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<document ID-DOI="http://doi.org/10.5281/zenodo.3789718" ID-GBIF-Dataset="5a91b4b1-6213-407b-8f3a-a9a71c44189c" ID-GBIF-Taxon="163749525" ID-ISSN="13132970" ID-Zenodo-Dep="3789718" approvalRequired="12" approvalRequired_for_taxonomicNames="1" approvalRequired_for_textStreams="11" checkinTime="1587974682905" checkinUser="plazi" docAuthor="Ratcliffe, Brett" docDate="2010" docId="03D587E5FFFFFFE8FF4FE5F5549DFBCB" docLanguage="en" docName="ZK_article_2141.pdf" docOrigin="ZooKeys 34 (34)" docStyle="DocumentStylede.uka.ipd.idaho.easyIO.settings.Settings@75a6cdb" docStyleName="zookeys.2008.journal_article" docTitle="Halffterinetis violetae Moron &amp; Nogueira 2007" docType="treatment" docVersion="4" lastPageId="18" lastPageNumber="123" masterDocId="FFECFF9DFFF3FFFAFFC3E1755657FFFA" masterDocTitle="A review of the Blaesiina (Coleoptera, Scarabaeidae, Cetoniinae, Gymnetini)" masterLastPageNumber="128" masterPageNumber="105" pageId="12" pageNumber="117" updateTime="1643562158794" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-3.0">
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<mods:titleInfo>
<mods:title>A review of the Blaesiina (Coleoptera, Scarabaeidae, Cetoniinae, Gymnetini)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ratcliffe, Brett</mods:namePart>
<mods:affiliation>University of Nebraska, Lincoln, NE, United States of America</mods:affiliation>
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<mods:title>ZooKeys</mods:title>
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<mods:part>
<mods:date>2010</mods:date>
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<mods:number>2010-01-28</mods:number>
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<mods:detail type="volume">
<mods:number>34</mods:number>
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<mods:number>34</mods:number>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.3789718" ID-GBIF-Taxon="163749525" ID-Zenodo-Dep="3789718" LSID="urn:lsid:plazi:treatment:03D587E5FFFFFFE8FF4FE5F5549DFBCB" httpUri="http://treatment.plazi.org/id/03D587E5FFFFFFE8FF4FE5F5549DFBCB" lastPageId="18" lastPageNumber="123" pageId="12" pageNumber="117">
<subSubSection box="[140,713,1151,1179]" pageId="12" pageNumber="117" type="nomenclature">
<paragraph blockId="12.[140,713,1151,1214]" box="[140,713,1151,1179]" pageId="12" pageNumber="117">
<heading box="[140,713,1151,1179]" fontSize="11" level="2" pageId="12" pageNumber="117" reason="3">
<taxonomicName authority="Moron &amp; Nogueira, 2007" authorityName="Moron &amp; Nogueira" authorityYear="2007" box="[140,713,1151,1179]" class="Insecta" family="Cetoniidae" genus="Halffterinetis" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="117" phylum="Arthropoda" rank="species" species="violetae">
<emphasis bold="true" box="[140,713,1151,1179]" pageId="12" pageNumber="117">
<emphasis bold="true" box="[140,398,1152,1179]" italics="true" pageId="12" pageNumber="117">Halffterinetis violetae</emphasis>
<bibRefCitation author="Moron MA &amp; Nogueira G" box="[405,713,1151,1178]" pageId="12" pageNumber="117" pagination="51 - 61" refId="ref9457" refString="Moron MA, Nogueira G (2007) Halffterinetis, nuevo genero mexicano de Cetoniidae (Coleoptera: Scarabaeoidea). In: Zunino M, Melic A (Eds) Escarabajos, Diversidad y Conservacion Biologica. Sociedad Entomologica Aragonesa, Monografias 3 ercer Milenio M 3 M, Volume 7: 51 - 61." type="journal article" year="2007">Morón &amp; Nogueira, 2007</bibRefCitation>
</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection box="[140,250,1187,1214]" pageId="12" pageNumber="117" type="description">
<paragraph blockId="12.[140,713,1151,1214]" box="[140,250,1187,1214]" pageId="12" pageNumber="117">
<figureCitation box="[140,208,1187,1214]" captionStart="Figure 5" captionStartId="8.[139,211,1654,1676]" captionText="Figure 5. Distribution map of Blaesia and Halffterinetis species. Open symbol is state record only." pageId="12" pageNumber="117">Figs 5</figureCitation>
, 11
</paragraph>
</subSubSection>
<subSubSection box="[140,719,1258,1285]" pageId="12" pageNumber="117" type="reference_group">
<paragraph blockId="12.[140,719,1258,1285]" box="[140,719,1258,1285]" pageId="12" pageNumber="117">
<treatmentCitationGroup box="[140,719,1258,1285]" pageId="12" pageNumber="117">
<treatmentCitation author="Moron MA &amp; Nogueira G" box="[140,715,1258,1285]" page="56" pageId="12" pageNumber="117" year="2007">
<taxonomicName authority="Moron &amp; Nogueira 2007: 56" authorityName="Moron &amp; Nogueira" authorityPageNumber="56" authorityYear="2007" box="[140,715,1258,1285]" class="Insecta" family="Cetoniidae" genus="Halffterinetis" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="117" phylum="Arthropoda" rank="species" species="violetae">
<emphasis box="[140,371,1258,1284]" italics="true" pageId="12" pageNumber="117">Halffterinetis violetae</emphasis>
<bibRefCitation author="Moron MA &amp; Nogueira G" box="[378,715,1258,1285]" pageId="12" pageNumber="117" pagination="51 - 61" refId="ref9457" refString="Moron MA, Nogueira G (2007) Halffterinetis, nuevo genero mexicano de Cetoniidae (Coleoptera: Scarabaeoidea). In: Zunino M, Melic A (Eds) Escarabajos, Diversidad y Conservacion Biologica. Sociedad Entomologica Aragonesa, Monografias 3 ercer Milenio M 3 M, Volume 7: 51 - 61." type="journal article" year="2007">Morón &amp; Nogueira 2007: 56</bibRefCitation>
</taxonomicName>
</treatmentCitation>
.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection pageId="12" pageNumber="117" type="materials_examined">
<paragraph blockId="12.[140,1108,1328,1672]" pageId="12" pageNumber="117">
<materialsCitation ID-GBIF-Occurrence="2608851043" collectingDate="1961-06-18" collectionCode="CMNC, MXAL" collectorName="D. H. Janzen." country="Mexico" location="Hwy" municipality="Types" pageId="12" pageNumber="117" specimenCount="2" stateProvince="Durango" typeStatus="holotype">
<typeStatus box="[140,247,1328,1354]" pageId="12" pageNumber="117">Holotype</typeStatus>
male at
<collectionCode box="[343,434,1328,1354]" pageId="12" pageNumber="117">CMNC</collectionCode>
and one
<typeStatus box="[539,636,1329,1355]" pageId="12" pageNumber="117">paratype</typeStatus>
male at
<collectionCode box="[732,816,1328,1354]" pageId="12" pageNumber="117">MXAL</collectionCode>
.
<collectingMunicipality box="[825,890,1329,1355]" pageId="12" pageNumber="117">Types</collectingMunicipality>
labeled
<collectingCountry box="[984,1104,1328,1354]" name="Mexico" pageId="12" pageNumber="117">MEXICO</collectingCountry>
:
<collectingRegion box="[140,245,1364,1390]" country="Mexico" name="Durango" pageId="12" pageNumber="117">Durango</collectingRegion>
,
<locationDeviation box="[256,474,1363,1390]" location="Torreon" pageId="12" pageNumber="117">
<quantity box="[256,324,1363,1390]" metricMagnitude="4" metricUnit="m" metricValue="6.437376" pageId="12" pageNumber="117" unit="mi" value="40.0">40 mi</quantity>
SW Torreón
</locationDeviation>
,
<location LSID="urn:lsid:plazi:treatment:03D587E5FFFFFFE8FF4FE5F5549DFBCB:8EA36028FFFFFFF6FE26E421544AFA94" box="[485,541,1364,1390]" country="Mexico" municipality="Types" name="Hwy" pageId="12" pageNumber="117" stateProvince="Durango">Hwy</location>
40,
<date box="[592,732,1363,1389]" pageId="12" pageNumber="117" value="1961-06-18">
<collectingDate box="[592,732,1363,1389]" pageId="12" pageNumber="117" value="1961-06-18">18-VI-1961</collectingDate>
</date>
,
<collectorName box="[743,901,1364,1390]" pageId="12" pageNumber="117">D. H. Janzen.</collectorName>
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection lastPageId="13" lastPageNumber="118" pageId="12" pageNumber="117" type="description">
<paragraph blockId="12.[140,1108,1328,1672]" lastBlockId="13.[140,1108,167,1074]" lastPageId="13" lastPageNumber="118" pageId="12" pageNumber="117">
<emphasis bold="true" box="[185,340,1398,1424]" pageId="12" pageNumber="117">
<emphasis box="[185,335,1398,1424]" inLineHeading="true" pageId="12" pageNumber="117" reason="1">Description</emphasis>
.
</emphasis>
Male (female unknown). Length
<quantity box="[745,925,1398,1425]" metricMagnitude="-2" metricUnit="m" metricValue="1.4949999999999999" metricValueMax="1.53" metricValueMin="1.46" pageId="12" pageNumber="117" unit="mm" value="14.95" valueMax="15.3" valueMin="14.6">14.615.3 mm</quantity>
; width
<quantity metricMagnitude="-3" metricUnit="m" metricValue="8.15" metricValueMax="8.4" metricValueMin="7.9" pageId="12" pageNumber="117" unit="mm" value="8.15" valueMax="8.4" valueMin="7.9">7.98.4 mm</quantity>
. Color black, shining, lacking cretaceous marks.
<emphasis box="[772,832,1434,1460]" italics="true" pageId="12" pageNumber="117">Head</emphasis>
: Surface densely punctate to rugopunctate; punctures small to large, deep. Frontoclypeal region lacking transverse ridge. Frons with short, moderately dense, black setae. Clypeus with apex broadly truncate, strongly emarginate at middle, slightly reflexed, subapex distinctly wider than base, surface weakly concave. Interocular width equals 3.9 transverse eye diameters. Antenna black, with 10 segments, club almost twice as long as antennomeres 27.
<emphasis box="[464,576,1646,1671]" italics="true" pageId="12" pageNumber="117">Pronotum</emphasis>
: Surface densely punctate; punctures moder- ate to large, deep, round to slightly transverse, punctures becoming larger to rugopunctate on sides, and with short, tawny setae. Sides margined, bead reduced in anterior fifth. Mesepimeron completely punctate, with sparse, black setae.
<emphasis box="[1033,1100,237,263]" italics="true" pageId="13" pageNumber="118">Elytra</emphasis>
: Surface superficially and irregularly striate, rugopunctate, punctures moderate to large, round to mostly ∩-shaped, setigerous; setae short, sparse, black. Bead present on lateral margin. Apical umbone pronounced. Apices nearly right-angled.
