holland M &amp; Jennings AE &amp; Wolff E" box="[1025,1237,1117,1139]" pageId="10" pageNumber="934" pagination="1757 - 1778" refId="ref11202" refString="Polyak L, Alley RB, Andrews JT, Brigham-Grette J, Cronin TM, Darby DA, Dyke AS, Fitzpatrick JJ, Funder S, Holland M, Jennings AE, Wolff E. 2010. History of sea ice in the Arctic. Quaternary Science Reviews 29: 1757 - 1778." type="journal article" year="2010">
Polyak
<emphasis id="8DFDEAB8FFF19120FBEB3F6512E17B2E" box="[1111,1168,1117,1139]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2010
</bibRefCitation>
). This would lead to the emergence of savannah in the lowlands between the mountains (
<bibRefCitation id="DB184B5BFFF19120FC493FA112807BEC" author="Sepulchre P &amp; Ramstein G &amp; Fluteau F &amp; Schuster M &amp; Tiercelin J-J &amp; Brunet M" box="[1013,1265,1178,1200]" pageId="10" pageNumber="934" pagination="1419 - 1423" refId="ref11473" refString="Sepulchre P, Ramstein G, Fluteau F, Schuster M, Tiercelin J-J, Brunet M. 2006. Tectonic uplift and Eastern Africa aridification. Science 313: 1419 - 1423." type="journal article" year="2006">
Sepulchre
<emphasis id="8DFDEAB8FFF19120FBCD3FA012DA7BF3" box="[1137,1195,1178,1200]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2006
</bibRefCitation>
;
<bibRefCitation id="DB184B5BFFF19120FB433FA115057B92" author="Segalen L &amp; Lee-Thorp JA &amp; Cerling T" pageId="10" pageNumber="934" pagination="549 - 559" refId="ref11442" refString="Segalen L, Lee-Thorp JA, Cerling T. 2007. Timing of C 4 grass expansion across sub-Saharan Africa. Journal of Human Evolution 53: 549 - 559." type="journal article" year="2007">
Ségalen
<emphasis id="8DFDEAB8FFF19120FADF3FA013EC7BF3" box="[1379,1437,1178,1200]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2007
</bibRefCitation>
;
<bibRefCitation id="DB184B5BFFF19120FC3F3F8212F37B92" author="Couvreur TLP &amp; Chatrou LW &amp; Sosef MSM &amp; Richardson JE" box="[899,1154,1209,1231]" pageId="10" pageNumber="934" pagination="54" refId="ref9512" refString="Couvreur TLP, Chatrou LW, Sosef MSM, Richardson JE. 2008. Molecular phylogenetics reveal multiple tertiary vicariance origins of the African rain forest trees. BMC Biology 6: 54." type="journal article" year="2008">
Couvreur
<emphasis id="8DFDEAB8FFF19120FC413F81124B7B92" box="[1021,1082,1209,1231]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2008
</bibRefCitation>
), isolating the montane forest and limiting migration between populations of forest-restricted species.
