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<mods:title id="E794B3A0803DC2073576064402CC450E">Is the grass greener on the other side? Phylogeny and biogeography of grass-feeding plant bugs of the genus Collaria Provancher (Hemiptera: Miridae: Mirinae)</mods:title>
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<mods:namePart id="9C61E5079CF50143707B0D6EBEE334A0">Forero, Dimitri</mods:namePart>
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<mods:date id="9147A422857CCFAF23B5A93F53984B57">2025</mods:date>
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<heading id="D0CD8199FFB99A161B87F9AD0CB1F9EB" bold="true" box="[160,721,1610,1636]" centered="true" fontSize="11" level="2" pageId="19" pageNumber="530" reason="3">
<emphasis id="B94EEAE7FFB99A161B87F9AD0CB1F9EB" bold="true" box="[160,721,1610,1636]" pageId="19" pageNumber="530">
<taxonomicName id="4C3A4D76FFB99A161B87F9AD0F60F9EB" ID-CoL="9G339" box="[160,256,1610,1636]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="530" phylum="Arthropoda" rank="genus">Collaria</taxonomicName>
<emphasis id="B94EEAE7FFB99A161A2FF9AD0C09F9EB" bold="true" box="[264,617,1610,1636]" italics="true" pageId="19" pageNumber="530">group and the monophyly of</emphasis>
<taxonomicName id="4C3A4D76FFB99A161956F9AD0CB1F9EB" box="[625,721,1610,1636]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="530" phylum="Arthropoda" rank="genus">Collaria</taxonomicName>
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<paragraph id="8B8536F5FFB99A101B87F9990FCCFDC8" blockId="19.[160,1156,1662,1722]" lastBlockId="21.[160,1157,142,965]" lastPageId="21" lastPageNumber="532" pageId="19" pageNumber="530">
<bibRefCitation id="EFAB4B04FFB99A161B87F9990F01F918" author="Schwartz MD" box="[160,353,1662,1687]" pageId="19" pageNumber="530" pagination="1111 - 1201" refId="ref17986" refString="Schwartz MD. 2008. Revision of the Stenodemini with a review of the included genera (Hemiptera: Heteroptera: Miridae: Mirinae). Proc Entomol Soc Wash. 110 (4): 1111-1201. doi:10.4289/0013- 8797 - 110.4.1111." type="journal article" year="2008">Schwartz (2008)</bibRefCitation>
, in his phylogenetic hypotheses for the tribe
<taxonomicName id="4C3A4D76FFB99A16188CF9990A22F918" box="[939,1090,1662,1687]" pageId="19" pageNumber="530" rank="tribe" tribe="Stenodemini">Stenodemini</taxonomicName>
, proposed the
<taxonomicName id="4C3A4D76FFB99A161A00F9460F1EF935" box="[295,382,1697,1722]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="530" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFB99A161A00F9460F1EF935" box="[295,382,1697,1722]" italics="true" pageId="19" pageNumber="530">Collaria</emphasis>
</taxonomicName>
group as composed of
<taxonomicName id="4C3A4D76FFB99A16198CF9460D62F935" box="[683,770,1697,1722]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="530" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFB99A16198CF9460D62F935" box="[683,770,1697,1722]" italics="true" pageId="19" pageNumber="530">Collaria</emphasis>
</taxonomicName>
and
<taxonomicName id="4C3A4D76FFB99A161860F9460DAFF935" authorityName="Poppius" authorityYear="1914" box="[839,975,1697,1722]" class="Insecta" family="Miridae" genus="Nabidomiris" kingdom="Animalia" order="Hemiptera" pageId="19" pageNumber="530" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFB99A161860F9460DAFF935" box="[839,975,1697,1722]" italics="true" pageId="19" pageNumber="530">Nabidomiris</emphasis>
</taxonomicName>
, based on the following non-homoplastic characters: eye ovoid and located in the medial portion of the head; male genitalia with dorsal and ventral ornamentation on the secondary gonopore; membrane of the vesica with a small spinose or filamentous sclerite basal to the secondary gonopore; and female genitalia with the dorsal surface of the posterior wall with a large process. In addition, two homoplastic characters also supported the
<taxonomicName id="4C3A4D76FFBF9A101AB3FF560F8BFF45" box="[404,491,177,202]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A101AB3FF560F8BFF45" box="[404,491,177,202]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
</taxonomicName>
group: the rounded humeral angle of the pronotum, and the proepisternum strongly rounded. In our study, we recovered a sister-group relationship between
<taxonomicName id="4C3A4D76FFBF9A101A8AFF110C64FE80" box="[429,516,246,271]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A101A8AFF110C64FE80" box="[429,516,246,271]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
</taxonomicName>
+
<taxonomicName id="4C3A4D76FFBF9A101917FF110CD8FE80" authorityName="Poppius" authorityYear="1914" box="[560,696,246,271]" class="Insecta" family="Miridae" genus="Nabidomiris" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A101917FF110CD8FE80" box="[560,696,246,271]" italics="true" pageId="21" pageNumber="532">Nabidomiris</emphasis>
</taxonomicName>
supported by two of the characters originally proposed by
<bibRefCitation id="EFAB4B04FFBF9A101A99FEFE0C1DFEBD" author="Schwartz MD" box="[446,637,281,306]" pageId="21" pageNumber="532" pagination="1111 - 1201" refId="ref17986" refString="Schwartz MD. 2008. Revision of the Stenodemini with a review of the included genera (Hemiptera: Heteroptera: Miridae: Mirinae). Proc Entomol Soc Wash. 110 (4): 1111-1201. doi:10.4289/0013- 8797 - 110.4.1111." type="journal article" year="2008">Schwartz (2008)</bibRefCitation>
: the postocular region elongated and the basal region of the endosoma with a ribbon-like sclerite. The eye located on the medial portion of the head is used by
<bibRefCitation id="EFAB4B04FFBF9A1019B6FEB90D30FEF8" author="Schwartz MD" box="[657,848,350,375]" pageId="21" pageNumber="532" pagination="1111 - 1201" refId="ref17986" refString="Schwartz MD. 2008. Revision of the Stenodemini with a review of the included genera (Hemiptera: Heteroptera: Miridae: Mirinae). Proc Entomol Soc Wash. 110 (4): 1111-1201. doi:10.4289/0013- 8797 - 110.4.1111." type="journal article" year="2008">Schwartz (2008)</bibRefCitation>
as one non-homoplastic character to support the
<taxonomicName id="4C3A4D76FFBF9A101AC7FE660C57FE15" box="[480,567,385,410]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A101AC7FE660C57FE15" box="[480,567,385,410]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
</taxonomicName>
group clade; however, the eye position corresponds to the relative anteocular and postocular lengths, and not to the eye position in the head
<emphasis id="B94EEAE7FFBF9A101A8BFE210F95FE50" box="[428,501,454,479]" italics="true" pageId="21" pageNumber="532">per se</emphasis>
. Therefore, we coded the anteocular and postocular structure as separate characters. Additionally, we recovered in our analysis that the dorsal surface of the pronotum clearly divided into lobes supports the clade
<taxonomicName id="4C3A4D76FFBF9A101B86FDC90E98FDC8" box="[161,248,558,583]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A101B86FDC90E98FDC8" box="[161,248,558,583]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
</taxonomicName>
+
<taxonomicName id="4C3A4D76FFBF9A101A38FDC90FC7FDC8" authorityName="Poppius" authorityYear="1914" box="[287,423,558,583]" class="Insecta" family="Miridae" genus="Nabidomiris" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A101A38FDC90FC7FDC8" box="[287,423,558,583]" italics="true" pageId="21" pageNumber="532">Nabidomiris</emphasis>
</taxonomicName>
.
</paragraph>
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<paragraph id="8B8536F5FFBE9A111B87FAAB0F2FFA79" blockId="20.[160,1157,1356,1526]" pageId="20" pageNumber="531">
<emphasis id="B94EEAE7FFBE9A111B87FAAB0E9AFAEC" bold="true" box="[160,250,1356,1379]" pageId="20" pageNumber="531">Figure 9.</emphasis>
Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 815. Symbols at the nodes of the phylogeny are: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares.
</paragraph>
</caption>
<paragraph id="8B8536F5FFBF9A101BE7FDB60CF5FC4A" blockId="21.[160,1157,142,965]" pageId="21" pageNumber="532">
Schwartzs hypothesis (2008) used genera as terminals, assuming
<emphasis id="B94EEAE7FFBF9A101881FDB60D95FDE5" box="[934,1013,593,618]" italics="true" pageId="21" pageNumber="532">a priori</emphasis>
their monophyly, and thus disregarding species in the analyses. Nine species of
<taxonomicName id="4C3A4D76FFBF9A1018CDFD930A20FD02" box="[1002,1088,628,653]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A1018CDFD930A20FD02" box="[1002,1088,628,653]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
</taxonomicName>
were analysed by
<bibRefCitation id="EFAB4B04FFBF9A101A14FD710F8AFD20" author="Schwartz MD" box="[307,490,662,687]" pageId="21" pageNumber="532" pagination="1111 - 1201" refId="ref17986" refString="Schwartz MD. 2008. Revision of the Stenodemini with a review of the included genera (Hemiptera: Heteroptera: Miridae: Mirinae). Proc Entomol Soc Wash. 110 (4): 1111-1201. doi:10.4289/0013- 8797 - 110.4.1111." type="journal article" year="2008">Schwartz (2008)</bibRefCitation>
:
<emphasis id="B94EEAE7FFBF9A101ADFFD710F8BFD5D" italics="true" pageId="21" pageNumber="532">
<taxonomicName id="4C3A4D76FFBF9A101ADFFD710CF7FD20" authorityName="Carvalho &amp; Fontes" authorityYear="1981" box="[504,663,662,687]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="guaraniana">C. guaraniana</taxonomicName>
,
<taxonomicName id="4C3A4D76FFBF9A101981FD710D51FD20" authorityName="Reuter" authorityYear="1893" box="[678,817,662,687]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="improvisa">C. improvisa</taxonomicName>
,
<taxonomicName id="4C3A4D76FFBF9A101818FD710DA2FD20" authorityName="Provancher" authorityYear="1872" box="[831,962,662,687]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="meilleurii">C. meilleurii</taxonomicName>
,
<taxonomicName id="4C3A4D76FFBF9A1018F6FD710A1FFD20" authorityName="Poppius" authorityYear="1910" box="[977,1151,662,687]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="obscuricornis">C. obscuricornis</taxonomicName>
,
<taxonomicName id="4C3A4D76FFBF9A101B87FD5E0F6DFD5D" baseAuthorityName="Reuter" baseAuthorityYear="1876" box="[160,269,697,722]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="oculata">C. oculata</taxonomicName>
,
<taxonomicName id="4C3A4D76FFBF9A101A30FD5E0F19FD5D" baseAuthorityName="Distant" baseAuthorityYear="1883" box="[279,377,697,722]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="oleosa">C. oleosa</taxonomicName>
,
<taxonomicName id="4C3A4D76FFBF9A101AA5FD5E0F8BFD5D" baseAuthorityName="Stal" baseAuthorityYear="1859" box="[386,491,697,722]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="scenica">C. scenica</taxonomicName>
</emphasis>
and two undescribed species, all coded as a single terminal,
<taxonomicName id="4C3A4D76FFBF9A101B87FD3B0E93FD7A" box="[160,243,732,757]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A101B87FD3B0E93FD7A" box="[160,243,732,757]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
</taxonomicName>
. Considering the diagnosis of
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<emphasis id="B94EEAE7FFBF9A10197FFD3B0CCBFD7A" box="[600,683,732,757]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
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offered by
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, our analysis agrees with one synapomorphy: the mandibular plates produced. Additionally, the generic diagnosis of
<taxonomicName id="4C3A4D76FFBF9A101A41FCC60FD9FCB5" box="[358,441,801,826]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A101A41FCC60FD9FCB5" box="[358,441,801,826]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
</taxonomicName>
did not consider the homoplastic characters recovered in his analyses: the setae on the first and second antennal segments long and erect (character 141;
<bibRefCitation id="EFAB4B04FFBF9A101BC3FC810FE8FCF0" author="Schwartz MD" box="[228,392,870,895]" pageId="21" pageNumber="532" pagination="1111 - 1201" refId="ref17986" refString="Schwartz MD. 2008. Revision of the Stenodemini with a review of the included genera (Hemiptera: Heteroptera: Miridae: Mirinae). Proc Entomol Soc Wash. 110 (4): 1111-1201. doi:10.4289/0013- 8797 - 110.4.1111." type="journal article" year="2008">Schwartz 2008</bibRefCitation>
), and the metatibia with long erect setae (character 181;
<bibRefCitation id="EFAB4B04FFBF9A101F39FC810EB8FC2D" author="Schwartz MD" pageId="21" pageNumber="532" pagination="1111 - 1201" refId="ref17986" refString="Schwartz MD. 2008. Revision of the Stenodemini with a review of the included genera (Hemiptera: Heteroptera: Miridae: Mirinae). Proc Entomol Soc Wash. 110 (4): 1111-1201. doi:10.4289/0013- 8797 - 110.4.1111." type="journal article" year="2008">Schwartz 2008</bibRefCitation>
). Having erect setae on the first antennal segment was recovered as a synapomorphy for
<taxonomicName id="4C3A4D76FFBF9A101AACFC4B0FBEFC4A" box="[395,478,940,965]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A101AACFC4B0FBEFC4A" box="[395,478,940,965]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
</taxonomicName>
in our analyses.