<taxonomicName authority=": Surface" authorityName="Surface" authorityYear="1834" class="Actinopterygii" family="Trichomycteridae" genus="Pygidium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Siluriformes" pageId="13" pageNumber="118" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="13" pageNumber="118">Pygidium</emphasis>
: Surface
</taxonomicName>
with oval punctures or with short, transverse strigae (often reduced) to transversely strigose. Base usually with sparse, short, black setae. In lateral view surface weakly convex.
<emphasis box="[419,490,449,474]" italics="true" pageId="13" pageNumber="118">Venter</emphasis>
: Setae black. Mesometasternal process short, nearly obsolete, flat, apex rounded. Abdominal sternites with transverse, irregular field of large punctures; punctures sparser in central third, mostly large, shallow, with short, black setae.
<emphasis box="[356,403,555,580]" italics="true" pageId="13" pageNumber="118">Legs</emphasis>
: Femora and tibiae with sparse fringe of mostly short, black setae on median surface. Protibia tridentate, apical tooth longer, slightly narrower. Metafemur normal, not enlarged. Metatibia at apex with 2 broad lobes and with 2 long, articulated apical spurs with apices rounded. Metatrochanter triangular, elongate, flush with posterior margin of metafemur, acuminate apex not projecting perpendicularly.
<emphasis box="[336,449,731,756]" italics="true" pageId="13" pageNumber="118">Parameres</emphasis>
: Fig.11.
</paragraph>
</subSubSection>
<subSubSection box="[185,1108,765,792]" pageId="13" pageNumber="118" type="distribution">
<paragraph blockId="13.[140,1108,167,1074]" box="[185,1108,765,792]" pageId="13" pageNumber="118">
<emphasis bold="true" box="[185,336,765,791]" inLineHeading="true" pageId="13" pageNumber="118" reason="1">Distribution</emphasis>
(
<figureCitation box="[350,416,765,792]" captionStart="Figure 5" captionStartId="8.[139,211,1654,1676]" captionText="Figure 5. Distribution map of Blaesia and Halffterinetis species. Open symbol is state record only." pageId="13" pageNumber="118">Fig. 5</figureCitation>
). Two specimens recorded from
<bibRefCitation author="Moron MA &amp; Nogueira G" box="[778,1104,765,791]" pageId="13" pageNumber="118" pagination="51 - 61" refId="ref9457" refString="Moron MA, Nogueira G (2007) Halffterinetis, nuevo genero mexicano de Cetoniidae (Coleoptera: Scarabaeoidea). In: Zunino M, Melic A (Eds) Escarabajos, Diversidad y Conservacion Biologica. Sociedad Entomologica Aragonesa, Monografias 3 ercer Milenio M 3 M, Volume 7: 51 - 61." type="journal article" year="2007">Morón and Nogueira (2007)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="13" pageNumber="118" type="materials_examined">
<paragraph blockId="13.[140,1108,167,1074]" box="[185,940,800,827]" pageId="13" pageNumber="118">
<emphasis bold="true" box="[185,302,800,826]" pageId="13" pageNumber="118">
<collectingCountry box="[185,302,800,826]" name="Mexico" pageId="13" pageNumber="118">MEXICO</collectingCountry>
</emphasis>
(2):
<collectingRegion box="[358,508,800,826]" country="Mexico" name="Durango" pageId="13" pageNumber="118">DURANGO</collectingRegion>
(2): Torreón (
<quantity box="[668,737,800,826]" metricMagnitude="4" metricUnit="m" metricValue="6.437376" pageId="13" pageNumber="118" unit="mi" value="40.0">40 mi</quantity>
SW on Hwy 40).
</paragraph>
<paragraph blockId="13.[140,1108,167,1074]" box="[185,572,835,862]" pageId="13" pageNumber="118">
<emphasis bold="true" box="[185,466,835,861]" pageId="13" pageNumber="118">
<emphasis box="[185,299,835,861]" inLineHeading="true" pageId="13" pageNumber="118" reason="1">Temporal</emphasis>
Distribution.
</emphasis>
June (2).
</paragraph>
</subSubSection>
<subSubSection lastPageId="18" lastPageNumber="123" pageId="13" pageNumber="118" type="biology_ecology">
<paragraph blockId="13.[140,1108,167,1074]" pageId="13" pageNumber="118">
<emphasis bold="true" box="[185,283,870,896]" pageId="13" pageNumber="118">
<emphasis box="[185,278,870,896]" inLineHeading="true" pageId="13" pageNumber="118" reason="1">Biology</emphasis>
.
</emphasis>
<bibRefCitation author="Moron MA &amp; Nogueira G" box="[292,603,871,897]" pageId="13" pageNumber="118" pagination="51 - 61" refId="ref9457" refString="Moron MA, Nogueira G (2007) Halffterinetis, nuevo genero mexicano de Cetoniidae (Coleoptera: Scarabaeoidea). In: Zunino M, Melic A (Eds) Escarabajos, Diversidad y Conservacion Biologica. Sociedad Entomologica Aragonesa, Monografias 3 ercer Milenio M 3 M, Volume 7: 51 - 61." type="journal article" year="2007">Morón &amp; Nogueira (2007)</bibRefCitation>
indicated the habitat where this species was collected is semiarid with an average annual temperature of 1218°C and located at 12871300 meters above sea level. Dominant plants consisted of
<taxonomicName box="[921,1050,941,967]" class="Magnoliopsida" family="Asteraceae" genus="Parthenium" kingdom="Plantae" order="Asterales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="genus">
<emphasis box="[921,1050,941,967]" italics="true" pageId="13" pageNumber="118">Parthenium</emphasis>
</taxonomicName>
species (
<taxonomicName box="[197,321,976,1002]" class="Magnoliopsida" family="Asteraceae" kingdom="Plantae" order="Asterales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="family">Asteraceae</taxonomicName>
),
<taxonomicName box="[341,460,977,1002]" class="Magnoliopsida" family="Fouquieriaceae" genus="Fouquieria" kingdom="Plantae" order="Ericales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="genus">
<emphasis box="[341,460,977,1002]" italics="true" pageId="13" pageNumber="118">Fouquieria</emphasis>
</taxonomicName>
species (
<taxonomicName box="[562,738,976,1003]" class="Magnoliopsida" family="Fouquieriaceae" kingdom="Plantae" order="Ericales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="family">Fouquieriaceae</taxonomicName>
),
<taxonomicName box="[758,830,977,1002]" class="Magnoliopsida" family="Zygophyllaceae" genus="Larrea" kingdom="Plantae" order="Zygophyllales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="genus">
<emphasis box="[758,830,977,1002]" italics="true" pageId="13" pageNumber="118">Larrea</emphasis>
</taxonomicName>
species (Zygophiliacae),
<taxonomicName authorityName="Klotzsch" authorityYear="1835" box="[140,225,1011,1037]" class="Liliopsida" family="Bromeliaceae" genus="Hechtia" kingdom="Plantae" order="Poales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="genus">
<emphasis box="[140,225,1011,1037]" italics="true" pageId="13" pageNumber="118">Hechtia</emphasis>
</taxonomicName>
species (
<taxonomicName box="[329,486,1011,1038]" class="Liliopsida" family="Bromeliaceae" kingdom="Plantae" order="Poales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="family">Bromeliaceae</taxonomicName>
),
<taxonomicName box="[506,599,1012,1038]" class="Magnoliopsida" family="Cactaceae" genus="Opuntia" kingdom="Plantae" order="Caryophyllales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="genus">
<emphasis box="[506,599,1012,1038]" italics="true" pageId="13" pageNumber="118">Opuntia</emphasis>
</taxonomicName>
species (
<taxonomicName authorityName="de Jussieu" authorityYear="1789" box="[703,821,1012,1038]" class="Magnoliopsida" family="Cactaceae" kingdom="Plantae" order="Caryophyllales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="family">Cactaceae</taxonomicName>
),
<taxonomicName box="[843,907,1012,1038]" class="Liliopsida" family="Asparagaceae" genus="Agave" kingdom="Plantae" order="Asparagales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="genus">
<emphasis box="[843,907,1012,1038]" italics="true" pageId="13" pageNumber="118">Agave</emphasis>
</taxonomicName>
species (Amarilidaceae), and
<taxonomicName box="[287,401,1047,1073]" class="Magnoliopsida" family="Euphorbiaceae" genus="Euphorbia" kingdom="Plantae" order="Malpighiales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="genus">
<emphasis box="[287,401,1047,1073]" italics="true" pageId="13" pageNumber="118">Euphorbia</emphasis>
</taxonomicName>
species (
<taxonomicName box="[500,674,1047,1074]" class="Magnoliopsida" family="Euphorbiaceae" kingdom="Plantae" order="Malpighiales" pageId="13" pageNumber="118" phylum="Tracheophyta" rank="family">Euphorbiaceae</taxonomicName>
).
</paragraph>
<paragraph blockId="13.[140,326,1167,1194]" box="[140,326,1167,1194]" pageId="13" pageNumber="118">
<emphasis bold="true" box="[140,326,1167,1194]" pageId="13" pageNumber="118">Biogeography</emphasis>
</paragraph>
<paragraph blockId="13.[140,1108,1222,1672]" pageId="13" pageNumber="118">
The distribution of blaesiine species (
<figureCitation box="[565,633,1222,1249]" captionStart="Figure 5" captionStartId="8.[139,211,1654,1676]" captionText="Figure 5. Distribution map of Blaesia and Halffterinetis species. Open symbol is state record only." pageId="13" pageNumber="118">Fig. 5</figureCitation>
) nearly mirrors that of
<taxonomicName authorityName="Martinez" authorityYear="1949" box="[907,1054,1223,1249]" class="Insecta" family="Cetoniidae" genus="Hologymnetis" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="118" phylum="Arthropoda" rank="genus">
<emphasis box="[907,1054,1223,1249]" italics="true" pageId="13" pageNumber="118">Hologymnetis</emphasis>
</taxonomicName>
species, another genus within the
<taxonomicName authorityName="Kirby" authorityYear="1827" box="[492,620,1258,1284]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="118" phylum="Arthropoda" rank="tribe" tribe="Gymnetini">Gymnetini</taxonomicName>
. The biogeography of
<taxonomicName authorityName="Martinez" authorityYear="1949" box="[876,1023,1258,1284]" class="Insecta" family="Cetoniidae" genus="Hologymnetis" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="118" phylum="Arthropoda" rank="genus">
<emphasis box="[876,1023,1258,1284]" italics="true" pageId="13" pageNumber="118">Hologymnetis</emphasis>
</taxonomicName>
species was reviewed by
<bibRefCitation author="Ratcliffe BC &amp; Deloya AC" box="[333,653,1293,1319]" pageId="13" pageNumber="118" pagination="161 - 202" refId="ref9772" refString="Ratcliffe BC, Deloya AC (1992) The biogeography and phylogeny of Hologymnetis (Coleoptera: Scrabaeidae: Cetoniinae) with a revision of the genus. Coleopterists Bulletin 46: 161 - 202." type="journal article" year="1992">Ratcliffe and Deloya (1992)</bibRefCitation>
and is reviewed here to understand the similar distribution of the
<taxonomicName authorityName="Schoch" authorityYear="1895" box="[447,551,1328,1354]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="118" phylum="Arthropoda" rank="subTribe" subTribe="Blaesiina">Blaesiina</taxonomicName>
. Lacking fossil evidence for the
<taxonomicName authorityName="Kirby" authorityYear="1827" box="[923,1050,1328,1354]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="118" phylum="Arthropoda" rank="tribe" tribe="Gymnetini">Gymnetini</taxonomicName>
, it is necessary to rely upon data from plate tectonics, present and paleodistribution of other plants and animals, and ecological factors to formulate a hypothesis to best explain the current distribution of these insects.
