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<mods:title id="3BEFA816E11A87D16080F6E111D3219A">A taxonomic revision and a review of the biology of the North American seed-harvester ant genus Veromessor (Hymenoptera: Formicidae: Myrmicinae)</mods:title>
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<paragraph id="8B8536EBFF87FFAD7FC5FE1E5DD7FE01" blockId="98.[151,425,402,429]" box="[151,425,402,429]" pageId="98" pageNumber="99">
<emphasis id="B94EEAF9FF87FFAD7FC5FE1E5DD7FE01" bold="true" box="[151,425,402,429]" pageId="98" pageNumber="99">
Biology of 
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<emphasis id="B94EEAF9FF87FFAD7E4FFE1F5DD7FE01" bold="true" box="[285,425,403,429]" italics="true" pageId="98" pageNumber="99">Veromessor</emphasis>
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<paragraph id="8B8536EBFF87FFAD7FC5FE57587AFCF1" blockId="98.[151,1437,475,861]" pageId="98" pageNumber="99">
<taxonomicName id="4C3A4D68FF87FFAD7FC5FE575D67FE58" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[151,281,475,500]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF87FFAD7FC5FE575D67FE58" box="[151,281,475,500]" italics="true" pageId="98" pageNumber="99">Veromessor</emphasis>
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is an interesting genus because it contains relatively few species, yet notable interspecific variation occurs in several morphological and ecological traits. Morphological variation occurs in degree of psammophore development, degree of worker polymorphism and worker body size, size of the propodeal spines, and eye size and structure; ecological variation occurs in colony size and associated foraging method, seasonality of mating flights, and queen size and colony founding strategy. We also discuss mating frequency given that queens of 
<taxonomicName id="4C3A4D68FF87FFAD7A5DFDE759E3FD29" box="[1295,1437,619,645]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF87FFAD7A5DFDE759E3FD29" box="[1295,1437,619,645]" italics="true" pageId="98" pageNumber="99">V. pergandei</emphasis>
</taxonomicName>
mate with multiple males. Interspecific variation in each character is discussed below, while a companion paper maps these traits onto a molecular phylogeny (M.L. Borowiec, unpub. data). Moreover, species of 
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<emphasis id="B94EEAF9FF87FFAD7BA3FD3F590DFD60" box="[1265,1395,691,716]" italics="true" pageId="98" pageNumber="99">Veromessor</emphasis>
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are used in ecological, sociobiological, and behavioral studies such that it is important to understand if trait evolution results from convergent evolution or common ancestry. Distribution patterns also are discussed because some species display parapatric distributions, while other species are broadly sympatric but may or may not occur at the same locales or in the same habitats (see also 
<bibRefCitation id="EFAB4B1AFF87FFAD7DCAFCCF5F47FCF1" author="Bennett, M." box="[664,825,835,861]" pageId="98" pageNumber="99" refId="ref66035" refString="Bennett, M. (2000) Systematics of the Nearctic granivore ant genus Veromessor. PhD dissertation, University of California, Davis, California, 114 pp." type="book" year="2000">Bennett, 2000</bibRefCitation>
; 
<bibRefCitation id="EFAB4B1AFF87FFAD7C16FCCF5F86FCF1" author="Johnson, R. A." box="[836,1016,835,861]" pageId="98" pageNumber="99" pagination="89 - 122" refId="ref69343" refString="Johnson, R. A. (2000 b) Seed-harvester ants (Hymenoptera: Formicidae) of North America: an overview of ecology and biogeography. Sociobiology, 36 (1), 89 - 122." type="journal article" year="2000">Johnson, 2000b</bibRefCitation>
).
</paragraph>
<paragraph id="8B8536EBFF87FFAD7FC5FC235D8DFC65" blockId="98.[151,499,943,969]" box="[151,499,943,969]" pageId="98" pageNumber="99">
<emphasis id="B94EEAF9FF87FFAD7FC5FC235D8DFC65" bold="true" box="[151,499,943,969]" pageId="98" pageNumber="99">Psammophore development</emphasis>
</paragraph>
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The psammophore consists of numerous long hairs that extend from the lateroventral margin of the head. The function of these hairs is unknown, but it has been suggested that they are used to carry sand or soil (
<bibRefCitation id="EFAB4B1AFF87FFAD7BEAFB975CACFBF5" author="Porter, S. D. &amp; Jorgensen, C. D." pageId="98" pageNumber="99" pagination="138 - 149" refId="ref71655" refString="Porter, S. D. &amp; Jorgensen, C. D. (1990) Psammophores: do harvester ants (Hymenoptera: Formicidae) use these pouches to transport seeds? Journal of the Kansas Entomological Society, 63 (1), 138 - 149." type="journal article" year="1990">Porter &amp; Jorgensen, 1990</bibRefCitation>
; 
<bibRefCitation id="EFAB4B1AFF87FFAD7FB3FBB35E38FBF5" author="Spangler, H. G. &amp; Rettenmeyer, C. W." box="[225,582,1087,1113]" pageId="98" pageNumber="99" pagination="739 - 745" refId="ref72817" refString="Spangler, H. G. &amp; Rettenmeyer, C. W. (1966) The function of the ammochaetae or psammophores of harvester ants, Pogonomyrmex spp. Journal of the Kansas Entomological Society, 39 (4), 739 - 745." type="journal article" year="1966">Spangler &amp; Rettenmeyer, 1966</bibRefCitation>
; 
<bibRefCitation id="EFAB4B1AFF87FFAD7D06FBB35E80FBF5" author="Wheeler, W. M." box="[596,766,1087,1113]" pageId="98" pageNumber="99" pagination="185 - 202" refId="ref73735" refString="Wheeler, W. M. (1907) On certain modified hairs peculiar to the ants of arid regions. Biological Bulletin, 13 (4), 185 - 202. https: // doi. org / 10.2307 / 1535694" type="journal article" year="1907">Wheeler, 1907</bibRefCitation>
). Psammophores can also be used to carry seeds. 
<bibRefCitation id="EFAB4B1AFF87FFAD7A13FBB35C9CFBD1" author="Johnson, R. A." pageId="98" pageNumber="99" pagination="1408 - 1419" refId="ref69142" refString="Johnson, R. A. (1991) Learning, memory, and foraging efficiency in two species of desert seed-harvester ants. Ecology, 72 (4), 1408 - 1419. https: // doi. org / 10.2307 / 1941112" type="journal article" year="1991">Johnson (1991)</bibRefCitation>
observed 
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<emphasis id="B94EEAF9FF87FFAD7E07FBEF5E1BFBD0" box="[341,613,1123,1148]" italics="true" pageId="98" pageNumber="99">Pogonomyrmex rugosus</emphasis>
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workers carry multiple seeds in their psammophore when presented with a patch of novel seeds, whereas 
<taxonomicName id="4C3A4D68FF87FFAD7EA8FB0B5EFBFB0D" box="[506,645,1159,1185]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF87FFAD7EA8FB0B5EFBFB0D" box="[506,645,1159,1185]" italics="true" pageId="98" pageNumber="99">V. pergandei</emphasis>
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did not use their psammophore to carry multiple seeds. Psammophores are rare in ants, but they are common in species that inhabit arid areas, with up to one-third of the species in some desert areas possessing psammophores (
<bibRefCitation id="EFAB4B1AFF87FFAD7D0BFB435E80FB45" author="Bernard, F." box="[601,766,1231,1257]" pageId="98" pageNumber="99" pagination="87 - 200" refId="ref66065" refString="Bernard, F. (1948) Les insectes sociaux du Fezzan. Comportement et biogeographie. In: Bernard, F. &amp; de Peyerimhoff, P. (Eds.), Mission scientifique du Fezzan (1944 - 1945). Tome V. Zoologie. Institut de Recherches Sahariennes de l'Universite d'Alger, Alger, pp. 87 - 200." type="book chapter" year="1948">Bernard, 1948</bibRefCitation>
).
</paragraph>
<paragraph id="8B8536EBFF87FFAD7F95FB7F5815F9A5" blockId="98.[151,1437,1015,1905]" pageId="98" pageNumber="99">
The psammophore ranges from poorly to well developed across species of 
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<emphasis id="B94EEAF9FF87FFAD7B69FB7F58C3FAA0" box="[1083,1213,1267,1292]" italics="true" pageId="98" pageNumber="99">Veromessor</emphasis>
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, with development diagnosed by number and distribution of J-shaped hairs on the ventral surface of the head capsule (=hypostomal region). As discussed by 
<bibRefCitation id="EFAB4B1AFF87FFAD7E91FAB75E0DFAF9" author="Bennett, M." box="[451,627,1339,1365]" pageId="98" pageNumber="99" refId="ref66035" refString="Bennett, M. (2000) Systematics of the Nearctic granivore ant genus Veromessor. PhD dissertation, University of California, Davis, California, 114 pp." type="book" year="2000">Bennett (2000)</bibRefCitation>
, the psammophore in species of 
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<emphasis id="B94EEAF9FF87FFAD7CAEFAB75800FAF8" box="[1020,1150,1339,1364]" italics="true" pageId="98" pageNumber="99">Veromessor</emphasis>
</taxonomicName>
can be placed into four categories of increasing development: (1) J-shaped hairs mostly absent, with scattered straight or evenly curved hairs, (2) J-shaped hairs present but not arranged in a distinct row, usually mixed with straight or evenly curved hairs, (3) J-shaped hairs arranged in a V-shaped row which does not reach the posterior part of the lateroventral margin of the head capsule, usually mixed with straight or evenly curved hairs, and (4) J-shaped hairs arranged in a distinct row around the lateroventral margin of the the head capsule (
<tableCitation id="C6B80350FF87FFAD7CC8FA635F91F9A5" box="[922,1007,1519,1545]" captionStart="TABLE 1" captionStartId="99.[151,244,643,669]" captionTargetPageId="99" captionText="TABLE 1. Summary of morphological and ecological traits for species of Veromessor." httpUri="http://table.plazi.org/id/DF456663FF86FFAC7FC5FD0F5847FD31" pageId="98" pageNumber="99" tableUuid="DF456663FF86FFAC7FC5FD0F5847FD31">Table 1</tableCitation>
, 
<figureCitation id="13012A6EFF87FFAD7CAFFA635823F9A5" box="[1021,1117,1519,1545]" captionStart="FIGURE 6" captionStartId="21.[151,250,1642,1666]" captionTargetBox="[151,1436,181,1618]" captionTargetId="figure-18@21.[151,1436,181,1618]" captionTargetPageId="21" captionText="FIGURE 6. Photographs of the four categories of increasing psammophore development for species of Veromessor based on number and distribution of long J-shaped hairs on the ventral surface of the head capsule (= hypostomal region) (see text). Photograph of: (A) V. chicoensis (CASENT0923125)—J-shaped hairs mostly absent with scattered straight or evenly curved hairs, (B) V. andrei (CASENT0923140)—J-shaped hairs present but not arranged in a distinct row, usually mixed with straight or evenly curved hairs, (C) V. chamberlini (CASENT0761101) and (D) V. smithi (CASENT0923131)—J-shaped hairs present, arranged in a V-shaped row which does not reach the posterior part of the lateroventral margin of head capsule, usually mixed with straight or evenly curved hairs, and (E) V. pergandei (CASENT0923124)—J-shaped hairs present, many long J-shaped hairs arranged in a distinct row around the outer margin of the ventral region of the head capsule. Photographs by Wade Lee from www.AntWeb.org." figureDoi="http://doi.org/10.5281/zenodo.7318842" httpUri="https://zenodo.org/record/7318842/files/figure.png" pageId="98" pageNumber="99">Figure 6</figureCitation>
).
</paragraph>
<paragraph id="8B8536EBFF87FFAD7F95F99F5865F8DD" blockId="98.[151,1437,1015,1905]" pageId="98" pageNumber="99">
Across species of 
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<emphasis id="B94EEAF9FF87FFAD7EC1F99F5E6BF980" box="[403,533,1555,1580]" italics="true" pageId="98" pageNumber="99">Veromessor</emphasis>
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, psammophore development loosely corresponds with habitat. The three species with the most poorly developed psammophores (
<taxonomicName id="4C3A4D68FF87FFAD7D9AF9BB5F50F9FD" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[712,814,1591,1617]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF87FFAD7D9AF9BB5F50F9FD" box="[712,814,1591,1617]" italics="true" pageId="98" pageNumber="99">V. andrei</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF87FFAD7C6CF9BB5FAFF9FD" authorityName="Smith" authorityYear="1956" box="[830,977,1591,1617]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF87FFAD7C6CF9BB5FAFF9FD" box="[830,977,1591,1617]" italics="true" pageId="98" pageNumber="99">V. chicoensis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF87FFAD7CB3F9BB5817F9FD" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[993,1129,1591,1617]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF87FFAD7CB3F9BB5817F9FD" box="[993,1129,1591,1617]" italics="true" pageId="98" pageNumber="99">V. stoddardi</emphasis>
</taxonomicName>
) are restricted to the more mesic 
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coastal ranges, the 
<collectingRegion id="49FEF809FF87FFAD7D78F9D75EE2F9D9" box="[554,668,1627,1653]" country="United States of America" name="California" pageId="98" pageNumber="99">California</collectingRegion>
Central Valley, and Sierra 
<collectingRegion id="49FEF809FF87FFAD7C94F9D75862F9D9" box="[966,1052,1627,1653]" country="United States of America" name="Nevada" pageId="98" pageNumber="99">Nevada</collectingRegion>
foothills. 
<taxonomicName id="4C3A4D68FF87FFAD7BDFF9D759E2F9D9" box="[1165,1436,1627,1653]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="chamberlini">
<emphasis id="B94EEAF9FF87FFAD7BDFF9D759E2F9D9" box="[1165,1436,1627,1653]" italics="true" pageId="98" pageNumber="99">Veromessor chamberlini</emphasis>
</taxonomicName>
has a moderately well developed psammophore, and it is also restricted to more mesic southern 
<collectingRegion id="49FEF809FF87FFAD7B87F9F35939F935" box="[1237,1351,1663,1689]" country="United States of America" name="California" pageId="98" pageNumber="99">California</collectingRegion>
coastal habitats. The two other species with a moderately well developed psammophore (
<taxonomicName id="4C3A4D68FF87FFAD7B18F92F588CF911" baseAuthorityName="Andrews" baseAuthorityYear="1916" box="[1098,1266,1699,1725]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="lobognathus">
<emphasis id="B94EEAF9FF87FFAD7B18F92F588CF911" box="[1098,1266,1699,1725]" italics="true" pageId="98" pageNumber="99">V. lobognathus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF87FFAD7A56F92F5918F911" authorityName="Cole" authorityYear="1963" box="[1284,1382,1699,1725]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF87FFAD7A56F92F5918F911" box="[1284,1382,1699,1725]" italics="true" pageId="98" pageNumber="99">V. smithi</emphasis>
</taxonomicName>
) are widespread and inhabit cooler, higher elevation areas such as pinyon pine-juniper habitats. Two species with well developed psammophores (
<taxonomicName id="4C3A4D68FF87FFAD7E83F9675E34F8A9" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[465,586,1771,1797]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF87FFAD7E83F9675E34F8A9" box="[465,586,1771,1797]" italics="true" pageId="98" pageNumber="99">V. julianus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF87FFAD7D09F9675E95F8A9" box="[603,747,1771,1797]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF87FFAD7D09F9675E95F8A9" box="[603,747,1771,1797]" italics="true" pageId="98" pageNumber="99">V. pergandei</emphasis>
</taxonomicName>
) inhabit low elevation, hot desert environs, while the other two species (
<taxonomicName id="4C3A4D68FF87FFAD7E7EF8835DD6F885" box="[300,424,1807,1833]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF87FFAD7E7EF8835DD6F885" box="[300,424,1807,1833]" italics="true" pageId="98" pageNumber="99">V. lariversi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF87FFAD7EEAF8835EFCF885" box="[440,642,1807,1833]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="98" pageNumber="99" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF87FFAD7EEAF8835EFCF885" box="[440,642,1807,1833]" italics="true" pageId="98" pageNumber="99">V. pseudolariversi</emphasis>
</taxonomicName>
) inhabit both low elevation, hot desert and mid-elevation, cool desert environs; all four species often occur in sandy habitats (see also 
<bibRefCitation id="EFAB4B1AFF87FFAD7C04F8BF5F82F8E1" author="Wheeler, W. M." box="[854,1020,1843,1869]" pageId="98" pageNumber="99" pagination="185 - 202" refId="ref73735" refString="Wheeler, W. M. (1907) On certain modified hairs peculiar to the ants of arid regions. Biological Bulletin, 13 (4), 185 - 202. https: // doi. org / 10.2307 / 1535694" type="journal article" year="1907">Wheeler, 1907</bibRefCitation>
). This interspecific variation suggests that degree of psammophore development is subject to environmental pressures.