</paragraph>
<paragraph id="8B8536F5FFBF9A101B87FBFE0F82FBBC" blockId="21.[160,482,1049,1075]" box="[160,482,1049,1075]" pageId="21" pageNumber="532">
<heading id="D0CD8199FFBF9A101B87FBFE0F82FBBC" bold="true" box="[160,482,1049,1075]" centered="true" fontSize="11" level="2" pageId="21" pageNumber="532" reason="3">
<emphasis id="B94EEAE7FFBF9A101B87FBFE0F82FBBC" bold="true" box="[160,482,1049,1075]" pageId="21" pageNumber="532">
<taxonomicName id="4C3A4D76FFBF9A101B87FBFE0F3FFBBC" box="[160,351,1049,1075]" pageId="21" pageNumber="532">
Collaria
<emphasis id="B94EEAE7FFBF9A101A2FFBFE0F3FFBBC" bold="true" box="[264,351,1049,1075]" italics="true" pageId="21" pageNumber="532">species</emphasis>
</taxonomicName>
<emphasis id="B94EEAE7FFBF9A101A41FBFE0F82FBBC" bold="true" box="[358,482,1049,1075]" italics="true" pageId="21" pageNumber="532">groupings</emphasis>
</emphasis>
</heading>
</paragraph>
<paragraph id="8B8536F5FFBF9A101B87FBA90FB2FA10" blockId="21.[160,1156,1102,1717]" pageId="21" pageNumber="532">
Missing data had an important effect on the species grouping hypothesis within
<taxonomicName id="4C3A4D76FFBF9A101F0DFBA90A1DFBE8" box="[1066,1149,1102,1127]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A101F0DFBA90A1DFBE8" box="[1066,1149,1102,1127]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
</taxonomicName>
. In our analysis of the complete matrix, the recently described species
<taxonomicName id="4C3A4D76FFBF9A101892FB960A4BFB05" authorityName="Matocq" authorityYear="2022" box="[949,1067,1137,1162]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="boulardi">
<emphasis id="B94EEAE7FFBF9A101892FB960A4BFB05" box="[949,1067,1137,1162]" italics="true" pageId="21" pageNumber="532">C. boulardi</emphasis>
</taxonomicName>
(
<collectingCountry id="F32D7665FFBF9A101F1EFB960A1AFB05" box="[1081,1146,1137,1162]" name="Brazil" pageId="21" pageNumber="532">Brazil</collectingCountry>
),
<taxonomicName id="4C3A4D76FFBF9A101B87FB730E92FB22" authorityName="Matocq" authorityYear="2021" box="[160,242,1172,1197]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="elsae">
<emphasis id="B94EEAE7FFBF9A101B87FB730E92FB22" box="[160,242,1172,1197]" italics="true" pageId="21" pageNumber="532">C. elsae</emphasis>
</taxonomicName>
(
<collectingCountry id="F32D7665FFBF9A101A27FB730FEDFB22" box="[256,397,1172,1197]" name="Madagascar" pageId="21" pageNumber="532">Madagascar</collectingCountry>
), and
<taxonomicName id="4C3A4D76FFBF9A101AE8FB730C59FB22" authorityName="Matocq" authorityYear="2022" box="[463,569,1172,1197]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="minuta">
<emphasis id="B94EEAE7FFBF9A101AE8FB730C59FB22" box="[463,569,1172,1197]" italics="true" pageId="21" pageNumber="532">C. minuta</emphasis>
</taxonomicName>
(
<collectingCountry id="F32D7665FFBF9A101960FB730C8EFB22" box="[583,750,1172,1197]" name="French Guiana" pageId="21" pageNumber="532">French Guiana</collectingCountry>
) were recovered as sister species to the remaining species of the genus, whereas
<taxonomicName id="4C3A4D76FFBF9A1019BFFB510C97FB40" authorityName="Linnavuori" authorityYear="1974" box="[664,759,1206,1231]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="danae">
<emphasis id="B94EEAE7FFBF9A1019BFFB510C97FB40" box="[664,759,1206,1231]" italics="true" pageId="21" pageNumber="532">C. danae</emphasis>
</taxonomicName>
(
<collectingCountry id="F32D7665FFBF9A101823FB510DE3FB40" box="[772,899,1206,1231]" name="Ivory Coast" pageId="21" pageNumber="532">Ivory Coast</collectingCountry>
) was recovered as part of the otherwise Neotropical species clade. The phylogenetic position of
<emphasis id="B94EEAE7FFBF9A101F24FB3E0F04FA9A" italics="true" pageId="21" pageNumber="532">
<taxonomicName id="4C3A4D76FFBF9A101F24FB3E0A1FFB7D" authorityName="Matocq" authorityYear="2022" box="[1027,1151,1241,1266]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="boulardi">C. boulardi</taxonomicName>
,
<taxonomicName id="4C3A4D76FFBF9A101B87FB1B0F63FA9A" authorityName="Linnavuori" authorityYear="1974" box="[160,259,1276,1301]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="danae">C. danae</taxonomicName>
,
<taxonomicName id="4C3A4D76FFBF9A101A37FB1B0F04FA9A" authorityName="Matocq" authorityYear="2021" box="[272,356,1276,1301]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="elsae">C. elsae</taxonomicName>
</emphasis>
, and
<taxonomicName id="4C3A4D76FFBF9A101A81FB1B0C72FA9A" authorityName="Matocq" authorityYear="2022" box="[422,530,1276,1301]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="minuta">
<emphasis id="B94EEAE7FFBF9A101A81FB1B0C72FA9A" box="[422,530,1276,1301]" italics="true" pageId="21" pageNumber="532">C. minuta</emphasis>
</taxonomicName>
derived from the complete matrix is doubtful. We did not examine specimens of these species, and because they are known only from females, the percentage of missing data for these species in our complete matrix surpasses 60%. Additional material for these species could contribute to more robust phylogenetic and biogeographic hypotheses.
</paragraph>
<paragraph id="8B8536F5FFBF9A101BE7FA4E0FA2F9A5" blockId="21.[160,1156,1102,1717]" pageId="21" pageNumber="532">
In our analyses with the reduced matrix,
<taxonomicName id="4C3A4D76FFBF9A1019ADFA4E0CBDFA4D" box="[650,733,1449,1474]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A1019ADFA4E0CBDFA4D" box="[650,733,1449,1474]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
</taxonomicName>
is composed of two main clades that correspond to the biogeographic distribution of the species: an Afrotropical species clade and a clade of Nearctic species +
<taxonomicName id="4C3A4D76FFBF9A101932FA090C38F988" box="[533,600,1518,1543]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="royi">
<emphasis id="B94EEAE7FFBF9A101932FA090C38F988" box="[533,600,1518,1543]" italics="true" pageId="21" pageNumber="532">C. royi</emphasis>
</taxonomicName>
with a Neotropical species clade (
<figureCitation id="13012A70FFBF9A1018F1FA090A51F988" box="[982,1073,1518,1543]" captionStart="Figure 7" captionStartId="17.[160,225,1207,1230]" captionTargetBox="[164,1152,144,1174]" captionTargetId="figure-110@17.[163,1153,142,1175]" captionTargetPageId="17" captionText="Figure 7. Single most parsimonious tree obtained with the implied weighting (IW) analysis and using k = 30. Main apomorphies mapped on the tree, non-homoplasious changes as black polygons, homoplasious changes as white polygons. Support values in the branches are within square brackets (Bremer support, absolute symmetric resampling and “Group present/Contradicted” (GC) frequencies, respectively). Clades A to F. Geographic distribution: Nearctic region (blue), African region (marine blue), Neotropical region (green)." figureDoi="http://doi.org/10.5281/zenodo.14970750" httpUri="https://zenodo.org/record/14970750/files/figure.png" pageId="21" pageNumber="532">Figure 7</figureCitation>
, clades C and E + F, respectively).
</paragraph>
<paragraph id="8B8536F5FFBF9A131BE7F9D30A00FF45" blockId="21.[160,1156,1102,1717]" lastBlockId="22.[160,1156,142,445]" lastPageId="22" lastPageNumber="533" pageId="21" pageNumber="532">
The Afrotropical clade (clade E) is supported by genital characters, as well as most of the species groupings. Similarly, the Neotropical species clade is supported by genitalic characters, but the groupings within the clade are weakly supported, except for the clade formed by
<taxonomicName id="4C3A4D76FFBF9A101A3CF97B0FE6F93A" baseAuthorityName="Stal" baseAuthorityYear="1859" box="[283,390,1692,1717]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="scenica">
<emphasis id="B94EEAE7FFBF9A101A3CF97B0FE6F93A" box="[283,390,1692,1717]" italics="true" pageId="21" pageNumber="532">C. scenica</emphasis>
</taxonomicName>
and
<taxonomicName id="4C3A4D76FFBF9A101A99F97B0C7FF93A" baseAuthorityName="Distant" baseAuthorityYear="1883" box="[446,543,1692,1717]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="species" species="oleosa">
<emphasis id="B94EEAE7FFBF9A101A99F97B0C7FF93A" box="[446,543,1692,1717]" italics="true" pageId="21" pageNumber="532">C. oleosa</emphasis>
</taxonomicName>
. The genitalia in
<taxonomicName id="4C3A4D76FFBF9A1019F8F97B0D52F93A" box="[735,818,1692,1717]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="21" pageNumber="532" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBF9A1019F8F97B0D52F93A" box="[735,818,1692,1717]" italics="true" pageId="21" pageNumber="532">Collaria</emphasis>
</taxonomicName>
are asymmetrical and include various sclerites and processes on the male endosoma and the female posterior wall. Thus, genitalic structures were informative in our analyses, supporting several clades.
</paragraph>
<paragraph id="8B8536F5FFBC9A131BE7FF330D6BFE32" blockId="22.[160,1156,142,445]" pageId="22" pageNumber="533">
The Nearctic species +
<taxonomicName id="4C3A4D76FFBC9A131AECFF330C70FF62" box="[459,528,212,237]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="species" species="royi">
<emphasis id="B94EEAE7FFBC9A131AECFF330C70FF62" box="[459,528,212,237]" italics="true" pageId="22" pageNumber="533">C. royi</emphasis>
</taxonomicName>
were recovered as the sister group of the Neotropical species, supported by two homoplastic characters.
<taxonomicName id="4C3A4D76FFBC9A1319CCFF110DCBFE80" authorityName="Provancher" authorityYear="1872" box="[747,939,246,271]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="species" species="meilleurii">
<emphasis id="B94EEAE7FFBC9A1319CCFF110DCBFE80" box="[747,939,246,271]" italics="true" pageId="22" pageNumber="533">Collaria meilleurii</emphasis>
</taxonomicName>
and
<taxonomicName id="4C3A4D76FFBC9A1318CFFF110A38FE80" baseAuthorityName="Reuter" baseAuthorityYear="1876" box="[1000,1112,246,271]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="species" species="oculata">
<emphasis id="B94EEAE7FFBC9A1318CFFF110A38FE80" box="[1000,1112,246,271]" italics="true" pageId="22" pageNumber="533">C. oculata</emphasis>
</taxonomicName>
are similar in endosomal structure in having a long sclerite and the medial left sclerite with a semicircular shape; whereas they are distinguished by the colouration patterns and the shape of the ventral right sclerite of the endosoma (
<bibRefCitation id="EFAB4B04FFBC9A1319CCFEB90DA0FEF8" author="Morales I &amp; Ferreira PSF &amp; Forero D" box="[747,960,350,375]" pageId="22" pageNumber="533" pagination="201 - 246" refId="ref17129" refString="Morales I, Ferreira PSF, Forero D. 2016. Taxonomic revision of Collaria Provancher, 1872 (Hemiptera: Miridae) with the description of a new species from the Afrotropical region. Zootaxa. 4138 (2): 201-246. doi:10.11646/zootaxa.4138.2.1." type="journal article" year="2016">
Morales
<emphasis id="B94EEAE7FFBC9A13186BFEB90D1CFEF8" box="[844,892,350,375]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2016
</bibRefCitation>
). Colour patterns are not included as evidence in our analyses, because
<taxonomicName id="4C3A4D76FFBC9A1319D9FE660DCAFE15" box="[766,938,385,410]" pageId="22" pageNumber="533">
<emphasis id="B94EEAE7FFBC9A1319D9FE660D31FE15" box="[766,849,385,410]" italics="true" pageId="22" pageNumber="533">Collaria</emphasis>
species
</taxonomicName>
are usually entirely brown or pale brown with some interspecific variation.