</paragraph>
<paragraph blockId="13.[140,1108,1222,1672]" lastBlockId="14.[140,1108,167,1672]" lastPageId="14" lastPageNumber="119" pageId="13" pageNumber="118">
As with most other genera of New World
<taxonomicName authorityName="Kirby" authorityYear="1827" box="[661,788,1469,1495]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="118" phylum="Arthropoda" rank="tribe" tribe="Gymnetini">Gymnetini</taxonomicName>
, the ancestral
<taxonomicName authorityName="Schoch" authorityYear="1895" box="[948,1048,1469,1495]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="118" phylum="Arthropoda" rank="subTribe" subTribe="Blaesiina">Blaesiina</taxonomicName>
were present in South America prior to the establishment of the Panamanian land bridge in the Miocene. Given the current distribution and ecological requirements of
<taxonomicName authorityName="Schoch" authorityYear="1895" box="[1008,1108,1539,1565]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="118" phylum="Arthropoda" rank="subTribe" subTribe="Blaesiina">Blaesiina</taxonomicName>
species, it is assumed that they evolved in drier habitats. Drier habitats have been present in South America since middle Tertiary time (
<bibRefCitation author="Webb SD" box="[663,803,1610,1636]" pageId="13" pageNumber="118" pagination="393 - 426" refId="ref10424" refString="Webb SD (1978) A history of savanna vertebrates in the New World, part 11: South America and the great interchange. Annual Review of Ecology and Systematics 9: 393 - 426." type="journal article" year="1978">Webb 1978</bibRefCitation>
). The Quaternary (
<emphasis box="[1023,1064,1611,1636]" italics="true" pageId="13" pageNumber="118">i.e.,</emphasis>
the last two million years) is characterized by great environmental instability (
<bibRefCitation author="Bigarella JJ &amp; Andrade-Lima D" lastPageId="14" lastPageNumber="119" pageId="13" pageNumber="118" pagination="27 - 40" refId="ref8368" refString="Bigarella JJ, Andrade-Lima D (1982) Paleoenvironmental changes in Brazil In: Prance, GT (Ed) Biological diversification in the tropics. Columbia University Press, New York, NY 27 - 40." type="book chapter" year="1982">Bigarella and Andrade-Lima 1982</bibRefCitation>
;
<bibRefCitation author="Whitmore TC &amp; Prance GT" box="[390,700,167,193]" pageId="14" pageNumber="119" refId="ref10544" refString="Whitmore TC, Prance GT (Eds) (1987) Biogeography and Quaternary History in Tropical America. Clarendon Press, Oxford, United Kingdom, 224 pp." type="book" year="1987">Whitmore &amp; Prance 1987</bibRefCitation>
). These climatic changes caused, in relatively short geologic time, successive expansions and contractions of either forest or open, drier vegetation. Forest and nonforest biomes broke up into isolated blocks or expanded and coalesced depending on varying humid or arid climatic conditions (
<bibRefCitation author="Haffer J" box="[147,290,307,333]" pageId="14" pageNumber="119" pagination="131 - 137" refId="ref8866" refString="Haffer J (1969) Speciation in Amazonian forest birds. Science 165: 131 - 137." type="journal article" year="1969">Haffer 1969</bibRefCitation>
,
<bibRefCitation author="Haffer J" box="[302,363,308,334]" pageId="14" pageNumber="119" pagination="6 - 24" refId="ref8884" refString="Haffer J (1982) General aspects of the refuge theory. In: Prance GT (Ed) Biological Diversification in the Tropics. Columbia University Press, New York, NY, 6 - 24." type="book chapter" year="1982">1982</bibRefCitation>
;
<bibRefCitation author="Vuilleumier BS" box="[374,575,307,334]" pageId="14" pageNumber="119" pagination="771 - 780" refId="ref10401" refString="Vuilleumier BS (1971) Pleistocene changes in the fauna and flora of South America. Science 173: 771 - 780." type="journal article" year="1971">Vuilleumier 1971</bibRefCitation>
;
<bibRefCitation author="Muller P" box="[587,731,307,334]" pageId="14" pageNumber="119" refId="ref9591" refString="Muller P (1973) The dispersal centers of terrestrial vertebrates in the Neotropical realm. Biogeographica 11. W. Junk, The Hague, Netherlands." type="book" year="1973">Müller 1973</bibRefCitation>
;
<bibRefCitation author="Prance GT" box="[743,886,308,334]" pageId="14" pageNumber="119" pagination="5 - 28" refId="ref9619" refString="Prance GT (1973) Phytogeographic support for the theory of Pleistocene forest refuges in the Amazon Basin, based on evidence from distribution patterns in Caryocaraceae, Chrysobalanaceae, Dichapetalaceae, and Lecythidaceae. Acta Amazonica 3: 5 - 28." type="journal article" year="1973">Prance 1973</bibRefCitation>
,
<bibRefCitation author="Prance GT" box="[898,959,308,334]" pageId="14" pageNumber="119" refId="ref9662" refString="Prance GT (Ed) (1982) Biological Diversification in the Tropics. Columbia University Press, New York, NY, 714 pp." type="book" year="1982">1982</bibRefCitation>
;
<bibRefCitation author="Brown KS &amp; Shepard PM &amp; Turner JRG" pageId="14" pageNumber="119" pagination="369 - 378" refId="ref8534" refString="Brown KS, Shepard PM, Turner JRG (1974) Quaternary refugia in tropical America: evidence from race formation in Heliconius butterflies. Proceedings of the Royal Society of London (B) 187: 369 - 378." type="journal article" year="1974">Brown et al. 1974</bibRefCitation>
;
<bibRefCitation author="Brown KS" box="[215,360,343,369]" pageId="14" pageNumber="119" pagination="75 - 137" refId="ref8496" refString="Brown KS (1977) Centros de evolucao refugias quaternarios e conservaca de patrimonias geneticos na regao neotropical: padroes de diferencao em Ithomiinae (Lepidoptera: Nymphalidae). Acta Amazonica 7: 75 - 137." type="journal article" year="1977">Brown 1977</bibRefCitation>
;
<bibRefCitation author="Tricart J" box="[372,517,343,369]" pageId="14" pageNumber="119" pagination="145 - 158" refId="ref10352" refString="Tricart J (1974) Existence de periodes seches au Quaternaire en Amazonie et dans les regions voisines. Revue de Geomorphologie Dynamique 4: 145 - 158." type="journal article" year="1974">Tricart 1974</bibRefCitation>
;
<bibRefCitation author="Simpson BB &amp; Haffer J" box="[531,834,343,369]" pageId="14" pageNumber="119" pagination="497 - 518" refId="ref10100" refString="Simpson BB, Haffer J (1978) Speciation patterns in the Amazonian forest biota. Annual Review of Ecology and Systematics 9: 497 - 518." type="journal article" year="1978">Simpson and Haffer 1978</bibRefCitation>
). With the reduction of forest vegetation during drier periods, there was a corresponding increase in nonforest formations that penetrated into the Amazon region from both north and south. Such formations consisted of large blocks or corridors connecting the open vegetation associations of the Central Brazilian Plateau with those of
<collectingCountry box="[796,908,483,509]" name="Venezuela" pageId="14" pageNumber="119">Venezuela</collectingCountry>
and the Guianas (
<bibRefCitation author="Eden MJ" box="[148,276,519,545]" pageId="14" pageNumber="119" pagination="95 - 109" refId="ref8750" refString="Eden MJ (1974) Paleoclimatic influences and the development of savanna in southern Venezuela. Journal of Biogeography 1: 95 - 109." type="journal article" year="1974">Eden 1974</bibRefCitation>
;
<bibRefCitation author="Sarmiento G" box="[288,473,518,545]" pageId="14" pageNumber="119" pagination="233 - 251" refId="ref9967" refString="Sarmiento G (1975) The dry plant formations of South America and their floristic connections. Journal of Biogeography 2: 233 - 251." type="journal article" year="1975">Sarmiento 1975</bibRefCitation>
;
<bibRefCitation author="Bigarella JJ &amp; Andrade-Lima D" box="[485,872,519,545]" pageId="14" pageNumber="119" pagination="27 - 40" refId="ref8368" refString="Bigarella JJ, Andrade-Lima D (1982) Paleoenvironmental changes in Brazil In: Prance, GT (Ed) Biological diversification in the tropics. Columbia University Press, New York, NY 27 - 40." type="book chapter" year="1982">Bigarella and Andrade-Lima 1982</bibRefCitation>
;
<bibRefCitation author="Marshall G" box="[884,1053,518,545]" pageId="14" pageNumber="119" pagination="49 - 85" refId="ref9242" refString="Marshall G (1985) Geochronology and land-mammal biochronology of the transamerican faunal interchange. In: Stehli FG, Webb SD (Eds) The Great American Biotic Interchange. Plenum Press, New York, NY, 49 - 85." type="book chapter" year="1985">Marshall 1985</bibRefCitation>
).
</paragraph>
<paragraph blockId="14.[140,1108,167,1672]" pageId="14" pageNumber="119">
Present day
<taxonomicName authorityName="Burmeister" authorityYear="1842" box="[319,395,554,580]" class="Insecta" family="Cetoniidae" genus="Blaesia" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="119" phylum="Arthropoda" rank="genus">
<emphasis box="[319,395,554,580]" italics="true" pageId="14" pageNumber="119">Blaesia</emphasis>
</taxonomicName>
species inhabit the mesic to semiarid, relatively open vegetation habitats of
<collectingCountry box="[274,356,589,615]" name="Bolivia" pageId="14" pageNumber="119">Bolivia</collectingCountry>
,
<collectingCountry box="[371,473,590,616]" name="Paraguay" pageId="14" pageNumber="119">Paraguay</collectingCountry>
,
<collectingCountry box="[488,587,590,616]" name="Uruguay" pageId="14" pageNumber="119">Uruguay</collectingCountry>
,
<collectingCountry box="[603,718,589,615]" name="Argentina" pageId="14" pageNumber="119">Argentina</collectingCountry>
, and southern
<collectingCountry box="[897,966,589,615]" name="Brazil" pageId="14" pageNumber="119">Brazil</collectingCountry>
. The broad, continuous band of present-day Amazonian rainforest is a barrier preventing further dispersal northward. Ancestral
<taxonomicName authorityName="Schoch" authorityYear="1895" box="[487,587,659,685]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="119" phylum="Arthropoda" rank="subTribe" subTribe="Blaesiina">Blaesiina</taxonomicName>
would have been afforded suitable avenues for traversing previously inhospitable lowland forested regions in Amazonia by the extensions of savanna-like habitat (Fig. 12). With the establishment or re-establishment of rain forest in the Amazon basin, populations of ancestral
<taxonomicName authorityName="Schoch" authorityYear="1895" box="[817,917,765,791]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="119" phylum="Arthropoda" rank="subTribe" subTribe="Blaesiina">Blaesiina</taxonomicName>
became divided and isolated both to the north and south of the Amazon region. The northern lineage (todays
<taxonomicName authorityName="Moron &amp; Nogueira" authorityYear="2007" box="[236,378,835,861]" class="Insecta" family="Cetoniidae" genus="Halffterinetis" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="119" phylum="Arthropoda" rank="genus">
<emphasis box="[236,378,835,861]" italics="true" pageId="14" pageNumber="119">Halffterinetis</emphasis>
</taxonomicName>
species) ultimately dispersed to nuclear northern
<collectingCountry box="[963,1050,835,861]" name="Mexico" pageId="14" pageNumber="119">Mexico</collectingCountry>
, and the southern lineage (todays
<taxonomicName authorityName="Burmeister" authorityYear="1842" box="[473,549,871,897]" class="Insecta" family="Cetoniidae" genus="Blaesia" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="119" phylum="Arthropoda" rank="genus">
<emphasis box="[473,549,871,897]" italics="true" pageId="14" pageNumber="119">Blaesia</emphasis>
</taxonomicName>
species) became isolated in the woodland savannas south of the Amazon basin. As habitats changed through time, ancestral
<taxonomicName authorityName="Schoch" authorityYear="1895" box="[1008,1108,906,932]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="119" phylum="Arthropoda" rank="subTribe" subTribe="Blaesiina">Blaesiina</taxonomicName>
disappeared entirely from between northern
<collectingCountry box="[643,727,941,967]" name="Mexico" pageId="14" pageNumber="119">Mexico</collectingCountry>
and southern South America.