</paragraph>
<caption id="DF456663FF86FFAC7FC5FD0F5847FD31" ID-Table-UUID="DF456663FF86FFAC7FC5FD0F5847FD31" box="[151,1081,643,669]" httpUri="http://table.plazi.org/id/DF456663FF86FFAC7FC5FD0F5847FD31" pageId="99" pageNumber="100" startId="99.[151,244,643,669]" targetBox="[165,2026,682,1296]" targetIsTable="true" targetPageId="99">
<paragraph id="8B8536EBFF86FFAC7FC5FD0F5847FD31" blockId="99.[151,1081,643,669]" box="[151,1081,643,669]" pageId="99" pageNumber="100">
<emphasis id="B94EEAF9FF86FFAC7FC5FD0F5D70FD31" bold="true" box="[151,270,643,669]" pageId="99" pageNumber="100">TABLE 1.</emphasis>
Summary of morphological and ecological traits for species of 
<taxonomicName id="4C3A4D68FF86FFAC7CE4FD0F584FFD31" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[950,1073,643,669]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF86FFAC7CE4FD0F584FFD31" box="[950,1073,643,669]" italics="true" pageId="99" pageNumber="100">Veromessor</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8B8536EBFF86FFAC7FF7FD205A9AFB40" pageId="99" pageNumber="100">
<table id="F93AC44BFF8600307FF7FD265B94FABC" box="[165,2026,682,1296]" gridcols="7" gridrows="12" pageId="99" pageNumber="100">
<tr id="350A34A9FF8600307FF7FD265B94FD45" box="[165,2026,682,745]" gridrow="0" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FD265D31FD45" box="[165,335,682,745]" gridcol="0" gridrow="0" pageId="99" pageNumber="100">Species&amp;</th>
<th id="76DB5DD5FF8600307EFFFD265E37FD45" box="[429,585,682,745]" gridcol="1" gridrow="0" pageId="99" pageNumber="100">
<emphasis id="B94EEAF9FF86FFAC7EFFFD205E3AFD45" bold="true" box="[429,585,684,745]" pageId="99" pageNumber="100">Psammophore development*</emphasis>
</th>
<th id="76DB5DD5FF8600307DFDFD265FD1FD45" box="[687,943,682,745]" gridcol="2" gridrow="0" pageId="99" pageNumber="100">
<emphasis id="B94EEAF9FF86FFAC7DFDFD205FD9FD69" bold="true" box="[687,935,684,709]" pageId="99" pageNumber="100">Propodeal spine length</emphasis>
<emphasis id="B94EEAF9FF86FFAC7CF5FD265FD1FD15" bold="true" box="[935,943,682,697]" pageId="99" pageNumber="100">+</emphasis>
</th>
<th id="76DB5DD5FF8600307C81FD2658DAFD45" box="[979,1188,682,745]" gridcol="3" gridrow="0" pageId="99" pageNumber="100">
<emphasis id="B94EEAF9FF86FFAC7C81FD20580FFD45" bold="true" pageId="99" pageNumber="100">Degree of worker polymorphism</emphasis>
</th>
<th id="76DB5DD5FF8600307B87FD2659C4FD45" box="[1237,1466,682,745]" gridcol="4" gridrow="0" pageId="99" pageNumber="100">
<emphasis id="B94EEAF9FF86FFAC7B87FD205952FD45" bold="true" pageId="99" pageNumber="100">Colony size; foraging strategy</emphasis>
</th>
<th id="76DB5DD5FF8600307A85FD265AEAFD45" box="[1495,1684,682,745]" gridcol="5" gridrow="0" pageId="99" pageNumber="100">
<emphasis id="B94EEAF9FF86FFAC7A85FD205A65FD45" bold="true" pageId="99" pageNumber="100">Timing of mating flights</emphasis>
</th>
<th id="76DB5DD5FF860030798BFD265B94FD45" box="[1753,2026,682,745]" gridcol="6" gridrow="0" pageId="99" pageNumber="100">
<emphasis id="B94EEAF9FF86FFAC798BFD205B94FD69" bold="true" box="[1753,2026,684,709]" pageId="99" pageNumber="100">Colony founding strategy</emphasis>
</th>
</tr>
<tr id="350A34A9FF8600307FF7FD745B94FC99" box="[165,2026,760,821]" gridrow="1" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FD745D31FC99" box="[165,335,760,821]" gridcol="0" gridrow="1" pageId="99" pageNumber="100">
<taxonomicName id="4C3A4D68FF86FFAC7FF7FD745C98FCBD" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[165,230,760,785]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF86FFAC7FF7FD745C98FCBD" box="[165,230,760,785]" italics="true" pageId="99" pageNumber="100">andrei</emphasis>
</taxonomicName>
</th>
<td id="76DB5DD5FF8600307EFFFD745E37FC99" box="[429,585,760,821]" gridcol="1" gridrow="1" pageId="99" pageNumber="100">2</td>
<td id="76DB5DD5FF8600307DFDFD745FD1FC99" box="[687,943,760,821]" gridcol="2" gridrow="1" pageId="99" pageNumber="100">&gt; 3.0</td>
<td id="76DB5DD5FF8600307C81FD7458DAFC99" box="[979,1188,760,821]" gridcol="3" gridrow="1" pageId="99" pageNumber="100">monomorphic</td>
<td id="76DB5DD5FF8600307B87FD7459C4FC99" box="[1237,1466,760,821]" gridcol="4" gridrow="1" pageId="99" pageNumber="100">large; column</td>
<td id="76DB5DD5FF8600307A85FD745AEAFC99" box="[1495,1684,760,821]" gridcol="5" gridrow="1" pageId="99" pageNumber="100">summer</td>
<td id="76DB5DD5FF860030798BFD745B94FC99" box="[1753,2026,760,821]" gridcol="6" gridrow="1" pageId="99" pageNumber="100">semi-claustral; obligate forager</td>
</tr>
<tr id="350A34A9FF8600307FF7FCC85B94FCF1" box="[165,2026,836,861]" gridrow="2" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FCC85D31FCF1" box="[165,335,836,861]" gridcol="0" gridrow="2" pageId="99" pageNumber="100">
<taxonomicName id="4C3A4D68FF86FFAC7FF7FCC85D61FCF1" box="[165,287,836,861]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="chamberlini">
<emphasis id="B94EEAF9FF86FFAC7FF7FCC85D61FCF1" box="[165,287,836,861]" italics="true" pageId="99" pageNumber="100">chamberlini</emphasis>
</taxonomicName>
</th>
<td id="76DB5DD5FF8600307EFFFCC85E37FCF1" box="[429,585,836,861]" gridcol="1" gridrow="2" pageId="99" pageNumber="100">3</td>
<td id="76DB5DD5FF8600307DFDFCC85FD1FCF1" box="[687,943,836,861]" gridcol="2" gridrow="2" pageId="99" pageNumber="100">&gt; 3.0</td>
<td id="76DB5DD5FF8600307C81FCC858DAFCF1" box="[979,1188,836,861]" gridcol="3" gridrow="2" pageId="99" pageNumber="100">monomorphic</td>
<td id="76DB5DD5FF8600307B87FCC859C4FCF1" box="[1237,1466,836,861]" gridcol="4" gridrow="2" pageId="99" pageNumber="100">small; solitary</td>
<td id="76DB5DD5FF8600307A85FCC85AEAFCF1" box="[1495,1684,836,861]" gridcol="5" gridrow="2" pageId="99" pageNumber="100">summer</td>
<td id="76DB5DD5FF860030798BFCC85B94FCF1" box="[1753,2026,836,861]" gridcol="6" gridrow="2" pageId="99" pageNumber="100">?</td>
</tr>
<tr id="350A34A9FF8600307FF7FCE15B94FC06" box="[165,2026,877,938]" gridrow="3" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FCE15D31FC06" box="[165,335,877,938]" gridcol="0" gridrow="3" pageId="99" pageNumber="100">
<taxonomicName id="4C3A4D68FF86FFAC7FF7FCE15D71FC2A" authorityName="Smith" authorityYear="1956" box="[165,271,877,902]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF86FFAC7FF7FCE15D71FC2A" box="[165,271,877,902]" italics="true" pageId="99" pageNumber="100">chicoensis</emphasis>
</taxonomicName>
</th>
<td id="76DB5DD5FF8600307EFFFCE15E37FC06" box="[429,585,877,938]" gridcol="1" gridrow="3" pageId="99" pageNumber="100">1</td>
<td id="76DB5DD5FF8600307DFDFCE15FD1FC06" box="[687,943,877,938]" gridcol="2" gridrow="3" pageId="99" pageNumber="100">&lt;1</td>
<td id="76DB5DD5FF8600307C81FCE158DAFC06" box="[979,1188,877,938]" gridcol="3" gridrow="3" pageId="99" pageNumber="100">polymorphic</td>
<td id="76DB5DD5FF8600307B87FCE159C4FC06" box="[1237,1466,877,938]" gridcol="4" gridrow="3" pageId="99" pageNumber="100">small; solitary or with short column</td>
<td id="76DB5DD5FF8600307A85FCE15AEAFC06" box="[1495,1684,877,938]" gridcol="5" gridrow="3" pageId="99" pageNumber="100">summer</td>
<td id="76DB5DD5FF860030798BFCE15B94FC06" box="[1753,2026,877,938]" gridcol="6" gridrow="3" pageId="99" pageNumber="100">semi-claustral; obligate forager?</td>
</tr>
<tr id="350A34A9FF8600307FF7FC355B94FC7E" box="[165,2026,953,978]" gridrow="4" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FC355D31FC7E" box="[165,335,953,978]" gridcol="0" gridrow="4" pageId="99" pageNumber="100">
<taxonomicName id="4C3A4D68FF86FFAC7FF7FC355C89FC7E" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[165,247,953,978]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF86FFAC7FF7FC355C89FC7E" box="[165,247,953,978]" italics="true" pageId="99" pageNumber="100">julianus</emphasis>
</taxonomicName>
</th>
<td id="76DB5DD5FF8600307EFFFC355E37FC7E" box="[429,585,953,978]" gridcol="1" gridrow="4" pageId="99" pageNumber="100">4</td>
<td id="76DB5DD5FF8600307DFDFC355FD1FC7E" box="[687,943,953,978]" gridcol="2" gridrow="4" pageId="99" pageNumber="100">&gt; 2</td>
<td id="76DB5DD5FF8600307C81FC3558DAFC7E" box="[979,1188,953,978]" gridcol="3" gridrow="4" pageId="99" pageNumber="100">weakly polymorphic</td>
<td id="76DB5DD5FF8600307B87FC3559C4FC7E" box="[1237,1466,953,978]" gridcol="4" gridrow="4" pageId="99" pageNumber="100">large; column</td>
<td id="76DB5DD5FF8600307A85FC355AEAFC7E" box="[1495,1684,953,978]" gridcol="5" gridrow="4" pageId="99" pageNumber="100">late winter/spring</td>
<td id="76DB5DD5FF860030798BFC355B94FC7E" box="[1753,2026,953,978]" gridcol="6" gridrow="4" pageId="99" pageNumber="100">?</td>
</tr>
<tr id="350A34A9FF8600307FF7FC6D5B94FC56" box="[165,2026,993,1018]" gridrow="5" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FC6D5D31FC56" box="[165,335,993,1018]" gridcol="0" gridrow="5" pageId="99" pageNumber="100">
<taxonomicName id="4C3A4D68FF86FFAC7FF7FC6D5C85FC56" box="[165,251,993,1018]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF86FFAC7FF7FC6D5C85FC56" box="[165,251,993,1018]" italics="true" pageId="99" pageNumber="100">lariversi</emphasis>
</taxonomicName>
</th>
<td id="76DB5DD5FF8600307EFFFC6D5E37FC56" box="[429,585,993,1018]" gridcol="1" gridrow="5" pageId="99" pageNumber="100">4</td>
<td id="76DB5DD5FF8600307DFDFC6D5FD1FC56" box="[687,943,993,1018]" gridcol="2" gridrow="5" pageId="99" pageNumber="100">&lt;1</td>
<td id="76DB5DD5FF8600307C81FC6D58DAFC56" box="[979,1188,993,1018]" gridcol="3" gridrow="5" pageId="99" pageNumber="100">monomorphic</td>
<td id="76DB5DD5FF8600307B87FC6D59C4FC56" box="[1237,1466,993,1018]" gridcol="4" gridrow="5" pageId="99" pageNumber="100">small; solitary</td>
<td id="76DB5DD5FF8600307A85FC6D5AEAFC56" box="[1495,1684,993,1018]" gridcol="5" gridrow="5" pageId="99" pageNumber="100">summer</td>
<td id="76DB5DD5FF860030798BFC6D5B94FC56" box="[1753,2026,993,1018]" gridcol="6" gridrow="5" pageId="99" pageNumber="100">?</td>
</tr>
<tr id="350A34A9FF8600307FF7FB855B94FB8E" box="[165,2026,1033,1058]" gridrow="6" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FB855D31FB8E" box="[165,335,1033,1058]" gridcol="0" gridrow="6" pageId="99" pageNumber="100">
<taxonomicName id="4C3A4D68FF86FFAC7FF7FB855D5CFB8E" baseAuthorityName="Andrews" baseAuthorityYear="1916" box="[165,290,1033,1058]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="lobognathus">
<emphasis id="B94EEAF9FF86FFAC7FF7FB855D5CFB8E" box="[165,290,1033,1058]" italics="true" pageId="99" pageNumber="100">lobognathus</emphasis>
</taxonomicName>
</th>
<td id="76DB5DD5FF8600307EFFFB855E37FB8E" box="[429,585,1033,1058]" gridcol="1" gridrow="6" pageId="99" pageNumber="100">3</td>
<td id="76DB5DD5FF8600307DFDFB855FD1FB8E" box="[687,943,1033,1058]" gridcol="2" gridrow="6" pageId="99" pageNumber="100">1-1.5</td>
<td id="76DB5DD5FF8600307C81FB8558DAFB8E" box="[979,1188,1033,1058]" gridcol="3" gridrow="6" pageId="99" pageNumber="100">monomorphic</td>
<td id="76DB5DD5FF8600307B87FB8559C4FB8E" box="[1237,1466,1033,1058]" gridcol="4" gridrow="6" pageId="99" pageNumber="100">small; solitary</td>
<td id="76DB5DD5FF8600307A85FB855AEAFB8E" box="[1495,1684,1033,1058]" gridcol="5" gridrow="6" pageId="99" pageNumber="100">summer</td>
<td id="76DB5DD5FF860030798BFB855B94FB8E" box="[1753,2026,1033,1058]" gridcol="6" gridrow="6" pageId="99" pageNumber="100">?</td>
</tr>
<tr id="350A34A9FF8600307FF7FBBE5B94FBE7" box="[165,2026,1074,1099]" gridrow="7" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FBBE5D31FBE7" box="[165,335,1074,1099]" gridcol="0" gridrow="7" pageId="99" pageNumber="100">
<taxonomicName id="4C3A4D68FF86FFAC7FF7FBBE5D75FBE7" box="[165,267,1074,1099]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF86FFAC7FF7FBBE5D75FBE7" box="[165,267,1074,1099]" italics="true" pageId="99" pageNumber="100">pergandei</emphasis>
</taxonomicName>
</th>
<td id="76DB5DD5FF8600307EFFFBBE5E37FBE7" box="[429,585,1074,1099]" gridcol="1" gridrow="7" pageId="99" pageNumber="100">4</td>
<td id="76DB5DD5FF8600307DFDFBBE5FD1FBE7" box="[687,943,1074,1099]" gridcol="2" gridrow="7" pageId="99" pageNumber="100">&lt;1</td>
<td id="76DB5DD5FF8600307C81FBBE58DAFBE7" box="[979,1188,1074,1099]" gridcol="3" gridrow="7" pageId="99" pageNumber="100">weakly polymorphic</td>
<td id="76DB5DD5FF8600307B87FBBE59C4FBE7" box="[1237,1466,1074,1099]" gridcol="4" gridrow="7" pageId="99" pageNumber="100">large; column</td>
<td id="76DB5DD5FF8600307A85FBBE5AEAFBE7" box="[1495,1684,1074,1099]" gridcol="5" gridrow="7" pageId="99" pageNumber="100">late winter/spring</td>
<td id="76DB5DD5FF860030798BFBBE5B94FBE7" box="[1753,2026,1074,1099]" gridcol="6" gridrow="7" pageId="99" pageNumber="100">fully claustral</td>
</tr>
<tr id="350A34A9FF8600307FF7FBD65B94FBDF" box="[165,2026,1114,1139]" gridrow="8" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FBD65D31FBDF" box="[165,335,1114,1139]" gridcol="0" gridrow="8" pageId="99" pageNumber="100">
sp. 
<taxonomicName id="4C3A4D68FF86FFAC7F9AFBD65D31FBDF" box="[200,335,1114,1139]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="pergandei">
cf. 
<emphasis id="B94EEAF9FF86FFAC7FBBFBD65D31FBDF" box="[233,335,1114,1139]" italics="true" pageId="99" pageNumber="100">pergandei</emphasis>
</taxonomicName>
</th>
<td id="76DB5DD5FF8600307EFFFBD65E37FBDF" box="[429,585,1114,1139]" gridcol="1" gridrow="8" pageId="99" pageNumber="100">?</td>
<td id="76DB5DD5FF8600307DFDFBD65FD1FBDF" box="[687,943,1114,1139]" gridcol="2" gridrow="8" pageId="99" pageNumber="100">?</td>
<td id="76DB5DD5FF8600307C81FBD658DAFBDF" box="[979,1188,1114,1139]" gridcol="3" gridrow="8" pageId="99" pageNumber="100">?</td>
<td id="76DB5DD5FF8600307B87FBD659C4FBDF" box="[1237,1466,1114,1139]" gridcol="4" gridrow="8" pageId="99" pageNumber="100">small; solitary?</td>
<td id="76DB5DD5FF8600307A85FBD65AEAFBDF" box="[1495,1684,1114,1139]" gridcol="5" gridrow="8" pageId="99" pageNumber="100">late winter/spring?</td>
<td id="76DB5DD5FF860030798BFBD65B94FBDF" box="[1753,2026,1114,1139]" gridcol="6" gridrow="8" pageId="99" pageNumber="100">?</td>
</tr>
<tr id="350A34A9FF8600307FF7FB0E5B94FB37" box="[165,2026,1154,1179]" gridrow="9" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FB0E5D31FB37" box="[165,335,1154,1179]" gridcol="0" gridrow="9" pageId="99" pageNumber="100">
<taxonomicName id="4C3A4D68FF86FFAC7FF7FB0E5D3DFB37" box="[165,323,1154,1179]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF86FFAC7FF7FB0E5D3DFB37" box="[165,323,1154,1179]" italics="true" pageId="99" pageNumber="100">pseudolariversi</emphasis>
</taxonomicName>
</th>
<td id="76DB5DD5FF8600307EFFFB0E5E37FB37" box="[429,585,1154,1179]" gridcol="1" gridrow="9" pageId="99" pageNumber="100">4</td>
<td id="76DB5DD5FF8600307DFDFB0E5FD1FB37" box="[687,943,1154,1179]" gridcol="2" gridrow="9" pageId="99" pageNumber="100">&lt;1</td>
<td id="76DB5DD5FF8600307C81FB0E58DAFB37" box="[979,1188,1154,1179]" gridcol="3" gridrow="9" pageId="99" pageNumber="100">monomorphic</td>
<td id="76DB5DD5FF8600307B87FB0E59C4FB37" box="[1237,1466,1154,1179]" gridcol="4" gridrow="9" pageId="99" pageNumber="100">small; solitary</td>
<td id="76DB5DD5FF8600307A85FB0E5AEAFB37" box="[1495,1684,1154,1179]" gridcol="5" gridrow="9" pageId="99" pageNumber="100">summer</td>
<td id="76DB5DD5FF860030798BFB0E5B94FB37" box="[1753,2026,1154,1179]" gridcol="6" gridrow="9" pageId="99" pageNumber="100">?</td>
</tr>
<tr id="350A34A9FF8600307FF7FB265B94FB6F" box="[165,2026,1194,1219]" gridrow="10" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FB265D31FB6F" box="[165,335,1194,1219]" gridcol="0" gridrow="10" pageId="99" pageNumber="100">
<taxonomicName id="4C3A4D68FF86FFAC7FF7FB265C9DFB6F" authorityName="Cole" authorityYear="1963" box="[165,227,1194,1219]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF86FFAC7FF7FB265C9DFB6F" box="[165,227,1194,1219]" italics="true" pageId="99" pageNumber="100">smithi</emphasis>
</taxonomicName>
</th>
<td id="76DB5DD5FF8600307EFFFB265E37FB6F" box="[429,585,1194,1219]" gridcol="1" gridrow="10" pageId="99" pageNumber="100">3</td>
<td id="76DB5DD5FF8600307DFDFB265FD1FB6F" box="[687,943,1194,1219]" gridcol="2" gridrow="10" pageId="99" pageNumber="100">&lt;1</td>
<td id="76DB5DD5FF8600307C81FB2658DAFB6F" box="[979,1188,1194,1219]" gridcol="3" gridrow="10" pageId="99" pageNumber="100">monomorphic</td>
<td id="76DB5DD5FF8600307B87FB2659C4FB6F" box="[1237,1466,1194,1219]" gridcol="4" gridrow="10" pageId="99" pageNumber="100">small; solitary</td>
<td id="76DB5DD5FF8600307A85FB265AEAFB6F" box="[1495,1684,1194,1219]" gridcol="5" gridrow="10" pageId="99" pageNumber="100">summer</td>
<td id="76DB5DD5FF860030798BFB265B94FB6F" box="[1753,2026,1194,1219]" gridcol="6" gridrow="10" pageId="99" pageNumber="100">?</td>
</tr>
<tr id="350A34A9FF8600307FF7FB5F5B94FABC" box="[165,2026,1235,1296]" gridrow="11" pageId="99" pageNumber="100">
<th id="76DB5DD5FF8600307FF7FB5F5D31FABC" box="[165,335,1235,1296]" gridcol="0" gridrow="11" pageId="99" pageNumber="100">
<taxonomicName id="4C3A4D68FF86FFAC7FF7FB5F5D7BFB40" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[165,261,1235,1260]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF86FFAC7FF7FB5F5D7BFB40" box="[165,261,1235,1260]" italics="true" pageId="99" pageNumber="100">stoddardi</emphasis>
</taxonomicName>
</th>
<td id="76DB5DD5FF8600307EFFFB5F5E37FABC" box="[429,585,1235,1296]" gridcol="1" gridrow="11" pageId="99" pageNumber="100">1</td>
<td id="76DB5DD5FF8600307DFDFB5F5FD1FABC" box="[687,943,1235,1296]" gridcol="2" gridrow="11" pageId="99" pageNumber="100">&lt;1</td>
<td id="76DB5DD5FF8600307C81FB5F58DAFABC" box="[979,1188,1235,1296]" gridcol="3" gridrow="11" pageId="99" pageNumber="100">polymorphic</td>
<td id="76DB5DD5FF8600307B87FB5F59C4FABC" box="[1237,1466,1235,1296]" gridcol="4" gridrow="11" pageId="99" pageNumber="100">small; solitary or with short colum</td>
<td id="76DB5DD5FF8600307A85FB5F5AEAFABC" box="[1495,1684,1235,1296]" gridcol="5" gridrow="11" pageId="99" pageNumber="100">summer</td>
<td id="76DB5DD5FF860030798BFB5F5B94FABC" box="[1753,2026,1235,1296]" gridcol="6" gridrow="11" pageId="99" pageNumber="100">?</td>
</tr>
</table>
</paragraph>
<tableNote id="76DC3765FF86FFAC7FC5FA965F8BF9A1" pageId="99" pageNumber="100" targetBox="[165,2026,682,1296]" targetPageId="99">
<paragraph id="8B8536EBFF86FFAC7FC5FA965CA9FAF5" blockId="99.[151,2041,1306,1549]" pageId="99" pageNumber="100">
<superScript id="7C4F9BA3FF86FFAC7FC5FA965CDDFA85" attach="right" box="[151,163,1306,1321]" fontSize="6" pageId="99" pageNumber="100">&amp;</superScript>
species are listed in alphabetical order, except for sp. 
<taxonomicName id="4C3A4D68FF86FFAC7DBBFA975F06FA99" box="[745,888,1307,1333]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Hymenoptera" pageId="99" pageNumber="100" phylum="Arthropoda" rank="species" species="pergandei">
cf. 
<emphasis id="B94EEAF9FF86FFAC7C5FFA975F06FA99" box="[781,888,1307,1333]" italics="true" pageId="99" pageNumber="100">pergandei</emphasis>
</taxonomicName>
, which is an undescribed species in which known specimens are presumed lost. See text under Additional notes.
</paragraph>
<paragraph id="8B8536EBFF86FFAC7FC5FAEE5BE7FA45" blockId="99.[151,2041,1306,1549]" pageId="99" pageNumber="100">
<superScript id="7C4F9BA3FF86FFAC7FC5FAEE5CE1FADD" attach="right" box="[151,159,1378,1393]" fontSize="6" pageId="99" pageNumber="100">*</superScript>
variation in psammophore development with regard to number and distribution of long J-shaped hairs on the ventral surface of the head capsule (=hypostomal regions) (see text): (1) J-shaped hairs mostly absent with scattered straight or evenly curved hairs, (2) J- shaped hairs present but not arranged in a distinct row, usually mixed with straight of evenly curved hairs, (3) J-shaped hairs present arranged in a V-shaped row which does not reach the posterior part of the lateroventral margin of head capsule, usually mixed with straight or evenly curved hairs, (4) J-shaped hairs present, many long J-shaped hairs arranged in a distinct row around the outer margin of the ventral region of the head capsule.
</paragraph>
<paragraph id="8B8536EBFF86FFAC7FC5FA7E5F8BF9A1" blockId="99.[151,2041,1306,1549]" box="[151,1013,1522,1549]" pageId="99" pageNumber="100">
<superScript id="7C4F9BA3FF86FFAC7FC5FA7E5CDEF9AD" attach="right" box="[151,160,1522,1537]" fontSize="6" pageId="99" pageNumber="100">+</superScript>
approximate length of propodeal spines relative to distance between their bases.