</paragraph>
<paragraph id="8B8536F5FFBC9A131B87FE1D0FB4FD9B" blockId="22.[160,468,506,532]" box="[160,468,506,532]" pageId="22" pageNumber="533">
<heading id="D0CD8199FFBC9A131B87FE1D0FB4FD9B" bold="true" box="[160,468,506,532]" centered="true" fontSize="11" level="2" pageId="22" pageNumber="533" reason="3">
<emphasis id="B94EEAE7FFBC9A131B87FE1D0FB4FD9B" bold="true" box="[160,468,506,532]" pageId="22" pageNumber="533">
<emphasis id="B94EEAE7FFBC9A131B87FE1D0F0CFD9B" bold="true" box="[160,364,506,532]" italics="true" pageId="22" pageNumber="533">Biogeography of</emphasis>
<taxonomicName id="4C3A4D76FFBC9A131A53FE1D0FB4FD9B" ID-CoL="9G339" box="[372,468,506,532]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="genus">Collaria</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8B8536F5FFBC9A131B87FDC80C20FCD1" blockId="22.[160,1156,559,1728]" pageId="22" pageNumber="533">
Attempts to explain species distribution require an integrated approach, because the processes behind the distribution patterns involve diverse aspects of historical biogeography, ecology, and even biology and behaviour (
<bibRefCitation id="EFAB4B04FFBC9A1319C5FD920A79FD01" author="Wiens JJ &amp; Donoghue MJ" box="[738,1049,629,654]" pageId="22" pageNumber="533" pagination="639 - 644" refId="ref18665" refString="Wiens JJ, Donoghue MJ. 2004. Historical biogeography, ecology and species richness. Trends Ecol Evol. 19: 639-644. doi:10.1016/j.tree.2004.09.011" type="journal article" year="2004">Wiens and Donoghue 2004</bibRefCitation>
). Despite several limitations hindering our understanding of the biogeographic processes that have resulted in the observed distributional patterns of
<taxonomicName id="4C3A4D76FFBC9A1319D2FD5D0DC8FD5C" box="[757,936,698,723]" pageId="22" pageNumber="533">
<emphasis id="B94EEAE7FFBC9A1319D2FD5D0D28FD5C" box="[757,840,698,723]" italics="true" pageId="22" pageNumber="533">Collaria</emphasis>
species
</taxonomicName>
, the results of our biogeographic analyses provide a set of hypotheses that could be further tested with additional data. Our results on the biogeography of
<taxonomicName id="4C3A4D76FFBC9A1319C4FD180D56FC97" box="[739,822,767,792]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBC9A1319C4FD180D56FC97" box="[739,822,767,792]" italics="true" pageId="22" pageNumber="533">Collaria</emphasis>
</taxonomicName>
show a mix of geographically explicit events among the Afrotropical, Nearctic, and Neotropical areas, including vicariance, sympatry, and founder events.
</paragraph>
<paragraph id="8B8536F5FFBC9A131BE7FC800E81FAAF" blockId="22.[160,1156,559,1728]" pageId="22" pageNumber="533">
The disjunct distribution between the Afrotropical and the Nearctic (except
<taxonomicName id="4C3A4D76FFBC9A131F38FC800A03FC0F" box="[1055,1123,871,896]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="species" species="royi">
<emphasis id="B94EEAE7FFBC9A131F38FC800A03FC0F" box="[1055,1123,871,896]" italics="true" pageId="22" pageNumber="533">C. royi</emphasis>
</taxonomicName>
) + Neotropical clades was inferred as a vicariant event in our analysis (
<figureCitation id="13012A70FFBC9A1318A9FC6D0D89FC2C" box="[910,1001,906,931]" captionStart="Figure 8" captionStartId="19.[160,225,1329,1352]" captionTargetBox="[164,1151,143,1296]" captionTargetId="figure-39@19.[163,1153,142,1298]" captionTargetPageId="19" captionText="Figure 8. Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 17. Symbols at the nodes of the phylogeny are as follows: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares." figureDoi="http://doi.org/10.5281/zenodo.14970752" httpUri="https://zenodo.org/record/14970752/files/figure.png" pageId="22" pageNumber="533">Figure 8</figureCitation>
, node 1). This pattern needs alternative explanations besides being of Gondwanan origin, considering the age of diversification for plant bugs (159.7167.5 MYA,
<bibRefCitation id="EFAB4B04FFBC9A131819FC280A78FC67" author="Johnson KP &amp; Dietrich CH &amp; Friedrich F &amp; Beutel RG &amp; Wipfler B &amp; Peters RS &amp; Allen JM &amp; Petersen M &amp; Donath A &amp; Walden KKO" box="[830,1048,975,1000]" pageId="22" pageNumber="533" pagination="12775 - 12780" refId="ref16404" refString="Johnson KP, Dietrich CH, Friedrich F, Beutel RG, Wipfler B, Peters RS, Allen JM, Petersen M, Donath A, Walden KKO, et al. 2018. Phylogenomics and the evolution of hemipteroid insects. PNAS. 115 (50): 12775-12780. doi:10.1073/pnas.1815820115." type="journal article" year="2018">
Johnson
<emphasis id="B94EEAE7FFBC9A131884FC280DB3FC67" box="[931,979,975,1000]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2018
</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFBC9A131F02FC280EB7FB84" author="Oh M &amp; Kim S &amp; Lee S" pageId="22" pageNumber="533" refId="ref17258" refString="Oh M, Kim S, Lee S. 2023. Revisiting the phylogeny of the family Miridae (Heteroptera: Cimicomorpha), with updated insights into its origin and life history evolution. Mol Phylogen Evol. 184: 107796. doi:10.1016/j.ympev.2023.107796" type="journal volume" year="2023">
Oh
<emphasis id="B94EEAE7FFBC9A131F6AFC280A1EFC67" box="[1101,1150,975,1000]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2023
</bibRefCitation>
), and of
<taxonomicName id="4C3A4D76FFBC9A131A1CFC150FAAFB84" box="[315,458,1010,1035]" pageId="22" pageNumber="530" rank="tribe" tribe="Stenodemini">Stenodemini</taxonomicName>
in particular, which ranges between 75 and 80 MYA (
<bibRefCitation id="EFAB4B04FFBC9A131F03FC150EB7FBA1" author="Oh M &amp; Kim S &amp; Lee S" pageId="22" pageNumber="533" refId="ref17258" refString="Oh M, Kim S, Lee S. 2023. Revisiting the phylogeny of the family Miridae (Heteroptera: Cimicomorpha), with updated insights into its origin and life history evolution. Mol Phylogen Evol. 184: 107796. doi:10.1016/j.ympev.2023.107796" type="journal volume" year="2023">
Oh
<emphasis id="B94EEAE7FFBC9A131F69FC150A1EFB84" box="[1102,1150,1010,1035]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2023
</bibRefCitation>
). Gondwana started breaking up during the late Jurassic, between 150 and 165 MYA, but South America and Africa were still connected until 95110 MYA, when northern South America and Africa started separating (
<bibRefCitation id="EFAB4B04FFBC9A1319BFFBBD0D85FBFC" author="Sanmartin I &amp; Ronquist F" box="[664,997,1114,1139]" pageId="22" pageNumber="533" pagination="216 - 243" refId="ref17714" refString="Sanmartin I, Ronquist F. 2004. Southern Hemisphere biogeography inferred by event-based models: plant versus animal patterns. Syst Biol. 53: 216-243. doi:10.1080/10635150490423430" type="journal article" year="2004">Sanmartín and Ronquist 2004</bibRefCitation>
), much earlier than the origin of stenodemines. The emergence of several lineages of stenodemines, including
<taxonomicName id="4C3A4D76FFBC9A131A37FB780F03FB37" box="[272,355,1183,1208]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBC9A131A37FB780F03FB37" box="[272,355,1183,1208]" italics="true" pageId="22" pageNumber="533">Collaria</emphasis>
</taxonomicName>
, might thus have happened in the early Palaeogene or late Cretaceous, between 55 and 80 MYA (
<bibRefCitation id="EFAB4B04FFBC9A131AEBFB250C10FB54" author="Oh M &amp; Kim S &amp; Lee S" box="[460,624,1218,1243]" pageId="22" pageNumber="533" refId="ref17258" refString="Oh M, Kim S, Lee S. 2023. Revisiting the phylogeny of the family Miridae (Heteroptera: Cimicomorpha), with updated insights into its origin and life history evolution. Mol Phylogen Evol. 184: 107796. doi:10.1016/j.ympev.2023.107796" type="journal volume" year="2023">
Oh
<emphasis id="B94EEAE7FFBC9A131ADFFB250C4AFB54" box="[504,554,1218,1243]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2023
</bibRefCitation>
), coinciding with the diversification of grasses (
<bibRefCitation id="EFAB4B04FFBC9A131B81FB020C62FB71" author="Bouchenak-Khelladi Y &amp; Verboom AG &amp; Savolainen V &amp; Hodkinson TR" box="[166,514,1253,1278]" pageId="22" pageNumber="533" pagination="543 - 557" refId="ref14670" refString="Bouchenak-Khelladi Y, Verboom AG, Savolainen V, Hodkinson TR. 2010. Biogeography of the grasses (Poaceae): a phylogenetic approach to reveal evolutionary history in geographical space and geological time. Bot J Linn Soc. 162 (4): 543-557. doi:10.1111/j.1095-8339.2010.01041.x." type="journal article" year="2010">
Bouchenak-Khelladi
<emphasis id="B94EEAE7FFBC9A131AAAFB020FDDFB71" box="[397,445,1253,1278]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2010
</bibRefCitation>
), and after Africa and South America had already drifted apart.
</paragraph>
<paragraph id="8B8536F5FFBC9A121BE7FACD0FD2FF62" blockId="22.[160,1156,559,1728]" lastBlockId="23.[160,1156,142,1727]" lastPageId="23" lastPageNumber="534" pageId="22" pageNumber="533">
A transoceanic disjunction pattern is found in several insect groups (eg
<bibRefCitation id="EFAB4B04FFBC9A1318C0FACD0A1EFACC" author="Ye Z &amp; Zhen Y &amp; Zhou Y &amp; Bu W" box="[999,1150,1322,1347]" pageId="22" pageNumber="533" pagination="793 - 802" refId="ref18700" refString="Ye Z, Zhen Y, Zhou Y, Bu W. 2017. Out of Africa: biogeography and diversification of the pantropical pond skater genus Limnogonus Stal, 1868 (Hemiptera: Gerridae). Ecol Evol. 7: 793-802. doi: 10. 1002 / ece 3.2688" type="journal article" year="2017">
Ye
<emphasis id="B94EEAE7FFBC9A131F2CFACD0A5AFACC" box="[1035,1082,1322,1347]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2017
</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFBC9A131B87FAAA0FD1FAE9" author="Bourguignon T &amp; Tang Q &amp; Syw H" box="[160,433,1357,1382]" pageId="22" pageNumber="533" pagination="970 - 983" refId="ref14732" refString="Bourguignon T, Tang Q, Syw H. 2018. Transoceanic dispersal and plate tectonics shaped global cockroach distributions: evidence from mitochondrial phylogenomics. Mol Biol Evol. 35 (4): 970-983. doi:10.1093/molbev/msy013." type="journal article" year="2018">
Bourguignon
<emphasis id="B94EEAE7FFBC9A131A1BFAAA0F0CFAE9" box="[316,364,1357,1382]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2018
</bibRefCitation>
). Traditionally, the tectonically driven vicariance of Gondwanan origin has been the principal mechanism to explain the distribution of organisms with transoceanic disjunctions (
<bibRefCitation id="EFAB4B04FFBC9A131AEEFA750CC1FA24" author="Amorim DS &amp; Santos CMD &amp; Oliveira SSD" box="[457,673,1426,1451]" pageId="22" pageNumber="533" pagination="2 - 9" refId="ref14414" refString="Amorim DS, Santos CMD, Oliveira SSD. 2009. Allochronic taxa as an alternative model to explain circumantarctic disjunctions. Syst Entomol. 34: 2-9. doi:10.1111/j.1365-3113.2008.00448.x" type="journal article" year="2009">
Amorim
<emphasis id="B94EEAE7FFBC9A13190AFA750C3DFA24" box="[557,605,1426,1451]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2009
</bibRefCitation>
). However, the inclusion of dating analysis in phylogenetic approaches has allowed testing this scenario and providing alternative mechanisms at the same time. One of these is long-distance dispersal, which has arisen as a possible hypothesis when there are inconsistencies between the divergent times of the biological groups and the estimated timing of Gondwana break-up (
<bibRefCitation id="EFAB4B04FFBC9A131882F9FA0A16F9B9" author="Givnish TJ &amp; Millam KC &amp; Evans TM &amp; Hall JC &amp; Chris Pires J &amp; Berry PE &amp; Sytsma KJ" box="[933,1142,1565,1590]" pageId="22" pageNumber="533" refId="ref15560" refString="Givnish TJ, Millam KC, Evans TM, Hall JC, Chris Pires J, Berry PE, Sytsma KJ. 2004. Ancient vicariance or recent long-distance dispersal? Inferences about phylogeny and south american - african disjunctions in rapateaceae and bromeliaceae based on ndh F sequence data. Int J Plant Sci. 165 (4): S 35 - S 54. doi:10.1086/421067." type="journal volume" year="2004">