</paragraph>
<paragraph blockId="14.[140,1108,167,1672]" pageId="14" pageNumber="119">
Pre-Miocene dispersal of the biota between North and South America was probably rare, and a small amount of biotic interchange agrees with the geologic evidence suggesting a relatively wide separation of the Americas in Cretaceous through Oligocene times (
<bibRefCitation author="Raven PH &amp; Axelrod DI" box="[304,596,1082,1108]" pageId="14" pageNumber="119" pagination="539 - 673" refId="ref9873" refString="Raven PH, Axelrod DI (1974) Angiosperm biogeography and past continental movements. Annals of the Missouri Botanical Garden 61: 539 - 673." type="journal article" year="1974">Raven and Axelrod 1974</bibRefCitation>
;
<bibRefCitation author="Smith AG &amp; Briden JC" box="[610,886,1082,1108]" pageId="14" pageNumber="119" refId="ref10176" refString="Smith AG, Briden JC (1977) Mesozoic and Cenozoic paleocontinental maps. Cambridge University Press, Cambridge, United Kingdom, 64 pp." type="book" year="1977">Smith and Briden 1977</bibRefCitation>
;
<bibRefCitation author="Gose WA &amp; Scott GR &amp; Swartz DK" box="[900,1095,1082,1108]" pageId="14" pageNumber="119" pagination="51 - 54" refId="ref8775" refString="Gose WA, Scott GR, Swartz DK (1980) The aggregation of Mesoamerica: paleomagnetic evidence. In: Pilger RH (Ed) The Origin of the Gulf of Mexico and the Early Opening of the Central North Atlantic Ocean. Symposium Proceedings, Louisiana State University and Louisiana Geological Survey, Baton Rouge, LA. 51 - 54." type="proceedings paper" year="1980">Gose et al. 1980</bibRefCitation>
). Consideration of climate is important both before and after establishment of a land connection (
<bibRefCitation author="Stehli FG &amp; Webb SD" box="[282,559,1152,1179]" pageId="14" pageNumber="119" pagination="3 - 16" refId="ref10267" refString="Stehli FG, Webb SD (1985 b) A kaleidoscope of plates, faunal and floral dispersals, and sea level changes. In: Stehli FG, Webb SD (Eds) The Great American Biotic Interchange. Plenum Press, New York, NY, 3 - 16." type="book chapter" year="1985">Stehli and Webb 1985b</bibRefCitation>
). For example, the existence of clear evidence of mountain glaciation along the continental divide in
<collectingCountry box="[746,869,1187,1213]" name="Guatemala" pageId="14" pageNumber="119">Guatemala</collectingCountry>
suggests that simply extending present-day conditions back in time will not suffice to allow a real understanding of the physical nature of the link between the two Americas or of its effect on biotic interchange. After Mesoamerica coalesced during the Pliocene 3.0 MYBP (
<bibRefCitation author="Marshall G" pageId="14" pageNumber="119" pagination="380 - 388" refId="ref9285" refString="Marshall G (1988) Land mammals and the great American interchange. Science 76: 380 - 388." type="journal article" year="1988">Marshall 1988</bibRefCitation>
) to 5.7 MYBP (
<bibRefCitation author="Lloyd JJ" box="[450,587,1328,1354]" pageId="14" pageNumber="119" pagination="88 - 1001" refId="ref9145" refString="Lloyd JJ (1963) Tectonic history of the south central-American orogen. In: Childs OE, Beebe BW (Eds). Backbone of the Americas. American Association of Petroleum Geologists, Memoir No. 2, 88 - 1001." type="book chapter" year="1963">Lloyd 1963</bibRefCitation>
;
<bibRefCitation author="Kaneps AC" box="[601,756,1328,1355]" pageId="14" pageNumber="119" pagination="297 - 301" refId="ref9057" refString="Kaneps AC (1979) Gulf Stream: velocity fluctuations during the late Cenozoic. Science 204: 297 - 301." type="journal article" year="1979">Kaneps 1979</bibRefCitation>
), an extensive faunal exchange began (
<bibRefCitation author="Webb SD" box="[224,364,1363,1390]" pageId="14" pageNumber="119" pagination="393 - 426" refId="ref10424" refString="Webb SD (1978) A history of savanna vertebrates in the New World, part 11: South America and the great interchange. Annual Review of Ecology and Systematics 9: 393 - 426." type="journal article" year="1978">Webb 1978</bibRefCitation>
;
<bibRefCitation author="Stehli FG &amp; Webb SD" box="[379,639,1363,1390]" pageId="14" pageNumber="119" refId="ref10238" refString="Stehli FG, Webb SD (Eds) (1985 a) The Great American Biotic Interchange. Plenum Press, New York, NY, 532 pp." type="book" year="1985">Stehli &amp; Webb 1985a</bibRefCitation>
). Formation of the Panamanian isthmus dispersal route permitted separate invasions of plants and animals at widely separated periods when climates and topographic features were different than today.
</paragraph>
<paragraph blockId="14.[140,1108,167,1672]" pageId="14" pageNumber="119">
After the formation of the isthmus of
<collectingCountry box="[618,710,1470,1496]" name="Panama" pageId="14" pageNumber="119">Panama</collectingCountry>
, members of the
<taxonomicName authorityName="Kirby" authorityYear="1827" box="[909,1033,1469,1495]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="119" phylum="Arthropoda" rank="tribe" tribe="Gymnetini">Gymnetini</taxonomicName>
began their northward dispersal from South America into Central America,
<collectingCountry box="[924,1011,1504,1530]" name="Mexico" pageId="14" pageNumber="119">Mexico</collectingCountry>
, and the
<collectingCountry box="[140,296,1539,1566]" name="United States of America" pageId="14" pageNumber="119">United States</collectingCountry>
.
<bibRefCitation author="Webb SD" box="[305,450,1539,1566]" pageId="14" pageNumber="119" pagination="393 - 426" refId="ref10424" refString="Webb SD (1978) A history of savanna vertebrates in the New World, part 11: South America and the great interchange. Annual Review of Ecology and Systematics 9: 393 - 426." type="journal article" year="1978">Webb (1978</bibRefCitation>
, 1985) provided an excellent analysis of the interamerican biotic exchange, pertinent parts of which are described here. The interval from 2.5 1.5 MYBP shows an extensive movement of savanna-adapted mammal faunas from south temperate to north temperate latitudes and vice versa. All of the animals that
</paragraph>
<paragraph blockId="15.[140,1107,666,752]" pageId="15" pageNumber="120">
<emphasis bold="true" box="[140,250,666,688]" pageId="15" pageNumber="120">Figure ļ2.</emphasis>
Maps showing approximate distribution of savannas (gray areas) in South America at
<emphasis bold="true" box="[1036,1049,666,688]" pageId="15" pageNumber="120">a</emphasis>
about 4.0 MYBP,
<emphasis bold="true" box="[250,264,698,720]" pageId="15" pageNumber="120">b</emphasis>
during glacial maxima, and
<emphasis bold="true" box="[534,546,698,720]" pageId="15" pageNumber="120">c</emphasis>
today. Arrow in
<emphasis bold="true" box="[708,722,698,720]" pageId="15" pageNumber="120">b</emphasis>
shows most likely dispersal route of taxa living in savanna habitats (after
<bibRefCitation author="Marshall G" box="[434,572,730,752]" pageId="15" pageNumber="120" pagination="49 - 85" refId="ref9242" refString="Marshall G (1985) Geochronology and land-mammal biochronology of the transamerican faunal interchange. In: Stehli FG, Webb SD (Eds) The Great American Biotic Interchange. Plenum Press, New York, NY, 49 - 85." type="book chapter" year="1985">Marshall 1985</bibRefCitation>
).
</paragraph>
<paragraph blockId="15.[140,1108,800,1672]" pageId="15" pageNumber="120">
are known to have dispersed between the Americas in the late Tertiary were tolerant of, or specifically adapted to, savanna woodland habitats. The savanna elements were not incidental parts of the interchange but represent the vast majority of the taxa involved. Notable among them were horses, llamas, armadillos, and ground sloths. The extent of savanna adaptations among the land mammals of the interchange indicates the presence of a uniformly nonforested corridor or a moving mosaic of such habitats between South America and North America. The more arid conditions that must be postulated for the isthmian region during its early history probably supported seasonal forests grading into thorn scrub savannas. Similar habitats exist today in northern
<collectingCountry box="[140,252,1117,1143]" name="Venezuela" pageId="15" pageNumber="120">Venezuela</collectingCountry>
and eastern
<collectingCountry box="[402,515,1117,1143]" name="Colombia" pageId="15" pageNumber="120">Colombia</collectingCountry>
and on the Pacific slopes of Central America from western
<collectingCountry box="[234,323,1153,1179]" name="Panama" pageId="15" pageNumber="120">Panama</collectingCountry>
northward. Less mesic conditions in the isthmian corridor were a result of a combination of factors having to do with climatic fluctuations associated with northern hemisphere glaciations, lowering of sea levels (with a concomitant increase in land area), regional uplift with large-scale volcanic extrusion, and creation of rain shadow regions.