</paragraph>
</tableNote>
<paragraph id="8B8536EBFF81FFAB7FC5FF1B5D12FF1D" blockId="100.[151,364,151,177]" box="[151,364,151,177]" pageId="100" pageNumber="101">
<emphasis id="B94EEAF9FF81FFAB7FC5FF1B5D12FF1D" bold="true" box="[151,364,151,177]" pageId="100" pageNumber="101">Propodeal spines</emphasis>
</paragraph>
<paragraph id="8B8536EBFF81FFAB7FC5FF535D7DFE59" blockId="100.[151,1437,223,501]" pageId="100" pageNumber="101">
The function of propodeal spines is unclear, but the two primary hypotheses are that these spines function for defense or structural support (
<bibRefCitation id="EFAB4B1AFF81FFAB7EB7FE8F5F60FEB1" author="Blanchard, B. D. &amp; Moreau, C. S." box="[485,798,259,286]" pageId="100" pageNumber="101" pagination="315 - 328" refId="ref66130" refString="Blanchard, B. D. &amp; Moreau, C. S. (2017) Defensive traits exhibit an evolutionary trade-off and drive diversification in ants. Evolution, 71 (2), 315 - 328. https: // doi. org / 10.1111 / evo. 13117" type="journal article" year="2017">Blanchard &amp; Moreau, 2017</bibRefCitation>
; Sarnat, Friedman, Fischer, Lecroq-Bennet, &amp; Economo, 2017; R. Keller, pers. comm.). Interestingly, size of the propodeal spines varies greatly across the few species of 
<taxonomicName id="4C3A4D68FF81FFAB7FC5FEC05D67FEC9" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[151,281,332,357]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF81FFAB7FC5FEC05D67FEC9" box="[151,281,332,357]" italics="true" pageId="100" pageNumber="101">Veromessor</emphasis>
</taxonomicName>
. The spines are short (length less than to similar to distance between their bases) in 
<taxonomicName id="4C3A4D68FF81FFAB7BB4FEC05906FEC9" authorityName="Smith" authorityYear="1956" box="[1254,1400,331,357]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF81FFAB7BB4FEC05906FEC9" box="[1254,1400,331,357]" italics="true" pageId="100" pageNumber="101">V. chicoensis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7ADAFEC05C88FE25" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF81FFAB7ADAFEC05C88FE25" italics="true" pageId="100" pageNumber="101">V. lariversi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7E51FEFC5DF0FE25" box="[259,398,367,393]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF81FFAB7E51FEFC5DF0FE25" box="[259,398,367,393]" italics="true" pageId="100" pageNumber="101">V. pergandei</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7ECEFEFC5E1BFE25" box="[412,613,367,393]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF81FFAB7ECEFEFC5E1BFE25" box="[412,613,367,393]" italics="true" pageId="100" pageNumber="101">V. pseudolariversi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7D21FEFC5EB5FE25" box="[627,715,367,393]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="smith">
<emphasis id="B94EEAF9FF81FFAB7D21FEFC5EB5FE25" box="[627,715,367,393]" italics="true" pageId="100" pageNumber="101">V. smith</emphasis>
</taxonomicName>
, and 
<taxonomicName id="4C3A4D68FF81FFAB7C55FEFC5FF2FE25" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[775,908,367,393]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF81FFAB7C55FEFC5FF2FE25" box="[775,908,367,393]" italics="true" pageId="100" pageNumber="101">V. stoddardi</emphasis>
</taxonomicName>
, medium length (length about 1.0–1.5× as long as the distance between their bases) in 
<taxonomicName id="4C3A4D68FF81FFAB7D11FE185E96FE01" baseAuthorityName="Andrews" baseAuthorityYear="1916" box="[579,744,403,429]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="lobognathus">
<emphasis id="B94EEAF9FF81FFAB7D11FE185E96FE01" box="[579,744,403,429]" italics="true" pageId="100" pageNumber="101">V. lobognathus</emphasis>
</taxonomicName>
, long (length&gt; 2× as long as the distance between their bases) in 
<taxonomicName id="4C3A4D68FF81FFAB7FE1FE345D56FE7D" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[179,296,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF81FFAB7FE1FE345D56FE7D" box="[179,296,439,465]" italics="true" pageId="100" pageNumber="101">V. julianus</emphasis>
</taxonomicName>
, and very long (length&gt; 3× as long as the distance between their bases) in 
<taxonomicName id="4C3A4D68FF81FFAB7B31FE3458B8FE7D" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[1123,1222,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF81FFAB7B31FE3458B8FE7D" box="[1123,1222,439,465]" italics="true" pageId="100" pageNumber="101">V. andrei</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF81FFAB7BA8FE3459E2FE7D" box="[1274,1436,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="chamberlini">
<emphasis id="B94EEAF9FF81FFAB7BA8FE3459E2FE7D" box="[1274,1436,439,465]" italics="true" pageId="100" pageNumber="101">V. chamberlini</emphasis>
</taxonomicName>
(
<tableCitation id="C6B80350FF81FFAB7FCDFE575C8CFE59" box="[159,242,475,501]" captionStart="TABLE 1" captionStartId="99.[151,244,643,669]" captionTargetPageId="99" captionText="TABLE 1. Summary of morphological and ecological traits for species of Veromessor." httpUri="http://table.plazi.org/id/DF456663FF86FFAC7FC5FD0F5847FD31" pageId="100" pageNumber="101" tableUuid="DF456663FF86FFAC7FC5FD0F5847FD31">Table 1</tableCitation>
).
</paragraph>
<paragraph id="8B8536EBFF81FFAB7FC5FDCB5DD2FDCD" blockId="100.[151,428,583,609]" box="[151,428,583,609]" pageId="100" pageNumber="101">
<emphasis id="B94EEAF9FF81FFAB7FC5FDCB5DD2FDCD" bold="true" box="[151,428,583,609]" pageId="100" pageNumber="101">Eye size and structure</emphasis>
</paragraph>
<paragraph id="8B8536EBFF81FFAB7FC5FD035E3CFC95" blockId="100.[151,1437,655,1366]" pageId="100" pageNumber="101">The eyes of insects possess an incredible array of adaptations to enhance vision across the gamut of light levels that they experience. One commonly studied contrast relates to differences in eye structure between nocturnal and diurnal species, with nocturnal species typically having larger eyes and larger eye facets. Cuticular pigment level also often is correlated with level of ambient light, with species having little or no cuticular pigment typically living in dim light to lightless environments.</paragraph>
<paragraph id="8B8536EBFF81FFAB7F95FCC8593CFB99" blockId="100.[151,1437,655,1366]" pageId="100" pageNumber="101">
Eye structure and pigment level varied between nocturnal and diurnal species of 
<taxonomicName id="4C3A4D68FF81FFAB7B24FCC85886FCF1" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[1142,1272,836,861]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF81FFAB7B24FCC85886FCF1" box="[1142,1272,836,861]" italics="true" pageId="100" pageNumber="101">Veromessor</emphasis>
</taxonomicName>
, as well as in three other ant genera (
<taxonomicName id="4C3A4D68FF81FFAB7EF1FCE45E2FFC2D" authorityName="Wesmael" authorityYear="1838" box="[419,593,872,897]" class="Insecta" family="Formicidae" genus="Myrmecocystus" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF81FFAB7EF1FCE45E2FFC2D" box="[419,593,872,897]" italics="true" pageId="100" pageNumber="101">Myrmecocystus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7D30FCEB5F72FC2D" baseAuthorityName="Demarco &amp; Cognato" baseAuthorityYear="2015" box="[610,780,871,897]" class="Insecta" family="Formicidae" genus="Aphaenogaster" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF81FFAB7D30FCEB5F72FC2D" box="[610,780,871,897]" italics="true" pageId="100" pageNumber="101">Aphaenogaster</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7C49FCEB5FD0FC2D" authorityName="Mayr" authorityYear="1861" box="[795,942,871,897]" class="Insecta" family="Formicidae" genus="Temnothorax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF81FFAB7C49FCEB5FD0FC2D" box="[795,942,871,897]" italics="true" pageId="100" pageNumber="101">Temnothorax</emphasis>
</taxonomicName>
) (
<bibRefCitation id="EFAB4B1AFF81FFAB7C9AFCEB5978FC2D" author="Johnson, R. A. &amp; Rutowski, R. L." box="[968,1286,871,897]" pageId="100" pageNumber="101" pagination="0257779" refId="ref69936" refString="Johnson, R. A. &amp; Rutowski, R. L. (2022) Color, activity period, and eye structure in four lineages of ants: pale, nocturnal species have evolved larger eyes and larger facets than their dark, diurnal congeners. PLoS ONE, 17 (9), e 0257779. https: // doi. org / 10.1371 / journal. pone. 0257779" type="journal article" year="2022">Johnson &amp; Rutowski, 2022</bibRefCitation>
). 
<taxonomicName id="4C3A4D68FF81FFAB7A49FCE459E3FC2D" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[1307,1437,872,897]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF81FFAB7A49FCE459E3FC2D" box="[1307,1437,872,897]" italics="true" pageId="100" pageNumber="101">Veromessor</emphasis>
</taxonomicName>
contains two species (
<taxonomicName id="4C3A4D68FF81FFAB7EC1FC005E6CFC09" box="[403,530,907,934]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF81FFAB7EC1FC005DDAFC09" box="[403,420,908,933]" italics="true" pageId="100" pageNumber="101">V</emphasis>
. 
<emphasis id="B94EEAF9FF81FFAB7EE1FC075E6CFC09" box="[435,530,907,933]" italics="true" pageId="100" pageNumber="101">lariversi</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF81FFAB7D1EFC005F69FC09" box="[588,791,907,933]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF81FFAB7D1EFC005F69FC09" box="[588,791,907,933]" italics="true" pageId="100" pageNumber="101">V. pseudolariversi</emphasis>
</taxonomicName>
) that are pale in color, i.e., light yellowish to yellowishorange, while all other congeners are darker (
<bibRefCitation id="EFAB4B1AFF81FFAB7DC5FC235FAEFC65" author="Johnson, R. A. &amp; Rutowski, R. L." box="[663,976,943,969]" pageId="100" pageNumber="101" pagination="0257779" refId="ref69936" refString="Johnson, R. A. &amp; Rutowski, R. L. (2022) Color, activity period, and eye structure in four lineages of ants: pale, nocturnal species have evolved larger eyes and larger facets than their dark, diurnal congeners. PLoS ONE, 17 (9), e 0257779. https: // doi. org / 10.1371 / journal. pone. 0257779" type="journal article" year="2022">Johnson &amp; Rutowski, 2022</bibRefCitation>
). Differences in pigment level correlated with foraging time—pale species mostly are strict nocturnal foragers whereas dark species usually forage diurnally, or foraging activity is temperature dependent (diurnal during cool seasons, nocturnal during hottest seasons), and several species are largely matinal-crepuscular-nocturnal foragers (
<taxonomicName id="4C3A4D68FF81FFAB7CD5FB905F82FB99" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[903,1020,1051,1077]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF81FFAB7CD5FB905F82FB99" box="[903,1020,1051,1077]" italics="true" pageId="100" pageNumber="101">V. julianus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7B58FB90580EFB99" authorityName="Cole" authorityYear="1963" box="[1034,1136,1051,1078]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF81FFAB7B58FB905865FB99" box="[1034,1051,1052,1077]" italics="true" pageId="100" pageNumber="101">V</emphasis>
. 
<emphasis id="B94EEAF9FF81FFAB7B7BFB97580EFB99" box="[1065,1136,1051,1077]" italics="true" pageId="100" pageNumber="101">smithi</emphasis>
</taxonomicName>
<emphasis id="B94EEAF9FF81FFAB7B23FB97580BFB99" box="[1137,1141,1051,1077]" italics="true" pageId="100" pageNumber="101">,</emphasis>
and 
<taxonomicName id="4C3A4D68FF81FFAB7BF9FB90594FFB99" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[1195,1329,1051,1077]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF81FFAB7BF9FB90594FFB99" box="[1195,1329,1051,1077]" italics="true" pageId="100" pageNumber="101">V. stoddardi</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="8B8536EBFF81FFAB7F95FBB359E3FAF9" blockId="100.[151,1437,655,1366]" pageId="100" pageNumber="101">
As predicted, both pale species had large eyes, but they displayed a different eye structure. 
<taxonomicName id="4C3A4D68FF81FFAB7A49FBCC5D3BFBD1" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF81FFAB7A49FBCC5D3BFBD1" italics="true" pageId="100" pageNumber="101">Veromessor pseudolariversi</emphasis>
</taxonomicName>
had larger eyes and more eye facets than 
<taxonomicName id="4C3A4D68FF81FFAB7C5EFBE85FFBFBD1" box="[780,901,1123,1149]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF81FFAB7C5EFBE85FFBFBD1" box="[780,901,1123,1149]" italics="true" pageId="100" pageNumber="101">V. lariversi</emphasis>
</taxonomicName>
, but mean facet diameter was similar for the two species (
<figureCitation id="13012A6EFF81FFAB7FA6FB0B5D14FB0D" box="[244,362,1159,1185]" captionStart="FIGURE 30" captionStartId="56.[151,250,1944,1968]" captionTargetBox="[393,1199,196,1904]" captionTargetId="figure-26@56.[376,1212,181,1920]" captionTargetPageId="56" captionText="FIGURE 30. Bivariate plots for workers of V. lariversi and V. pseudolariversi: (A) mesosoma length versus number of eye facets, and (B) mesosoma length versus eye length. Sample size for each species is in parentheses." figureDoi="http://doi.org/10.5281/zenodo.7318900" httpUri="https://zenodo.org/record/7318900/files/figure.png" pageId="100" pageNumber="101">Figures 30</figureCitation>
&amp; 
<figureCitation id="13012A6EFF81FFAB7EDBFB0B5DD6FB0E" box="[393,424,1159,1186]" captionStart="FIGURE 53" captionStartId="102.[151,250,1802,1826]" captionTargetBox="[376,1218,194,1775]" captionTargetId="figure-26@102.[365,1223,181,1778]" captionTargetPageId="102" captionText="FIGURE 53. Eye area (mm2) (A), facet number (B), and mean facet diameter (μm) (C) for pale and dark colored species of Veromessor. Two species are pale (V. lariversi, V. pseudolariversi—open symbols and regular font), while the other eight species are dark (filled symbols and bold font). For each species, number of workers examined and number of colonies they were derived from is given in parentheses. Significant differences (P &lt;0.05) among species are denoted after each species name by the letters a–g: a&gt; b&gt; c&gt; d&gt; e&gt; f&gt; g; the three sets of letters for each species correspond to panels A, B, and C, respectively. Groupings are based on univariate F tests within MANCOVA followed by pairwise comparisons using a least significant differences test (see also Johnson &amp; Rutowki, 2022)." figureDoi="http://doi.org/10.5281/zenodo.7318948" httpUri="https://zenodo.org/record/7318948/files/figure.png" pageId="100" pageNumber="101">53</figureCitation>
), despite their being sister species.All three variables (eye size, facet number, facet diameter) were larger for these two species than for all other dark congeners except for 
<taxonomicName id="4C3A4D68FF81FFAB7C81FB20584EFB69" authorityName="Cole" authorityYear="1963" box="[979,1072,1195,1221]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF81FFAB7C81FB20584EFB69" box="[979,1072,1195,1221]" italics="true" pageId="100" pageNumber="101">V. smithi</emphasis>
</taxonomicName>
. Compared to 
<taxonomicName id="4C3A4D68FF81FFAB7B9CFB2059EBFB69" box="[1230,1429,1195,1221]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF81FFAB7B9CFB2059EBFB69" box="[1230,1429,1195,1221]" italics="true" pageId="100" pageNumber="101">V.pseudolariversi</emphasis>
</taxonomicName>
, the eyes of 
<taxonomicName id="4C3A4D68FF81FFAB7E4EFB5C5D03FB45" authorityName="Cole" authorityYear="1963" box="[284,381,1231,1257]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF81FFAB7E4EFB5C5D03FB45" box="[284,381,1231,1257]" italics="true" pageId="100" pageNumber="101">V. smithi</emphasis>
</taxonomicName>
were smaller and had fewer facets, but facet diameter was similar. Compared to 
<taxonomicName id="4C3A4D68FF81FFAB7A4BFB5C59EBFB45" box="[1305,1429,1231,1257]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF81FFAB7A4BFB5C59EBFB45" box="[1305,1429,1231,1257]" italics="true" pageId="100" pageNumber="101">V. lariversi</emphasis>
</taxonomicName>
, the eyes of 
<taxonomicName id="4C3A4D68FF81FFAB7E44FB785D0BFAA1" authorityName="Cole" authorityYear="1963" box="[278,373,1267,1293]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF81FFAB7E44FB785D0BFAA1" box="[278,373,1267,1293]" italics="true" pageId="100" pageNumber="101">V. smithi</emphasis>
</taxonomicName>
were larger and facet diameter was larger, but facet number was similar (
<figureCitation id="13012A6EFF81FFAB7BF6FB7F596AFAA2" box="[1188,1300,1267,1294]" captionStart="FIGURE 53" captionStartId="102.[151,250,1802,1826]" captionTargetBox="[376,1218,194,1775]" captionTargetId="figure-26@102.[365,1223,181,1778]" captionTargetPageId="102" captionText="FIGURE 53. Eye area (mm2) (A), facet number (B), and mean facet diameter (μm) (C) for pale and dark colored species of Veromessor. Two species are pale (V. lariversi, V. pseudolariversi—open symbols and regular font), while the other eight species are dark (filled symbols and bold font). For each species, number of workers examined and number of colonies they were derived from is given in parentheses. Significant differences (P &lt;0.05) among species are denoted after each species name by the letters a–g: a&gt; b&gt; c&gt; d&gt; e&gt; f&gt; g; the three sets of letters for each species correspond to panels A, B, and C, respectively. Groupings are based on univariate F tests within MANCOVA followed by pairwise comparisons using a least significant differences test (see also Johnson &amp; Rutowki, 2022)." figureDoi="http://doi.org/10.5281/zenodo.7318948" httpUri="https://zenodo.org/record/7318948/files/figure.png" pageId="100" pageNumber="101">Figure 53</figureCitation>
). Moreover, eyes of the two pale species of 
<taxonomicName id="4C3A4D68FF81FFAB7D50FA945EFAFA9D" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[514,644,1304,1329]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF81FFAB7D50FA945EFAFA9D" box="[514,644,1304,1329]" italics="true" pageId="100" pageNumber="101">Veromessor</emphasis>
</taxonomicName>
evolved to maximize sensitivity over resolution, which is the pattern typical for most nocturnal insects with apposition eyes (
<bibRefCitation id="EFAB4B1AFF81FFAB7DAEFAB75FE5FAF9" author="Greiner, B." box="[764,923,1339,1365]" pageId="100" pageNumber="101" pagination="1 - 46" refId="ref68406" refString="Greiner, B. (2006) Adaptations for nocturnal vision in insect apposition eyes. International Review of Cytology, 250, 1 - 46. https: // doi. org / 10.1016 / S 0074 - 7696 (06) 50001 - 4" type="journal article" year="2006">Greiner, 2006</bibRefCitation>
; Stöckl, Smolka, O’Carroll, &amp; Warrant, 2017).