Givnish
<emphasis id="B94EEAE7FFBC9A131F25F9FA0A52F9B9" box="[1026,1074,1565,1590]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2004
</bibRefCitation>
). Usually, a long-distance dispersal mechanism is commonly invoked in many grass-feeding insects; for example,
<taxonomicName id="4C3A4D76FFBC9A131AB9F9850F93F9F4" box="[414,499,1634,1659]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="tribe" tribe="Satyrini">Satyrini</taxonomicName>
butterflies (
<taxonomicName id="4C3A4D76FFBC9A1319A2F9850D41F9F4" box="[645,801,1634,1659]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="family">Nymphalidae</taxonomicName>
:
<taxonomicName id="4C3A4D76FFBC9A131808F9850DFAF9F4" box="[815,922,1634,1659]" class="Insecta" family="Nymphalidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="subFamily" subFamily="Satyrinae">Satyrinae</taxonomicName>
) (
<bibRefCitation id="EFAB4B04FFBC9A131895F9850A15F9F4" author="Pena C &amp; Nylin S &amp; Wahlberg N" box="[946,1141,1634,1659]" pageId="22" pageNumber="533" pagination="64 - 87" refId="ref17427" refString="Pena C, Nylin S, Wahlberg N. 2011. The radiation of Satyrini butterflies (Nymphalidae: Satyrinae): a challenge for phylogenetic methods. Zool J Linn Soc. 161: 64-87. doi:10.1111/j.1096-3642.2009. 00627. x." type="journal article" year="2011">
Peña
<emphasis id="B94EEAE7FFBC9A1318DFF9850A4DF9F4" box="[1016,1069,1634,1659]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2011
</bibRefCitation>
),
<taxonomicName id="4C3A4D76FFBC9A131B87F9620F7DF911" authorityName="Guenee" authorityYear="1852" box="[160,285,1669,1694]" class="Insecta" family="Noctuidae" genus="Spodoptera" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBC9A131B87F9620F7DF911" box="[160,285,1669,1694]" italics="true" pageId="22" pageNumber="533">Spodoptera</emphasis>
</taxonomicName>
moths (
<taxonomicName id="4C3A4D76FFBC9A131A5EF9620C65F911" authorityName="Linnaeus" authorityYear="1758" box="[377,517,1669,1694]" class="Insecta" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="order">Lepidoptera</taxonomicName>
:
<taxonomicName id="4C3A4D76FFBC9A131937F9620CE5F911" authorityName="Latreille" authorityYear="1809" box="[528,645,1669,1694]" class="Insecta" family="Noctuidae" kingdom="Animalia" order="Lepidoptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="family">Noctuidae</taxonomicName>
) (
<bibRefCitation id="EFAB4B04FFBC9A1319BFF9620D0DF911" author="Kergoat GJ &amp; Prowell DP &amp; Le Ru BP &amp; Mitchell A &amp; Dumas P &amp; Clamens A-L &amp; Condamine FL &amp; Silvain J-F" box="[664,877,1669,1694]" pageId="22" pageNumber="533" pagination="855 - 870" refId="ref16466" refString="Kergoat GJ, Prowell DP, Le Ru BP, Mitchell A, Dumas P, Clamens A-L, Condamine FL, Silvain J-F. 2012. Disentangling dispersal, vicariance and adaptive radiation patterns: a case study using armyworms in the pest genus Spodoptera (Lepidoptera: Noctuidae). Mol Phylogen Evol. 65: 855-870. doi:10.1016/j.ympev.2012.08.006." type="journal article" year="2012">
Kergoat
<emphasis id="B94EEAE7FFBC9A1319DEF9620D49F911" box="[761,809,1669,1694]" italics="true" pageId="22" pageNumber="533">et al</emphasis>
. 2012
</bibRefCitation>
), and grassland leafhoppers of the tribe
<taxonomicName id="4C3A4D76FFBC9A131A56F9400F80F94F" box="[369,480,1703,1728]" class="Insecta" family="Cicadellidae" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="tribe" tribe="Chiasmini">Chiasmini</taxonomicName>
(
<taxonomicName id="4C3A4D76FFBC9A131AD5F9400C0EF94F" authorityName="Linnaeus" authorityYear="1758" box="[498,622,1703,1728]" class="Insecta" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="order">Hemiptera</taxonomicName>
:
<taxonomicName id="4C3A4D76FFBC9A13195BF9400D68F94F" authorityName="Latreille" authorityYear="1825" box="[636,776,1703,1728]" class="Insecta" family="Cicadellidae" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="family">Cicadellidae</taxonomicName>
:
<taxonomicName id="4C3A4D76FFBC9A131830F9400DB2F94F" box="[791,978,1703,1728]" class="Insecta" family="Cicadellidae" kingdom="Animalia" order="Hemiptera" pageId="22" pageNumber="533" phylum="Arthropoda" rank="subFamily" subFamily="Deltocephalinae">Deltocephalinae</taxonomicName>
) (
<bibRefCitation id="EFAB4B04FFBC9A1218CEF9400F56FF28" author="Zahniser JN &amp; Dietrich CH" lastPageId="23" lastPageNumber="534" pageId="22" pageNumber="533" pagination="473 - 495" refId="ref18754" refString="Zahniser JN, Dietrich CH. 2015. Phylogeny, evolution, and historical biogeography of the grassland leafhopper tribe Chiasmini (Hemiptera: Cicadellidae: Deltocephalinae). Zool J Linn Soc. 175 (3): 473-495. doi:10.1111/zoj.12292." type="journal article" year="2015">Zahniser and Dietrich 2015</bibRefCitation>
). Nonetheless, 50 MYA the distance between Africa and South America was about
<quantity id="4CC29B10FFBD9A121BCFFF560F27FF45" box="[232,327,177,202]" metricMagnitude="6" metricUnit="m" metricValue="1.0" pageId="23" pageNumber="534" unit="km" value="1000.0">1000 km</quantity>
on their closest path (
<bibRefCitation id="EFAB4B04FFBD9A12191EFF560D65FF45" author="Oliveira FB &amp; Cassola Molina E &amp; Marroig G" box="[569,773,177,202]" pageId="23" pageNumber="534" pagination="55 - 68" refId="ref17310" refString="Oliveira FB, Cassola Molina E, Marroig G. 2010. Paleogeography of the South Atlantic: a route for primates and rodents into the New World? In: Garber PA, Estrada A, Bicca-Marques JC, Heymann EW, Strier KB, editors. South American primates, developments in primatology: progress and prospects. Houten (The Netherlands): Springer Science; p. 55-68." type="book chapter" year="2010">
Oliveira
<emphasis id="B94EEAE7FFBD9A1219B2FF560CA3FF45" box="[661,707,177,202]" italics="true" pageId="23" pageNumber="534">et al</emphasis>
. 2010
</bibRefCitation>
), making a long-distance dispersal mechanism challenging.
</paragraph>
<paragraph id="8B8536F5FFBD9A121BE7FF110F2FFDE5" blockId="23.[160,1156,142,1727]" pageId="23" pageNumber="534">
Even though mirids are usually regarded as poor dispersers at long distances because of their small size and limited flight power (
<bibRefCitation id="EFAB4B04FFBD9A121983FEFE0DBFFEBD" author="Schuh RT &amp; Stonedahl GM" box="[676,991,281,306]" pageId="23" pageNumber="534" pagination="337 - 355" refId="ref17857" refString="Schuh RT, Stonedahl GM. 1986. Historical biogeography in the Indo - Pacific: a cladistic approach. Cladistics. 2: 337-355. doi:10.1111/j.1096-0031.1986.tb00456.x" type="journal article" year="1986">Schuh and Stonedahl 1986</bibRefCitation>
), various data might suggest otherwise.
<taxonomicName id="4C3A4D76FFBD9A121AE1FEDB0C7AFEDA" box="[454,538,316,341]" class="Insecta" family="Miridae" kingdom="Animalia" order="Hemiptera" pageId="23" pageNumber="534" phylum="Arthropoda" rank="family">Miridae</taxonomicName>
are actually good flyers, better than other heteropterans studied (
<bibRefCitation id="EFAB4B04FFBD9A121A15FEB90F96FEF8" author="Southwood TRE" box="[306,502,350,375]" pageId="23" pageNumber="534" pagination="173 - 220" refId="ref18239" refString="Southwood TRE. 1960. The flight activity of Heteroptera. Trans R Entomol Soc Lond. 112: 173-220. doi:10.1111/j.1365-2311.1960.tb00498.x" type="journal article" year="1960">Southwood 1960</bibRefCitation>
). For instance,
<taxonomicName id="4C3A4D76FFBD9A121982FEB90DD8FEF8" authority="(Stal)" baseAuthorityName="Stal" baseAuthorityYear="1859" box="[677,952,350,375]" class="Insecta" family="Miridae" genus="Creontiades" kingdom="Animalia" order="Hemiptera" pageId="23" pageNumber="534" phylum="Arthropoda" rank="species" species="dilutus">
<emphasis id="B94EEAE7FFBD9A121982FEB90D17FEF8" box="[677,887,350,375]" italics="true" pageId="23" pageNumber="534">Creontiades dilutus</emphasis>
(Stål)
</taxonomicName>
can migrate over long distances over land, up to
<quantity id="4CC29B10FFBD9A12192DFE660C0EFE15" box="[522,622,385,410]" metricMagnitude="6" metricUnit="m" metricValue="1.5" pageId="23" pageNumber="534" unit="km" value="1500.0">1500 km</quantity>
, and can potentially disperse across a range of
<quantity id="4CC29B10FFBD9A121B87FE430F61FE32" box="[160,257,420,445]" metricMagnitude="6" metricUnit="m" metricValue="5.0" pageId="23" pageNumber="534" unit="km" value="5000.0">5000 km</quantity>
(
<bibRefCitation id="EFAB4B04FFBD9A121A2AFE430F96FE32" author="Hereward JP &amp; Walter GH &amp; DeBarro PJ &amp; Lowe AJ &amp; Riginos C" box="[269,502,420,445]" pageId="23" pageNumber="534" pagination="807 - 821" refId="ref16110" refString="Hereward JP, Walter GH, DeBarro PJ, Lowe AJ, Riginos C. 2013. Gene flow in the green mirid, Creontiades dilutus (Hemiptera: Miridae), across arid and agricultural environments with different host plant species. Ecol Evol. 3: 807-821. doi:10.1002/ece3.510" type="journal article" year="2013">
Hereward
<emphasis id="B94EEAE7FFBD9A121AA4FE430FD2FE32" box="[387,434,420,445]" italics="true" pageId="23" pageNumber="534">et al</emphasis>
. 2013
</bibRefCitation>
); this species can also disperse from mainland
<collectingCountry id="F32D7665FFBD9A121F24FE430A04FE32" box="[1027,1124,420,445]" name="Australia" pageId="23" pageNumber="534">Australia</collectingCountry>
to
<collectingRegion id="49FEF817FFBD9A121B87FE210F6AFE50" box="[160,266,454,479]" country="Australia" name="Tasmania" pageId="23" pageNumber="534">Tasmania</collectingRegion>
(
<bibRefCitation id="EFAB4B04FFBD9A121A30FE210F1AFE50" author="Hill L" box="[279,378,454,479]" pageId="23" pageNumber="534" pagination="211 - 220" refId="ref16168" refString="Hill L. 2017. Migration of green mirid, Creontiades dilutus (Stal) and residence of potato bug, Closterotomus norwegicus (Gmelin) in Tasmania (Hemiptera: Miridae: Mirinae: Mirini). Crop Prot. 96: 211-220. doi:10.1016/j.cropro.2017.02.006" type="journal article" year="2017">Hill 2017</bibRefCitation>
). It also has been noted that mirids can be found at sea far from land (about
<quantity id="4CC29B10FFBD9A121BD2FE0E0F2EFD8D" box="[245,334,489,514]" metricMagnitude="5" metricUnit="m" metricValue="6.0" pageId="23" pageNumber="534" unit="km" value="600.0">600 km</quantity>
) (
<bibRefCitation id="EFAB4B04FFBD9A121A41FE0E0C66FD8D" author="Wheeler AG Jr." box="[358,518,489,514]" pageId="23" pageNumber="534" refId="ref18579" refString="Wheeler AG Jr. 2001. Biology of the plant bugs (Hemiptera: Miridae). Pests, Predators, Opportunists. New York: Comstock Publishing Associates, Cornell University Press. Ithaca; p. 558." type="journal volume" year="2001">Wheeler 2001</bibRefCitation>
) which is congruent with data on the colonisation of oceanic islands by mirids (eg
<bibRefCitation id="EFAB4B04FFBD9A121AD7FDEB0C1AFDAA" author="Leston D" box="[496,634,524,549]" pageId="23" pageNumber="534" pagination="41 - 46" refId="ref16681" refString="Leston D. 1957. Spread potential and the colonisation of islands. Syst Zool. 6: 41-46. doi: 10.2307 / 2411707" type="journal article" year="1957">Leston 1957</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFBD9A1219AFFDEB0D7DFDAA" author="Asquith A" box="[648,797,524,549]" pageId="23" pageNumber="534" pagination="356 - 365" refId="ref14537" refString="Asquith A. 1997. Hawaiian Miridae (Hemiptera: Heteroptera): the evolution of bugs and thought. Pacific Sci. 51: 356-365." type="journal article" year="1997">Asquith 1997</bibRefCitation>
). These dispersal abilities probably can be explained by a combination of favourable wind conditions and flight abilities (
<bibRefCitation id="EFAB4B04FFBD9A121B81FDB60F22FDE5" author="Wheeler AG Jr." box="[166,322,593,618]" pageId="23" pageNumber="534" refId="ref18579" refString="Wheeler AG Jr. 2001. Biology of the plant bugs (Hemiptera: Miridae). Pests, Predators, Opportunists. New York: Comstock Publishing Associates, Cornell University Press. Ithaca; p. 558." type="journal volume" year="2001">Wheeler 2001</bibRefCitation>
).