</paragraph>
<paragraph blockId="15.[140,1108,800,1672]" lastBlockId="16.[140,1108,167,1671]" lastPageId="16" lastPageNumber="121" pageId="15" pageNumber="120">
The glacial maxima at and following the emergence of the Panamanian land bridge, combined with the presence of a north-south corridor over the bridge, occurred only twice in the late Tertiary (
<bibRefCitation author="Shackelton NJ &amp; Opdyke DN" box="[578,926,1399,1425]" pageId="15" pageNumber="120" pagination="216 - 219" refId="ref10074" refString="Shackelton NJ, Opdyke DN (1977) Oxygen isotope and paleomagnetic evidence for early northern hemisphere glaciation. Nature 270: 216 - 219." type="journal article" year="1977">Shackelton and Opdyke 1977</bibRefCitation>
;
<bibRefCitation author="Cronin TM" box="[941,1095,1399,1425]" pageId="15" pageNumber="120" pagination="812 - 833" refId="ref8651" refString="Cronin TM (1981) Rates and possible causes of neotectonic vertical crustal movements of the emerged southeastern United States Atlantic Coastal Plane. Bulletin of the Geolological Society of America 92: 812 - 833." type="journal article" year="1981">Cronin 1981</bibRefCitation>
). These times (2.5 and 1.8 MYBP) represent “optimal ecological windows” that permitted dispersal of taxa living in savanna habitats between the Americas (
<bibRefCitation author="Marshall G" box="[924,1095,1469,1496]" pageId="15" pageNumber="120" pagination="49 - 85" refId="ref9242" refString="Marshall G (1985) Geochronology and land-mammal biochronology of the transamerican faunal interchange. In: Stehli FG, Webb SD (Eds) The Great American Biotic Interchange. Plenum Press, New York, NY, 49 - 85." type="book chapter" year="1985">Marshall 1985</bibRefCitation>
). The earliest known South American mammals to disperse to North America across the Panamanian land bridge occur in rocks dated at 2.82.6 MYBP. This reciprocal event favoring savanna-adapted forms could not have occurred earlier due to absence of a suitable corridor, habitat, and climate. Subsequent opportunities did not exist until the next glacial maxima at about 2.01.9 MYBP (
<bibRefCitation author="Marshall G" box="[677,850,1645,1672]" pageId="15" pageNumber="120" pagination="49 - 85" refId="ref9242" refString="Marshall G (1985) Geochronology and land-mammal biochronology of the transamerican faunal interchange. In: Stehli FG, Webb SD (Eds) The Great American Biotic Interchange. Plenum Press, New York, NY, 49 - 85." type="book chapter" year="1985">Marshall 1985</bibRefCitation>
). “Thus, two synchro- nous and reciprocal dispersal events of late Tertiary age are recognized. The first event (2.82.6 Ma) included dispersal of
<taxonomicName authority="Cuvier" authorityName="Cuvier" box="[551,743,202,228]" class="Mammalia" family="Erethizontidae" genus="Erethizon" kingdom="Animalia" order="Rodentia" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[551,655,202,228]" italics="true" pageId="16" pageNumber="121">Erethizon</emphasis>
Cuvier
</taxonomicName>
,
<taxonomicName authority="Hay" authorityName="Hay" authorityYear="1926" box="[756,936,202,228]" class="Mammalia" family="Caviidae" genus="Neochoerus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[756,877,202,228]" italics="true" pageId="16" pageNumber="121">Neochoerus</emphasis>
Hay
</taxonomicName>
,
<taxonomicName authority="Osborn, Glossothrium Glossothrium Owen, Othrotheriops Hoffstetter" authorityName="Osborn, Glossothrium Glossothrium Owen, Othrotheriops Hoffstetter" authorityYear="1903" class="Mammalia" family="Glyptodontidae" genus="Glyptotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cingulata" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[949,1107,202,228]" italics="true" pageId="16" pageNumber="121">Glyptotherium</emphasis>
Osborn,
<emphasis box="[241,534,237,263]" italics="true" pageId="16" pageNumber="121">Glossothrium Glossothrium</emphasis>
Owen,
<emphasis box="[625,774,237,263]" italics="true" pageId="16" pageNumber="121">Othrotheriops</emphasis>
Hoffstetter
</taxonomicName>
,
<taxonomicName authority="Casatellanos" authorityName="Casatellanos" authorityYear="1927" class="Mammalia" family="Pampatheriidae" genus="Kraglievichia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cingulata" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[916,1059,237,263]" italics="true" pageId="16" pageNumber="121">Kraglievichia</emphasis>
Casatellanos
</taxonomicName>
, and
<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[312,400,273,298]" class="Mammalia" family="Dasypodidae" genus="Dasypus" kingdom="Animalia" order="Cingulata" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[312,400,273,298]" italics="true" pageId="16" pageNumber="121">Dasypus</emphasis>
</taxonomicName>
L. (and the ground bird
<taxonomicName authority="Brodkorb" authorityName="Brodkorb" authorityYear="1963" box="[686,882,272,299]" class="Reptilia" family="Phorusrhacidae" genus="Titanis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[686,763,273,298]" italics="true" pageId="16" pageNumber="121">Titanis</emphasis>
Brodkorb
</taxonomicName>
) to North America, and
<taxonomicName authority="Gray" authorityName="Gray" authorityYear="1837" box="[190,366,308,334]" class="Mammalia" family="Mephitidae" genus="Conepatus" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[190,302,308,334]" italics="true" pageId="16" pageNumber="121">Conepatus</emphasis>
Gray
</taxonomicName>
,
<taxonomicName authority="Roth" authorityName="Roth" box="[378,559,307,333]" class="Mammalia" family="Equidae" genus="Hippidion" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[378,491,307,333]" italics="true" pageId="16" pageNumber="121">Hippidion</emphasis>
Roth
</taxonomicName>
, and
<taxonomicName authority="LeConte" authorityName="LeConte" box="[621,843,307,334]" class="Mammalia" family="Tayassuidae" genus="Platygonus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[621,736,307,333]" italics="true" pageId="16" pageNumber="121">Platygonus</emphasis>
LeConte
</taxonomicName>
to South America. The second event (2.01.9 Ma) included dispersal of
<taxonomicName authority="Brisson" authorityName="Brisson" authorityYear="1762" box="[699,936,343,369]" class="Mammalia" family="Caviidae" genus="Hydrochoerus" kingdom="Animalia" order="Rodentia" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[699,845,343,369]" italics="true" pageId="16" pageNumber="121">Hydrochoerus</emphasis>
Brisson
</taxonomicName>
,
<taxonomicName authority="Spillmann" authorityName="Spillmann" authorityYear="1948" class="Mammalia" family="Megatheriidae" genus="Eremotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pilosa" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[952,1107,343,369]" italics="true" pageId="16" pageNumber="121">Eremotherium</emphasis>
Spillmann
</taxonomicName>
, and
<taxonomicName authority="Simpson" authorityName="Simpson" box="[326,552,378,404]" class="Mammalia" family="Dasypodidae" genus="Holmesina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cingulata" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[326,442,378,404]" italics="true" pageId="16" pageNumber="121">Holmesina</emphasis>
Simpson
</taxonomicName>
to North America, and
<taxonomicName authority="Leidy" authorityName="Leidy" authorityYear="1854" box="[841,1007,378,404]" class="Mammalia" family="Ursidae" genus="Arctodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[841,934,378,404]" italics="true" pageId="16" pageNumber="121">Arctodus</emphasis>
Leidy
</taxonomicName>
,
<taxonomicName authority="Bell" authorityName="Bell" authorityYear="1826" class="Mammalia" family="Mustelidae" genus="Galictis" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[1025,1107,378,404]" italics="true" pageId="16" pageNumber="121">Galictis</emphasis>
Bell
</taxonomicName>
,
<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[197,245,413,439]" class="Mammalia" family="Felidae" genus="Felis" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[197,245,413,439]" italics="true" pageId="16" pageNumber="121">Felis</emphasis>
</taxonomicName>
L.,
<taxonomicName authority="Lund" authorityName="Lund" authorityYear="1842" box="[288,460,413,439]" class="Mammalia" family="Felidae" genus="Smilodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[288,390,413,439]" italics="true" pageId="16" pageNumber="121">Smilodon</emphasis>
Lund
</taxonomicName>
,
<taxonomicName authority="Brunnich" authorityName="Brunnich" authorityYear="1771" box="[469,668,413,439]" class="Mammalia" family="Tapiridae" genus="Tapirus" kingdom="Animalia" order="Perissodactyla" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[469,551,414,439]" italics="true" pageId="16" pageNumber="121">Tapirus</emphasis>
Brünnich
</taxonomicName>
,
<taxonomicName authority="Gervais and Ameghino" authorityName="Gervais and Ameghino" box="[678,1100,413,439]" class="Mammalia" family="Camelidae" genus="Hemiauchenia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[678,837,413,439]" italics="true" pageId="16" pageNumber="121">Hemiauchenia</emphasis>
Gervais and Ameghino
</taxonomicName>
,
<taxonomicName authority="Moreno" authorityName="Moreno" authorityYear="1891" box="[140,403,448,475]" class="Mammalia" family="Equidae" genus="Onohippidium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[140,302,448,474]" italics="true" pageId="16" pageNumber="121">Onohippidium</emphasis>
Moreno
</taxonomicName>
, and
<taxonomicName authority="Osborn" authorityName="Osborn" authorityYear="1923" box="[462,686,448,475]" class="Mammalia" family="Gomphotheriidae" genus="Cuvieronius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proboscidea" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[462,593,449,475]" italics="true" pageId="16" pageNumber="121">Cuvieronius</emphasis>
Osborn
</taxonomicName>
(and possibly
<taxonomicName authority="Reig" authorityName="Reig" authorityYear="1956" box="[847,1035,448,475]" class="Mammalia" family="Mustelidae" genus="Stipanicicia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[847,975,449,475]" italics="true" pageId="16" pageNumber="121">Stipanicicia</emphasis>
Reig
</taxonomicName>
,
<taxonomicName authority="Smith" authorityName="Smith" class="Mammalia" family="Canidae" genus="Dusicyon" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="16" pageNumber="121">Dusicyon</emphasis>
Smith
</taxonomicName>
, and
<taxonomicName authority="Giebel" authorityName="Giebel" authorityYear="1855" box="[326,513,483,510]" class="Mammalia" family="Canidae" genus="Protocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[326,431,484,509]" italics="true" pageId="16" pageNumber="121">Protocyon</emphasis>
Giebel
</taxonomicName>
) to South America. Only one dispersal event of early Pleistocene age is evident, and this occurred at about 1.4 Ma. It corresponds to the earliest of the Pleistocene glacial maxima recognized by
<bibRefCitation author="Cronin TM" box="[763,930,554,580]" pageId="16" pageNumber="121" pagination="812 - 833" refId="ref8651" refString="Cronin TM (1981) Rates and possible causes of neotectonic vertical crustal movements of the emerged southeastern United States Atlantic Coastal Plane. Bulletin of the Geolological Society of America 92: 812 - 833." type="journal article" year="1981">Cronin (1981)</bibRefCitation>
and follows the one at 2.01.9 Ma. During this event,
<taxonomicName box="[582,644,590,616]" class="Mammalia" family="Canidae" genus="Canis" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[582,644,590,616]" italics="true" pageId="16" pageNumber="121">Canis</emphasis>