</paragraph>
<paragraph id="8B8536EBFF81FFAB7FC5FA2B5EB4FA6D" blockId="100.[151,714,1447,1473]" box="[151,714,1447,1473]" pageId="100" pageNumber="101">
<emphasis id="B94EEAF9FF81FFAB7FC5FA2B5EB4FA6D" bold="true" box="[151,714,1447,1473]" pageId="100" pageNumber="101">Worker polymorphism and worker body size</emphasis>
</paragraph>
<paragraph id="8B8536EBFF81FFAA7FC5FA635D8DFF79" blockId="100.[151,1437,1519,2013]" lastBlockId="101.[151,1437,151,717]" lastPageId="101" lastPageNumber="102" pageId="100" pageNumber="101">
The genus 
<taxonomicName id="4C3A4D68FF81FFAB7E45FA7C5DE7F9A5" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[279,409,1520,1545]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF81FFAB7E45FA7C5DE7F9A5" box="[279,409,1520,1545]" italics="true" pageId="100" pageNumber="101">Veromessor</emphasis>
</taxonomicName>
is interesting because workers range from monomorphic to polymorphic (
<tableCitation id="C6B80350FF81FFAB7BA0FA635936F9A5" box="[1266,1352,1519,1545]" captionStart="TABLE 1" captionStartId="99.[151,244,643,669]" captionTargetPageId="99" captionText="TABLE 1. Summary of morphological and ecological traits for species of Veromessor." httpUri="http://table.plazi.org/id/DF456663FF86FFAC7FC5FD0F5847FD31" pageId="100" pageNumber="101" tableUuid="DF456663FF86FFAC7FC5FD0F5847FD31">Table 1</tableCitation>
). Most species are monomorphic (
<taxonomicName id="4C3A4D68FF81FFAB7E95F9985E54F981" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[455,554,1555,1581]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF81FFAB7E95F9985E54F981" box="[455,554,1555,1581]" italics="true" pageId="100" pageNumber="101">V. andrei</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7D6BF9985EA2F981" box="[569,732,1555,1581]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="chamberlini">
<emphasis id="B94EEAF9FF81FFAB7D6BF9985EA2F981" box="[569,732,1555,1581]" italics="true" pageId="100" pageNumber="101">V. chamberlini</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7DB8F9985F1BF981" box="[746,869,1555,1581]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF81FFAB7DB8F9985F1BF981" box="[746,869,1555,1581]" italics="true" pageId="100" pageNumber="101">V. lariversi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7C21F9985867F981" baseAuthorityName="Andrews" baseAuthorityYear="1916" box="[883,1049,1555,1581]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="lobognathus">
<emphasis id="B94EEAF9FF81FFAB7C21F9985867F981" box="[883,1049,1555,1581]" italics="true" pageId="100" pageNumber="101">V. lobognathus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7B7AF998588CF981" box="[1064,1266,1555,1581]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF81FFAB7B7AF998588CF981" box="[1064,1266,1555,1581]" italics="true" pageId="100" pageNumber="101">V. pseudolariversi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7A52F998591EF981" authorityName="Cole" authorityYear="1963" box="[1280,1376,1555,1581]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF81FFAB7A52F998591EF981" box="[1280,1376,1555,1581]" italics="true" pageId="100" pageNumber="101">V. smithi</emphasis>
</taxonomicName>
), but two species are weakly polymorphic (
<taxonomicName id="4C3A4D68FF81FFAB7D11F9B45EC7F9FD" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[579,697,1591,1617]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF81FFAB7D11F9B45EC7F9FD" box="[579,697,1591,1617]" italics="true" pageId="100" pageNumber="101">V. julianus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7D9AF9B45F2AF9FD" box="[712,852,1591,1617]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF81FFAB7D9AF9B45F2AF9FD" box="[712,852,1591,1617]" italics="true" pageId="100" pageNumber="101">V. pergandei</emphasis>
</taxonomicName>
) and two species are polymorphic (
<taxonomicName id="4C3A4D68FF81FFAB7BBBF9B45904F9FD" authorityName="Smith" authorityYear="1956" box="[1257,1402,1591,1617]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF81FFAB7BBBF9B45904F9FD" box="[1257,1402,1591,1617]" italics="true" pageId="100" pageNumber="101">V. chicoensis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF81FFAB7ADAF9B45D7FF9D9" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF81FFAB7ADAF9B45D7FF9D9" italics="true" pageId="100" pageNumber="101">V. stoddardi</emphasis>
</taxonomicName>
). All species of 
<taxonomicName id="4C3A4D68FF81FFAB7EE0F9D05E4AF9D9" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[434,564,1628,1653]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF81FFAB7EE0F9D05E4AF9D9" box="[434,564,1628,1653]" italics="true" pageId="100" pageNumber="101">Veromessor</emphasis>
</taxonomicName>
that display some degree of polymorphism show monophasic allometry, which is the most primitive form of polymorphism in which growth is nonisometric (Wills, Powell, Rivera, &amp; Suarez, 2018; 
<bibRefCitation id="EFAB4B1AFF81FFAB7FC5F92F5D4CF912" author="Wilson, E. O." box="[151,306,1699,1726]" pageId="100" pageNumber="101" pagination="136 - 156" refId="ref74082" refString="Wilson, E. O. (1953) The origin and evolution of polymorphism in ants. Quarterly Review of Biology, 28 (2), 136 - 156. https: // doi. org / 10.1086 / 399512" type="journal article" year="1953">Wilson, 1953</bibRefCitation>
). Worker polymorphism has been examined in detail only in the weakly polymorphic 
<taxonomicName id="4C3A4D68FF81FFAB7A5AF92859EBF911" box="[1288,1429,1699,1725]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF81FFAB7A5AF92859EBF911" box="[1288,1429,1699,1725]" italics="true" pageId="100" pageNumber="101">V. pergandei</emphasis>
</taxonomicName>
, where workers display an annual cycle of mean body size with the largest workers present during winter months (
<bibRefCitation id="EFAB4B1AFF81FFAB7FCDF9675D3FF8A9" author="Gordon, S. H." box="[159,321,1771,1797]" pageId="100" pageNumber="101" refId="ref68321" refString="Gordon, S. H. (1978) Food and foraging ecology of a desert harvester ant, Veromessor pergandei (Mayr). PhD dissertation, University of California, Berkeley, California, 186 pp." type="book" year="1978">Gordon, 1978</bibRefCitation>
; Rissing, 1987). Additionally, some tasks are related to worker size in 
<taxonomicName id="4C3A4D68FF81FFAB7B38F9605889F8A9" box="[1130,1271,1771,1797]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF81FFAB7B38F9605889F8A9" box="[1130,1271,1771,1797]" italics="true" pageId="100" pageNumber="101">V. pergandei</emphasis>
</taxonomicName>
. For example, 
<taxonomicName id="4C3A4D68FF81FFAB7FC5F8835C87F885" authorityName="Sundevall" authorityYear="1833" box="[151,249,1807,1833]" class="Arachnida" family="Theridiidae" genus="Steatoda" kingdom="Animalia" order="Araneae" pageId="100" pageNumber="101" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF81FFAB7FC5F8835C87F885" box="[151,249,1807,1833]" italics="true" pageId="100" pageNumber="101">Steatoda</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF81FFAB7E63F8835DEFF885" authorityName="Sundevall" authorityYear="1833" box="[305,401,1807,1833]" class="Arachnida" family="Theridiidae" genus="Asagena" kingdom="Animalia" order="Araneae" pageId="100" pageNumber="101" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF81FFAB7E63F8835DEFF885" box="[305,401,1807,1833]" italics="true" pageId="100" pageNumber="101">Asagena</emphasis>
</taxonomicName>
spiders prey on workers of 
<taxonomicName id="4C3A4D68FF81FFAB7D9DF89C5F25F885" box="[719,859,1807,1833]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="100" pageNumber="101" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF81FFAB7D9DF89C5F25F885" box="[719,859,1807,1833]" italics="true" pageId="100" pageNumber="101">V. pergandei</emphasis>
</taxonomicName>
, but only the largest 30% of workers remove webs of these species from the foraging column, and only the largest workers retrieve nestmates that are ensnared in webs (
<bibRefCitation id="EFAB4B1AFF81FFAB7FCDF8DB5D92F8DD" author="Kwapich, C. L. &amp; Holldobler, B." box="[159,492,1879,1905]" pageId="100" pageNumber="101" pagination="395 - 404" refId="ref70304" refString="Kwapich, C. L. &amp; Holldobler, B. (2019) Destruction of spider webs and rescue of ensnared nestmates by a granivorous desert ant (Veromessor pergandei). American Naturalist, 194 (3), 395 - 404. https: // doi. org / 10.1086 / 704338" type="journal article" year="2019">Kwapich &amp; Hölldobler, 2019</bibRefCitation>
). Groups of polymorphic workers also dig longer nests with more branching than do groups of monomorphic workers, and larger workers build tunnels with a larger diameter (
<bibRefCitation id="EFAB4B1AFF81FFAB7BCEF8F759EFF839" author="Kwapich, C. L. &amp; Valentini, G. &amp; Holldobler, B." box="[1180,1425,1915,1942]" pageId="100" pageNumber="101" pagination="20170235" refId="ref70359" refString="Kwapich, C. L., Valentini, G. &amp; Holldobler, B. (2018) The non-additive effects of body size on nest architecture in a polymorphic ant. Philosophical Transactions of the Royal Society, Series B, 373, 20170235. https: // doi. org / 10.1098 / rstb. 2017.0235" type="journal article" year="2018">
Kwapich 
<emphasis id="B94EEAF9FF81FFAB7A59F8F0593BF839" box="[1291,1349,1915,1941]" italics="true" pageId="100" pageNumber="101">et al.</emphasis>
, 2018
</bibRefCitation>
). Further studies should examine distribution of body sizes within and across colonies for all species. These studies should include seasonal and geographic variation in worker body size given the annual cycle displayed by colonies of 
<taxonomicName id="4C3A4D68FF80FFAA7FE4FF1B5D86FF1D" authority="(Rissing, 1987)" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[182,504,151,177]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF80FFAA7FE4FF1B5D3CFF1D" box="[182,322,151,177]" italics="true" pageId="101" pageNumber="102">V. pergandei</emphasis>
(Rissing, 1987)
</taxonomicName>
, along with the size-frequency distribution for foragers within colonies, as graphed by 
<bibRefCitation id="EFAB4B1AFF80FFAA7FE8FF375D91FF79" author="Rissing, S. W. &amp; Pollock, G. B." box="[186,495,187,213]" pageId="101" pageNumber="102" pagination="121 - 126" refId="ref72032" refString="Rissing, S. W. &amp; Pollock, G. B. (1984) Worker size variability and foraging efficiency in Veromessor pergandei (Hymenoptera: Formicidae). Behavioral Ecology and Sociobiology, 15 (2), 121 - 126. https: // doi. org / 10.1007 / BF 00299379" type="journal article" year="1984">Rissing and Pollock (1984)</bibRefCitation>
.
</paragraph>
<paragraph id="8B8536EBFF80FFAA7F95FF535D63FDB5" blockId="101.[151,1437,151,717]" pageId="101" pageNumber="102">
Species of 
<taxonomicName id="4C3A4D68FF80FFAA7E18FF535DB2FF54" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[330,460,223,248]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF80FFAA7E18FF535DB2FF54" box="[330,460,223,248]" italics="true" pageId="101" pageNumber="102">Veromessor</emphasis>
</taxonomicName>
also show morphological similarities and differences within groups of ecologically similar species. The three large-colony species (
<taxonomicName id="4C3A4D68FF80FFAA7DE2FE8F5F6DFEB1" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[688,787,259,285]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF80FFAA7DE2FE8F5F6DFEB1" box="[688,787,259,285]" italics="true" pageId="101" pageNumber="102">V. andrei</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF80FFAA7C73FE8F5FE9FEB1" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[801,919,259,285]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF80FFAA7C73FE8F5FE9FEB1" box="[801,919,259,285]" italics="true" pageId="101" pageNumber="102">V. julianus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF80FFAA7CF7FE8F584EFEB1" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[933,1072,259,285]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF80FFAA7CF7FE8F584EFEB1" box="[933,1072,259,285]" italics="true" pageId="101" pageNumber="102">V. pergandei</emphasis>
</taxonomicName>
) form one group of ecologically similar species, while small-colony species form three groups of ecologically similar species: (1) 
<taxonomicName id="4C3A4D68FF80FFAA7BBDFEAB5915FEED" box="[1263,1387,295,321]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF80FFAA7BBDFEAB5915FEED" box="[1263,1387,295,321]" italics="true" pageId="101" pageNumber="102">V. lariversi</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF80FFAA7FC5FEC75D1FFEC9" box="[151,353,331,357]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF80FFAA7FC5FEC75D1FFEC9" box="[151,353,331,357]" italics="true" pageId="101" pageNumber="102">V. pseudolariversi</emphasis>
</taxonomicName>
, (2) 
<taxonomicName id="4C3A4D68FF80FFAA7EC4FEC75E45FEC9" baseAuthorityName="Andrews" baseAuthorityYear="1916" box="[406,571,331,357]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="lobognathus">
<emphasis id="B94EEAF9FF80FFAA7EC4FEC75E45FEC9" box="[406,571,331,357]" italics="true" pageId="101" pageNumber="102">V. lobognathus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF80FFAA7D20FEC75EAFFEC9" authorityName="Cole" authorityYear="1963" box="[626,721,331,357]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF80FFAA7D20FEC75EAFFEC9" box="[626,721,331,357]" italics="true" pageId="101" pageNumber="102">V. smithi</emphasis>
</taxonomicName>
, and (3) 
<taxonomicName id="4C3A4D68FF80FFAA7C65FEC75FB9FEC9" authorityName="Smith" authorityYear="1956" box="[823,967,331,357]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF80FFAA7C65FEC75FB9FEC9" box="[823,967,331,357]" italics="true" pageId="101" pageNumber="102">V. chicoensis</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF80FFAA7CAFFEC758FCFEC9" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[1021,1154,331,357]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF80FFAA7CAFFEC758FCFEC9" box="[1021,1154,331,357]" italics="true" pageId="101" pageNumber="102">V. stoddardi</emphasis>
</taxonomicName>
. Two of the large-colony species, 
<taxonomicName id="4C3A4D68FF80FFAA7FAAFEE35D0EFE25" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[248,368,367,393]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF80FFAA7FAAFEE35D0EFE25" box="[248,368,367,393]" italics="true" pageId="101" pageNumber="102">V. julianus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF80FFAA7EF8FEE35E46FE25" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[426,568,367,393]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF80FFAA7EF8FEE35E46FE25" box="[426,568,367,393]" italics="true" pageId="101" pageNumber="102">V. pergandei</emphasis>
</taxonomicName>
, have a similar allometry for head width versus head length and mesosoma length versus pronotum width. For 
<taxonomicName id="4C3A4D68FF80FFAA7D4FFE1F5E01FE01" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[541,639,403,429]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF80FFAA7D4FFE1F5E01FE01" box="[541,639,403,429]" italics="true" pageId="101" pageNumber="102">V. andrei</emphasis>
</taxonomicName>
, the allometry for mesosoma length versus pronotum width is similar to that of 
<taxonomicName id="4C3A4D68FF80FFAA7FB7FE3B5D24FE7D" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[229,346,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF80FFAA7FB7FE3B5D24FE7D" box="[229,346,439,465]" italics="true" pageId="101" pageNumber="102">V. julianus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF80FFAA7EDDFE3B5E67FE7D" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[399,537,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF80FFAA7EDDFE3B5E67FE7D" box="[399,537,439,465]" italics="true" pageId="101" pageNumber="102">V. pergandei</emphasis>
</taxonomicName>
, but head length in 
<taxonomicName id="4C3A4D68FF80FFAA7DA6FE3B5F29FE7D" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[756,855,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF80FFAA7DA6FE3B5F29FE7D" box="[756,855,439,465]" italics="true" pageId="101" pageNumber="102">V. andrei</emphasis>
</taxonomicName>
is greater than head width for any given worker size in the other two species. Workers of 
<taxonomicName id="4C3A4D68FF80FFAA7D7EFE575EF1FE59" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[556,655,475,501]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF80FFAA7D7EFE575EF1FE59" box="[556,655,475,501]" italics="true" pageId="101" pageNumber="102">V. andrei</emphasis>
</taxonomicName>
are also less variable in size than those of 
<taxonomicName id="4C3A4D68FF80FFAA7B35FE5758A5FE59" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[1127,1243,475,501]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF80FFAA7B35FE5758A5FE59" box="[1127,1243,475,501]" italics="true" pageId="101" pageNumber="102">V. julianus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF80FFAA7A43FE5759E3FE59" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[1297,1437,475,501]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF80FFAA7A43FE5759E3FE59" box="[1297,1437,475,501]" italics="true" pageId="101" pageNumber="102">V. pergandei</emphasis>
</taxonomicName>
(
<figureCitation id="13012A6EFF80FFAA7FCDFE735D6FFDB5" box="[159,273,510,537]" captionStart="FIGURE 54" captionStartId="103.[151,250,1926,1950]" captionTargetBox="[383,1206,200,1885]" captionTargetId="figure-18@103.[368,1219,181,1903]" captionTargetPageId="103" captionText="FIGURE 54. Bivariate plots for workers showing head width versus head length (A) and mesosoma length versus pronotum width (B) for the three large colony species of Veromessor (V. andrei, V. julianus, V. pergandei). Sample size for each species is given in parentheses." figureDoi="http://doi.org/10.5281/zenodo.7318950" httpUri="https://zenodo.org/record/7318950/files/figure.png" pageId="101" pageNumber="102">Figure 54</figureCitation>
).
</paragraph>
<paragraph id="8B8536EBFF80FFAA7F95FDAF5E5AFD61" blockId="101.[151,1437,151,717]" pageId="101" pageNumber="102">
The ecologically similar pairs of small-colony species display two patterns. First, body size varies across groups with there being a group of smaller species (
<taxonomicName id="4C3A4D68FF80FFAA7DBEFDCB5F16FDCD" box="[748,872,583,609]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF80FFAA7DBEFDCB5F16FDCD" box="[748,872,583,609]" italics="true" pageId="101" pageNumber="102">V. lariversi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF80FFAA7C2AFDCB583DFDCD" box="[888,1091,583,609]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF80FFAA7C2AFDCB583DFDCD" box="[888,1091,583,609]" italics="true" pageId="101" pageNumber="102">V. pseudolariversi</emphasis>
</taxonomicName>
), a group of larger species (
<emphasis id="B94EEAF9FF80FFAA7ADAFDCB5DECFD29" italics="true" pageId="101" pageNumber="102">
<taxonomicName id="4C3A4D68FF80FFAA7ADAFDCB5D5AFD29" baseAuthorityName="Andrews" baseAuthorityYear="1916" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="lobognathus">V. lobognathus</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF80FFAA7E63FDE75DECFD29" authorityName="Cole" authorityYear="1963" box="[305,402,619,645]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="smithi">V. smithi</taxonomicName>
</emphasis>
), and a group of polymorphic species (
<taxonomicName id="4C3A4D68FF80FFAA7C07FDE75F98FD29" authorityName="Smith" authorityYear="1956" box="[853,998,619,645]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF80FFAA7C07FDE75F98FD29" box="[853,998,619,645]" italics="true" pageId="101" pageNumber="102">V. chicoensis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF80FFAA7CA4FDE75802FD29" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[1014,1148,619,645]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF80FFAA7CA4FDE75802FD29" box="[1014,1148,619,645]" italics="true" pageId="101" pageNumber="102">V. stoddardi</emphasis>
</taxonomicName>
). The former two groups also have one species that has greater variation in body size (
<figureCitation id="13012A6EFF80FFAA7C1BFD035FC5FD05" box="[841,955,654,681]" captionStart="FIGURE 55" captionStartId="104.[151,250,1872,1896]" captionTargetBox="[227,1370,195,1836]" captionTargetId="figure-26@104.[206,1381,181,1848]" captionTargetPageId="104" captionText="FIGURE 55. Bivariate plots for workers showing head width versus head length (left panels) and mesosoma length versus pronotum width (right panels) for ecologically similar pairs of small colony species of Veromessor: V. lariversi and V. pseudolariversi (A–B), V. lobognathus and V. smithi (C–D), V. chicoensis and V. stoddardi (E–F). Note that the x-axis scale is the same for all panels on the left and for all panels on the right so that sizes can be compared directly across species groups. Sample size for each species is given in parentheses." figureDoi="http://doi.org/10.5281/zenodo.7318952" httpUri="https://zenodo.org/record/7318952/files/figure.png" pageId="101" pageNumber="102">Figure 55</figureCitation>
). 
<taxonomicName id="4C3A4D68FF80FFAA7C82FD03589EFD05" box="[976,1248,655,681]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="chamberlini">
<emphasis id="B94EEAF9FF80FFAA7C82FD03589EFD05" box="[976,1248,655,681]" italics="true" pageId="101" pageNumber="102">Veromessor chamberlini</emphasis>
</taxonomicName>
does not readily fit into any of these species groups.
</paragraph>
</subSubSection>
<subSubSection id="C3206560FF80FFA67FC5FC925F40F911" lastPageId="105" lastPageNumber="106" pageId="101" pageNumber="102" type="biology_ecology">
<paragraph id="8B8536EBFF80FFAA7FC5FC925E4AFC95" blockId="101.[151,564,798,825]" box="[151,564,798,825]" pageId="101" pageNumber="102">
<emphasis id="B94EEAF9FF80FFAA7FC5FC925E4AFC95" bold="true" box="[151,564,798,825]" pageId="101" pageNumber="102">Colony size and foraging method</emphasis>
</paragraph>
<paragraph id="8B8536EBFF80FFAA7FC5FCEB5ECBFBF5" blockId="101.[151,1437,871,1329]" pageId="101" pageNumber="102">
Species of 
<taxonomicName id="4C3A4D68FF80FFAA7E40FCEB5DEAFC2C" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[274,404,871,896]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF80FFAA7E40FCEB5DEAFC2C" box="[274,404,871,896]" italics="true" pageId="101" pageNumber="102">Veromessor</emphasis>
</taxonomicName>
display a dimorphism in colony size with three species (
<taxonomicName id="4C3A4D68FF80FFAA7B5DFCEB580CFC2D" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[1039,1138,871,897]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF80FFAA7B5DFCEB580CFC2D" box="[1039,1138,871,897]" italics="true" pageId="101" pageNumber="102">V. andrei</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF80FFAA7BD2FCEB5887FC2D" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[1152,1273,871,897]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF80FFAA7BD2FCEB58EFFC2C" box="[1152,1169,871,896]" italics="true" pageId="101" pageNumber="102">V</emphasis>
. 
<emphasis id="B94EEAF9FF80FFAA7BCDFCEB5887FC2D" box="[1183,1273,871,897]" italics="true" pageId="101" pageNumber="102">julianus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF80FFAA7A55FCEB59EDFC2D" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[1287,1427,871,897]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF80FFAA7A55FCEB59EDFC2D" box="[1287,1427,871,897]" italics="true" pageId="101" pageNumber="102">V. pergandei</emphasis>
</taxonomicName>
) having colonies that probably greatly exceed 50,000 workers, while the other seven species have colonies that typically range from several hundred to 
<geoCoordinate id="EE0E502CFF80FFAA7D3EFC235ED4FC65" box="[620,682,943,969]" degrees="1,000" direction="north" orientation="latitude" pageId="101" pageNumber="102" precision="55" value="1.0">1,000</geoCoordinate>
<geoCoordinate id="EE0E502CFF80FFAA7DF8FC235E86FC65" box="[682,760,943,969]" degrees="2,000" direction="west" orientation="longitude" pageId="101" pageNumber="102" precision="55" value="-2.0">–2,000</geoCoordinate>
or more workers. All three large-colony species forage in columns that extend 
<quantity id="4CC29B0EFF80FFAA7E2FFC5F5DA1FC41" box="[381,479,979,1005]" metricMagnitude="1" metricUnit="m" metricValue="2.5" metricValueMax="4.0" metricValueMin="1.0" pageId="101" pageNumber="102" unit="m" value="25.0" valueMax="40.0" valueMin="10.0">10–40 m</quantity>
from the nest, with workers dispersing into a fan to forage individually near the end of the column (for a review see Plowes, Johnson, &amp; Hölldobler, 2012). In contrast, all small-colony species use solitary foraging, although 
<taxonomicName id="4C3A4D68FF80FFAA7E3CFB975E6FFB99" box="[366,529,1051,1077]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="chamberlini">
<emphasis id="B94EEAF9FF80FFAA7E3CFB975E6FFB99" box="[366,529,1051,1077]" italics="true" pageId="101" pageNumber="102">V. chamberlini</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF80FFAA7D72FB975ECCFB99" authorityName="Smith" authorityYear="1956" box="[544,690,1051,1077]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF80FFAA7D72FB975ECCFB99" box="[544,690,1051,1077]" italics="true" pageId="101" pageNumber="102">V. chicoensis</emphasis>
</taxonomicName>
, and 
<taxonomicName id="4C3A4D68FF80FFAA7DA2FB975F08FB99" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[752,886,1051,1077]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF80FFAA7DA2FB975F08FB99" box="[752,886,1051,1077]" italics="true" pageId="101" pageNumber="102">V. stoddardi</emphasis>
</taxonomicName>
sometimes forage in short columns (
<tableCitation id="C6B80350FF80FFAA7A4AFB975910FB99" box="[1304,1390,1051,1077]" captionStart="TABLE 1" captionStartId="99.[151,244,643,669]" captionTargetPageId="99" captionText="TABLE 1. Summary of morphological and ecological traits for species of Veromessor." httpUri="http://table.plazi.org/id/DF456663FF86FFAC7FC5FD0F5847FD31" pageId="101" pageNumber="102" tableUuid="DF456663FF86FFAC7FC5FD0F5847FD31">Table 1</tableCitation>
; M. Bennett, pers. comm.; R.A. Johnson, pers. obs.).
</paragraph>
<paragraph id="8B8536EBFF80FFAA7F95FBEF5F88FA9D" blockId="101.[151,1437,871,1329]" pageId="101" pageNumber="102">
Colony size and foraging strategy also correlate with development of the pygidial gland. The three large-colony, column foraging species possess a well developed pygidial gland that consists of large paired clusters of glandular cells, large paired reservoir sacks, and a pair of distinct textured cuticle structures associated with the gland. In contrast, the pygidial gland reservoir and textured tergal cuticle structures are lacking or rudimentary in six of the seven small-colony species (not examined in 
<taxonomicName id="4C3A4D68FF80FFAA7DF3FB7F5F63FAA1" box="[673,797,1267,1293]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF80FFAA7DF3FB7F5F63FAA1" box="[673,797,1267,1293]" italics="true" pageId="101" pageNumber="102">V. lariversi</emphasis>
</taxonomicName>
) (
<bibRefCitation id="EFAB4B1AFF80FFAA7C65FB7F583CFAA1" author="Holldobler, B. &amp; Plowes, N. J. R. &amp; Johnson, R. A. &amp; Nishshanka, U. &amp; Liu, C. &amp; Attygalle, A. B." box="[823,1090,1267,1293]" pageId="101" pageNumber="102" pagination="863 - 869" refId="ref69008" refString="Holldobler, B., Plowes, N. J. R., Johnson, R. A., Nishshanka, U., Liu, C. &amp; Attygalle, A. B. (2013) Pygidial gland chemistry and potential alarm-recruitment function in column foraging, but not solitary, Nearctic Messor harvesting ants (Hymenoptera: Formicidae: Myrmicinae). Journal of Insect Physiology, 59 (9), 863 - 869. https: // doi. org / 10.1016 / j. jinsphys. 2013.06.006" type="journal article" year="2013">
Hölldobler 
<emphasis id="B94EEAF9FF80FFAA7CE9FB7F5F88FAA1" box="[955,1014,1267,1293]" italics="true" pageId="101" pageNumber="102">et al.</emphasis>
, 2013
</bibRefCitation>
). These two groups of species should be examined for additional differences in glands and gland chemistry.