</paragraph>
<paragraph id="8B8536F5FFBD9A121BE7FD930C5CFBA2" blockId="23.[160,1156,142,1727]" pageId="23" pageNumber="534">
An alternative to the long-distance dispersal mechanism to explain the observed vicariant pattern of species of
<taxonomicName id="4C3A4D76FFBD9A121904FD710C19FD20" box="[547,633,662,687]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="23" pageNumber="534" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBD9A121904FD710C19FD20" box="[547,633,662,687]" italics="true" pageId="23" pageNumber="534">Collaria</emphasis>
</taxonomicName>
, distributed between the Neotropical and Nearctic + Afrotropical clades, is geodispersal through a series of large dry land extensions found in the South Atlantic at 50 MYA (Brigs 2003;
<bibRefCitation id="EFAB4B04FFBD9A1219E9FD3B0DF9FD7A" author="Oliveira FB &amp; Cassola Molina E &amp; Marroig G" box="[718,921,732,757]" pageId="23" pageNumber="534" pagination="55 - 68" refId="ref17310" refString="Oliveira FB, Cassola Molina E, Marroig G. 2010. Paleogeography of the South Atlantic: a route for primates and rodents into the New World? In: Garber PA, Estrada A, Bicca-Marques JC, Heymann EW, Strier KB, editors. South American primates, developments in primatology: progress and prospects. Houten (The Netherlands): Springer Science; p. 55-68." type="book chapter" year="2010">
Oliveira
<emphasis id="B94EEAE7FFBD9A12180FFD3B0D37FD7A" box="[808,855,732,757]" italics="true" pageId="23" pageNumber="534">et al</emphasis>
. 2010
</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFBD9A121883FD3B0EB7FC98" author="Ezcurra MD &amp; Agnolin FL" pageId="23" pageNumber="534" pagination="553 - 566" refId="ref15368" refString="Ezcurra MD, Agnolin FL. 2012. A new global palaeobiogeographical model for the late Mesozoic and early tertiary. Syst Biol. 61: 553-566. doi:10.1093/sysbio/syr115" type="journal article" year="2012">Ezcurra and Agnolín 2012</bibRefCitation>
). Between 80 and 50 MYA, there was a series of large islands that connected Africa and South America in the South Atlantic (
<bibRefCitation id="EFAB4B04FFBD9A12194DFCC60D58FCB5" author="Oliveira FB &amp; Cassola Molina E &amp; Marroig G" box="[618,824,801,826]" pageId="23" pageNumber="534" pagination="55 - 68" refId="ref17310" refString="Oliveira FB, Cassola Molina E, Marroig G. 2010. Paleogeography of the South Atlantic: a route for primates and rodents into the New World? In: Garber PA, Estrada A, Bicca-Marques JC, Heymann EW, Strier KB, editors. South American primates, developments in primatology: progress and prospects. Houten (The Netherlands): Springer Science; p. 55-68." type="book chapter" year="2010">
Oliveira
<emphasis id="B94EEAE7FFBD9A1219E0FCC60C96FCB5" box="[711,758,801,826]" italics="true" pageId="23" pageNumber="534">et al</emphasis>
. 2010
</bibRefCitation>
). Evidence of a South Atlantic dispersal pattern has been shown by some groups of plants and vertebrates (eg
<bibRefCitation id="EFAB4B04FFBD9A121F1AFCA30EB7FCF0" author="Briggs JC" pageId="23" pageNumber="534" pagination="548 - 553" refId="ref14815" refString="Briggs JC. 2003. Fishes and birds: gondwana life rafts reconsidered. Syst Biol. 52: 548-553. doi: 10. 1080 / 10635150390218385" type="journal article" year="2003">Briggs 2003</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFBD9A121BC2FC810F13FCF0" author="Renner S" box="[229,371,870,895]" pageId="23" pageNumber="534" refId="ref17517" refString="Renner S. 2004. Plant dispersal across the Tropical Atlantic by wind and sea currents. Int J Plant Sci. 165 (4): S 23 - S 33. doi:10.1086/383334." type="journal volume" year="2004">Renner 2004</bibRefCitation>
). Predominant palaeocurrents and palaeowinds from Africa to South America (
<bibRefCitation id="EFAB4B04FFBD9A121A2EFC6E0FBAFC2D" author="Oliveira FB &amp; Cassola Molina E &amp; Marroig G" box="[265,474,905,930]" pageId="23" pageNumber="534" pagination="55 - 68" refId="ref17310" refString="Oliveira FB, Cassola Molina E, Marroig G. 2010. Paleogeography of the South Atlantic: a route for primates and rodents into the New World? In: Garber PA, Estrada A, Bicca-Marques JC, Heymann EW, Strier KB, editors. South American primates, developments in primatology: progress and prospects. Houten (The Netherlands): Springer Science; p. 55-68." type="book chapter" year="2010">
Oliveira
<emphasis id="B94EEAE7FFBD9A121A41FC6E0FF6FC2D" box="[358,406,905,930]" italics="true" pageId="23" pageNumber="534">et al</emphasis>
. 2010
</bibRefCitation>
) lead us to favour the idea of a geodispersal of
<taxonomicName id="4C3A4D76FFBD9A1218D4FC6E0A26FC2D" box="[1011,1094,905,930]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="23" pageNumber="534" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBD9A1218D4FC6E0A26FC2D" box="[1011,1094,905,930]" italics="true" pageId="23" pageNumber="534">Collaria</emphasis>
</taxonomicName>
from Africa to South America through this island chain between 40 and 50 MYA. These islands were non-existent by 20 MYA (
<bibRefCitation id="EFAB4B04FFBD9A121ADFFC290CAAFC68" author="Oliveira FB &amp; Cassola Molina E &amp; Marroig G" box="[504,714,974,999]" pageId="23" pageNumber="534" pagination="55 - 68" refId="ref17310" refString="Oliveira FB, Cassola Molina E, Marroig G. 2010. Paleogeography of the South Atlantic: a route for primates and rodents into the New World? In: Garber PA, Estrada A, Bicca-Marques JC, Heymann EW, Strier KB, editors. South American primates, developments in primatology: progress and prospects. Houten (The Netherlands): Springer Science; p. 55-68." type="book chapter" year="2010">
Oliveira
<emphasis id="B94EEAE7FFBD9A121971FC290CE6FC68" box="[598,646,974,999]" italics="true" pageId="23" pageNumber="534">et al</emphasis>
. 2010
</bibRefCitation>
), severing the connection between the two continents, and, thus, probably creating the vicariant pattern observed between South America and Africa in
<taxonomicName id="4C3A4D76FFBD9A121AC4FBF30C56FBA2" box="[483,566,1044,1069]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="23" pageNumber="534" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBD9A121AC4FBF30C56FBA2" box="[483,566,1044,1069]" italics="true" pageId="23" pageNumber="534">Collaria</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B8536F5FFBD9A121BE7FBD10CF7F9BA" blockId="23.[160,1156,142,1727]" pageId="23" pageNumber="534">
The African clade was reconstructed as having three unambiguous vicariant events. The first one is between
<taxonomicName id="4C3A4D76FFBD9A121A98FBBE0C23FBFD" box="[447,579,1113,1138]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="23" pageNumber="534" phylum="Arthropoda" rank="species" species="schwartzi">
<emphasis id="B94EEAE7FFBD9A121A98FBBE0C23FBFD" box="[447,579,1113,1138]" italics="true" pageId="23" pageNumber="534">C. schwartzi</emphasis>
</taxonomicName>
, distributed in Central and Eastern Africa, and the remaining species found in Africa, which have a more south-eastern and Madagascan distribution (
<figureCitation id="13012A70FFBD9A121A11FB790FF8FB38" box="[310,408,1182,1207]" captionStart="Figure 8" captionStartId="19.[160,225,1329,1352]" captionTargetBox="[164,1151,143,1296]" captionTargetId="figure-39@19.[163,1153,142,1298]" captionTargetPageId="19" captionText="Figure 8. Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 17. Symbols at the nodes of the phylogeny are as follows: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares." figureDoi="http://doi.org/10.5281/zenodo.14970752" httpUri="https://zenodo.org/record/14970752/files/figure.png" pageId="23" pageNumber="534">Figure 8</figureCitation>
, node 2). The observed pattern could have formed during the Miocene (823 MYA). Initially, the rainforest stretched from coast to coast, but around the mid Miocene
<emphasis id="B94EEAE7FFBD9A121A46FB030FBCFB72" bold="true" box="[353,476,1252,1277]" pageId="23" pageNumber="534">(~17 MYA)</emphasis>
the uplift of eastern Africa started to separate the eastern and western rainforests (
<bibRefCitation id="EFAB4B04FFBD9A121AA5FAE10C76FA90" author="Dijkstra KDB" box="[386,534,1286,1311]" pageId="23" pageNumber="534" pagination="143 - 187" refId="ref15291" refString="Dijkstra KDB. 2007. Demise and rise: the biogeography and taxonomy of the Odonata of tropical Africa [PhD Thesis]. Leiden University, 143-187." type="book chapter" year="2007">Dijkstra 2007</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFBD9A121905FAE10C98FA90" author="Wichura H &amp; Bousquet R &amp; Oberhansli R &amp; Strecker MR &amp; Trauth MH" box="[546,760,1286,1311]" pageId="23" pageNumber="534" pagination="543 - 546" refId="ref18619" refString="Wichura H, Bousquet R, Oberhansli R, Strecker MR, Trauth MH. 2010. Evidence for middle Miocene uplift of the East African Plateau. Geology. 38 (6): 543-546. doi:10.1130/G31022.1." type="journal article" year="2010">
Wichura
<emphasis id="B94EEAE7FFBD9A1219A2FAE10CD5FA90" box="[645,693,1286,1311]" italics="true" pageId="23" pageNumber="534">et al</emphasis>
. 2010
</bibRefCitation>
). The consequence was the expansion of the grass-dominated biome between the mid and late Miocene (816 MYA) (
<bibRefCitation id="EFAB4B04FFBD9A121B81FAAB0F52FAEA" author="Jacobs BF" box="[166,306,1356,1381]" pageId="23" pageNumber="534" pagination="1573 - 1583" refId="ref16362" refString="Jacobs BF. 2004. Palaeobotanical studies from tropical Africa: relevance to the evolution of forest, woodland and Savannah biomes. R Soc London Philos Trans. 359: 1573-1583. doi:10.1098/rstb. 2004.1533" type="journal article" year="2004">Jacobs 2004</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFBD9A121A18FAAB0CC6FAEA" author="Utescher T &amp; Mosbrugger V" box="[319,678,1356,1381]" pageId="23" pageNumber="534" pagination="243 - 271" refId="ref18413" refString="Utescher T, Mosbrugger V. 2007. Eocene vegetation patterns reconstructed from plant diversity - A global perspective. Palaeogeograph Palaeoclimatol Palaeoecol. 247: 243-271. doi:10.1016/j. palaeo. 2006.10.022" type="book chapter" year="2007">Utescher and Mosbrugger 2007</bibRefCitation>
), which could have had an impact on the diversification of grass-associated
<taxonomicName id="4C3A4D76FFBD9A12190AFA890CE1FA08" box="[557,641,1390,1415]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="23" pageNumber="534" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFBD9A12190AFA890CE1FA08" box="[557,641,1390,1415]" italics="true" pageId="23" pageNumber="534">Collaria</emphasis>
</taxonomicName>
. During the Pliocene and early Pleistocene, a major uplift also created the Great Rift Valley and shaped the
<collectingCountry id="F32D7665FFBD9A1218B9FA760D88FA25" box="[926,1000,1425,1450]" name="Democratic Republic of the Congo" pageId="23" pageNumber="534">Congo</collectingCountry>
Basin, which could have divided the continent into eastern and western regions (
<bibRefCitation id="EFAB4B04FFBD9A1218C6FA530A1EFA42" author="Dijkstra KDB" box="[993,1150,1460,1485]" pageId="23" pageNumber="534" pagination="143 - 187" refId="ref15291" refString="Dijkstra KDB. 2007. Demise and rise: the biogeography and taxonomy of the Odonata of tropical Africa [PhD Thesis]. Leiden University, 143-187." type="book chapter" year="2007">Dijkstra 2007</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFBD9A121B87FA310FE2FA60" author="Wichura H &amp; Bousquet R &amp; Oberhansli R &amp; Strecker MR &amp; Trauth MH" box="[160,386,1494,1519]" pageId="23" pageNumber="534" pagination="543 - 546" refId="ref18619" refString="Wichura H, Bousquet R, Oberhansli R, Strecker MR, Trauth MH. 2010. Evidence for middle Miocene uplift of the East African Plateau. Geology. 38 (6): 543-546. doi:10.1130/G31022.1." type="journal article" year="2010">
Wichura
<emphasis id="B94EEAE7FFBD9A121A20FA310F5AFA60" box="[263,314,1494,1519]" italics="true" pageId="23" pageNumber="534">et al</emphasis>
. 2010
</bibRefCitation>
). These complex expansions and contractions of grass-dominated biomes, coupled with changes in temperature (
<bibRefCitation id="EFAB4B04FFBD9A121988FA1E0DF0F99D" author="Couvreur TLP &amp; Dauby G &amp; Blach-Overgaard A" box="[687,912,1529,1554]" pageId="23" pageNumber="534" pagination="16 - 51" refId="ref15099" refString="Couvreur TLP, Dauby G, Blach-Overgaard A. 2021. Tectonics, climate and the diversification of the tropical African terrestrial flora and fauna. Biol Rev. 96: 16-51. doi:10.1111/brv.12644." type="journal article" year="2021">
Couvreur
<emphasis id="B94EEAE7FFBD9A121839FA1E0D2DF99D" box="[798,845,1529,1554]" italics="true" pageId="23" pageNumber="534">et al</emphasis>
. 2021
</bibRefCitation>
), might have affected the diversification of African
<taxonomicName id="4C3A4D76FFBD9A121AC3F9FB0CF7F9BA" box="[484,663,1564,1589]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="23" pageNumber="534" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="B94EEAE7FFBD9A121AC3F9FB0C57F9BA" box="[484,567,1564,1589]" italics="true" pageId="23" pageNumber="534">Collaria</emphasis>
species.