</taxonomicName>
L.,
<taxonomicName authority="Brisson" authorityName="Brisson" authorityYear="1762" box="[691,846,589,615]" class="Mammalia" family="Mustelidae" genus="Lutra" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[691,752,590,615]" italics="true" pageId="16" pageNumber="121">Lutra</emphasis>
Brisson
</taxonomicName>
,
<taxonomicName box="[858,979,589,615]" class="Mammalia" family="Canidae" genus="Chrysocyon" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[858,979,589,615]" italics="true" pageId="16" pageNumber="121">Chrysocyon</emphasis>
</taxonomicName>
Hamilton- Smith,
<taxonomicName authority="Hamilton-Smith, Leo" authorityName="Hamilton-Smith, Leo" box="[231,593,624,650]" class="Mammalia" family="Canidae" genus="Cerdocyon" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[231,342,624,650]" italics="true" pageId="16" pageNumber="121">Cerdocyon</emphasis>
Hamilton-Smith,
<emphasis box="[555,593,625,650]" italics="true" pageId="16" pageNumber="121">Leo</emphasis>
</taxonomicName>
(
<emphasis box="[610,724,624,650]" italics="true" pageId="16" pageNumber="121">=Panthera</emphasis>
Oken), and
<taxonomicName authority="Pohlig" authorityName="Pohlig" authorityYear="1912" box="[870,1108,624,650]" class="Mammalia" family="Gomphotheriidae" genus="Stegomastodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proboscidea" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[870,1028,624,650]" italics="true" pageId="16" pageNumber="121">Stegomastodon</emphasis>
Pohlig
</taxonomicName>
dispersed to South America, and
<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[534,638,659,685]" class="Mammalia" family="Didelphidae" genus="Didelphis" kingdom="Animalia" order="Didelphimorphia" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[534,638,659,685]" italics="true" pageId="16" pageNumber="121">Didelphis</emphasis>
</taxonomicName>
L. and
<taxonomicName authority="Gervais" authorityName="Gervais" box="[737,958,659,685]" class="Mammalia" family="Camelidae" genus="Palaeolama" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="16" pageNumber="121" phylum="Chordata" rank="genus">
<emphasis box="[737,862,659,685]" italics="true" pageId="16" pageNumber="121">Palaeolama</emphasis>
Gervais
</taxonomicName>
dispersed to North America” (
<bibRefCitation author="Marshall G" box="[342,516,694,721]" pageId="16" pageNumber="121" pagination="49 - 85" refId="ref9242" refString="Marshall G (1985) Geochronology and land-mammal biochronology of the transamerican faunal interchange. In: Stehli FG, Webb SD (Eds) The Great American Biotic Interchange. Plenum Press, New York, NY, 49 - 85." type="book chapter" year="1985">Marshall 1985</bibRefCitation>
). The last glacial maximum permitting dispersal of savanna biotas over the land bridge occurred
<geoCoordinate box="[641,722,730,756]" degrees="12,000" direction="north" orientation="latitude" pageId="16" pageNumber="121" precision="55" value="12.0">12,000</geoCoordinate>
<geoCoordinate box="[721,802,730,756]" degrees="1,000" direction="west" orientation="longitude" pageId="16" pageNumber="121" precision="55" value="-1.0">1,000</geoCoordinate>
years B.P. (
<bibRefCitation author="Bradbury JP" box="[927,1102,730,756]" pageId="16" pageNumber="121" pagination="46 - 48" refId="ref8445" refString="Bradbury JP (1982) Holocene chronostratigraphy of Mexico and Central America. In: Mangerud J, Birks, HJB, Jager JD (Eds) Chronostratigraphic Subdivision of the Holocene. Striae (Societas Upsaliensis pro Geologia Quaternaria) 16, Uppsala, Sweden, 46 - 48." type="book chapter" year="1982">Bradbury 1982</bibRefCitation>
;
<bibRefCitation author="Markgraf V &amp; Bradbury JP" box="[140,475,765,791]" pageId="16" pageNumber="121" pagination="40 - 45" refId="ref9189" refString="Markgraf V, Bradbury JP (1982) Holocene climatic history of South America. In: Mangerud J, Birks, HJB, Jager JD (Eds) Chronostratigraphic Subdivision of the Holocene. Striae (Societas Upsaliensis pro Geologia Quaternaria) 16, Uppsala, Sweden, 40 - 45." type="book chapter" year="1982">Markgraf and Bradbury 1982</bibRefCitation>
). A savanna corridor formed along the eastern side of the Andes connecting the now disjunct habitats in South America (Fig. 12).
</paragraph>
<paragraph blockId="16.[140,1108,167,1671]" lastBlockId="17.[140,1108,167,1672]" lastPageId="17" lastPageNumber="122" pageId="16" pageNumber="121">
The major obstacles to such dispersal events were distance and potential competitive exclusion. Given that cetoniines are capable of such powerful flight, distance may not have been such a deterrent to long distance dispersal. The Japanese beetle (
<taxonomicName authority="Newman" authorityName="Newman" class="Insecta" family="Rutelidae" genus="Popillia" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="121" phylum="Arthropoda" rank="species" species="japonica">
<emphasis italics="true" pageId="16" pageNumber="121">Popillia japonica</emphasis>
Newman
</taxonomicName>
) (
<taxonomicName authorityName="Latreille" authorityYear="1802" box="[372,522,941,967]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="121" phylum="Arthropoda" rank="family">Scarabaeidae</taxonomicName>
:
<taxonomicName authorityName="MacLeay" authorityYear="1819" box="[536,649,941,968]" class="Insecta" family="Rutelinae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="121" phylum="Arthropoda" rank="family">Rutelinae</taxonomicName>
), for example, has spread from the east coast of North America (where it was introduced) to the central states (
<quantity box="[992,1107,976,1002]" metricMagnitude="6" metricUnit="m" metricValue="1.9" pageId="16" pageNumber="121" unit="km" value="1900.0">1,900 km</quantity>
away) in only 70 years; that averages
<quantity box="[564,639,1011,1038]" metricMagnitude="4" metricUnit="m" metricValue="2.7" pageId="16" pageNumber="121" unit="km" value="27.0">27 km</quantity>
/year.
<taxonomicName baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[708,925,1011,1038]" class="Insecta" family="Aphodiidae" genus="Aphodius" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="121" phylum="Arthropoda" rank="species" species="fimetarius">
<emphasis box="[708,925,1011,1038]" italics="true" pageId="16" pageNumber="121">Aphodius fimetarius</emphasis>
</taxonomicName>
(L.) (
<taxonomicName authorityName="Latreille" authorityYear="1802" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="121" phylum="Arthropoda" rank="family">Scarabaeidae</taxonomicName>
:
<taxonomicName authorityName="Leach" authorityYear="1815" box="[195,331,1047,1074]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="121" phylum="Arthropoda" rank="subFamily" subFamily="Aphodiinae">Aphodiinae</taxonomicName>
), introduced into North America from Europe probably in colonial times, is now found over much of the continent.
<taxonomicName authority="(Fabr.)" baseAuthorityName="Fabr." box="[705,1063,1082,1109]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="121" phylum="Arthropoda" rank="species" species="gazella">
<emphasis box="[705,979,1082,1108]" italics="true" pageId="16" pageNumber="121">Digitonthophagus gazella</emphasis>
(Fabr.)
</taxonomicName>
has dispersed from 43 to 808 (!) km/year in
<collectingCountry box="[617,701,1117,1143]" name="Mexico" pageId="16" pageNumber="121">Mexico</collectingCountry>
and the
<collectingCountry box="[809,859,1117,1143]" name="United States of America" pageId="16" pageNumber="121">U.S.</collectingCountry>
(Barbero and López- Guerrero 1992). The tussock moth, monarch butterfly, European corn borer, and honeybee all represent contemporary examples of long distance dispersal by insects in short periods of time. The Africanized honeybee,
<taxonomicName authority="Lepeletier" authorityName="Lepeletier" authorityYear="1836" box="[729,1104,1223,1249]" class="Insecta" family="Apidae" genus="Apis" kingdom="Animalia" order="Hymenoptera" pageId="16" pageNumber="121" phylum="Arthropoda" rank="subSpecies" species="mellifera" subSpecies="scutellata">
<emphasis box="[729,983,1223,1249]" italics="true" pageId="16" pageNumber="121">Apis mellifera scutellata</emphasis>
Lepeletier
</taxonomicName>
, has dispersed
<quantity box="[297,444,1258,1285]" metricMagnitude="5" metricUnit="m" metricValue="4.0" metricValueMax="5.0" metricValueMin="3.0" pageId="16" pageNumber="121" unit="km" value="400.0" valueMax="500.0" valueMin="300.0">300500 km</quantity>
per year from southern
<collectingCountry box="[722,787,1258,1284]" name="Brazil" pageId="16" pageNumber="121">Brazil</collectingCountry>
to northern
<collectingCountry box="[935,1019,1258,1284]" name="Mexico" pageId="16" pageNumber="121">Mexico</collectingCountry>
in only 30 years (
<bibRefCitation author="Camazine S &amp; Morse RA" box="[254,560,1293,1320]" pageId="16" pageNumber="121" pagination="465 - 471" refId="ref8602" refString="Camazine S, Morse RA (1988) The Africanized honeybee. American Scientist 76: 465 - 471." type="journal article" year="1988">Camazine &amp; Morse 1988</bibRefCitation>
). The rapid and historically near-instantaneous colonization of the Australian continent by the European hare,
<taxonomicName authority="Pallas" authorityName="Pallas" authorityYear="1778" box="[856,1104,1328,1354]" class="Mammalia" family="Leporidae" genus="Lepus" kingdom="Animalia" order="Lagomorpha" pageId="16" pageNumber="121" phylum="Chordata" rank="species" species="europaeus">
<emphasis box="[856,1030,1329,1354]" italics="true" pageId="16" pageNumber="121">Lepus europaeus</emphasis>
Pallas
</taxonomicName>
, highlights the phenomenal dispersal ability of a small mammal (
<bibRefCitation author="Marshall G" box="[872,1044,1363,1390]" pageId="16" pageNumber="121" pagination="49 - 85" refId="ref9242" refString="Marshall G (1985) Geochronology and land-mammal biochronology of the transamerican faunal interchange. In: Stehli FG, Webb SD (Eds) The Great American Biotic Interchange. Plenum Press, New York, NY, 49 - 85." type="book chapter" year="1985">Marshall 1985</bibRefCitation>
). The opossum,
<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[257,495,1399,1425]" class="Mammalia" family="Didelphidae" genus="Didelphis" kingdom="Animalia" order="Didelphimorphia" pageId="16" pageNumber="121" phylum="Chordata" rank="species" species="marsupialis">
<emphasis box="[257,495,1399,1425]" italics="true" pageId="16" pageNumber="121">Didelphis marsupialis</emphasis>
</taxonomicName>
L., had an average dispersal rate of
<quantity box="[917,992,1398,1425]" metricMagnitude="4" metricUnit="m" metricValue="5.0" pageId="16" pageNumber="121" unit="km" value="50.0">50 km</quantity>
/year during the 26 years following its introduction into California (
<bibRefCitation author="Tyndale-Biscoe H" box="[814,1060,1434,1460]" pageId="16" pageNumber="121" refId="ref10381" refString="Tyndale-Biscoe H (1973) Life of marsupials. Edward Arnold, London, United Kingdom. 442 pp." type="book" year="1973">Tyndale-Biscoe 1973</bibRefCitation>
). At such a rate this species could extend its range
<quantity box="[675,803,1469,1496]" metricMagnitude="7" metricUnit="m" metricValue="2.5" pageId="16" pageNumber="121" unit="km" value="25000.0">25,000 km</quantity>
in only 500 years (
<bibRefCitation author="Savage DE &amp; Russell DE" pageId="16" pageNumber="121" refId="ref10031" refString="Savage DE, Russell DE (1983) Mammalian Paleofaunas of the World. Addison-Wesley, London, United Kingdom." type="book" year="1983">Savage and Russell 1983</bibRefCitation>
).