</paragraph>
<paragraph id="8B8536EBFF80FFAA7FC5FA0E5D87FA30" blockId="101.[151,505,1410,1436]" box="[151,505,1410,1436]" pageId="101" pageNumber="102">
<emphasis id="B94EEAF9FF80FFAA7FC5FA0E5D87FA30" bold="true" box="[151,505,1410,1436]" pageId="101" pageNumber="102">Seasonality of mating flights</emphasis>
</paragraph>
<paragraph id="8B8536EBFF80FFAA7FC5FA475D93F815" blockId="101.[151,1437,1483,1977]" pageId="101" pageNumber="102">
Most desert ants have summer mating flights that are triggered by rain (
<bibRefCitation id="EFAB4B1AFF80FFAA7C8CFA4758EBFA49" author="Johnson, R. A." box="[990,1173,1483,1509]" pageId="101" pageNumber="102" pagination="89 - 122" refId="ref69343" refString="Johnson, R. A. (2000 b) Seed-harvester ants (Hymenoptera: Formicidae) of North America: an overview of ecology and biogeography. Sociobiology, 36 (1), 89 - 122." type="journal article" year="2000">Johnson, 2000b</bibRefCitation>
, 
<bibRefCitation id="EFAB4B1AFF80FFAA7BF1FA4758A1FA49" author="Johnson, R. A." box="[1187,1247,1483,1509]" pageId="101" pageNumber="102" pagination="1 - 29" refId="ref69435" refString="Johnson, R. A. (2001) Biogeography and community structure of North American seed-harvester ants. Annual Review of Entomology, 46, 1 - 29. https: // doi. org / 10.1146 / annurev. ento. 46.1.1" type="journal article" year="2001">2001</bibRefCitation>
). In contrast, all species of 
<taxonomicName id="4C3A4D68FF80FFAA7E42FA635DECF9A4" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[272,402,1519,1544]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF80FFAA7E42FA635DECF9A4" box="[272,402,1519,1544]" italics="true" pageId="101" pageNumber="102">Veromessor</emphasis>
</taxonomicName>
have mating flights that appear to be triggered by photoperiod, i.e., flights do not occur in response to rains. All species of 
<taxonomicName id="4C3A4D68FF80FFAA7D59F99F5EF3F980" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[523,653,1555,1580]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF80FFAA7D59F99F5EF3F980" box="[523,653,1555,1580]" italics="true" pageId="101" pageNumber="102">Veromessor</emphasis>
</taxonomicName>
appear to release few sexuals per day over a period of up to several weeks. Photoperiod-triggered mating flights appear to be a derived trait given that species of 
<taxonomicName id="4C3A4D68FF80FFAA7BC0F9BB5962F9FC" baseAuthorityName="Donisthorpe" baseAuthorityYear="1943" box="[1170,1308,1591,1616]" class="Insecta" family="Formicidae" genus="Novomessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF80FFAA7BC0F9BB5962F9FC" box="[1170,1308,1591,1616]" italics="true" pageId="101" pageNumber="102">Novomessor</emphasis>
</taxonomicName>
, the mostly closely related genus (
<bibRefCitation id="EFAB4B1AFF80FFAA7EC7F9D75E1FF9D9" author="Ward, P. S. &amp; Brady, S. G. &amp; Fisher, B. L. &amp; Schultz, T. R." box="[405,609,1626,1653]" pageId="101" pageNumber="102" pagination="61 - 81" refId="ref73206" refString="Ward, P. S., Brady, S. G., Fisher, B. L. &amp; Schultz, T. R. (2015) The evolution of myrmicine ants: phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae). Systematic Entomology, 40 (1), 61 - 81. https: // doi. org / 10.1111 / syen. 12090" type="journal article" year="2015">
Ward 
<emphasis id="B94EEAF9FF80FFAA7E8EF9D75E68F9D9" box="[476,534,1627,1653]" italics="true" pageId="101" pageNumber="102">et al.</emphasis>
, 2015
</bibRefCitation>
), have mating flights triggered by summer rains (
<bibRefCitation id="EFAB4B1AFF80FFAA7BC5F9D75930F9D9" author="Johnson, R. A." box="[1175,1358,1627,1653]" pageId="101" pageNumber="102" pagination="89 - 122" refId="ref69343" refString="Johnson, R. A. (2000 b) Seed-harvester ants (Hymenoptera: Formicidae) of North America: an overview of ecology and biogeography. Sociobiology, 36 (1), 89 - 122." type="journal article" year="2000">Johnson, 2000b</bibRefCitation>
). Most species of 
<taxonomicName id="4C3A4D68FF80FFAA7E42F9F35DECF934" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[272,402,1663,1688]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF80FFAA7E42F9F35DECF934" box="[272,402,1663,1688]" italics="true" pageId="101" pageNumber="102">Veromessor</emphasis>
</taxonomicName>
have summer mating flights (June–August), whereas those of 
<taxonomicName id="4C3A4D68FF80FFAA7B09F9F358ACF935" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[1115,1234,1663,1689]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF80FFAA7B09F9F358ACF935" box="[1115,1234,1663,1689]" italics="true" pageId="101" pageNumber="102">V. julianus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF80FFAA7A59F9F359E3F935" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[1291,1437,1663,1689]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF80FFAA7A59F9F35962F934" box="[1291,1308,1663,1688]" italics="true" pageId="101" pageNumber="102">V</emphasis>
. 
<emphasis id="B94EEAF9FF80FFAA7A7EF9F359E3F935" box="[1324,1437,1663,1689]" italics="true" pageId="101" pageNumber="102">pergandei</emphasis>
</taxonomicName>
occur during late winter–early spring (
<bibRefCitation id="EFAB4B1AFF80FFAA7D6CF92F5E8CF911" author="Johnson, R. A." box="[574,754,1699,1725]" pageId="101" pageNumber="102" pagination="377 - 384" refId="ref69291" refString="Johnson, R. A. (2000 a) Reproductive biology of the seed-harvester ants Messor julianus (Pergande) and Messor pergandei (Mayr) (Hymenoptera: Formicidae) in Baja California, Mexico. Journal of Hymenoptera Research, 9 (2), 377 - 384." type="journal article" year="2000">Johnson, 2000a</bibRefCitation>
). The late winter–spring flights for these two species correlate with their being the only two species of 
<taxonomicName id="4C3A4D68FF80FFAA7D33F94B5E9DF94C" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[609,739,1735,1760]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF80FFAA7D33F94B5E9DF94C" box="[609,739,1735,1760]" italics="true" pageId="101" pageNumber="102">Veromessor</emphasis>
</taxonomicName>
that are restricted to hot desert habitats (
<bibRefCitation id="EFAB4B1AFF80FFAA7BE3F94B5CADF8A9" author="Wheeler, G. C. &amp; Wheeler, J." pageId="101" pageNumber="102" refId="ref73592" refString="Wheeler, G. C. &amp; Wheeler, J. (1973) Ants of Deep Canyon. University of California Press, Berkeley, California, 162 pp." type="book" year="1973">Wheeler &amp; Wheeler, 1973</bibRefCitation>
). Queens of both species have poor heat tolerance compared to species of 
<taxonomicName id="4C3A4D68FF80FFAA7B7AF96758CCF8A8" baseAuthorityName="Donisthorpe" baseAuthorityYear="1943" box="[1064,1202,1771,1796]" class="Insecta" family="Formicidae" genus="Novomessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF80FFAA7B7AF96758CCF8A8" box="[1064,1202,1771,1796]" italics="true" pageId="101" pageNumber="102">Novomessor</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF80FFAA7BBEF96759E2F8A8" authorityName="Mayr" authorityYear="1868" box="[1260,1436,1771,1796]" class="Insecta" family="Formicidae" genus="Pogonomyrmex" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF80FFAA7BBEF96759E2F8A8" box="[1260,1436,1771,1796]" italics="true" pageId="101" pageNumber="102">Pogonomyrmex</emphasis>
</taxonomicName>
(see 
<bibRefCitation id="EFAB4B1AFF80FFAA7F9DF8835DFBF885" author="Johnson, R. A." box="[207,389,1807,1833]" pageId="101" pageNumber="102" pagination="377 - 384" refId="ref69291" refString="Johnson, R. A. (2000 a) Reproductive biology of the seed-harvester ants Messor julianus (Pergande) and Messor pergandei (Mayr) (Hymenoptera: Formicidae) in Baja California, Mexico. Journal of Hymenoptera Research, 9 (2), 377 - 384." type="journal article" year="2000">Johnson, 2000a</bibRefCitation>
), suggesting that 
<taxonomicName id="4C3A4D68FF80FFAA7D01F8835EABF884" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[595,725,1807,1832]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF80FFAA7D01F8835EABF884" box="[595,725,1807,1832]" italics="true" pageId="101" pageNumber="102">Veromessor</emphasis>
</taxonomicName>
queens could not survive hot summer soil temperatures (see also Rissing, Johnson, &amp; Pollock, 1986). Moreover, it seems probable that shifting mating flights from summer to late winter–early spring was prerequisite to these two species invading hot desert habitats (
<bibRefCitation id="EFAB4B1AFF80FFAA7BC7F8DB5936F8DD" author="Johnson, R. A." box="[1173,1352,1879,1905]" pageId="101" pageNumber="102" pagination="377 - 384" refId="ref69291" refString="Johnson, R. A. (2000 a) Reproductive biology of the seed-harvester ants Messor julianus (Pergande) and Messor pergandei (Mayr) (Hymenoptera: Formicidae) in Baja California, Mexico. Journal of Hymenoptera Research, 9 (2), 377 - 384." type="journal article" year="2000">
Johnson, 2000 
<bibRefCitation id="EFAB4B1AFF80FFAA7A6EF8DB5936F8DD" author="Johnson, R. A." box="[1340,1352,1879,1905]" pageId="101" pageNumber="102" pagination="1 - 29" refId="ref69435" refString="Johnson, R. A. (2001) Biogeography and community structure of North American seed-harvester ants. Annual Review of Entomology, 46, 1 - 29. https: // doi. org / 10.1146 / annurev. ento. 46.1.1" type="journal article" year="2001">a</bibRefCitation>
</bibRefCitation>
, 2001). Based on these patterns, we predict that mating flights of 
<emphasis id="B94EEAF9FF80FFAA7C71F8F75F4AF838" box="[803,820,1915,1940]" italics="true" pageId="101" pageNumber="102">V</emphasis>
. sp. 
<taxonomicName id="4C3A4D68FF80FFAA7C38F8F7587EF839" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[874,1024,1915,1941]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Hymenoptera" pageId="101" pageNumber="102" phylum="Arthropoda" rank="species" species="pergandei">
cf. 
<emphasis id="B94EEAF9FF80FFAA7CDDF8F7587EF839" box="[911,1024,1915,1941]" italics="true" pageId="101" pageNumber="102">pergandei</emphasis>
</taxonomicName>
(sexuals are unknown) occur during late winter or spring (
<tableCitation id="C6B80350FF80FFAA7EDAF8135DA2F815" box="[392,476,1951,1977]" captionStart="TABLE 1" captionStartId="99.[151,244,643,669]" captionTargetPageId="99" captionText="TABLE 1. Summary of morphological and ecological traits for species of Veromessor." httpUri="http://table.plazi.org/id/DF456663FF86FFAC7FC5FD0F5847FD31" pageId="101" pageNumber="102" tableUuid="DF456663FF86FFAC7FC5FD0F5847FD31">Table 1</tableCitation>
).
</paragraph>
<caption id="DF456663FF83FFA97FC5F8865E71F856" ID-DOI="http://doi.org/10.5281/zenodo.7318948" ID-Zenodo-Dep="7318948" httpUri="https://zenodo.org/record/7318948/files/figure.png" pageId="102" pageNumber="103" startId="102.[151,250,1802,1826]" targetBox="[376,1218,194,1775]" targetPageId="102">
<paragraph id="8B8536EBFF83FFA97FC5F8865E71F856" blockId="102.[151,1436,1801,2042]" pageId="102" pageNumber="103">
<emphasis id="B94EEAF9FF83FFA97FC5F8865D5FF88E" bold="true" box="[151,289,1802,1826]" pageId="102" pageNumber="103">FIGURE 53.</emphasis>
Eye area (mm 
<superScript id="7C4F9BA3FF83FFA97EE8F8855DBFF8BB" attach="left" box="[442,449,1801,1815]" fontSize="6" pageId="102" pageNumber="103">2</superScript>
) (A), facet number (B), and mean facet diameter (μm) (C) for pale and dark colored species of 
<taxonomicName id="4C3A4D68FF83FFA97FC5F8A35D72F8EA" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[151,268,1839,1862]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="102" pageNumber="103" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF83FFA97FC5F8A35D72F8EA" box="[151,268,1839,1862]" italics="true" pageId="102" pageNumber="103">Veromessor</emphasis>
</taxonomicName>
. Two species are pale (
<taxonomicName id="4C3A4D68FF83FFA97EA7F8A35E1DF8EA" box="[501,611,1838,1862]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="102" pageNumber="103" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF83FFA97EA7F8A35E1DF8EA" box="[501,611,1838,1862]" italics="true" pageId="102" pageNumber="103">V. lariversi</emphasis>
</taxonomicName>
, 
<emphasis id="B94EEAF9FF83FFA97D3CF8A35F44F8EA" box="[622,826,1838,1862]" italics="true" pageId="102" pageNumber="103">
<taxonomicName id="4C3A4D68FF83FFA97D3CF8A35F5EF8EA" box="[622,800,1838,1862]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="102" pageNumber="103" phylum="Arthropoda" rank="species" species="pseudolariversi">V. pseudolariversi</taxonomicName>
—
</emphasis>
open symbols and regular font), while the other eight species are dark (filled symbols and bold font). For each species, number of workers examined and number of colonies they were derived from is given in parentheses. Significant differences (
<emphasis id="B94EEAF9FF83FFA97DE2F8FB5EC1F822" box="[688,703,1911,1934]" italics="true" pageId="102" pageNumber="103">P</emphasis>
&lt;0.05) among species are denoted after each species name by the letters 
<emphasis id="B94EEAF9FF83FFA97FC5F8175CC3F81E" box="[151,189,1947,1970]" italics="true" pageId="102" pageNumber="103">a–g</emphasis>
: 
<emphasis id="B94EEAF9FF83FFA97F98F8175CA9F81E" box="[202,215,1947,1970]" italics="true" pageId="102" pageNumber="103">a</emphasis>
&gt; 
<emphasis id="B94EEAF9FF83FFA97FA3F8165C80F81E" box="[241,254,1946,1970]" italics="true" pageId="102" pageNumber="103">b</emphasis>
&gt; 
<emphasis id="B94EEAF9FF83FFA97E4AF8175D5DF81E" box="[280,291,1947,1970]" italics="true" pageId="102" pageNumber="103">c</emphasis>
&gt; 
<emphasis id="B94EEAF9FF83FFA97E6FF8165D34F81E" box="[317,330,1946,1970]" italics="true" pageId="102" pageNumber="103">d</emphasis>
&gt; 
<emphasis id="B94EEAF9FF83FFA97E36F8175D11F81E" box="[356,367,1947,1970]" italics="true" pageId="102" pageNumber="103">e</emphasis>
&gt; 
<emphasis id="B94EEAF9FF83FFA97EDBF8165DEEF81E" box="[393,400,1946,1970]" italics="true" pageId="102" pageNumber="103">f</emphasis>
&gt; 
<emphasis id="B94EEAF9FF83FFA97EF9F8175DC6F81E" box="[427,440,1947,1970]" italics="true" pageId="102" pageNumber="103">g</emphasis>
; the three sets of letters for each species correspond to panels A, B, and C, respectively. Groupings are based on univariate F tests within MANCOVA followed by pairwise comparisons using a least significant differences test (see also Johnson &amp; Rutowki, 2022).
</paragraph>
</caption>
<caption id="DF456663FF82FFA87FC5F80A5D13F84B" ID-DOI="http://doi.org/10.5281/zenodo.7318950" ID-Zenodo-Dep="7318950" httpUri="https://zenodo.org/record/7318950/files/figure.png" pageId="103" pageNumber="104" startId="103.[151,250,1926,1950]" targetBox="[383,1206,200,1885]" targetPageId="103">
<paragraph id="8B8536EBFF82FFA87FC5F80A5D13F84B" blockId="103.[151,1437,1926,2023]" pageId="103" pageNumber="104">
<emphasis id="B94EEAF9FF82FFA87FC5F80A5D5FF832" bold="true" box="[151,289,1926,1950]" pageId="103" pageNumber="104">FIGURE 54.</emphasis>
Bivariate plots for workers showing head width versus head length (A) and mesosoma length versus pronotum width (B) for the three large colony species of 
<taxonomicName id="4C3A4D68FF82FFA87D3DF8275E9AF86E" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[623,740,1963,1986]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF82FFA87D3DF8275E9AF86E" box="[623,740,1963,1986]" italics="true" pageId="103" pageNumber="104">Veromessor</emphasis>
</taxonomicName>
(
<taxonomicName id="4C3A4D68FF82FFA87DA1F8275F33F86F" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[755,845,1963,1987]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF82FFA87DA1F8275F33F86F" box="[755,845,1963,1987]" italics="true" pageId="103" pageNumber="104">V. andrei</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF82FFA87C0BF8275FBAF86F" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[857,964,1963,1987]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF82FFA87C0BF8275FBAF86F" box="[857,964,1963,1987]" italics="true" pageId="103" pageNumber="104">V. julianus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF82FFA87C82F8275831F86F" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[976,1103,1963,1987]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="103" pageNumber="104" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF82FFA87C82F8275831F86F" box="[976,1103,1963,1987]" italics="true" pageId="103" pageNumber="104">V. pergandei</emphasis>
</taxonomicName>
). Sample size for each species is given in parentheses.
</paragraph>
</caption>
<caption id="DF456663FF8DFFA77FC5F8DC5ED3F854" ID-DOI="http://doi.org/10.5281/zenodo.7318952" ID-Zenodo-Dep="7318952" httpUri="https://zenodo.org/record/7318952/files/figure.png" pageId="104" pageNumber="105" startId="104.[151,250,1872,1896]" targetBox="[227,1370,195,1836]" targetPageId="104">
<paragraph id="8B8536EBFF8DFFA77FC5F8DC5ED3F854" blockId="104.[151,1437,1872,2040]" pageId="104" pageNumber="105">
<emphasis id="B94EEAF9FF8DFFA77FC5F8DC5D5CF8C5" bold="true" box="[151,290,1872,1897]" pageId="104" pageNumber="105">FIGURE 55.</emphasis>
Bivariate plots for workers showing head width versus head length (left panels) and mesosoma length versus pronotum width (right panels) for ecologically similar pairs of small colony species of 
<taxonomicName id="4C3A4D68FF8DFFA77B03F8F958B8F820" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[1105,1222,1909,1932]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="104" pageNumber="105" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF8DFFA77B03F8F958B8F820" box="[1105,1222,1909,1932]" italics="true" pageId="104" pageNumber="105">Veromessor</emphasis>
</taxonomicName>
: 
<taxonomicName id="4C3A4D68FF8DFFA77B8BF8F95933F820" box="[1241,1357,1908,1932]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="104" pageNumber="105" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF8DFFA77B8BF8F95933F820" box="[1241,1357,1908,1932]" italics="true" pageId="104" pageNumber="105">V. lariversi</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF8DFFA77AD8F8F95D4BF81C" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="104" pageNumber="105" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF8DFFA77AD8F8F95D4BF81C" italics="true" pageId="104" pageNumber="105">V. pseudolariversi</emphasis>
</taxonomicName>
(A–B), 
<taxonomicName id="4C3A4D68FF8DFFA77ED8F8155E5FF81C" baseAuthorityName="Andrews" baseAuthorityYear="1916" box="[394,545,1944,1968]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="104" pageNumber="105" phylum="Arthropoda" rank="species" species="lobognathus">
<emphasis id="B94EEAF9FF8DFFA77ED8F8155E5FF81C" box="[394,545,1944,1968]" italics="true" pageId="104" pageNumber="105">V. lobognathus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF8DFFA77D06F8155ED2F81C" authorityName="Cole" authorityYear="1963" box="[596,684,1944,1968]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="104" pageNumber="105" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF8DFFA77D06F8155ED2F81C" box="[596,684,1944,1968]" italics="true" pageId="104" pageNumber="105">V. smithi</emphasis>
</taxonomicName>
(C–D), 
<taxonomicName id="4C3A4D68FF8DFFA77C53F8155FFBF81C" authorityName="Smith" authorityYear="1956" box="[769,901,1944,1968]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="104" pageNumber="105" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF8DFFA77C53F8155FFBF81C" box="[769,901,1944,1968]" italics="true" pageId="104" pageNumber="105">V. chicoensis</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF8DFFA77CE5F815584FF81C" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[951,1073,1944,1968]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="104" pageNumber="105" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF8DFFA77CE5F815584FF81C" box="[951,1073,1944,1968]" italics="true" pageId="104" pageNumber="105">V. stoddardi</emphasis>
</taxonomicName>
(E–F). Note that the x-axis scale is the same for all panels on the left and for all panels on the right so that sizes can be compared directly across species groups. Sample size for each species is given in parentheses.