</paragraph>
<paragraph id="BF3636AAFFF19121FCEF3E2D14557E44" blockId="10.[824,1444,197,1906]" lastBlockId="11.[144,760,197,280]" lastPageId="11" lastPageNumber="935" pageId="10" pageNumber="934">
We emphasize that we have neither good fossil records nor mtDNA mutation rates estimated specifically for millipedes, which is why the split times of our millipede phylogenetic tree should be interpreted with caution. Moreover, comparable studies with dated phylogenies of Eastern Arc species are sparse, hence it is difficult to compare the split times we have estimated with those of other species in the region. Examining two separate studies of chameleons (
<taxonomicName id="78894D29FFF19120FC723D1113D27963" authority="Tilbury, Tolley &amp; Branch, 2006" authorityName="Tilbury, Tolley &amp; Branch" authorityYear="2006" box="[974,1443,1577,1599]" class="Reptilia" family="Chamaeleonidae" genus="Kinyongia" kingdom="Animalia" order="Squamata" pageId="10" pageNumber="934" phylum="Chordata" rank="genus">
<emphasis id="8DFDEAB8FFF19120FC723D1112327963" box="[974,1091,1578,1599]" italics="true" pageId="10" pageNumber="934">Kinyongia</emphasis>
Tilbury, Tolley &amp; Branch, 2006
</taxonomicName>
and
<taxonomicName id="78894D29FFF19120FCD03D7312FD7901" authority="Swainson, 1839" authorityName="Swainson" authorityYear="1839" box="[876,1164,1608,1630]" class="Reptilia" family="Chamaeleonidae" genus="Trioceros" kingdom="Animalia" order="Squamata" pageId="10" pageNumber="934" phylum="Chordata" rank="genus">
<emphasis id="8DFDEAB8FFF19120FCD03D7315A27901" box="[876,979,1608,1629]" italics="true" pageId="10" pageNumber="934">Trioceros</emphasis>
Swainson, 1839
</taxonomicName>
) with dated phylogenies based on both mitochondrial and nuclear markers did show some correspondence with our dated splits (
<bibRefCitation id="DB184B5BFFF19120FCFF3D9F127A79E5" author="Tolley KA &amp; Tilbury CR &amp; Measey GJ &amp; Menegon M &amp; Branch WR &amp; Matthee CA" box="[835,1035,1700,1722]" pageId="10" pageNumber="934" pagination="1748 - 1760" refId="ref11779" refString="Tolley KA, Tilbury CR, Measey GJ, Menegon M, Branch WR, Matthee CA. 2011. Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hotspot. Journal of Biogeography 3 8: 1748 - 1760." type="journal article" year="2011">
Tolley
<emphasis id="8DFDEAB8FFF19120FC333D9E15B679E5" box="[911,967,1700,1722]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2011
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;
<bibRefCitation id="DB184B5BFFF19120FBAB3D9F137A79E5" author="Ceccarelli FS &amp; Menegon M &amp; Tolley KA &amp; Tilbury CR &amp; Gower DJ &amp; Laserna MH &amp; Kasahun R &amp; Prieto A &amp; Hagmann R &amp; Loader SP" box="[1047,1291,1700,1722]" pageId="10" pageNumber="934" pagination="125 - 136" refId="ref9424" refString="Ceccarelli FS, Menegon M, Tolley KA, Tilbury CR, Gower DJ, Laserna MH, Kasahun R, Rodriguez- Prieto A, Hagmann R, Loader SP. 2014. Evolutionary relationships, species delimitation and biogeography of Eastern Afromontane horned chameleons (Chamaeleonidae: Trioceros). Molecular Phylogenetics and Evolution 80: 125 - 136." type="journal article" year="2014">
Ceccarelli
<emphasis id="8DFDEAB8FFF19120FB333D9E12B679E5" box="[1167,1223,1700,1722]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
, 2014
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).
<bibRefCitation id="DB184B5BFFF19120FAA33D9F15F57984" author="Tolley KA &amp; Tilbury CR &amp; Measey GJ &amp; Menegon M &amp; Branch WR &amp; Matthee CA" pageId="10" pageNumber="934" pagination="1748 - 1760" refId="ref11779" refString="Tolley KA, Tilbury CR, Measey GJ, Menegon M, Branch WR, Matthee CA. 2011. Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hotspot. Journal of Biogeography 3 8: 1748 - 1760." type="journal article" year="2011">
Tolley
<emphasis id="8DFDEAB8FFF19120FAD73D9E13D279E5" box="[1387,1443,1700,1722]" italics="true" pageId="10" pageNumber="934">et al.</emphasis>
(2011)
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dated the earliest split between the northern and southern Eastern Arc species to ~28 Mya, and both studies show several radiation events between 5 and 20 Mya, corresponding to the same overall time frame that we are discussing for the millipedes. In contrast, the chameleons display fewer speciation events during the last 5 Myr than the millipedes, perhaps suggesting that the latter have been more susceptible to vicariance during more recent climatic events.
</paragraph>
</subSubSection>
</treatment>
</document>
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