</taxonomicName>
</paragraph>
<paragraph id="8B8536F5FFBD9A1D1BE7F9D90C9BFDC8" blockId="23.[160,1156,142,1727]" lastBlockId="24.[160,1157,142,1727]" lastPageId="24" lastPageNumber="535" pageId="23" pageNumber="534">
The second vicariant event was between
<taxonomicName id="4C3A4D76FFBD9A1219B4F9D90C88F9D8" authorityName="Linnavuori" authorityYear="1975" box="[659,744,1598,1623]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="23" pageNumber="534" phylum="Arthropoda" rank="species" species="nigra">
<emphasis id="B94EEAE7FFBD9A1219B4F9D90C88F9D8" box="[659,744,1598,1623]" italics="true" pageId="23" pageNumber="534">C. nigra</emphasis>
</taxonomicName>
, distributed into East Africa, and the remaining African species, distributed in
<collectingCountry id="F32D7665FFBD9A1219A1F9860D70F9F5" box="[646,784,1633,1658]" name="Madagascar" pageId="23" pageNumber="534">Madagascar</collectingCountry>
, West Africa and Eastern Africa (
<figureCitation id="13012A70FFBD9A121B8FF9630F69F912" box="[168,265,1668,1693]" captionStart="Figure 8" captionStartId="19.[160,225,1329,1352]" captionTargetBox="[164,1151,143,1296]" captionTargetId="figure-39@19.[163,1153,142,1298]" captionTargetPageId="19" captionText="Figure 8. Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 17. Symbols at the nodes of the phylogeny are as follows: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares." figureDoi="http://doi.org/10.5281/zenodo.14970752" httpUri="https://zenodo.org/record/14970752/files/figure.png" pageId="23" pageNumber="534">Figure 8</figureCitation>
, node 4). This event could similarly be explained by the uplift of the East African plateau. The third vicariant event was between the African species distributed in the continent and species found in
<collectingCountry id="F32D7665FFB29A1D191AFF690CA6FF28" box="[573,710,142,167]" name="Madagascar" pageId="24" pageNumber="535">Madagascar</collectingCountry>
, and the
<collectingCountry id="F32D7665FFB29A1D1813FF690DF1FF28" box="[820,913,142,167]" name="Réunion" pageId="24" pageNumber="535">Réunion</collectingCountry>
and Grande Comore Islands (
<figureCitation id="13012A70FFB29A1D1BD9FF560F3EFF45" box="[254,350,177,202]" captionStart="Figure 8" captionStartId="19.[160,225,1329,1352]" captionTargetBox="[164,1151,143,1296]" captionTargetId="figure-39@19.[163,1153,142,1298]" captionTargetPageId="19" captionText="Figure 8. Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 17. Symbols at the nodes of the phylogeny are as follows: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares." figureDoi="http://doi.org/10.5281/zenodo.14970752" httpUri="https://zenodo.org/record/14970752/files/figure.png" pageId="24" pageNumber="535">Figure 8</figureCitation>
, node 5).
<collectingCountry id="F32D7665FFB29A1D1AF0FF560C3DFF45" box="[471,605,177,202]" name="Madagascar" pageId="24" pageNumber="535">Madagascar</collectingCountry>
and Africa separated as part of the Gondwana break-up into eastern and western areas, about 155170 MYA (
<bibRefCitation id="EFAB4B04FFB29A1D18B3FF330A16FF62" author="Masters JC &amp; de Wit MJ &amp; Asher RJ" box="[916,1142,212,237]" pageId="24" pageNumber="535" pagination="399 - 418" refId="ref16810" refString="Masters JC, de Wit MJ, Asher RJ. 2006. Reconciling the origins of Africa, India and Madagascar with vertebrate dispersal scenarios. Folia Primatologica. 77: 399-418. doi:10.1159/000095388" type="journal article" year="2006">
Masters
<emphasis id="B94EEAE7FFB29A1D18DEFF330A4DFF62" box="[1017,1069,212,237]" italics="true" pageId="24" pageNumber="535">et al</emphasis>
. 2006
</bibRefCitation>
).
<collectingCountry id="F32D7665FFB29A1D1B87FF110F2DFE80" box="[160,333,246,271]" name="Reunion Island" pageId="24" pageNumber="535">Réunion Island</collectingCountry>
is located about
<quantity id="4CC29B10FFB29A1D1901FF110C1EFE80" box="[550,638,246,271]" metricMagnitude="5" metricUnit="m" metricValue="6.8" pageId="24" pageNumber="535" unit="km" value="680.0">680 km</quantity>
east of
<collectingCountry id="F32D7665FFB29A1D19C0FF110D0FFE80" box="[743,879,246,271]" name="Madagascar" pageId="24" pageNumber="535">Madagascar</collectingCountry>
on the western Indian Ocean, and it is composed of three shield volcanoes (
<bibRefCitation id="EFAB4B04FFB29A1D19CEFEFE0DD4FEBD" author="Michon L &amp; Saint-Ange F &amp; Bachelery P &amp; Villeneuve N &amp; Staudacher T" box="[745,948,281,306]" pageId="24" pageNumber="535" refId="ref17071" refString="Michon L, Saint-Ange F, Bachelery P, Villeneuve N, Staudacher T. 2007. Role of the structural inheritance of the oceanic lithosphere in the magmato-tectonic evolution of Piton de la Fournaise volcano (La Reunion Island). J Geophys Res. 112: B 04205. doi:10.1029/2006JB004598" type="journal volume" year="2007">
Michon
<emphasis id="B94EEAE7FFB29A1D1864FEFE0D11FEBD" box="[835,881,281,306]" italics="true" pageId="24" pageNumber="535">et al</emphasis>
. 2007
</bibRefCitation>
). The archipelagos around
<collectingCountry id="F32D7665FFB29A1D1BDCFEDB0FE1FEDA" box="[251,385,316,341]" name="Madagascar" pageId="24" pageNumber="535">Madagascar</collectingCountry>
vary widely in origin, age, and structure, although their formation could respond to Gondwana separating from Africa (
<bibRefCitation id="EFAB4B04FFB29A1D1835FEB90A7DFEF8" author="Ali JR &amp; Aitchison JC" box="[786,1053,350,375]" pageId="24" pageNumber="535" pagination="145 - 166" refId="ref14351" refString="Ali JR, Aitchison JC. 2008. Gondwana to Asia: plate tectonics, paleogeography and the biological connectivity of the Indian sub-continent from the Middle Jurassic through latest Eocene (166 - 35 Ma). Earth Sci Rev. 88 (3 - 4): 145-166. doi:10.1016/j.earscirev.2008.01.007." type="journal article" year="2008">Ali and Aitchison 2008</bibRefCitation>
). Recent studies propose that faunal composition, high endemic biodiversity, and the presence of unique lineages in these islands could be the result of dispersal and in situ diversification processes (
<bibRefCitation id="EFAB4B04FFB29A1D1A68FE210C66FE50" author="Strijk JS &amp; Noyes RD &amp; Strasberg D &amp; Cruaud C &amp; Gavory F &amp; Chase MW &amp; Abbott RJ &amp; Thebaud C" box="[335,518,454,479]" pageId="24" pageNumber="535" refId="ref18340" refString="Strijk JS, Noyes RD, Strasberg D, Cruaud C, Gavory F, Chase MW, Abbott RJ, Thebaud C. 2012. In and out of Madagascar: dispersal to peripheral islands, insular speciation and diversification of Indian Ocean daisy trees (Psiadia, Asteraceae). PLOS ONE. 7 (8): e 42932. doi:10.1371/journal.pone. 0042932." type="book" year="2012">
Strijk
<emphasis id="B94EEAE7FFB29A1D1AB6FE210FA2FE50" box="[401,450,454,479]" italics="true" pageId="24" pageNumber="535">et al</emphasis>
. 2012
</bibRefCitation>
). If we accept that the previous vicariant event in Africa probably occurred in the mid-Miocene, the present distributional pattern between Africa and
<collectingCountry id="F32D7665FFB29A1D1BF6FDEB0F37FDAA" box="[209,343,524,549]" name="Madagascar" pageId="24" pageNumber="535">Madagascar</collectingCountry>
and related areas is likely not the product of a vicariant event, but could instead correspond to more recent dispersion events.
</paragraph>
<paragraph id="8B8536F5FFB29A1D1BE7FDB60CD6FACD" blockId="24.[160,1157,142,1727]" pageId="24" pageNumber="535">
The event between the North American and the African species
<taxonomicName id="4C3A4D76FFB29A1D18C7FDB60A4CFDE5" box="[992,1068,593,618]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="24" pageNumber="535" phylum="Arthropoda" rank="species" species="royi">
<emphasis id="B94EEAE7FFB29A1D18C7FDB60A4CFDE5" box="[992,1068,593,618]" italics="true" pageId="24" pageNumber="535">C. royi</emphasis>
</taxonomicName>
in our analysis was reconstructed as a founder event (
<figureCitation id="13012A70FFB29A1D183AFD930DE9FD02" box="[797,905,628,653]" captionStart="Figure 9" captionStartId="20.[160,225,1356,1379]" captionTargetBox="[164,1151,143,1322]" captionTargetId="figure-54@20.[163,1153,142,1324]" captionTargetPageId="20" captionText="Figure 9. Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 815. Symbols at the nodes of the phylogeny are: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares." figureDoi="http://doi.org/10.5281/zenodo.14970754" httpUri="https://zenodo.org/record/14970754/files/figure.png" pageId="24" pageNumber="535">Figure 9</figureCitation>
, node 14). Several
<taxonomicName id="4C3A4D76FFB29A1D1B86FD710F53FD20" box="[161,307,662,687]" pageId="24" pageNumber="530" rank="tribe" tribe="Stenodemini">Stenodemini</taxonomicName>
lineages probably originated between the latest Cretaceous and the early Palaeogene, from 55 to 80 MYA (
<bibRefCitation id="EFAB4B04FFB29A1D1949FD5E0D78FD5D" author="Oh M &amp; Kim S &amp; Lee S" box="[622,792,697,722]" pageId="24" pageNumber="535" refId="ref17258" refString="Oh M, Kim S, Lee S. 2023. Revisiting the phylogeny of the family Miridae (Heteroptera: Cimicomorpha), with updated insights into its origin and life history evolution. Mol Phylogen Evol. 184: 107796. doi:10.1016/j.ympev.2023.107796" type="journal volume" year="2023">
Oh
<emphasis id="B94EEAE7FFB29A1D19BBFD5E0CB0FD5D" box="[668,720,697,722]" italics="true" pageId="24" pageNumber="535">et al</emphasis>
. 2023
</bibRefCitation>
). A biogeographic relationship between these two areas has rarely been documented or explained.