</paragraph>
</caption>
<paragraph id="8B8536EBFF8CFFA67FC5FF1B5D09FF1D" blockId="105.[151,375,151,177]" box="[151,375,151,177]" pageId="105" pageNumber="106">
<emphasis id="B94EEAF9FF8CFFA67FC5FF1B5D09FF1D" bold="true" box="[151,375,151,177]" pageId="105" pageNumber="106">Mating frequency</emphasis>
</paragraph>
<paragraph id="8B8536EBFF8CFFA67FC5FF535970FE59" blockId="105.[151,1437,223,502]" pageId="105" pageNumber="106">
Obligate multiple mating (effective queen mating frequency&gt; 2) is rare in ants (but see Denny, Franks, Powell, &amp; Edwards, 2004; Overson, Fewell, &amp; Gadau, 2016; Villesen, Murakami, Schultz, &amp; Boomsma, 2002; Wiernasz, Perroni, &amp; Cole, 2004). Little is known about mating frequency in species of 
<taxonomicName id="4C3A4D68FF8CFFA67CA8FEA45802FEED" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[1018,1148,296,321]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF8CFFA67CA8FEA45802FEED" box="[1018,1148,296,321]" italics="true" pageId="105" pageNumber="106">Veromessor</emphasis>
</taxonomicName>
, but we include this topic to highlight the need for data so as to determine if species variation correlates with life history traits such as colony size. Mating frequency is known only for 
<taxonomicName id="4C3A4D68FF8CFFA67D3AFEFC5E8DFE25" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[616,755,367,393]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8CFFA67D3AFEFC5E8DFE25" box="[616,755,367,393]" italics="true" pageId="105" pageNumber="106">V. pergandei</emphasis>
</taxonomicName>
, in which queens mate with multiple males; patriline number ranges from one to seven (mean = 3.80, 
<emphasis id="B94EEAF9FF8CFFA67D01FE185E1FFE01" box="[595,609,404,429]" italics="true" pageId="105" pageNumber="106">n</emphasis>
= 9 colonies) with an effective mating frequency (
<emphasis id="B94EEAF9FF8CFFA67BC3FE1858D1FE18" box="[1169,1199,404,436]" italics="true" pageId="105" pageNumber="106">
M 
<subScript id="17BE34AEFF8CFFA67BFAFE2A58D1FE18" attach="left" box="[1192,1199,422,436]" fontSize="6" pageId="105" pageNumber="106">e</subScript>
</emphasis>
) of 2.56 (
<bibRefCitation id="EFAB4B1AFF8CFFA67A49FE1F5C83FE7D" author="Kwapich, C. L. &amp; Gadau, J. &amp; Holldobler, B." pageId="105" pageNumber="106" pagination="110" refId="ref70239" refString="Kwapich, C. L., Gadau, J. &amp; Holldobler, B. (2017) The ecological and genetic basis of annual worker production in the desert seed harvesting ant, Veromessor pergandei. Behavioral Ecology and Sociobiology, 71, Article 110. https: // doi. org / 10.1007 / s 00265 - 017 - 2333 - 1" type="journal article" year="2017">
Kwapich 
<emphasis id="B94EEAF9FF8CFFA67ADAFE185CCAFE7D" italics="true" pageId="105" pageNumber="106">et al.</emphasis>
, 2017
</bibRefCitation>
; 
<bibRefCitation id="EFAB4B1AFF8CFFA67E5AFE3B5D86FE7D" author="Ode, P. J. &amp; Rissing, S. W." box="[264,504,439,465]" pageId="105" pageNumber="106" pagination="548 - 556" refId="ref70724" refString="Ode, P. J. &amp; Rissing, S. W. (2002) Resource abundance and sex allocation by queen and workers in the harvester ant, Messor pergandei. Behavioral Ecology and Sociobiology, 51 (6), 548 - 556. https: // doi. org / 10.1007 / s 00265 - 002 - 0462 - 6" type="journal article" year="2002">Ode &amp; Rissing, 2002</bibRefCitation>
). Additionally, the most productive colonies of 
<taxonomicName id="4C3A4D68FF8CFFA67B5BFE3458EBFE7D" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[1033,1173,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8CFFA67B5BFE3458EBFE7D" box="[1033,1173,439,465]" italics="true" pageId="105" pageNumber="106">V. pergandei</emphasis>
</taxonomicName>
had significantly fewer patrilines, a larger peak forager population, and a larger annual foraging range (
<bibRefCitation id="EFAB4B1AFF8CFFA67B43FE57597CFE59" author="Kwapich, C. L. &amp; Gadau, J. &amp; Holldobler, B." box="[1041,1282,475,502]" pageId="105" pageNumber="106" pagination="110" refId="ref70239" refString="Kwapich, C. L., Gadau, J. &amp; Holldobler, B. (2017) The ecological and genetic basis of annual worker production in the desert seed harvesting ant, Veromessor pergandei. Behavioral Ecology and Sociobiology, 71, Article 110. https: // doi. org / 10.1007 / s 00265 - 017 - 2333 - 1" type="journal article" year="2017">
Kwapich 
<emphasis id="B94EEAF9FF8CFFA67B2DFE5058C9FE59" box="[1151,1207,475,501]" italics="true" pageId="105" pageNumber="106">et al.</emphasis>
, 2017
</bibRefCitation>
).
</paragraph>
<paragraph id="8B8536EBFF8CFFA67FC5FDCB5EEFFDCD" blockId="105.[151,657,583,609]" box="[151,657,583,609]" pageId="105" pageNumber="106">
<emphasis id="B94EEAF9FF8CFFA67FC5FDCB5EEFFDCD" bold="true" box="[151,657,583,609]" pageId="105" pageNumber="106">Queen size and colony founding strategy</emphasis>
</paragraph>
<paragraph id="8B8536EBFF8CFFA67FC5FD035E15FBF6" blockId="105.[151,1437,655,1726]" pageId="105" pageNumber="106">
The evolution of queen size and method of colony founding is a promising research area given the wide variation in known queen founding strategies combined with the large variation in queen body size across species of 
<taxonomicName id="4C3A4D68FF8CFFA67A49FD3859E3FD61" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[1307,1437,692,717]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF8CFFA67A49FD3859E3FD61" box="[1307,1437,692,717]" italics="true" pageId="105" pageNumber="106">Veromessor</emphasis>
</taxonomicName>
(
<figureCitation id="13012A6EFF8CFFA67FCDFD5B5D6FFD5E" box="[159,273,727,754]" captionStart="FIGURE 56" captionStartId="106.[151,250,1727,1751]" captionTargetBox="[173,1422,538,1690]" captionTargetId="figure-211@106.[157,1430,525,1704]" captionTargetPageId="106" captionText="FIGURE 56. Queen body size across species of Veromessor based on average head width, average head length, and average mesosoma length. Species are ordered (left to right) from shortest to longest mesosoma length." figureDoi="http://doi.org/10.5281/zenodo.7318956" httpUri="https://zenodo.org/record/7318956/files/figure.png" pageId="105" pageNumber="106">Figure 56</figureCitation>
). The three large-colony, column foraging species have the largest queens in the genus, with an average dry mass of 
<quantity id="4CC29B0EFF8CFFA67E70FD775D01FCBA" box="[290,383,763,790]" metricMagnitude="-5" metricUnit="kg" metricValue="2.05" pageId="105" pageNumber="106" unit="mg" value="20.5">20.5 mg</quantity>
for 
<taxonomicName id="4C3A4D68FF8CFFA67EFCFD705E45FCB9" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[430,571,763,789]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8CFFA67EFCFD705E45FCB9" box="[430,571,763,789]" italics="true" pageId="105" pageNumber="106">V. pergandei</emphasis>
</taxonomicName>
(see also 
<bibRefCitation id="EFAB4B1AFF8CFFA67DF9FD775FC8FCBA" author="Cahan, S. &amp; Rissing, S. W." box="[683,950,763,790]" pageId="105" pageNumber="106" pagination="84 - 88" refId="ref67033" refString="Cahan, S. &amp; Rissing, S. W. (2005) Variation in queen size across a behavioral transition zone in the ant Messor pergandei. Insectes Sociaux, 52 (1), 84 - 88. https: // doi. org / 10.1007 / s 00040 - 004 - 0776 - 7" type="journal article" year="2005">Cahan &amp; Rissing, 2005</bibRefCitation>
), 
<quantity id="4CC29B0EFF8CFFA67C98FD775858FCBA" box="[970,1062,763,790]" metricMagnitude="-5" metricUnit="kg" metricValue="1.1099999999999999" pageId="105" pageNumber="106" unit="mg" value="11.1">11.1 mg</quantity>
for 
<taxonomicName id="4C3A4D68FF8CFFA67B07FD7058B5FCB9" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[1109,1227,763,789]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF8CFFA67B07FD7058B5FCB9" box="[1109,1227,763,789]" italics="true" pageId="105" pageNumber="106">V. julianus</emphasis>
</taxonomicName>
, and 
<quantity id="4CC29B0EFF8CFFA67A58FD775926FCBA" box="[1290,1368,763,790]" metricMagnitude="-6" metricUnit="kg" metricValue="7.5" pageId="105" pageNumber="106" unit="mg" value="7.5">7.5 mg</quantity>
for 
<taxonomicName id="4C3A4D68FF8CFFA67ADAFD705CA1FC95" baseAuthorityName="Mayr" baseAuthorityYear="1886" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF8CFFA67ADAFD705CA1FC95" italics="true" pageId="105" pageNumber="106">V. andrei</emphasis>
</taxonomicName>
(see above). The queens of 
<taxonomicName id="4C3A4D68FF8CFFA67D47FCAC5EF4FC95" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[533,650,799,825]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF8CFFA67D47FCAC5EF4FC95" box="[533,650,799,825]" italics="true" pageId="105" pageNumber="106">V. julianus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF8CFFA67DEDFCAC5F34FC95" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[703,842,799,825]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8CFFA67DEDFCAC5F34FC95" box="[703,842,799,825]" italics="true" pageId="105" pageNumber="106">V. pergandei</emphasis>
</taxonomicName>
are the largest queens in the genus, and these also are the only two species that are restricted to hot desert habitats (
<bibRefCitation id="EFAB4B1AFF8CFFA67C69FCCF581AFCF1" author="Wheeler, G. C. &amp; Wheeler, J." box="[827,1124,835,861]" pageId="105" pageNumber="106" refId="ref73592" refString="Wheeler, G. C. &amp; Wheeler, J. (1973) Ants of Deep Canyon. University of California Press, Berkeley, California, 162 pp." type="book" year="1973">Wheeler &amp; Wheeler, 1973</bibRefCitation>
), where increased body size can increase longevity under desiccating conditions (
<bibRefCitation id="EFAB4B1AFF8CFFA67DB7FCEB5FF2FC2D" author="Johnson, R. A." box="[741,908,871,897]" pageId="105" pageNumber="106" pagination="255 - 266" refId="ref69237" refString="Johnson, R. A. (1998) Foundress survival and brood production in the desert seed-harvester ants Pogonomyrmex rugosus and P. barbatus (Hymenoptera, Formicidae). Insectes Sociaux, 45 (3), 255 - 266. https: // doi. org / 10.1007 / s 000400050086" type="journal article" year="1998">Johnson, 1998</bibRefCitation>
, 
<bibRefCitation id="EFAB4B1AFF8CFFA67CC4FCEB5FAFFC2D" author="Johnson, R. A." box="[918,977,871,897]" pageId="105" pageNumber="106" pagination="294 - 308" refId="ref69639" refString="Johnson, R. A. (2021) Dessication limits recruitment in the pleometrotic desert seed-harvester ant Veromessor pergandei. Ecology and Evolution, 11 (1), 294 - 308. https: // doi. org / 10.1002 / ece 3.7039" type="journal article" year="2021">2021</bibRefCitation>
; 
<bibRefCitation id="EFAB4B1AFF8CFFA67C8EFCEB5899FC2D" author="Wiernasz, D. C. &amp; Cole, B. J." box="[988,1255,871,897]" pageId="105" pageNumber="106" pagination="2179 - 2183" refId="ref73898" refString="Wiernasz, D. C. &amp; Cole, B. J. (2003) Queen size mediates queen survival and colony fitness in harvester ants. Evolution, 57 (9), 2179 - 2183. https: // doi. org / 10.1111 / j. 0014 - 3820.2003. tb 00396. x" type="journal article" year="2003">Wiernasz &amp; Cole, 2003</bibRefCitation>
). At the low end of the scale, size of 
<taxonomicName id="4C3A4D68FF8CFFA67E21FC005E7DFC09" authorityName="Smith" authorityYear="1956" box="[371,515,907,933]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF8CFFA67E21FC005E7DFC09" box="[371,515,907,933]" italics="true" pageId="105" pageNumber="106">V. chicoensis</emphasis>
</taxonomicName>
queens is similar to that of their workers (
<figureCitation id="13012A6EFF8CFFA67C85FC075838FC09" box="[983,1094,907,933]" captionStart="FIGURE 19" captionStartId="38.[151,250,1949,1973]" captionTargetBox="[217,1373,196,1910]" captionTargetId="figure-26@38.[203,1384,181,1925]" captionTargetPageId="38" captionText="FIGURE 19. Bivariate plots for workers, queens, and males of Veromessor chicoensis and V. stoddardi: (A) head width versus head length and (B) head width versus mesosoma length. Sample size for each caste is in parentheses." figureDoi="http://doi.org/10.5281/zenodo.7318868" httpUri="https://zenodo.org/record/7318868/files/figure.png" pageId="105" pageNumber="106">Figure 19</figureCitation>
), and undoubtedly this species has the lowest queen to worker dimorphism in the genus. The queens of 
<taxonomicName id="4C3A4D68FF8CFFA67C8EFC3C583FFC65" authorityName="Cole" authorityYear="1963" box="[988,1089,943,970]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF8CFFA67C8EFC3C5F93FC65" box="[988,1005,944,969]" italics="true" pageId="105" pageNumber="106">V</emphasis>
. 
<emphasis id="B94EEAF9FF8CFFA67CAFFC23583FFC65" box="[1021,1089,943,969]" italics="true" pageId="105" pageNumber="106">smithi</emphasis>
</taxonomicName>
are also relatively small, with a size similar to or only slightly larger than that of workers (workers: HW = 
<quantity id="4CC29B0EFF8CFFA67CAEFC5F58DFFC42" box="[1020,1185,979,1006]" metricMagnitude="-3" metricUnit="m" metricValue="1.47" metricValueMax="1.61" metricValueMin="1.33" pageId="105" pageNumber="106" unit="mm" value="1.47" valueMax="1.61" valueMin="1.33">1.33–1.61 mm</quantity>
, HL = 
<quantity id="4CC29B0EFF8CFFA67BA0FC5F59E6FC42" box="[1266,1432,979,1006]" metricMagnitude="-3" metricUnit="m" metricValue="1.455" metricValueMax="1.58" metricValueMin="1.33" pageId="105" pageNumber="106" unit="mm" value="1.455" valueMax="1.58" valueMin="1.33">1.33–1.58 mm</quantity>
, ML = 
<quantity id="4CC29B0EFF8CFFA67FB2FC7B5DF9FBBE" box="[224,391,1015,1042]" metricMagnitude="-3" metricUnit="m" metricValue="1.915" metricValueMax="2.07" metricValueMin="1.76" pageId="105" pageNumber="106" unit="mm" value="1.915" valueMax="2.07" valueMin="1.76">1.76–2.07 mm</quantity>
; queens: HL = 
<quantity id="4CC29B0EFF8CFFA67D64FC7B5EA2FBBE" box="[566,732,1015,1042]" metricMagnitude="-3" metricUnit="m" metricValue="1.515" metricValueMax="1.63" metricValueMin="1.4" pageId="105" pageNumber="106" unit="mm" value="1.515" valueMax="1.63" valueMin="1.4">1.40–1.63 mm</quantity>
, HW = 
<quantity id="4CC29B0EFF8CFFA67C65FC7B5FA3FBBE" box="[823,989,1015,1042]" metricMagnitude="-3" metricUnit="m" metricValue="1.515" metricValueMax="1.63" metricValueMin="1.4" pageId="105" pageNumber="106" unit="mm" value="1.515" valueMax="1.63" valueMin="1.4">1.40–1.63 mm</quantity>
, ML = 
<quantity id="4CC29B0EFF8CFFA67B61FC7B58A5FBBD" box="[1075,1243,1015,1041]" metricMagnitude="-3" metricUnit="m" metricValue="2.2199999999999998" metricValueMax="2.38" metricValueMin="2.06" pageId="105" pageNumber="106" unit="mm" value="2.2199999999999998" valueMax="2.38" valueMin="2.06">2.06–2.38 mm</quantity>
). A similar range of queen body size also occurs in 
<taxonomicName id="4C3A4D68FF8CFFA67D4BFB905EB7FB99" authorityName="Mayr" authorityYear="1868" box="[537,713,1052,1077]" class="Insecta" family="Formicidae" genus="Pogonomyrmex" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF8CFFA67D4BFB905EB7FB99" box="[537,713,1052,1077]" italics="true" pageId="105" pageNumber="106">Pogonomyrmex</emphasis>
</taxonomicName>
, where queens of 
<taxonomicName id="4C3A4D68FF8CFFA67CC8FB905871FB99" authorityName="Forel" authorityYear="1901" box="[922,1039,1051,1077]" class="Insecta" family="Formicidae" genus="Pogonomyrmex" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="schmitti">
<emphasis id="B94EEAF9FF8CFFA67CC8FB905871FB99" box="[922,1039,1051,1077]" italics="true" pageId="105" pageNumber="106">P. schmitti</emphasis>
</taxonomicName>
are similar in size to their workers (Johnson, 2015; 
<bibRefCitation id="EFAB4B1AFF8CFFA67E03FBB35E21FBF6" author="Johnson, R. A. &amp; Cover, S. P." box="[337,607,1087,1114]" pageId="105" pageNumber="106" pagination="231 - 249" refId="ref69684" refString="Johnson, R. A. &amp; Cover, S. P. (2015) A taxonomic revision of the seed-harvester ant genus Pogonomyrmex (Hymenoptera: Formicidae) on Hispaniola. Zootaxa, 3972 (2), 231 - 249. https: // doi. org / 10.11646 / zootaxa. 3972.2.5" type="journal article" year="2015">Johnson &amp; Cover, 2015</bibRefCitation>
).
</paragraph>
<paragraph id="8B8536EBFF8CFFA67F95FBEF5F40F911" blockId="105.[151,1437,655,1726]" pageId="105" pageNumber="106">
Little is known about colony founding strategies for species of 
<taxonomicName id="4C3A4D68FF8CFFA67CD6FBE85878FBD1" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[900,1030,1124,1149]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF8CFFA67CD6FBE85878FBD1" box="[900,1030,1124,1149]" italics="true" pageId="105" pageNumber="106">Veromessor</emphasis>
</taxonomicName>
. All species of 
<taxonomicName id="4C3A4D68FF8CFFA67BE3FBE8594DFBD1" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[1201,1331,1124,1149]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF8CFFA67BE3FBE8594DFBD1" box="[1201,1331,1124,1149]" italics="true" pageId="105" pageNumber="106">Veromessor</emphasis>
</taxonomicName>
probably have independent colony founding, but this has only been documented in 
<taxonomicName id="4C3A4D68FF8CFFA67C8CFB04583CFB0D" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[990,1090,1159,1185]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF8CFFA67C8CFB04583CFB0D" box="[990,1090,1159,1185]" italics="true" pageId="105" pageNumber="106">V. andrei</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF8CFFA67B03FB0458B6FB0D" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[1105,1224,1159,1185]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF8CFFA67B03FB0458B6FB0D" box="[1105,1224,1159,1185]" italics="true" pageId="105" pageNumber="106">V. julianus</emphasis>
</taxonomicName>
, and 
<taxonomicName id="4C3A4D68FF8CFFA67A5AFB0459EBFB0D" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[1288,1429,1159,1185]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8CFFA67A5AFB0459EBFB0D" box="[1288,1429,1159,1185]" italics="true" pageId="105" pageNumber="106">V. pergandei</emphasis>
</taxonomicName>
. 