<bibRefCitation author="Martin PS" box="[363,529,1504,1530]" pageId="16" pageNumber="121" pagination="969 - 974" refId="ref9305" refString="Martin PS (1973) The discovery of America. Science 179: 969 - 974." type="journal article" year="1973">Martin (1973)</bibRefCitation>
noted that a conservative dispersal rate of
<quantity box="[1022,1099,1504,1530]" metricMagnitude="4" metricUnit="m" metricValue="1.6" pageId="16" pageNumber="121" unit="km" value="16.0">16 km</quantity>
/ year would have permitted prehistoric humans to spread from
<collectingCountry box="[868,954,1539,1565]" name="Canada" pageId="16" pageNumber="121">Canada</collectingCountry>
to Tierra del Fuego in less than 1,000 years. The dispersal of insects between the mid-continental regions of North and South America may have occurred in only a few thousand years with the availability of suitable habitat. The vertebrate fossil evidence clearly indicates that dispersal of savanna-adapted animals occurred twice in the late Tertiary. South American ancestral
<taxonomicName authorityName="Schoch" authorityYear="1895" box="[361,461,202,228]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="122" phylum="Arthropoda" rank="subTribe" subTribe="Blaesiina">Blaesiina</taxonomicName>
, adapted to dry habitats, was part of that dispersal.
<bibRefCitation author="Webb SD" pageId="17" pageNumber="122" pagination="393 - 426" refId="ref10424" refString="Webb SD (1978) A history of savanna vertebrates in the New World, part 11: South America and the great interchange. Annual Review of Ecology and Systematics 9: 393 - 426." type="journal article" year="1978">Webb (1978)</bibRefCitation>
observed that it may be difficult for some biologists to accept so short a time scale for such evolutionary change, but the paleontological record of the interamerican interchange demonstrates that two or three million years is sufficient time to produce fundamental evolutionary reorganization of a major biota.
</paragraph>
<paragraph blockId="17.[140,1108,167,1672]" pageId="17" pageNumber="122">
The late Pleistocene shift to more humid conditions in lower Central America produced a major set of savanna disjunctions spanning the isthmian gap (
<bibRefCitation author="Webb SD" pageId="17" pageNumber="122" pagination="393 - 426" refId="ref10424" refString="Webb SD (1978) A history of savanna vertebrates in the New World, part 11: South America and the great interchange. Annual Review of Ecology and Systematics 9: 393 - 426." type="journal article" year="1978">Webb 1978</bibRefCitation>
). The disjunct distribution across the American tropics shared by many presentday organisms provides additional evidence of a previous woodland savanna corridor. Within the temperate to subtropical
<taxonomicName authorityName="Burmeister" authorityYear="1844" box="[570,676,519,545]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="122" phylum="Arthropoda" rank="subTribe" subTribe="Areodina">Areodina</taxonomicName>
(
<taxonomicName authorityName="Latreille" authorityYear="1802" box="[692,846,519,545]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="122" phylum="Arthropoda" rank="family">Scarabaeidae</taxonomicName>
:
<taxonomicName authorityName="MacLeay" authorityYear="1819" box="[857,973,519,546]" class="Insecta" family="Rutelinae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="122" phylum="Arthropoda" rank="family">Rutelinae</taxonomicName>
), six genera are found ranging from the
<collectingCountry box="[463,617,554,580]" name="United States of America" pageId="17" pageNumber="122">United States</collectingCountry>
to
<collectingCountry box="[657,786,554,580]" name="Guatemala" pageId="17" pageNumber="122">Guatemala</collectingCountry>
, and three genera are found in South America (
<bibRefCitation author="Jameson ML" box="[361,533,589,615]" pageId="17" pageNumber="122" pagination="377 - 422" refId="ref9021" refString="Jameson ML (1990) Revision, phylogeny and biogeography of the genera Parabyrsopolis Ohaus and Viridimicus, new genus (Coleoptera: Scarabaeidae: Rutelinae). Coleopterists Bulletin 44: 377 - 422." type="journal article" year="1990">Jameson 1990</bibRefCitation>
). None of these genera occur in the remainder of Central America, which, for the most part, has been historically covered by tropical rainforest. This Central American gap might seem like a paradox until, noticing its occurrence in other groups, we recognize a pattern. Many birds adapted to savanna or thorn scrub show a wide interamerican disjunction. These include the Green Jay, Military Macaw, Melodious Blackbird, Homed Lark, Vermillion Flycatcher, small woodpeckers, and the Grasshopper Sparrow (
<bibRefCitation author="Griscom L" box="[656,826,800,827]" pageId="17" pageNumber="122" pagination="341 - 382" refId="ref8839" refString="Griscom L (1950) Distribution and origin of the birds of Mexico. Bulletin of the Museum of Comparative Zoology 103: 341 - 382." type="journal article" year="1950">Griscom 1950</bibRefCitation>
;
<bibRefCitation author="Mengel RM" box="[837,994,800,826]" pageId="17" pageNumber="122" pagination="279 - 340" refId="ref9354" refString="Mengel RM (1970) The North American central plains as an isolating agent in bird speciation. In: Dort W, Jones JK (Eds) Pleistocene and Recent Environments of the Central Great Plains. University of Kansas Press, Lawrence, KS, 279 - 340." type="book chapter" year="1970">Mengel 1970</bibRefCitation>
). Cricetid rodents such as
<taxonomicName authority="Giglioli" authorityName="Giglioli" authorityYear="1874" box="[322,600,835,861]" class="Mammalia" family="Muridae" genus="Reithrodontomys" kingdom="Animalia" order="Rodentia" pageId="17" pageNumber="122" phylum="Chordata" rank="genus">
<emphasis box="[322,506,835,861]" italics="true" pageId="17" pageNumber="122">Reithrodontomys</emphasis>
Giglioli
</taxonomicName>
skip from semiarid habitats in
<collectingCountry box="[959,1077,835,861]" name="Nicaragua" pageId="17" pageNumber="122">Nicaragua</collectingCountry>
to similar habitats in the Andes, and
<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[539,632,871,897]" class="Reptilia" family="Viperidae" genus="Crotalus" kingdom="Animalia" order="Squamata" pageId="17" pageNumber="122" phylum="Chordata" rank="genus">
<emphasis box="[539,632,871,897]" italics="true" pageId="17" pageNumber="122">Crotalus</emphasis>
</taxonomicName>
L. vipers (preferring scrub habitats) now have a large gap across the rainforest of the isthmus (
<bibRefCitation author="Webb SD" box="[739,880,906,933]" pageId="17" pageNumber="122" pagination="357 - 386" refId="ref10461" refString="Webb SD (1985) Late Cenozoic mammal dispersals between the Americas. In: Stehli FG, Webb SD (Eds) The Great American Biotic Interchange. Plenum Press, New York, NY, 357 - 386." type="book chapter" year="1985">Webb 1985</bibRefCitation>
). The distributional gap across the isthmian region is well known for many plants as well as many bees that specialize on these plants (
<bibRefCitation author="Raven PH" box="[503,648,976,1002]" pageId="17" pageNumber="122" pagination="151 - 177" refId="ref9847" refString="Raven PH (1963) Amphitropical relationships in the floras of North and South America. Quarterly Review of Biology 38: 151 - 177." type="journal article" year="1963">Raven 1963</bibRefCitation>
;
<bibRefCitation author="Solbrig OT" box="[661,816,976,1002]" pageId="17" pageNumber="122" pagination="218 - 223" refId="ref10202" refString="Solbrig OT (1972) The floristic disjunctions between the Monte in Argentina and the Sonoran Desert in Mexico and the United States. Annals of the Missouri Botanical Garden 59: 218 - 223." type="journal article" year="1972">Solbrig 1972</bibRefCitation>
;
<bibRefCitation author="Rzedowski JL" box="[829,1024,976,1002]" pageId="17" pageNumber="122" pagination="61 - 72" refId="ref9926" refString="Rzedowski JL (1973) Geographic relationships of the flora of Mexican dry regions. In: Graham A (Ed) Vegetation and Vegetational History of Northern Latin America. Elsevier, Amsterdam, Netherlands, 61 - 72." type="book chapter" year="1973">Rzedowski 1973</bibRefCitation>
;
<bibRefCitation author="Simpson BB &amp; Neff JL" pageId="17" pageNumber="122" pagination="427 - 452" refId="ref10128" refString="Simpson BB, Neff JL (1985) Plants, their pollinating bees, and the great American interchange. In: Stehli FG, Webb SD (Eds) The Great American Biotic Interchange. Plenum Press, New York, NY, 427 - 452." type="book chapter" year="1985">Simpson and Neff 1985</bibRefCitation>
).
<bibRefCitation author="Webb SD" box="[380,535,1011,1038]" pageId="17" pageNumber="122" pagination="357 - 386" refId="ref10461" refString="Webb SD (1985) Late Cenozoic mammal dispersals between the Americas. In: Stehli FG, Webb SD (Eds) The Great American Biotic Interchange. Plenum Press, New York, NY, 357 - 386." type="book chapter" year="1985">Webb (1985)</bibRefCitation>
observed that one of the most convincing indications of former continuity is a string of relict populations of
<taxonomicName authority="Cav." authorityName="Cav." box="[834,967,1047,1073]" class="Magnoliopsida" family="Zygophyllaceae" genus="Larrea" kingdom="Plantae" order="Zygophyllales" pageId="17" pageNumber="122" phylum="Tracheophyta" rank="genus">
<emphasis box="[834,907,1047,1072]" italics="true" pageId="17" pageNumber="122">Larrea</emphasis>
Cav.