<bibRefCitation id="EFAB4B04FFB29A1D18E2FD3B0A1FFD7A" author="Hembree DI" box="[965,1151,732,757]" pageId="24" pageNumber="535" pagination="340 - 354" refId="ref15971" refString="Hembree DI. 2006. Amphisbaenian paleobiogeography: evidence of vicariance and geodispersal patterns. Palaeogeograph Palaeoclimatol Palaeoecol. 235 (4): 340-354. doi:10.1016/j.palaeo.2005. 11.006." type="journal article" year="2006">Hembree (2006)</bibRefCitation>
, in a biogeographic analysis of amphisbaenians, showed with a molecular data set that African areas had complex relationships with both Asia and North America, rather than with South America. In his analysis, North America was closely related to both Western Asia and Northern Africa, a relationship which was not satisfactorily explained. Certain Scarabaeinae dung beetle taxa have been shown to have a North AmericanAfrican relationship (Phillips 2016). Africa was connected to Eurasia during the Miocene (
<bibRefCitation id="EFAB4B04FFB29A1D1B81FC290F8EFC68" author="Steinthorsdottir M &amp; Coxall HK &amp; De Boer AM &amp; Huber M &amp; Barbolini N &amp; Bradshaw CD &amp; Burls NJ &amp; Feakins SJ &amp; Gasson E &amp; Henderiks J" box="[166,494,974,999]" pageId="24" pageNumber="535" refId="ref18276" refString="Steinthorsdottir M, Coxall HK, De Boer AM, Huber M, Barbolini N, Bradshaw CD, Burls NJ, Feakins SJ, Gasson E, Henderiks J, et al. 2021. The Miocene: the future of the past. Paleoceanograp Paleoclimatol. 36 (4): e 2020. doi:10.1029/2020PA004037." type="journal volume" year="2021">
Steinthorsdottir
<emphasis id="B94EEAE7FFB29A1D1A4CFC290FC2FC68" box="[363,418,974,999]" italics="true" pageId="24" pageNumber="535">et al</emphasis>
. 2021
</bibRefCitation>
), and an exchange of fauna with North America was hypothesised to have occurred from the Miocene up to the Pleistocene via tectonic or sea level changes (
<bibRefCitation id="EFAB4B04FFB29A1D1AB9FBF30C00FBA2" author="Davis ALV &amp; Scholtz CH &amp; Philips TK" box="[414,608,1044,1069]" pageId="24" pageNumber="535" pagination="1217 - 1256" refId="ref15142" refString="Davis ALV, Scholtz CH, Philips TK. 2002. Historical biogeography of scarabaeine dung beetles. J Biogeograph. 29 (9): 1217-1256. doi:10.1046/j.1365-2699.2002.00776.x." type="journal article" year="2002">
Davis
<emphasis id="B94EEAE7FFB29A1D1AC0FBF30C79FBA2" box="[487,537,1044,1069]" italics="true" pageId="24" pageNumber="535">et al</emphasis>
. 2002
</bibRefCitation>
). Boreotropical migration has been invoked to explain the current distribution of certain groups of plants (
<bibRefCitation id="EFAB4B04FFB29A1D1875FBD10EB7FBFD" author="Smedmark JE &amp; Anderberg AA" pageId="24" pageNumber="535" pagination="1491 - 1505" refId="ref18192" refString="Smedmark JE, Anderberg AA. 2007. Boreotropical migration explains hybridization between geographically distant lineages in the pantropical clade Sideroxyleae (Sapotaceae). Am. J. Bot 94 (9): 1491-1505. doi:10.3732/ajb.94.9.1491." type="journal article" year="2007">Smedmark and Anderberg 2007</bibRefCitation>
;
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Wei
<emphasis id="B94EEAE7FFB29A1D1A38FBBE0F34FBFD" box="[287,340,1113,1138]" italics="true" pageId="24" pageNumber="535">et al</emphasis>
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;
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Huang
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) and animals (
<bibRefCitation id="EFAB4B04FFB29A1D1812FBBE0A6FFBFD" author="Brunke AJ &amp; Chatzimanolis S &amp; Metscher BD &amp; Wolf-Schwenninger K &amp; Solodovnikov A" box="[821,1039,1113,1138]" pageId="24" pageNumber="535" refId="ref14845" refString="Brunke AJ, Chatzimanolis S, Metscher BD, Wolf-Schwenninger K, Solodovnikov A. 2017. Dispersal of thermophilic beetles across the intercontinental Arctic forest belt during the early Eocene. Sci Rep. 7: 12972. doi:10.1038/s41598-017-13207-4" type="journal volume" year="2017">
Brunke
<emphasis id="B94EEAE7FFB29A1D18B5FBBE0DA7FBFD" box="[914,967,1113,1138]" italics="true" pageId="24" pageNumber="535">et al</emphasis>
. 2017
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;
<bibRefCitation id="EFAB4B04FFB29A1D1F06FBBE0EB7FB1A" author="Ye Z &amp; Zhen Y &amp; Zhou Y &amp; Bu W" pageId="24" pageNumber="535" pagination="793 - 802" refId="ref18700" refString="Ye Z, Zhen Y, Zhou Y, Bu W. 2017. Out of Africa: biogeography and diversification of the pantropical pond skater genus Limnogonus Stal, 1868 (Hemiptera: Gerridae). Ecol Evol. 7: 793-802. doi: 10. 1002 / ece 3.2688" type="journal article" year="2017">
Ye
<emphasis id="B94EEAE7FFB29A1D1F6EFBBE0A1DFBFD" box="[1097,1149,1113,1138]" italics="true" pageId="24" pageNumber="535">et al</emphasis>
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), and thus could be an explanation for the pattern of
<taxonomicName id="4C3A4D76FFB29A1D186FFB9B0D97FB1A" box="[840,1015,1148,1173]" pageId="24" pageNumber="535">
<emphasis id="B94EEAE7FFB29A1D186FFB9B0DFCFB1A" box="[840,924,1148,1173]" italics="true" pageId="24" pageNumber="535">Collaria</emphasis>
species
</taxonomicName>
distribution between North America and Africa. This would imply that the African and North American
<taxonomicName id="4C3A4D76FFB29A1D1A38FB260F16FB55" box="[287,374,1217,1242]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="24" pageNumber="535" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFB29A1D1A38FB260F16FB55" box="[287,374,1217,1242]" italics="true" pageId="24" pageNumber="535">Collaria</emphasis>
</taxonomicName>
clade diversified during and after the Miocene. Alternatively, because the phylogenetic support for the relationship between
<taxonomicName id="4C3A4D76FFB29A1D18E0FB030A74FB72" box="[967,1044,1252,1277]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="24" pageNumber="535" phylum="Arthropoda" rank="species" species="royi">
<emphasis id="B94EEAE7FFB29A1D18E0FB030A74FB72" box="[967,1044,1252,1277]" italics="true" pageId="24" pageNumber="535">C. royi</emphasis>
</taxonomicName>
and the Nearctic clade is weak (
<figureCitation id="13012A70FFB29A1D1AE4FAE10C46FA90" box="[451,550,1286,1311]" captionStart="Figure 7" captionStartId="17.[160,225,1207,1230]" captionTargetBox="[164,1152,144,1174]" captionTargetId="figure-110@17.[163,1153,142,1175]" captionTargetPageId="17" captionText="Figure 7. Single most parsimonious tree obtained with the implied weighting (IW) analysis and using k = 30. Main apomorphies mapped on the tree, non-homoplasious changes as black polygons, homoplasious changes as white polygons. Support values in the branches are within square brackets (Bremer support, absolute symmetric resampling and “Group present/Contradicted” (GC) frequencies, respectively). Clades A to F. Geographic distribution: Nearctic region (blue), African region (marine blue), Neotropical region (green)." figureDoi="http://doi.org/10.5281/zenodo.14970750" httpUri="https://zenodo.org/record/14970750/files/figure.png" pageId="24" pageNumber="535">Figure 7</figureCitation>
), the biogeographic pattern might be an artefact and, thus, its interpretation remains tentative.
</paragraph>
<paragraph id="8B8536F5FFB29A1C1BE7FAAB0C99FF62" blockId="24.[160,1157,142,1727]" lastBlockId="25.[160,1156,142,1173]" lastPageId="25" lastPageNumber="536" pageId="24" pageNumber="535">
The Nearctic Region comprises cold temperate and arid subtropical areas, shaped by two main events: the glaciation dynamic and aridity during the Holocene (
<bibRefCitation id="EFAB4B04FFB29A1D1F07FA890EB7FA25" author="Morrone J" pageId="24" pageNumber="535" refId="ref17213" refString="Morrone J. 2019. Regionalizacion biogeografica y evolucion biotica de Mexico: encrucijada de la de la biodiversidad del Nuevo Mundo. Rev Mex Biodivers. 90: e 902980. doi:10.22201/ib.20078706e. 2019.90. 2980." type="journal volume" year="2019">Morrone 2019</bibRefCitation>
). The Nearctic species of
<taxonomicName id="4C3A4D76FFB29A1D1925FA760C36FA25" box="[514,598,1425,1450]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="24" pageNumber="535" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFB29A1D1925FA760C36FA25" box="[514,598,1425,1450]" italics="true" pageId="24" pageNumber="535">Collaria</emphasis>
</taxonomicName>
occur in Eastern North America (
<figureCitation id="13012A70FFB29A1D18FCFA760A5AFA25" box="[987,1082,1425,1450]" captionStart="Figure 9" captionStartId="20.[160,225,1356,1379]" captionTargetBox="[164,1151,143,1322]" captionTargetId="figure-54@20.[163,1153,142,1324]" captionTargetPageId="20" captionText="Figure 9. Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 815. Symbols at the nodes of the phylogeny are: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares." figureDoi="http://doi.org/10.5281/zenodo.14970754" httpUri="https://zenodo.org/record/14970754/files/figure.png" pageId="24" pageNumber="535">Figure 9</figureCitation>
, node 15), which could be a response to glaciation dynamics during the Holocene, approximately 11,700 years ago. An Eastern North America distribution pattern is considered an endemic area, as shown by several biological groups (
<bibRefCitation id="EFAB4B04FFB29A1D185DFA1E0A1DF99D" author="Escalante T &amp; Rodriguez-Tapia G &amp; Szumik C &amp; Morrone JJ &amp; Rivas M" box="[890,1149,1529,1554]" pageId="24" pageNumber="535" pagination="1381 - 1388" refId="ref15322" refString="Escalante T, Rodriguez-Tapia G, Szumik C, Morrone JJ, Rivas M. 2010. Delimitation of the nearctic region according to mammalian distributional patterns. J Mammal. 91: 1381-1388. doi: 10.1644 / 10 - MAMM-A- 136.1." type="journal article" year="2010">
Escalante
<emphasis id="B94EEAE7FFB29A1D18DFFA1E0A51F99D" box="[1016,1073,1529,1554]" italics="true" pageId="24" pageNumber="535">et al</emphasis>
. 2010
</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFB29A1D1B86F9FB0FF2F9BA" author="Weirauch C &amp; Seltmann KC &amp; Schuh RT &amp; Schwartz MD &amp; Johnson C &amp; Feist MA &amp; Soltis PS" box="[161,402,1564,1589]" pageId="24" pageNumber="535" pagination="279 - 294" refId="ref18512" refString="Weirauch C, Seltmann KC, Schuh RT, Schwartz MD, Johnson C, Feist MA, Soltis PS. 2017. Areas of endemism in the Nearctic: a case study of 1339 species of Miridae (Insecta: Hemiptera) and their plant hosts. Cladistics. 33 (3): 279-294. doi:10.1111/cla.12169." type="journal article" year="2017">
Weirauch
<emphasis id="B94EEAE7FFB29A1D1A31F9FB0F2AF9BA" box="[278,330,1564,1589]" italics="true" pageId="24" pageNumber="535">et al</emphasis>
. 2017
</bibRefCitation>
). The north-eastern region in the
<collectingCountry id="F32D7665FFB29A1D180AF9FB0DA9F9BA" box="[813,969,1564,1589]" name="United States of America" pageId="24" pageNumber="535">United States</collectingCountry>
was influenced and shaped by the advance and retreat of these glaciers, forming glacial lakes in low areas between or in front of glaciers, and shaped its current topography, drainage patterns, and organism distribution (
<bibRefCitation id="EFAB4B04FFB29A1D196FF9630C92F912" author="Sandford KS" box="[584,754,1668,1693]" pageId="24" pageNumber="535" pagination="543 - 552" refId="ref17678" refString="Sandford KS. 1929. The glacial conditions and Quaternary history of North-East Land (Continued). Geog J. 74 (6): 543-552. doi:10.2307/1785162." type="journal article" year="1929">Sandford 1929</bibRefCitation>
).
<taxonomicName id="4C3A4D76FFB29A1D182EF9630DA5F912" authorityName="Akingbohungbe, Libby, &amp; Shenefelt" authorityYear="1973" box="[777,965,1668,1693]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="24" pageNumber="535" phylum="Arthropoda" rank="species" species="meilleuri">
<emphasis id="B94EEAE7FFB29A1D182EF9630DA5F912" box="[777,965,1668,1693]" italics="true" pageId="24" pageNumber="535">Collaria meilleuri</emphasis>
</taxonomicName>
is distributed in
<collectingCountry id="F32D7665FFB29A1D1B87F9410E98F930" box="[160,248,1702,1727]" name="Canada" pageId="24" pageNumber="535">Canada</collectingCountry>
, the north-eastern
<collectingCountry id="F32D7665FFB29A1D1AC0F9410CE7F930" box="[487,647,1702,1727]" name="United States of America" pageId="24" pageNumber="535">United States</collectingCountry>
and Appalachian regions, and
<taxonomicName id="4C3A4D76FFB29A1D1F2AF9410AE3F930" baseAuthorityName="Reuter" baseAuthorityYear="1876" box="[1037,1155,1702,1727]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="24" pageNumber="535" phylum="Arthropoda" rank="species" species="oculata">
<emphasis id="B94EEAE7FFB29A1D1F2AF9410AE3F930" box="[1037,1155,1702,1727]" italics="true" pageId="24" pageNumber="535">C. oculata</emphasis>
</taxonomicName>
mostly in the Southeastern Plains and Florida (Southeast Coastal Plain) (
<bibRefCitation id="EFAB4B04FFB39A1C18C0FF690EB7FF45" author="Morales I &amp; Ferreira PSF &amp; Forero D" pageId="25" pageNumber="536" pagination="201 - 246" refId="ref17129" refString="Morales I, Ferreira PSF, Forero D. 2016. Taxonomic revision of Collaria Provancher, 1872 (Hemiptera: Miridae) with the description of a new species from the Afrotropical region. Zootaxa. 4138 (2): 201-246. doi:10.11646/zootaxa.4138.2.1." type="journal article" year="2016">
Morales
<emphasis id="B94EEAE7FFB39A1C1F6CFF690A1DFF28" box="[1099,1149,142,167]" italics="true" pageId="25" pageNumber="536">et al</emphasis>
. 2016
</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFB39A1C1BCFFF560FB9FF45" author="Weirauch C &amp; Seltmann KC &amp; Schuh RT &amp; Schwartz MD &amp; Johnson C &amp; Feist MA &amp; Soltis PS" box="[232,473,177,202]" pageId="25" pageNumber="536" pagination="279 - 294" refId="ref18512" refString="Weirauch C, Seltmann KC, Schuh RT, Schwartz MD, Johnson C, Feist MA, Soltis PS. 2017. Areas of endemism in the Nearctic: a case study of 1339 species of Miridae (Insecta: Hemiptera) and their plant hosts. Cladistics. 33 (3): 279-294. doi:10.1111/cla.12169." type="journal article" year="2017">
Weirauch
<emphasis id="B94EEAE7FFB39A1C1A79FF560FF1FF45" box="[350,401,177,202]" italics="true" pageId="25" pageNumber="536">et al</emphasis>
. 2017
</bibRefCitation>
), with a broad overlapping area between the two, which could be a response to similar glaciation processes.