<taxonomicName id="4C3A4D68FF8CFFA67FC5FB205D00FB69" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[151,382,1195,1221]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF8CFFA67FC5FB205D00FB69" box="[151,382,1195,1221]" italics="true" pageId="105" pageNumber="106">Veromessor julianus</emphasis>
</taxonomicName>
is haplometrotic (
<bibRefCitation id="EFAB4B1AFF8CFFA67D04FB275F70FB69" author="Johnson, R. A." box="[598,782,1195,1221]" pageId="105" pageNumber="106" pagination="377 - 384" refId="ref69291" refString="Johnson, R. A. (2000 a) Reproductive biology of the seed-harvester ants Messor julianus (Pergande) and Messor pergandei (Mayr) (Hymenoptera: Formicidae) in Baja California, Mexico. Journal of Hymenoptera Research, 9 (2), 377 - 384." type="journal article" year="2000">Johnson, 2000a</bibRefCitation>
), whereas 
<taxonomicName id="4C3A4D68FF8CFFA67CDFFB205860FB69" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[909,1054,1195,1221]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8CFFA67CDFFB205860FB69" box="[909,1054,1195,1221]" italics="true" pageId="105" pageNumber="106">V. pergandei</emphasis>
</taxonomicName>
displays geographic variation in founding strategy (see above). The larger queens of 
<taxonomicName id="4C3A4D68FF8CFFA67C52FB5C5FEFFB45" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[768,913,1231,1257]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8CFFA67C52FB5C5FEFFB45" box="[768,913,1231,1257]" italics="true" pageId="105" pageNumber="106">V. pergandei</emphasis>
</taxonomicName>
are fully claustral, while the smaller queens of 
<taxonomicName id="4C3A4D68FF8CFFA67FEAFB785D60FAA1" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[184,286,1267,1293]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF8CFFA67FEAFB785D60FAA1" box="[184,286,1267,1293]" italics="true" pageId="105" pageNumber="106">V. andrei</emphasis>
</taxonomicName>
appear to be semi-claustral with obligate queen foraging based on their absence of storage proteins (
<bibRefCitation id="EFAB4B1AFF8CFFA67FCDFA9B5D39FA9D" author="Brown, M. J. F." box="[159,327,1303,1329]" pageId="105" pageNumber="106" pagination="194 - 195" refId="ref66700" refString="Brown, M. J. F. (1999 b) Semi-claustral founding and worker behaviour in gynes of Messor andrei. Insectes Sociaux, 46 (2), 194 - 195. https: // doi. org / 10.1007 / s 000400050133" type="journal article" year="1999">Brown, 1999b</bibRefCitation>
). Nothing is known about colony founding for queens of small-colony species. However, if body size correlates with colony founding strategy, then queens for most to all small-colony species are predicted to be semi-claustral obligate foragers. For example, the very small size for queens of 
<taxonomicName id="4C3A4D68FF8CFFA67B44FAEC58D8FAD5" authorityName="Smith" authorityYear="1956" box="[1046,1190,1375,1401]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF8CFFA67B44FAEC58D8FAD5" box="[1046,1190,1375,1401]" italics="true" pageId="105" pageNumber="106">V. chicoensis</emphasis>
</taxonomicName>
, especially relative to worker size, suggests that this species is semi-claustral with obligate queen foraging. We also note that queens of 
<taxonomicName id="4C3A4D68FF8CFFA67ADAFA085D3BFA6D" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF8CFFA67ADAFA085D3BFA6D" italics="true" pageId="105" pageNumber="106">V. pseudolariversi</emphasis>
</taxonomicName>
are very small (
<figureCitation id="13012A6EFF8CFFA67EAEFA2B5E10FA6E" box="[508,622,1447,1474]" captionStart="FIGURE 56" captionStartId="106.[151,250,1727,1751]" captionTargetBox="[173,1422,538,1690]" captionTargetId="figure-211@106.[157,1430,525,1704]" captionTargetPageId="106" captionText="FIGURE 56. Queen body size across species of Veromessor based on average head width, average head length, and average mesosoma length. Species are ordered (left to right) from shortest to longest mesosoma length." figureDoi="http://doi.org/10.5281/zenodo.7318956" httpUri="https://zenodo.org/record/7318956/files/figure.png" pageId="105" pageNumber="106">Figure 56</figureCitation>
), but they are much larger than their workers compared to the queens of 
<taxonomicName id="4C3A4D68FF8CFFA67FC5FA405D40FA49" baseAuthorityName="Andrews" baseAuthorityYear="1916" box="[151,318,1483,1509]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="lobognathus">
<emphasis id="B94EEAF9FF8CFFA67FC5FA405D40FA49" box="[151,318,1483,1509]" italics="true" pageId="105" pageNumber="106">V. lobognathus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF8CFFA67E25FA405DA6FA49" authorityName="Cole" authorityYear="1963" box="[375,472,1483,1509]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF8CFFA67E25FA405DA6FA49" box="[375,472,1483,1509]" italics="true" pageId="105" pageNumber="106">V. smithi</emphasis>
</taxonomicName>
. Interestingly, 
<taxonomicName id="4C3A4D68FF8CFFA67DD1FA405E81FA49" box="[643,767,1483,1509]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF8CFFA67DD1FA405E81FA49" box="[643,767,1483,1509]" italics="true" pageId="105" pageNumber="106">V. lariversi</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF8CFFA67C65FA40587CFA49" box="[823,1026,1483,1509]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF8CFFA67C65FA40587CFA49" box="[823,1026,1483,1509]" italics="true" pageId="105" pageNumber="106">V. pseudolariversi</emphasis>
</taxonomicName>
have similar geographic ranges and are sympatric at several sites, but the queens differ in size with those of 
<taxonomicName id="4C3A4D68FF8CFFA67CEEFA7C58FBF9A5" box="[956,1157,1519,1545]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF8CFFA67CEEFA7C58FBF9A5" box="[956,1157,1519,1545]" italics="true" pageId="105" pageNumber="106">V. pseudolariversi</emphasis>
</taxonomicName>
being the smallest in the genus, while those of 
<taxonomicName id="4C3A4D68FF8CFFA67EDEF9985E78F981" box="[396,518,1555,1581]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF8CFFA67EDEF9985E78F981" box="[396,518,1555,1581]" italics="true" pageId="105" pageNumber="106">V. lariversi</emphasis>
</taxonomicName>
are the largest among small-colony species (
<figureCitation id="13012A6EFF8CFFA67CA9F99F5813F982" box="[1019,1133,1555,1582]" captionStart="FIGURE 56" captionStartId="106.[151,250,1727,1751]" captionTargetBox="[173,1422,538,1690]" captionTargetId="figure-211@106.[157,1430,525,1704]" captionTargetPageId="106" captionText="FIGURE 56. Queen body size across species of Veromessor based on average head width, average head length, and average mesosoma length. Species are ordered (left to right) from shortest to longest mesosoma length." figureDoi="http://doi.org/10.5281/zenodo.7318956" httpUri="https://zenodo.org/record/7318956/files/figure.png" pageId="105" pageNumber="106">Figure 56</figureCitation>
). Additional studies should examine colony founding strategies and variation in queen body size. For most of these species, it is doubtful that sufficient numbers of founding queens could be located to experimentally test founding strategy (e.g., Johnson, 2002, 2006). However, founding strategy might be inferred by comparing storage protein levels for alate queens across species (see Hahn, Johnson, Buck, &amp; Wheeler, 2004).
</paragraph>
</subSubSection>
<subSubSection id="C3206560FF8CFFA47FC5F8835DECFBBD" lastPageId="107" lastPageNumber="108" pageId="105" pageNumber="106" type="distribution">
<paragraph id="8B8536EBFF8CFFA67FC5F8835DDEF885" blockId="105.[151,416,1807,1833]" box="[151,416,1807,1833]" pageId="105" pageNumber="106">
<emphasis id="B94EEAF9FF8CFFA67FC5F8835DDEF885" bold="true" box="[151,416,1807,1833]" pageId="105" pageNumber="106">Distribution patterns</emphasis>
</paragraph>
<paragraph id="8B8536EBFF8CFFA57FC5F8D458DCFE59" blockId="105.[151,1437,1879,2014]" lastBlockId="106.[151,1437,151,501]" lastPageId="106" lastPageNumber="107" pageId="105" pageNumber="106">
<taxonomicName id="4C3A4D68FF8CFFA67FC5F8D45D67F8DD" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[151,281,1880,1905]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF8CFFA67FC5F8D45D67F8DD" box="[151,281,1880,1905]" italics="true" pageId="105" pageNumber="106">Veromessor</emphasis>
</taxonomicName>
is mostly restricted to western North America with species inhabiting arid and semi-arid habitats, grasslands, sclerophyll woodlands, and desert scrublands. Distribution patterns of 
<taxonomicName id="4C3A4D68FF8CFFA67B04F8F058A6F839" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[1110,1240,1916,1941]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF8CFFA67B04F8F058A6F839" box="[1110,1240,1916,1941]" italics="true" pageId="105" pageNumber="106">Veromessor</emphasis>
</taxonomicName>
are discussed in two groups that correspond to colony size—species that have large colonies (&gt;50,000 workers) and species that have small colonies (usually &lt;
<geoCoordinate id="EE0E502CFF8CFFA67EA6F84F5E4CF871" box="[500,562,1987,2013]" degrees="1,000" direction="north" orientation="latitude" pageId="105" pageNumber="106" precision="55" value="1.0">1,000</geoCoordinate>
<geoCoordinate id="EE0E502CFF8CFFA67D63F84F5EFEF871" box="[561,640,1987,2013]" degrees="2,000" direction="west" orientation="longitude" pageId="105" pageNumber="106" precision="55" value="-2.0">–2,000</geoCoordinate>
workers) (see also 
<bibRefCitation id="EFAB4B1AFF8CFFA67C08F84F5870F871" author="Johnson, R. A." box="[858,1038,1987,2013]" pageId="105" pageNumber="106" pagination="89 - 122" refId="ref69343" refString="Johnson, R. A. (2000 b) Seed-harvester ants (Hymenoptera: Formicidae) of North America: an overview of ecology and biogeography. Sociobiology, 36 (1), 89 - 122." type="journal article" year="2000">Johnson, 2000b</bibRefCitation>
). The three large-colony species (
<taxonomicName id="4C3A4D68FF8CFFA57ADAF8485CA1FF1D" baseAuthorityName="Mayr" baseAuthorityYear="1886" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" lastPageId="106" lastPageNumber="107" order="Hymenoptera" pageId="105" pageNumber="106" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF8CFFA57ADAF8485CA1FF1D" italics="true" lastPageId="106" lastPageNumber="107" pageId="105" pageNumber="106">V. andrei</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF8FFFA57FBCFF1B5D1DFF1D" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[238,355,151,177]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF8FFFA57FBCFF1B5D1DFF1D" box="[238,355,151,177]" italics="true" pageId="106" pageNumber="107">V. julianus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF8FFFA57E20FF1B5D80FF1D" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[370,510,151,177]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8FFFA57E20FF1B5D80FF1D" box="[370,510,151,177]" italics="true" pageId="106" pageNumber="107">V. pergandei</emphasis>
</taxonomicName>
) have mostly parapatric distribution patterns with their ranges showing little to no overlap across the narrow contact zones that separate the species (herein we define parapatric as species that occupy separate but adjoining areas, such that only a small fraction of individuals in each species encounter the other species) (see 
<figureCitation id="13012A6EFF8FFFA57E7CFE8F5DF2FEB1" box="[302,396,259,285]" captionStart="FIGURE 1" captionStartId="8.[151,250,1768,1792]" captionTargetBox="[378,1208,181,1742]" captionTargetId="figure-26@8.[378,1208,181,1744]" captionTargetPageId="8" captionText="FIGURE 1. Photographs of the metasternal process on males of Veromessor: (A) V. andrei (CASENT4010823), (B) V. chamberlini (UCRC_ENT00500152), (C)V.chicoensis (CASENT0869853), (D) V.julianus (LACMENT359792),(E)V.lariversi (CASENT0761204), (F) V. lobognathus (LACMENT363986), (G) V. pergandei (CASENT0869850), (H) V. pseudolariversi (CASENT0869851), (I) V. smithi (LACMENT364071), and (J) V. stoddardi (LACMENT364102). Photographs by Robert Johnson from www.AntWeb.org." figureDoi="http://doi.org/10.5281/zenodo.7318831" httpUri="https://zenodo.org/record/7318831/files/figure.png" pageId="106" pageNumber="107">Figure 1</figureCitation>
, pg 
<quantity id="4CC29B0EFF8FFFA57EEDFE8F5D86FEB1" box="[447,504,259,285]" metricMagnitude="0" metricUnit="m" metricValue="2.1082" pageId="106" pageNumber="107" unit="in" value="83.0">83 in</quantity>
<bibRefCitation id="EFAB4B1AFF8FFFA57D52FE8F5ECBFEB1" author="Johnson, R. A." box="[512,693,259,285]" pageId="106" pageNumber="107" pagination="89 - 122" refId="ref69343" refString="Johnson, R. A. (2000 b) Seed-harvester ants (Hymenoptera: Formicidae) of North America: an overview of ecology and biogeography. Sociobiology, 36 (1), 89 - 122." type="journal article" year="2000">Johnson, 2000b</bibRefCitation>
). 
<taxonomicName id="4C3A4D68FF8FFFA57D9AFE8F5FD5FEB1" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[712,939,259,285]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF8FFFA57D9AFE8F5FD5FEB1" box="[712,939,259,285]" italics="true" pageId="106" pageNumber="107">Veromessor julianus</emphasis>
</taxonomicName>
is restricted to central and southern parts of the 
<collectingRegion id="49FEF809FF8FFFA57F93FEAB5D11FEED" box="[193,367,295,321]" country="Mexico" name="Baja California" pageId="106" pageNumber="107">Baja California</collectingRegion>
peninsula in 
<collectingCountry id="F32D767BFF8FFFA57D5AFEAB5E1EFEED" box="[520,608,295,321]" name="Mexico" pageId="106" pageNumber="107">Mexico</collectingCountry>
, as well as a narrow band along cool Pacific coastal areas north to Bahía San Quintin (
<bibRefCitation id="EFAB4B1AFF8FFFA57E79FEC75DA2FEC9" author="Johnson, R. A." box="[299,476,331,357]" pageId="106" pageNumber="107" pagination="377 - 384" refId="ref69291" refString="Johnson, R. A. (2000 a) Reproductive biology of the seed-harvester ants Messor julianus (Pergande) and Messor pergandei (Mayr) (Hymenoptera: Formicidae) in Baja California, Mexico. Journal of Hymenoptera Research, 9 (2), 377 - 384." type="journal article" year="2000">
Johnson, 2000 
<bibRefCitation id="EFAB4B1AFF8FFFA57E82FEC75DA2FEC9" author="Johnson, R. A." box="[464,476,331,357]" pageId="106" pageNumber="107" pagination="89 - 122" refId="ref69343" refString="Johnson, R. A. (2000 b) Seed-harvester ants (Hymenoptera: Formicidae) of North America: an overview of ecology and biogeography. Sociobiology, 36 (1), 89 - 122." type="journal article" year="2000">a</bibRefCitation>
</bibRefCitation>
, 2000b) (
<figureCitation id="13012A6EFF8FFFA57D11FEC75EB7FEC9" box="[579,713,331,357]" captionStart="FIGURE 27" captionStartId="50.[151,250,1943,1967]" captionTargetBox="[417,1170,181,1920]" captionTargetId="figure-26@50.[417,1170,181,1920]" captionTargetPageId="50" captionText="FIGURE 27. Geographic distribution of: (A) Veromessor julianus, (B) V. lariversi, and (C) V. lobognathus. The larger black circle in each panel denotes the type locality." figureDoi="http://doi.org/10.5281/zenodo.7318888" httpUri="https://zenodo.org/record/7318888/files/figure.png" pageId="106" pageNumber="107">Figure 27A</figureCitation>
). Alternatively, 
<taxonomicName id="4C3A4D68FF8FFFA57C2AFEC7587AFEC9" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[888,1028,331,357]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8FFFA57C2AFEC7587AFEC9" box="[888,1028,331,357]" italics="true" pageId="106" pageNumber="107">V. pergandei</emphasis>
</taxonomicName>
occurs in the more xeric desert areas along the east coast of the peninsula south to northern 
<collectingRegion id="49FEF809FF8FFFA57DBFFEE35FB9FE25" box="[749,967,367,393]" country="Mexico" name="Baja California Sur" pageId="106" pageNumber="107">Baja California Sur</collectingRegion>
. Where these two species come in contact, 
<taxonomicName id="4C3A4D68FF8FFFA57FC5FE1F5D58FE01" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[151,294,403,429]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8FFFA57FC5FE1F5D58FE01" box="[151,294,403,429]" italics="true" pageId="106" pageNumber="107">V. pergandei</emphasis>
</taxonomicName>
occurs in more xeric microhabitats, i.e., in drier soils and/or at lower elevations (
<bibRefCitation id="EFAB4B1AFF8FFFA57B8BFE1F59EFFE01" author="Johnson, R. A." box="[1241,1425,403,429]" pageId="106" pageNumber="107" pagination="377 - 384" refId="ref69291" refString="Johnson, R. A. (2000 a) Reproductive biology of the seed-harvester ants Messor julianus (Pergande) and Messor pergandei (Mayr) (Hymenoptera: Formicidae) in Baja California, Mexico. Journal of Hymenoptera Research, 9 (2), 377 - 384." type="journal article" year="2000">Johnson, 2000a</bibRefCitation>
). The different micro- and macrohabitats occupied by 
<taxonomicName id="4C3A4D68FF8FFFA57DB4FE3B5F22FE7D" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[742,860,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF8FFFA57DB4FE3B5F22FE7D" box="[742,860,439,465]" italics="true" pageId="106" pageNumber="107">V. julianus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF8FFFA57CC1FE3B585EFE7D" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[915,1056,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8FFFA57CC1FE3B585EFE7D" box="[915,1056,439,465]" italics="true" pageId="106" pageNumber="107">V. pergandei</emphasis>
</taxonomicName>
correlate with queen body size as queens of 
<taxonomicName id="4C3A4D68FF8FFFA57E58FE575DE9FE59" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[266,407,475,501]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8FFFA57E58FE575DE9FE59" box="[266,407,475,501]" italics="true" pageId="106" pageNumber="107">V. pergandei</emphasis>
</taxonomicName>
are larger than those of 
<taxonomicName id="4C3A4D68FF8FFFA57DF4FE575F65FE59" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[678,795,475,501]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF8FFFA57DF4FE575F65FE59" box="[678,795,475,501]" italics="true" pageId="106" pageNumber="107">V. julianus</emphasis>
</taxonomicName>
(see also 
<bibRefCitation id="EFAB4B1AFF8FFFA57CD8FE5758E9FE59" author="Wiernasz, D. C. &amp; Cole, B. J." box="[906,1175,475,501]" pageId="106" pageNumber="107" pagination="2179 - 2183" refId="ref73898" refString="Wiernasz, D. C. &amp; Cole, B. J. (2003) Queen size mediates queen survival and colony fitness in harvester ants. Evolution, 57 (9), 2179 - 2183. https: // doi. org / 10.1111 / j. 0014 - 3820.2003. tb 00396. x" type="journal article" year="2003">Wiernasz &amp; Cole, 2003</bibRefCitation>
).
</paragraph>
<caption id="DF456663FF8FFFA57FC5F9335827F957" ID-DOI="http://doi.org/10.5281/zenodo.7318956" ID-Zenodo-Dep="7318956" httpUri="https://zenodo.org/record/7318956/files/figure.png" pageId="106" pageNumber="107" startId="106.[151,250,1727,1751]" targetBox="[173,1422,538,1690]" targetPageId="106">
<paragraph id="8B8536EBFF8FFFA57FC5F9335827F957" blockId="106.[151,1436,1727,1787]" pageId="106" pageNumber="107">
<emphasis id="B94EEAF9FF8FFFA57FC5F9335D5FF97B" bold="true" box="[151,289,1727,1751]" pageId="106" pageNumber="107">FIGURE 56.</emphasis>
Queen body size across species of 
<taxonomicName id="4C3A4D68FF8FFFA57DC3F94C5F78F97B" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[657,774,1728,1751]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF8FFFA57DC3F94C5F78F97B" box="[657,774,1728,1751]" italics="true" pageId="106" pageNumber="107">Veromessor</emphasis>
</taxonomicName>
based on average head width, average head length, and average mesosoma length. Species are ordered (left to right) from shortest to longest mesosoma length.
</paragraph>
</caption>
<paragraph id="8B8536EBFF8FFFA57F95F8A6581DF85B" blockId="106.[151,1437,1833,2040]" pageId="106" pageNumber="107">
<taxonomicName id="4C3A4D68FF8FFFA57F95F8A65DD2F8EF" baseAuthorityName="Pergande" baseAuthorityYear="1894" box="[199,428,1833,1859]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="julianus">
<emphasis id="B94EEAF9FF8FFFA57F95F8A65DD2F8EF" box="[199,428,1833,1859]" italics="true" pageId="106" pageNumber="107">Veromessor julianus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF8FFFA57EB5F8A65E33F8EF" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[487,589,1833,1859]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF8FFFA57EB5F8A65E33F8EF" box="[487,589,1833,1859]" italics="true" pageId="106" pageNumber="107">V. andrei</emphasis>
</taxonomicName>
contact each other only near Bahía San Quintin, 
<collectingRegion id="49FEF809FF8FFFA57BDAF8A55947F8EF" box="[1160,1337,1833,1859]" country="Mexico" name="Baja California" pageId="106" pageNumber="107">Baja California</collectingRegion>
, 
<collectingCountry id="F32D767BFF8FFFA57A15F8A559E3F8EF" box="[1351,1437,1833,1859]" name="Mexico" pageId="106" pageNumber="107">Mexico</collectingCountry>
(
<figureCitation id="13012A6EFF8FFFA57FCDF8C15D52F8CB" box="[159,300,1869,1895]" captionStart="FIGURE 10" captionStartId="25.[151,250,1943,1967]" captionTargetBox="[417,1169,181,1919]" captionTargetId="figure-18@25.[417,1170,181,1919]" captionTargetPageId="25" captionText="FIGURE 10. Geographic distribution of: (A) Veromessor andrei, (B) V. chamberlini, and (C) V. chicoensis. The larger black circle in each panel denotes the type locality. The northernmost locale for V. chamberlini was given only as Santa Clara County, and we have placed this locale near the center of the county." figureDoi="http://doi.org/10.5281/zenodo.7318850" httpUri="https://zenodo.org/record/7318850/files/figure.png" pageId="106" pageNumber="107">Figures 10A</figureCitation>
, 
<figureCitation id="13012A6EFF8FFFA57E65F8C15D12F8CB" box="[311,364,1869,1895]" captionStart="FIGURE 27" captionStartId="50.[151,250,1943,1967]" captionTargetBox="[417,1170,181,1920]" captionTargetId="figure-26@50.[417,1170,181,1920]" captionTargetPageId="50" captionText="FIGURE 27. Geographic distribution of: (A) Veromessor julianus, (B) V. lariversi, and (C) V. lobognathus. The larger black circle in each panel denotes the type locality." figureDoi="http://doi.org/10.5281/zenodo.7318888" httpUri="https://zenodo.org/record/7318888/files/figure.png" pageId="106" pageNumber="107">27A</figureCitation>
). 
<taxonomicName id="4C3A4D68FF8FFFA57E2EF8C25E0BF8CB" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[380,629,1869,1895]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8FFFA57E2EF8C25E0BF8CB" box="[380,629,1869,1895]" italics="true" pageId="106" pageNumber="107">Veromessor pergandei</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF8FFFA57DF5F8C25F77F8CB" baseAuthorityName="Mayr" baseAuthorityYear="1886" box="[679,777,1869,1895]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="andrei">
<emphasis id="B94EEAF9FF8FFFA57DF5F8C25F77F8CB" box="[679,777,1869,1895]" italics="true" pageId="106" pageNumber="107">V. andrei</emphasis>
</taxonomicName>
rarely coexist as the latter species inhabits the cool foothills of the California coastal range and inland valleys from northwestern 
<collectingRegion id="49FEF809FF8FFFA57CDFF8FD5846F827" box="[909,1080,1905,1931]" country="Mexico" name="Baja California" pageId="106" pageNumber="107">Baja California</collectingRegion>
to southern Oregon. In contrast, 
<taxonomicName id="4C3A4D68FF8FFFA57FC5F81A5D5BF803" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[151,293,1941,1967]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8FFFA57FC5F81A5D5BF803" box="[151,293,1941,1967]" italics="true" pageId="106" pageNumber="107">V. pergandei</emphasis>
</taxonomicName>
inhabits hot desert areas to the east and only occasionally penetrates to the adjacent coastal valleys, but rarely above elevations of 
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<geoCoordinate id="EE0E502CFF8FFFA57EBAF8355E59F87F" box="[488,551,1977,2003]" degrees="1,100" direction="north" orientation="latitude" pageId="106" pageNumber="107" precision="55" value="1.1">1,100</geoCoordinate>
<geoCoordinate id="EE0E502CFF8FFFA57D75F8355E0BF87F" box="[551,629,1977,2003]" degrees="1,200" direction="west" orientation="longitude" pageId="106" pageNumber="107" precision="55" value="-1.2">–1,200</geoCoordinate>
m
</quantity>
(see also 
<bibRefCitation id="EFAB4B1AFF8FFFA57DACF8355FD1F87F" author="Johnson, R. A." box="[766,943,1977,2003]" pageId="106" pageNumber="107" pagination="377 - 384" refId="ref69291" refString="Johnson, R. A. (2000 a) Reproductive biology of the seed-harvester ants Messor julianus (Pergande) and Messor pergandei (Mayr) (Hymenoptera: Formicidae) in Baja California, Mexico. Journal of Hymenoptera Research, 9 (2), 377 - 384." type="journal article" year="2000">
Johnson, 2000 
<bibRefCitation id="EFAB4B1AFF8FFFA57CF1F8355FD1F87F" author="Johnson, R. A." box="[931,943,1977,2003]" pageId="106" pageNumber="107" pagination="89 - 122" refId="ref69343" refString="Johnson, R. A. (2000 b) Seed-harvester ants (Hymenoptera: Formicidae) of North America: an overview of ecology and biogeography. Sociobiology, 36 (1), 89 - 122." type="journal article" year="2000">a</bibRefCitation>
</bibRefCitation>
, 2000b); there are also several records for 
<taxonomicName id="4C3A4D68FF8FFFA57ADAF8365D76F85B" baseAuthorityName="Rissing" baseAuthorityYear="1987" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="106" pageNumber="107" phylum="Arthropoda" rank="species" species="pergandei">
<emphasis id="B94EEAF9FF8FFFA57ADAF8365D76F85B" italics="true" pageId="106" pageNumber="107">V. pergandei</emphasis>
</taxonomicName>
in hot coastal areas of California such as in the Central Valley (
<figureCitation id="13012A6EFF8FFFA57C82F8515828F85B" box="[976,1110,2013,2039]" captionStart="FIGURE 41" captionStartId="78.[151,250,1943,1967]" captionTargetBox="[425,1163,181,1919]" captionTargetId="figure-26@78.[425,1163,181,1919]" captionTargetPageId="78" captionText="FIGURE 41. Geographic distribution of: (A) Veromessor pergandei, (B) V. pseudolariversi, and (C) V. smithi. The larger black circle in panels B &amp; C denotes the type locality." figureDoi="http://doi.org/10.5281/zenodo.7318924" httpUri="https://zenodo.org/record/7318924/files/figure.png" pageId="106" pageNumber="107">Figure 41A</figureCitation>
).