</taxonomicName>
in
<collectingCountry box="[1005,1057,1047,1073]" name="Peru" pageId="17" pageNumber="122">Peru</collectingCountry>
and
<collectingCountry box="[140,221,1082,1108]" name="Bolivia" pageId="17" pageNumber="122">Bolivia</collectingCountry>
partly connecting its main south temperate and north temperate ranges. This idea is strengthened by the fact that one of the principal foods of the extinct ground sloth (
<taxonomicName authority="Hoffstetter" authorityName="Hoffstetter" box="[215,514,1152,1178]" class="Mammalia" family="Nothrotheriidae" genus="Nothrotheriops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pilosa" pageId="17" pageNumber="122" phylum="Chordata" rank="genus">
<emphasis box="[215,379,1152,1178]" italics="true" pageId="17" pageNumber="122">Nothrotheriops</emphasis>
Hoffstetter
</taxonomicName>
species), as indicated by its dung, was
<taxonomicName box="[975,1048,1153,1178]" class="Magnoliopsida" family="Zygophyllaceae" genus="Larrea" kingdom="Plantae" order="Zygophyllales" pageId="17" pageNumber="122" phylum="Tracheophyta" rank="genus">
<emphasis box="[975,1048,1153,1178]" italics="true" pageId="17" pageNumber="122">Larrea</emphasis>
</taxonomicName>
, and that both genera clearly came to North America from temperate South America (
<bibRefCitation author="Martin PS &amp; Sabels B &amp; Shutler D" box="[148,362,1223,1249]" pageId="17" pageNumber="122" pagination="102 - 127" refId="ref9322" refString="Martin PS, Sabels B, Shutler D (1961) Rampart cave coprolite and ecology of the Shasta Ground Sloth. American Journal of Science 259: 102 - 127." type="journal article" year="1961">Martin et al. 1961</bibRefCitation>
;
<bibRefCitation author="Hunziker JH &amp; Palaciosa RA &amp; De Valesi G &amp; Poggio L" box="[373,614,1223,1249]" pageId="17" pageNumber="122" pagination="224 - 233" refId="ref8976" refString="Hunziker JH, Palaciosa RA, De Valesi G, Poggio L (1973) Species disjunctions in Larrea: evidence from morphology, cytogenetics, phenolic compounds, and seed albumins. Annals of the Missouri Botanical Garden 59: 224 - 233." type="journal article" year="1973">Hunziker et al. 1973</bibRefCitation>
). By the late Pleistocene, as now, woodland savanna taxa were excluded from the isthmian region due to the dissolution of savanna habitats and replacement by tropical rainforest. Late Pleistocene pollen samples from Lake Gatun in
<collectingCountry box="[384,475,1329,1355]" name="Panama" pageId="17" pageNumber="122">Panama</collectingCountry>
reveal a forest flora much like that of present lowland
<collectingCountry box="[140,231,1364,1390]" name="Panama" pageId="17" pageNumber="122">Panama</collectingCountry>
(
<bibRefCitation author="Webb SD" box="[246,388,1363,1390]" pageId="17" pageNumber="122" pagination="393 - 426" refId="ref10424" refString="Webb SD (1978) A history of savanna vertebrates in the New World, part 11: South America and the great interchange. Annual Review of Ecology and Systematics 9: 393 - 426." type="journal article" year="1978">Webb 1978</bibRefCitation>
). About
<quantity box="[488,601,1363,1390]" metricMagnitude="6" metricUnit="m" metricValue="1.7" pageId="17" pageNumber="122" unit="km" value="1700.0">1,700 km</quantity>
of tropical wet forest extending from
<collectingCountry name="Costa Rica" pageId="17" pageNumber="122">Costa Rica</collectingCountry>
and
<collectingCountry box="[257,348,1399,1425]" name="Panama" pageId="17" pageNumber="122">Panama</collectingCountry>
through northern
<collectingCountry box="[577,693,1399,1425]" name="Colombia" pageId="17" pageNumber="122">Colombia</collectingCountry>
now separates the nearest areas of savanna and thorn forest (
<bibRefCitation author="Sarmiento G" box="[445,638,1434,1461]" pageId="17" pageNumber="122" pagination="65 - 99" refId="ref9993" refString="Sarmiento G (1976) Evolution of arid vegetation in tropical America. In: Goodall DW (Ed) Evolution of Desert Biota. University of Texas Press, Austin, TX, 65 - 99." type="book chapter" year="1976">Sarmiento 1976</bibRefCitation>
). Consequently, ancestral
<taxonomicName authorityName="Schoch" authorityYear="1895" box="[943,1046,1434,1460]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="122" phylum="Arthropoda" rank="subTribe" subTribe="Blaesiina">Blaesiina</taxonomicName>
were also excluded from this region because they could not survive in tropical wet forests. Northern Central America retained a woodland savanna fauna as evidenced by the present biota and Pleistocene samples from
<collectingCountry box="[655,784,1539,1565]" name="Guatemala" pageId="17" pageNumber="122">Guatemala</collectingCountry>
,
<collectingCountry box="[798,914,1539,1565]" name="Honduras" pageId="17" pageNumber="122">Honduras</collectingCountry>
and
<collectingCountry box="[975,1107,1539,1565]" name="El Salvador" pageId="17" pageNumber="122">El Salvador</collectingCountry>
(
<bibRefCitation author="Stirton RA &amp; Gealey WK" box="[148,446,1575,1601]" pageId="17" pageNumber="122" pagination="1731 - 1754" refId="ref10319" refString="Stirton RA, Gealey WK (1949) Reconnaissance geology and vertebrate paleontology of El Salvador, Central America. Bulletin of the Geological Society of America 60: 1731 - 1754." type="journal article" year="1949">Stirton and Gealey 1949</bibRefCitation>
;
<bibRefCitation author="Carr A" box="[460,586,1574,1601]" pageId="17" pageNumber="122" pagination="563 - 594" refId="ref8622" refString="Carr A (1950) Outline for a classification of animal habitats in Honduras. Bulletin of the American Museum of Natural History 94: 563 - 594." type="journal article" year="1950">Carr 1950</bibRefCitation>
;
<bibRefCitation author="Duellman WE" box="[601,791,1575,1601]" pageId="17" pageNumber="122" pagination="700 - 719" refId="ref8729" refString="Duellman WE (1966) The Central American herpetofauna: an ecological perspective. Copeia 1966: 700 - 719." type="journal article" year="1966">Duellman 1966</bibRefCitation>
;
<bibRefCitation author="Savage JM" box="[805,955,1575,1601]" pageId="17" pageNumber="122" pagination="719 - 766" refId="ref10052" refString="Savage JM (1966) The origins and history of the Central American herpetofauna. Copeia 1966: 719 - 766." type="journal article" year="1966">Savage 1966</bibRefCitation>
;
<bibRefCitation author="Woodburne M" pageId="17" pageNumber="122" pagination="121 - 125" refId="ref10574" refString="Woodburne M (1969) A Late Pleistocene occurrence of the collared peccary, Dicotyles tajacu, in Guatemala. Journal of Mammalogy 50: 121 - 125." type="journal article" year="1969">Woodburne 1969</bibRefCitation>
;
<bibRefCitation author="Howell R" box="[214,370,1610,1637]" pageId="17" pageNumber="122" pagination="293 - 326" refId="ref8957" refString="Howell R (1969) Avian distribution in Central America. The Auk 86: 293 - 326." type="journal article" year="1969">Howell 1969</bibRefCitation>
). Species of
<taxonomicName authorityName="Moron &amp; Nogueira" authorityYear="2007" box="[510,656,1610,1636]" class="Insecta" family="Cetoniidae" genus="Halffterinetis" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="122" phylum="Arthropoda" rank="genus">
<emphasis box="[510,656,1610,1636]" italics="true" pageId="17" pageNumber="122">Halffterinetis</emphasis>
</taxonomicName>
are today found in mesic to xeric habitats in north central
<collectingCountry box="[378,467,1645,1671]" name="Mexico" pageId="17" pageNumber="122">Mexico</collectingCountry>
.
</paragraph>
<paragraph blockId="18.[140,1108,167,510]" pageId="18" pageNumber="123">
Based on his analysis of the entomofauna,
<bibRefCitation author="Halffter G" box="[702,882,167,194]" pageId="18" pageNumber="123" pagination="1 - 64" refId="ref8922" refString="Halffter G (1976) Distribucion de 1 os insectos en la Zona de Transicion Mexicana. Relaciones con la entomofauna de Norteamerica. Folia Entomologica Mexicana No. 35: 1 - 64." type="book chapter" year="1976">Halffter (1976)</bibRefCitation>
formulated several different dispersal patterns to explain the present distribution of taxa in the Mexican Transition Zone. The distribution of
<taxonomicName authorityName="Moron &amp; Nogueira" authorityYear="2007" box="[636,780,237,263]" class="Insecta" family="Cetoniidae" genus="Halffterinetis" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="123" phylum="Arthropoda" rank="genus">
<emphasis box="[636,780,237,263]" italics="true" pageId="18" pageNumber="123">Halffterinetis</emphasis>
</taxonomicName>
species coincides well with Halffters “Typical Neotropical Dispersal Pattern”. In this pattern, South American elements penetrated into the Mexican Transition Zone after the formation of the Panamanian land bridge and after most of the elevation of the Mexican Plateau. As ancestral
<taxonomicName authorityName="Schoch" authorityYear="1895" box="[250,351,378,404]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="123" phylum="Arthropoda" rank="subTribe" subTribe="Blaesiina">Blaesiina</taxonomicName>
spread northward, they used as their principal expansion route from Central America the mountains of
<collectingRegion box="[614,698,413,439]" country="Mexico" name="Oaxaca" pageId="18" pageNumber="123">Oaxaca</collectingRegion>
and the Sierra Madres, which funneled the dispersal of
<taxonomicName authorityName="Schoch" authorityYear="1895" box="[384,485,448,474]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="123" phylum="Arthropoda" rank="subTribe" subTribe="Blaesiina">Blaesiina</taxonomicName>
to the west and north, respectively, where
<taxonomicName authorityName="Moron &amp; Nogueira" authorityYear="2007" box="[963,1107,448,474]" class="Insecta" family="Cetoniidae" genus="Halffterinetis" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="123" phylum="Arthropoda" rank="genus">
<emphasis box="[963,1107,448,474]" italics="true" pageId="18" pageNumber="123">Halffterinetis</emphasis>
</taxonomicName>
species occur today.
</paragraph>
<paragraph blockId="18.[140,393,604,631]" box="[140,393,604,631]" pageId="18" pageNumber="123">
<emphasis bold="true" box="[140,393,604,631]" pageId="18" pageNumber="123">Acknowledgments</emphasis>
</paragraph>
<paragraph blockId="18.[140,1108,659,1073]" pageId="18" pageNumber="123">
I am grateful to the institutions, curators, and collection managers mentioned in the “Methods” section for their generous cooperation in either loaning or allowing me access to specimens under their care. I thank Mary Liz Jameson (then University of Nebraska State Museum) for her assistance in collating specimen data at the National Museum of Natural History (Naturalis) in Leiden. I am grateful to two anonymous reviewers for their constructive criticism of the manuscript. Miguel Morón (Instituto de Ecología, Xalapa,
<collectingCountry box="[377,463,871,898]" name="Mexico" pageId="18" pageNumber="123">Mexico</collectingCountry>
) is gratefully acknowledged for permission to use Figures 69, which originally appeared in his 2007 paper on
<taxonomicName authorityName="Moron &amp; Nogueira" authorityYear="2007" box="[744,886,906,932]" class="Insecta" family="Cetoniidae" genus="Halffterinetis" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="123" phylum="Arthropoda" rank="genus">
<emphasis box="[744,886,906,932]" italics="true" pageId="18" pageNumber="123">Halffterinetis</emphasis>
</taxonomicName>
. I thank Angie Fox (Scientific Illustrator, University of Nebraska State Museum) for making ready the figures, maps, and image files. This project was supported by an NSF/BS&amp;I grant (DEB 0716899) to B. C. Ratcliffe and R. D. Cave and an NSF Multiuser Equipment grant (DBI 0500767) to M. L. Jameson and F. Ocampo.
</paragraph>
</subSubSection>
</treatment>
</document>