</paragraph>
<paragraph id="8B8536F5FFB39A1C1BE7FF110D29FC98" blockId="25.[160,1156,142,1173]" pageId="25" pageNumber="536">
The Neotropical clade exhibits only one vicariant event between the north-western and the south-eastern species (
<figureCitation id="13012A70FFB39A1C1ADDFEFE0C35FEBD" box="[506,597,281,306]" captionStart="Figure 9" captionStartId="20.[160,225,1356,1379]" captionTargetBox="[164,1151,143,1322]" captionTargetId="figure-54@20.[163,1153,142,1324]" captionTargetPageId="20" captionText="Figure 9. Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 815. Symbols at the nodes of the phylogeny are: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares." figureDoi="http://doi.org/10.5281/zenodo.14970754" httpUri="https://zenodo.org/record/14970754/files/figure.png" pageId="25" pageNumber="536">Figure 9</figureCitation>
, node 10). Nonetheless, this distribution pattern is not always clear in the case of
<taxonomicName id="4C3A4D76FFB39A1C1AD3FEDB0C35FEDA" baseAuthorityName="Distant" baseAuthorityYear="1883" box="[500,597,316,341]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="25" pageNumber="536" phylum="Arthropoda" rank="species" species="oleosa">
<emphasis id="B94EEAE7FFB39A1C1AD3FEDB0C35FEDA" box="[500,597,316,341]" italics="true" pageId="25" pageNumber="536">C. oleosa</emphasis>
</taxonomicName>
and
<taxonomicName id="4C3A4D76FFB39A1C19ABFEDB0C96FEDA" baseAuthorityName="Stal" baseAuthorityYear="1859" box="[652,758,316,341]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="25" pageNumber="536" phylum="Arthropoda" rank="species" species="scenica">
<emphasis id="B94EEAE7FFB39A1C19ABFEDB0C96FEDA" box="[652,758,316,341]" italics="true" pageId="25" pageNumber="536">C. scenica</emphasis>
</taxonomicName>
, in which both are distributed from the southern portion of the Chacoan subregion of the Atlantic Forest to the north of the Mexican transition zone, including the Antillean and Brazilian subregion (
<bibRefCitation id="EFAB4B04FFB39A1C18F1FE660A16FE15" author="Morrone J" box="[982,1142,385,410]" pageId="25" pageNumber="536" pagination="81 - 90" refId="ref17183" refString="Morrone J. 2015. Biogeographical regionalisation of the world: a reappraisal. Aust Syst Bot. 28: 81-90. doi:10.1071/SB14042" type="journal article" year="2015">Morrone 2015</bibRefCitation>
). The disjunction between the north-west and south-east areas could be explained by the origin of the Atlantic Forest with open vegetation formations since the Miocene and its complete isolation since at least the Pliocene. These events have been invoked to explain similar distribution patterns in other biological groups (
<bibRefCitation id="EFAB4B04FFB39A1C1825FDEB0A7DFDAA" author="Sigrist MS &amp; Carvalho CJB" box="[770,1053,524,549]" pageId="25" pageNumber="536" pagination="79 - 91" refId="ref18113" refString="Sigrist MS, Carvalho CJB. 2009. Historical relationships among areas of endemism in the tropical South America using Brooks parsimony analysis (BPA). Biota Neotropica. 9: 79-91. doi: 10.1590 / S 1676 - 06032009000400009" type="journal article" year="2009">Sigrist and Carvalho 2009</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFB39A1C1F0FFDEB0F35FDC8" author="Batalha-Filho H &amp; Fjeldsa J &amp; Fabre PH &amp; Miyaki CY" pageId="25" pageNumber="536" pagination="41 - 50" refId="ref14566" refString="Batalha-Filho H, Fjeldsa J, Fabre PH, Miyaki CY. 2013. Connections between the Atlantic and the Amazonian forest avifaunas represent distinct historical events. J Ornitho. 154: 41-50. doi: 10. 1007 / s 10336 - 012 - 0866 - 7" type="journal article" year="2013">
Batalha-Filho
<emphasis id="B94EEAE7FFB39A1C1BF8FDC90F70FDC8" box="[223,272,558,583]" italics="true" pageId="25" pageNumber="536">et al</emphasis>
. 2013
</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFB39A1C1A44FDC90C5EFDC8" author="Silva M &amp; Noll FB" box="[355,574,558,583]" pageId="25" pageNumber="536" pagination="1 - 10" refId="ref18157" refString="Silva M, Noll FB. 2014. Biogeography of the social wasp genus Brachygastra (Hymenoptera: Vespidade: Polistinae). J Biogeograph. 18 (141): 1-10." type="journal article" year="2014">Silva and Noll 2014</bibRefCitation>
). The patterns of the clade containing
<emphasis id="B94EEAE7FFB39A1C18DBFDC90F77FDE5" italics="true" pageId="25" pageNumber="536">
<taxonomicName id="4C3A4D76FFB39A1C18DBFDC90A1FFDC8" authorityName="Carvalho" authorityYear="1990" box="[1020,1151,558,583]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="25" pageNumber="536" phylum="Arthropoda" rank="species" species="boliviana">C. boliviana</taxonomicName>
,
<taxonomicName id="4C3A4D76FFB39A1C1B87FDB60F77FDE5" box="[160,279,593,618]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="25" pageNumber="536" phylum="Arthropoda" rank="species" species="manoloi">C. manoloi</taxonomicName>
</emphasis>
,
<taxonomicName id="4C3A4D76FFB39A1C1A03FDB60FE7FDE5" baseAuthorityName="Distant" baseAuthorityYear="1883" box="[292,391,593,618]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="25" pageNumber="536" phylum="Arthropoda" rank="species" species="oleosa">
<emphasis id="B94EEAE7FFB39A1C1A03FDB60FE7FDE5" box="[292,391,593,618]" italics="true" pageId="25" pageNumber="536">C. oleosa</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C3A4D76FFB39A1C1AE0FDB60C52FDE5" baseAuthorityName="Stal" baseAuthorityYear="1859" box="[455,562,593,618]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="25" pageNumber="536" phylum="Arthropoda" rank="species" species="scenica">
<emphasis id="B94EEAE7FFB39A1C1AE0FDB60C52FDE5" box="[455,562,593,618]" italics="true" pageId="25" pageNumber="536">C. scenica</emphasis>
</taxonomicName>
are mostly explained by sympatric events (
<figureCitation id="13012A70FFB39A1C1F06FDB60A1EFDE5" box="[1057,1150,593,618]" captionStart="Figure 8" captionStartId="19.[160,225,1329,1352]" captionTargetBox="[164,1151,143,1296]" captionTargetId="figure-39@19.[163,1153,142,1298]" captionTargetPageId="19" captionText="Figure 8. Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 17. Symbols at the nodes of the phylogeny are as follows: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares." figureDoi="http://doi.org/10.5281/zenodo.14970752" httpUri="https://zenodo.org/record/14970752/files/figure.png" pageId="25" pageNumber="536">Figure 8</figureCitation>
, nodes 5 and 6). Given these sympatric patterns, besides geographical factors, other biological aspects such as intraspecific biochemical recognition could be at play (
<bibRefCitation id="EFAB4B04FFB39A1C1F62FD710F7BFD5D" author="Byers JA &amp; Fefer D &amp; Levi-Zada A" pageId="25" pageNumber="536" pagination="1115 - 1123" refId="ref14949" refString="Byers JA, Fefer D, Levi-Zada A. 2013. Sex pheromone component ratios and mating isolation among three Lygus plant bug species of North America. Naturwissenschaften. 100: 1115-1123. doi: 10. 1007 / s 00114 - 013 - 1113 - 7" type="journal article" year="2013">
Byers
<emphasis id="B94EEAE7FFB39A1C1B87FD5E0EB3FD5D" box="[160,211,697,722]" italics="true" pageId="25" pageNumber="536">et al</emphasis>
. 2013
</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFB39A1C1A0CFD5E0FB0FD5D" author="Sai Z &amp; Shuwei Y &amp; Zhixiang Z" box="[299,464,697,722]" pageId="25" pageNumber="536" refId="ref17632" refString="Sai Z, Shuwei Y, Zhixiang Z. 2021. Identification and functional characterization of sex pheromone receptors in mirid bugs (Heteroptera: Miridae). Insect Biochem Mol Biol. 136: 103621. doi: 10.1016 / j. ibmb. 2021.103621" type="journal volume" year="2021">
Sai
<emphasis id="B94EEAE7FFB39A1C1A71FD5E0FE9FD5D" box="[342,393,697,722]" italics="true" pageId="25" pageNumber="536">et al</emphasis>
. 2021
</bibRefCitation>
). The founder event between
<taxonomicName id="4C3A4D76FFB39A1C1810FD5E0DC5FD5D" authorityName="Carvalho" authorityYear="1955" box="[823,933,697,722]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="25" pageNumber="536" phylum="Arthropoda" rank="species" species="husseyi">
<emphasis id="B94EEAE7FFB39A1C1810FD5E0DC5FD5D" box="[823,933,697,722]" italics="true" pageId="25" pageNumber="536">C. husseyi</emphasis>
</taxonomicName>
and
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<emphasis id="B94EEAE7FFB39A1C18C2FD5E0AE3FD5D" box="[997,1155,697,722]" italics="true" pageId="25" pageNumber="536">C. guaraniana</emphasis>
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(
<figureCitation id="13012A70FFB39A1C1B8FFD3B0F64FD7A" box="[168,260,732,757]" captionStart="Figure 8" captionStartId="19.[160,225,1329,1352]" captionTargetBox="[164,1151,143,1296]" captionTargetId="figure-39@19.[163,1153,142,1298]" captionTargetPageId="19" captionText="Figure 8. Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 17. Symbols at the nodes of the phylogeny are as follows: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares." figureDoi="http://doi.org/10.5281/zenodo.14970752" httpUri="https://zenodo.org/record/14970752/files/figure.png" pageId="25" pageNumber="536">Figure 8</figureCitation>
, node 6) could be explained by the vegetation changes between the Araucaria Forest province and the Atlantic and
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Forests regions.
</paragraph>
<paragraph id="8B8536F5FFB39A1C1BE7FCC60C90FB1A" blockId="25.[160,1156,142,1173]" pageId="25" pageNumber="536">
Information on species distributions is scattered in natural history collections and various scientific publications, affecting biogeographical inferences through the incomplete taxonomic and distributional knowledge (
<bibRefCitation id="EFAB4B04FFB39A1C19E0FC810DE4FCF0" author="Arias" box="[711,900,870,895]" pageId="25" pageNumber="536" pagination="617 - 628" refId="ref14488" refString="Arias, JS, Szumik, CA, Goloboff, PA. 2011. Spatial analysis of vicariance: a method for using direct geographical information in historical biogeography. Cladistics. 27: 617-628. doi:10.1111/j.1096- 0031.2011.00353. x" type="journal article" year="2011">
Arias
<emphasis id="B94EEAE7FFB39A1C182DFC810D23FCF0" box="[778,835,870,895]" italics="true" pageId="25" pageNumber="536">et al.</emphasis>
2011
</bibRefCitation>
;
<bibRefCitation id="EFAB4B04FFB39A1C18B3FC810A16FCF0" author="Kreft H &amp; Jetz W" box="[916,1142,870,895]" pageId="25" pageNumber="536" pagination="2029 - 2053" refId="ref16586" refString="Kreft H, Jetz W. 2010. A framework for delineating biogeographical regions based on species distributions. J Biogeograph. 37 (11): 2029-2053. doi:10.1111/j.1365-2699.2010.02375.x." type="journal article" year="2010">Kreft and Jetz 2010</bibRefCitation>
). Therefore, it is necessary to collect data and specimens from areas with an obvious lack of information, which may enable more precise identification of possible events and patterns they have generated (
<bibRefCitation id="EFAB4B04FFB39A1C1AF2FC290CCEFC68" author="Silva M &amp; Noll FB" box="[469,686,974,999]" pageId="25" pageNumber="536" pagination="1 - 10" refId="ref18157" refString="Silva M, Noll FB. 2014. Biogeography of the social wasp genus Brachygastra (Hymenoptera: Vespidade: Polistinae). J Biogeograph. 18 (141): 1-10." type="journal article" year="2014">Silva and Noll 2014</bibRefCitation>
). In the case of
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<emphasis id="B94EEAE7FFB39A1C1878FC290DD2FC68" box="[863,946,974,999]" italics="true" pageId="25" pageNumber="536">Collaria</emphasis>
</taxonomicName>
, the most obvious gaps are found in Africa (
<figureCitation id="13012A70FFB39A1C1AE7FC160C47FB85" box="[448,551,1009,1034]" captionStart="Figure 8" captionStartId="19.[160,225,1329,1352]" captionTargetBox="[164,1151,143,1296]" captionTargetId="figure-39@19.[163,1153,142,1298]" captionTargetPageId="19" captionText="Figure 8. Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 17. Symbols at the nodes of the phylogeny are as follows: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares." figureDoi="http://doi.org/10.5281/zenodo.14970752" httpUri="https://zenodo.org/record/14970752/files/figure.png" pageId="25" pageNumber="536">Figures 8</figureCitation>
and
<figureCitation id="13012A70FFB39A1C1946FC160C0FFB85" box="[609,623,1009,1034]" captionStart="Figure 9" captionStartId="20.[160,225,1356,1379]" captionTargetBox="[164,1151,143,1322]" captionTargetId="figure-54@20.[163,1153,142,1324]" captionTargetPageId="20" captionText="Figure 9. Optimal reconstructions found for the geographical data and the phylogenetic hypothesis, nodes 815. Symbols at the nodes of the phylogeny are: black square, vicariance; white square, sympatry; white circle, point sympatry; and white triangle, founder event. In the maps, the ancestral range is drawn in green; the vicariance is drawn in red and blue squares to indicate the disjunction; regarding the founder event and point sympatry, the founder and point descendants are indicated by white squares." figureDoi="http://doi.org/10.5281/zenodo.14970754" httpUri="https://zenodo.org/record/14970754/files/figure.png" pageId="25" pageNumber="536">9</figureCitation>
). Despite these shortcomings, we were able to propose biogeographic hypotheses within
<taxonomicName id="4C3A4D76FFB39A1C19B7FBF30C83FBA2" box="[656,739,1044,1069]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="25" pageNumber="536" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFB39A1C19B7FBF30C83FBA2" box="[656,739,1044,1069]" italics="true" pageId="25" pageNumber="536">Collaria</emphasis>
</taxonomicName>
, which should be tested with additional data (eg molecular). New data, and refined phylogenetic hypotheses, could help date the nodes of the phylogeny, and thus help us test the critical biogeographical events identified here that shaped the evolution of
<taxonomicName id="4C3A4D76FFB39A1C19B1FB9B0C89FB1A" box="[662,745,1148,1173]" class="Insecta" family="Miridae" genus="Collaria" kingdom="Animalia" order="Hemiptera" pageId="25" pageNumber="536" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAE7FFB39A1C19B1FB9B0C89FB1A" box="[662,745,1148,1173]" italics="true" pageId="25" pageNumber="536">Collaria</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>

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@ -1,171 +1,173 @@
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