</paragraph>
<paragraph id="8B8536EBFF8EFFA47F95FF1B58F3FE01" blockId="107.[151,1437,151,1041]" pageId="107" pageNumber="108">
The other seven species of 
<taxonomicName id="4C3A4D68FF8EFFA47D50FF1B5EFAFF1C" baseAuthorityName="Brown" baseAuthorityYear="1949" box="[514,644,151,176]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="genus">
<emphasis id="B94EEAF9FF8EFFA47D50FF1B5EFAFF1C" box="[514,644,151,176]" italics="true" pageId="107" pageNumber="108">Veromessor</emphasis>
</taxonomicName>
have relatively small colonies with number of workers ranging from several hundred to 
<geoCoordinate id="EE0E502CFF8EFFA47E3EFF375DD4FF79" box="[364,426,187,213]" degrees="1,000" direction="north" orientation="latitude" pageId="107" pageNumber="108" precision="55" value="1.0">1,000</geoCoordinate>
<geoCoordinate id="EE0E502CFF8EFFA47EF8FF375D86FF79" box="[426,504,187,213]" degrees="2,000" direction="west" orientation="longitude" pageId="107" pageNumber="108" precision="55" value="-2.0">–2,000</geoCoordinate>
or more. Distribution patterns of these species can be divided into three groups: (1) three California species (
<taxonomicName id="4C3A4D68FF8EFFA47EFDFF535E2CFF55" box="[431,594,223,249]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="chamberlini">
<emphasis id="B94EEAF9FF8EFFA47EFDFF535E2CFF55" box="[431,594,223,249]" italics="true" pageId="107" pageNumber="108">V. chamberlini</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF8EFFA47D32FF535E8EFF55" authorityName="Smith" authorityYear="1956" box="[608,752,223,249]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF8EFFA47D32FF535E8EFF55" box="[608,752,223,249]" italics="true" pageId="107" pageNumber="108">V. chicoensis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF8EFFA47DACFF535FFDFF55" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[766,899,223,249]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF8EFFA47DACFF535FFDFF55" box="[766,899,223,249]" italics="true" pageId="107" pageNumber="108">V. stoddardi</emphasis>
</taxonomicName>
) that occur in non-desert areas of the California coastal range west of the Sierra Nevada from northern to southern California, with one species extending into central 
<collectingRegion id="49FEF809FF8EFFA47FC5FEAB5D38FEED" box="[151,326,295,321]" country="Mexico" name="Baja California" pageId="107" pageNumber="108">Baja California</collectingRegion>
, (2) four broadly sympatric species (
<taxonomicName id="4C3A4D68FF8EFFA47DB4FEAB5F1FFEED" box="[742,865,295,321]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF8EFFA47DB4FEAB5F1FFEED" box="[742,865,295,321]" italics="true" pageId="107" pageNumber="108">V. lariversi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF8EFFA47C3DFEAB586BFEED" baseAuthorityName="Andrews" baseAuthorityYear="1916" box="[879,1045,295,321]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="lobognathus">
<emphasis id="B94EEAF9FF8EFFA47C3DFEAB586BFEED" box="[879,1045,295,321]" italics="true" pageId="107" pageNumber="108">V. lobognathus</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF8EFFA47B76FEAB588CFEED" box="[1060,1266,295,321]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF8EFFA47B76FEAB584BFEEC" box="[1060,1077,295,320]" italics="true" pageId="107" pageNumber="108">V</emphasis>
. 
<emphasis id="B94EEAF9FF8EFFA47B16FEAB588CFEED" box="[1092,1266,295,321]" italics="true" pageId="107" pageNumber="108">pseudolariversi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF8EFFA47A53FEAB591FFEED" authorityName="Cole" authorityYear="1963" box="[1281,1377,295,321]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF8EFFA47A53FEAB591FFEED" box="[1281,1377,295,321]" italics="true" pageId="107" pageNumber="108">V. smithi</emphasis>
</taxonomicName>
) that occur east of the Sierra Nevada in California and are most common in Nevada, with ranges sometimes extending into Arizona and adjacent states, and (3) one species (
<emphasis id="B94EEAF9FF8EFFA47DBBFEE35E84FE24" box="[745,762,367,392]" italics="true" pageId="107" pageNumber="108">V</emphasis>
. sp. 
<taxonomicName id="4C3A4D68FF8EFFA47C7FFEE35FBEFE25" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[813,960,367,393]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="pergandei">
cf. 
<emphasis id="B94EEAF9FF8EFFA47C02FEE35FBEFE25" box="[848,960,367,393]" italics="true" pageId="107" pageNumber="108">pergandei</emphasis>
</taxonomicName>
) that appears to be restricted to sand dunes of hot desert habitats in southwestern Arizona and probably northwestern 
<collectingRegion id="49FEF809FF8EFFA47C81FE1F585BFE01" box="[979,1061,403,429]" country="Mexico" name="Sonora" pageId="107" pageNumber="108">Sonora</collectingRegion>
, 
<collectingCountry id="F32D767BFF8EFFA47B62FE1F58F7FE01" box="[1072,1161,403,429]" name="Mexico" pageId="107" pageNumber="108">Mexico</collectingCountry>
.
</paragraph>
<paragraph id="8B8536EBFF8EFFA47F95FE3B5914FD61" blockId="107.[151,1437,151,1041]" pageId="107" pageNumber="108">
The first group consists of three species (
<taxonomicName id="4C3A4D68FF8EFFA47DE0FE3B5F27FE7D" box="[690,857,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="chamberlini">
<emphasis id="B94EEAF9FF8EFFA47DE0FE3B5F27FE7D" box="[690,857,439,465]" italics="true" pageId="107" pageNumber="108">V. chamberlini</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF8EFFA47C39FE3B587EFE7D" authorityName="Smith" authorityYear="1956" box="[875,1024,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF8EFFA47C39FE3B587EFE7D" box="[875,1024,439,465]" italics="true" pageId="107" pageNumber="108">V. chicoensis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C3A4D68FF8EFFA47B40FE3B58E2FE7D" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[1042,1180,439,465]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF8EFFA47B40FE3B58E2FE7D" box="[1042,1180,439,465]" italics="true" pageId="107" pageNumber="108">V. stoddardi</emphasis>
</taxonomicName>
) that are restricted to 
<collectingRegion id="49FEF809FF8EFFA47FC5FE575D77FE59" box="[151,265,475,501]" country="United States of America" name="California" pageId="107" pageNumber="108">California</collectingRegion>
and 
<collectingRegion id="49FEF809FF8EFFA47E11FE575D8AFE59" box="[323,500,475,501]" country="Mexico" name="Baja California" pageId="107" pageNumber="108">Baja California</collectingRegion>
. 
<taxonomicName id="4C3A4D68FF8EFFA47D52FE575F7EFE59" authorityName="Smith" authorityYear="1956" box="[512,768,475,501]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF8EFFA47D52FE575F7EFE59" box="[512,768,475,501]" italics="true" pageId="107" pageNumber="108">Veromessor chicoensis</emphasis>
</taxonomicName>
is geographically isolated from the other two species as it is restricted to the Sacramento and San Joaquin Valleys of northern and central 
<collectingRegion id="49FEF809FF8EFFA47B7AFE7358E4FDB5" box="[1064,1178,511,537]" country="United States of America" name="California" pageId="107" pageNumber="108">California</collectingRegion>
(
<bibRefCitation id="EFAB4B1AFF8EFFA47BF8FE735CAFFD91" author="Hamm, C. A. &amp; Kamansky, B." pageId="107" pageNumber="108" pagination="543 - 547" refId="ref68677" refString="Hamm, C. A. &amp; Kamansky, B. (2009) New record of Messor chicoensis (Hymenoptera: Formicidae) from the San Joaquin Valley of California. Sociobiology, 53 (2), 543 - 547." type="journal article" year="2009">Hamm &amp; Kamansky, 2009</bibRefCitation>
) (
<figureCitation id="13012A6EFF8EFFA47FBBFDAF5D11FD91" box="[233,367,547,573]" captionStart="FIGURE 10" captionStartId="25.[151,250,1943,1967]" captionTargetBox="[417,1169,181,1919]" captionTargetId="figure-18@25.[417,1170,181,1919]" captionTargetPageId="25" captionText="FIGURE 10. Geographic distribution of: (A) Veromessor andrei, (B) V. chamberlini, and (C) V. chicoensis. The larger black circle in each panel denotes the type locality. The northernmost locale for V. chamberlini was given only as Santa Clara County, and we have placed this locale near the center of the county." figureDoi="http://doi.org/10.5281/zenodo.7318850" httpUri="https://zenodo.org/record/7318850/files/figure.png" pageId="107" pageNumber="108">Figure 10C</figureCitation>
). 
<taxonomicName id="4C3A4D68FF8EFFA47ED6FDAF5E06FD91" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[388,632,547,573]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF8EFFA47ED6FDAF5E06FD91" box="[388,632,547,573]" italics="true" pageId="107" pageNumber="108">Veromessor stoddardi</emphasis>
</taxonomicName>
is most closely related to 
<taxonomicName id="4C3A4D68FF8EFFA47CF5FDAF5847FD91" authorityName="Smith" authorityYear="1956" box="[935,1081,547,573]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF8EFFA47CF5FDAF5847FD91" box="[935,1081,547,573]" italics="true" pageId="107" pageNumber="108">V. chicoensis</emphasis>
</taxonomicName>
(M.L. Borowiec, unpub. data), and these two species have a mostly contiguous distribution with 
<taxonomicName id="4C3A4D68FF8EFFA47C38FDCB5F84FDCD" authorityName="Smith" authorityYear="1956" box="[874,1018,583,609]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="chicoensis">
<emphasis id="B94EEAF9FF8EFFA47C38FDCB5F84FDCD" box="[874,1018,583,609]" italics="true" pageId="107" pageNumber="108">V. chicoensis</emphasis>
</taxonomicName>
in northern coastal range areas, while 
<taxonomicName id="4C3A4D68FF8EFFA47FC5FDE75D62FD29" authorityName="Ward, Brady, Fisher, and Schultz" authorityYear="2015" box="[151,284,619,645]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="stoddardi">
<emphasis id="B94EEAF9FF8EFFA47FC5FDE75D62FD29" box="[151,284,619,645]" italics="true" pageId="107" pageNumber="108">V. stoddardi</emphasis>
</taxonomicName>
occurs in southern coastal range areas, with the nearest locales being about 
<quantity id="4CC29B0EFF8EFFA47B38FDE758C0FD29" box="[1130,1214,619,645]" metricMagnitude="5" metricUnit="m" metricValue="1.7" pageId="107" pageNumber="108" unit="km" value="170.0">170 km</quantity>
apart (
<figureCitation id="13012A6EFF8EFFA47A58FDE759E9FD29" box="[1290,1431,619,645]" captionStart="FIGURE 10" captionStartId="25.[151,250,1943,1967]" captionTargetBox="[417,1169,181,1919]" captionTargetId="figure-18@25.[417,1170,181,1919]" captionTargetPageId="25" captionText="FIGURE 10. Geographic distribution of: (A) Veromessor andrei, (B) V. chamberlini, and (C) V. chicoensis. The larger black circle in each panel denotes the type locality. The northernmost locale for V. chamberlini was given only as Santa Clara County, and we have placed this locale near the center of the county." figureDoi="http://doi.org/10.5281/zenodo.7318850" httpUri="https://zenodo.org/record/7318850/files/figure.png" pageId="107" pageNumber="108">Figures 10C</figureCitation>
, 
<figureCitation id="13012A6EFF8EFFA47FC5FD025CC9FD05" box="[151,183,654,681]" captionStart="FIGURE 52" captionStartId="97.[151,250,1652,1676]" captionTargetBox="[151,1434,633,1628]" captionTargetId="figure-213@97.[151,1435,633,1629]" captionTargetPageId="97" captionText="FIGURE 52. Geographic distribution of Veromessor stoddardi. The larger black circle denotes the type locality." figureDoi="http://doi.org/10.5281/zenodo.7318946" httpUri="https://zenodo.org/record/7318946/files/figure.png" pageId="107" pageNumber="108">52</figureCitation>
). 
<taxonomicName id="4C3A4D68FF8EFFA47F9EFD035DA0FD05" box="[204,478,655,681]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="chamberlini">
<emphasis id="B94EEAF9FF8EFFA47F9EFD035DA0FD05" box="[204,478,655,681]" italics="true" pageId="107" pageNumber="108">Veromessor chamberlini</emphasis>
</taxonomicName>
occurs on Santa Cruz and 
<taxonomicName id="4C3A4D68FF8EFFA47C46FD035F95FD05" authority="Rosa Islands" authorityName="Rosa Islands" box="[788,1003,655,681]" class="Insecta" family="Hesperiidae" genus="Santa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="genus">Santa Rosa Islands</taxonomicName>
and across locales along the southern base of the San Gabriel Mountains and in coastal areas north to near the San Francisco Bay area (
<figureCitation id="13012A6EFF8EFFA47B8BFD3F5923FD61" box="[1241,1373,691,717]" captionStart="FIGURE 10" captionStartId="25.[151,250,1943,1967]" captionTargetBox="[417,1169,181,1919]" captionTargetId="figure-18@25.[417,1170,181,1919]" captionTargetPageId="25" captionText="FIGURE 10. Geographic distribution of: (A) Veromessor andrei, (B) V. chamberlini, and (C) V. chicoensis. The larger black circle in each panel denotes the type locality. The northernmost locale for V. chamberlini was given only as Santa Clara County, and we have placed this locale near the center of the county." figureDoi="http://doi.org/10.5281/zenodo.7318850" httpUri="https://zenodo.org/record/7318850/files/figure.png" pageId="107" pageNumber="108">Figure 10B</figureCitation>
).
</paragraph>
<paragraph id="8B8536EBFF8EFFA47F95FD5B5FC9FC65" blockId="107.[151,1437,151,1041]" pageId="107" pageNumber="108">
The four species in the second group occur in 
<collectingRegion id="49FEF809FF8EFFA47D9FFD5B5F41FD5D" box="[717,831,727,753]" country="United States of America" name="California" pageId="107" pageNumber="108">California</collectingRegion>
east of the Sierra 
<collectingRegion id="49FEF809FF8EFFA47B5BFD5B5821FD5D" box="[1033,1119,727,753]" country="United States of America" name="Nevada" pageId="107" pageNumber="108">Nevada</collectingRegion>
and have broadly sympatric distributions. These species form two groups based on habitat of occurrence. 
<taxonomicName id="4C3A4D68FF8EFFA47CA6FD775978FCB9" baseAuthorityName="Andrews" baseAuthorityYear="1916" box="[1012,1286,763,789]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="lobognathus">
<emphasis id="B94EEAF9FF8EFFA47CA6FD775978FCB9" box="[1012,1286,763,789]" italics="true" pageId="107" pageNumber="108">Veromessor lobognathus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF8EFFA47A6FFD7759E2FCB9" authorityName="Cole" authorityYear="1963" box="[1341,1436,763,789]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="smithi">
<emphasis id="B94EEAF9FF8EFFA47A6FFD7759E2FCB9" box="[1341,1436,763,789]" italics="true" pageId="107" pageNumber="108">V. smithi</emphasis>
</taxonomicName>
typically inhabit mid-elevation, pinyon pine–juniper habitats, rocky bajadas, and buttes. Both species are sympatric at some locales (
<figureCitation id="13012A6EFF8EFFA47E1CFCCF5DA2FCF1" box="[334,476,835,861]" captionStart="FIGURE 27" captionStartId="50.[151,250,1943,1967]" captionTargetBox="[417,1170,181,1920]" captionTargetId="figure-26@50.[417,1170,181,1920]" captionTargetPageId="50" captionText="FIGURE 27. Geographic distribution of: (A) Veromessor julianus, (B) V. lariversi, and (C) V. lobognathus. The larger black circle in each panel denotes the type locality." figureDoi="http://doi.org/10.5281/zenodo.7318888" httpUri="https://zenodo.org/record/7318888/files/figure.png" pageId="107" pageNumber="108">Figures 27C</figureCitation>
, 
<figureCitation id="13012A6EFF8EFFA47EB4FCCF5E66FCF1" box="[486,536,835,861]" captionStart="FIGURE 41" captionStartId="78.[151,250,1943,1967]" captionTargetBox="[425,1163,181,1919]" captionTargetId="figure-26@78.[425,1163,181,1919]" captionTargetPageId="78" captionText="FIGURE 41. Geographic distribution of: (A) Veromessor pergandei, (B) V. pseudolariversi, and (C) V. smithi. The larger black circle in panels B &amp; C denotes the type locality." figureDoi="http://doi.org/10.5281/zenodo.7318924" httpUri="https://zenodo.org/record/7318924/files/figure.png" pageId="107" pageNumber="108">41C</figureCitation>
), but it is not known if these two species have different microhabitat preferences. Alternatively, 
<taxonomicName id="4C3A4D68FF8EFFA47E6BFCEB5DC7FC2D" box="[313,441,871,897]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="lariversi">
<emphasis id="B94EEAF9FF8EFFA47E6BFCEB5D34FC2C" box="[313,330,871,896]" italics="true" pageId="107" pageNumber="108">V</emphasis>
. 
<emphasis id="B94EEAF9FF8EFFA47E08FCEB5DC7FC2D" box="[346,441,871,897]" italics="true" pageId="107" pageNumber="108">lariversi</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C3A4D68FF8EFFA47EA0FCEB5EBFFC2D" box="[498,705,871,897]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="pseudolariversi">
<emphasis id="B94EEAF9FF8EFFA47EA0FCEB5E7DFC2C" box="[498,515,871,896]" italics="true" pageId="107" pageNumber="108">V</emphasis>
. 
<emphasis id="B94EEAF9FF8EFFA47D41FCEB5EBFFC2D" box="[531,705,871,897]" italics="true" pageId="107" pageNumber="108">pseudolariversi</emphasis>
</taxonomicName>
typically inhabit high desert habitats, often in sandy soils. Both species have a similar geographic distribution and both species are sympatric at some sites (
<figureCitation id="13012A6EFF8EFFA47BC6FC07595FFC09" box="[1172,1313,907,933]" captionStart="FIGURE 27" captionStartId="50.[151,250,1943,1967]" captionTargetBox="[417,1170,181,1920]" captionTargetId="figure-26@50.[417,1170,181,1920]" captionTargetPageId="50" captionText="FIGURE 27. Geographic distribution of: (A) Veromessor julianus, (B) V. lariversi, and (C) V. lobognathus. The larger black circle in each panel denotes the type locality." figureDoi="http://doi.org/10.5281/zenodo.7318888" httpUri="https://zenodo.org/record/7318888/files/figure.png" pageId="107" pageNumber="108">Figures 27B</figureCitation>
, 
<figureCitation id="13012A6EFF8EFFA47A7EFC075921FC09" box="[1324,1375,907,933]" captionStart="FIGURE 41" captionStartId="78.[151,250,1943,1967]" captionTargetBox="[425,1163,181,1919]" captionTargetId="figure-26@78.[425,1163,181,1919]" captionTargetPageId="78" captionText="FIGURE 41. Geographic distribution of: (A) Veromessor pergandei, (B) V. pseudolariversi, and (C) V. smithi. The larger black circle in panels B &amp; C denotes the type locality." figureDoi="http://doi.org/10.5281/zenodo.7318924" httpUri="https://zenodo.org/record/7318924/files/figure.png" pageId="107" pageNumber="108">41B</figureCitation>
). It is not known if these two species have different microhabitat preferences.
</paragraph>
<paragraph id="8B8536EBFF8EFFA47F95FC5F5DECFBBD" blockId="107.[151,1437,151,1041]" pageId="107" pageNumber="108">
The final species, 
<emphasis id="B94EEAF9FF8EFFA47ECCFC5F5DD1FC40" box="[414,431,979,1004]" italics="true" pageId="107" pageNumber="108">V</emphasis>
. sp. 
<taxonomicName id="4C3A4D68FF8EFFA47EB9FC5F5EFDFC41" baseAuthorityName="Rissing" baseAuthorityYear="1987" box="[491,643,979,1005]" class="Insecta" family="Formicidae" genus="Veromessor" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Hymenoptera" pageId="107" pageNumber="108" phylum="Arthropoda" rank="species" species="pergandei">
cf. 
<emphasis id="B94EEAF9FF8EFFA47D41FC5F5EFDFC41" box="[531,643,979,1005]" italics="true" pageId="107" pageNumber="108">pergandei</emphasis>
</taxonomicName>
, is only known from one hot desert habitat locale in sand dunes of southwestern 
<collectingRegion id="49FEF809FF8EFFA47E62FC7B5DF0FBBD" box="[304,398,1015,1041]" country="United States of America" name="Arizona" pageId="107" pageNumber="108">Arizona</collectingRegion>
.
</paragraph>
</subSubSection>
</treatment>
</document>