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<mods:title id="394428B67BC53BAED62FCE4427AC5766">Early Eocene snakeflies (Raphidioptera) of western North America from the Okanagan Highlands and Green River Formation</mods:title>
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<mods:affiliation id="AEFB29E8F7F91FDEC30258FBB6E2E0E5">Department of Biological Sciences, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia, V 5 A 1 S 6, Canada. https: // orcid. org / 0000 - 0002 - 4397 - 2497 &amp; Museum of Comparative Zoology, 26 Oxford Street, Cambridge, Massachusetts, 02138, United States of America &amp; Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia, V 8 W 9 W 2, Canada</mods:affiliation>
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<taxonomicName id="4C774D1BFFEBC453FF67F895CD2A874C" ID-CoL="6236X" ID-ENA="50482" authorityName="Navas" authorityYear="1916" box="[151,325,1842,1868]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="29" pageNumber="70" phylum="Arthropoda" rank="order">Raphidioptera</taxonomicName>
and the necessity of cold winter
</emphasis>
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It has been generally thought that extant snakeflies (
<taxonomicName id="4C774D1BFFEBC453FCF8F8F0CFE18771" authorityName="Navas" authorityYear="1913" box="[776,910,1879,1905]" class="Insecta" family="Inocelliidae" kingdom="Animalia" order="Raphidioptera" pageId="29" pageNumber="70" phylum="Arthropoda" rank="family">Inocelliidae</taxonomicName>
and
<taxonomicName id="4C774D1BFFEBC453FC33F8F0C8238772" authorityName="Latreille" authorityYear="1810" box="[963,1100,1879,1906]" class="Insecta" family="Raphidiidae" kingdom="Animalia" order="Raphidioptera" pageId="29" pageNumber="70" phylum="Arthropoda" rank="family">Raphidiidae</taxonomicName>
) require a cold interval to mature, explaining their restriction to mid- and higher latitudes of the Northern Hemisphere, reportedly in high, cooler elevations in the southern portions of their range (
<bibRefCitation id="EFE64B69FFEBC453FD36F838CF0887B9" author="Aspock, H." box="[710,871,1951,1977]" pageId="29" pageNumber="70" pagination="33 - 44" refId="ref20619" refString="Aspock, H. (1998) Distribution and biogeography of the order Raphidioptera: updated facts and a new hypothesis. Acta Zoologica Fennica, 209, 33 - 44." type="journal article" year="1998">Aspöck, 1998</bibRefCitation>
,
<bibRefCitation id="EFE64B69FFEBC453FC82F838CFC287B9" author="Aspock, H." box="[882,941,1951,1977]" pageId="29" pageNumber="70" pagination="223 - 233" refId="ref20651" refString="Aspock, H. (2000) Der endkreidezeitliche Impakt und das Uberleben der Raphidiopteren. Entomologica Basiliena, 22, 223 - 233." type="journal article" year="2000">2000</bibRefCitation>
,
<bibRefCitation id="EFE64B69FFEBC453FC48F838CF9B87B9" author="Aspock, H." box="[952,1012,1951,1977]" pageId="29" pageNumber="70" pagination="35 - 50" refId="ref20676" refString="Aspock, H. (2002) The biology of Raphidioptera: a review of present knowledge. In: G. Sziraki, G. (Ed.), Neuropterology 2000. Proceedings of the Seventh International Symposium on Neuropterology. Acta Zoologica Academiae Scientiarum Hungaricae, 48 (Supplement 2), 35 - 50." type="journal article" year="2002">2002</bibRefCitation>
; Aspöck &amp; Aspöck, 2009, 2014; Abbt
<emphasis id="B903EA8AFFEBC453FF67F864CCA587DD" box="[151,202,1987,2013]" italics="true" pageId="29" pageNumber="70">et al</emphasis>
., 2018).
<bibRefCitation id="EFE64B69FFEBC453FEC1F864CDB587DD" author="Aspock, H." box="[305,474,1987,2013]" pageId="29" pageNumber="70" pagination="223 - 233" refId="ref20651" refString="Aspock, H. (2000) Der endkreidezeitliche Impakt und das Uberleben der Raphidiopteren. Entomologica Basiliena, 22, 223 - 233." type="journal article" year="2000">Aspöck (2000)</bibRefCitation>
stated that the cold interval is necessary to induce pupation and develop the imago, which must be 0° C or below. Abbt
<emphasis id="B903EA8AFFE8C450FDDEFF30CE0F80B1" box="[558,608,151,177]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
. (2018) later found that at least some can mature in the laboratory with a cold interval of 4°C.
</paragraph>
<paragraph id="8BC83698FFE8C450FF37FF78C9398189" blockId="30.[151,1437,151,2013]" pageId="30" pageNumber="71">
This would account for their exclusion from the tropics. In North America, they are found west of the Rocky Mountains from about
<geoCoordinate id="EE43505FFFE8C450FE68FEA5CDB5811D" box="[408,474,258,285]" degrees="53" direction="north" orientation="latitude" pageId="30" pageNumber="71" precision="55555" value="53.0">53° N</geoCoordinate>
in southern British Columbia with some known from as far south as the Mexico-Guatemala border. In the Eastern Hemisphere, they range from
<collectingCountry id="F3607608FFE8C450FCC0FE80CFE58141" box="[816,906,295,321]" name="Norway" pageId="30" pageNumber="71">Norway</collectingCountry>
at
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as far south as parts of Mediterranean North Africa, with scattered records eastward through Asia to the Pacific, southward to the Himalayas, northern
<collectingCountry id="F3607608FFE8C450FF67FEC8CC948189" box="[151,251,367,393]" name="Thailand" pageId="30" pageNumber="71">Thailand</collectingCountry>
and
<collectingCountry id="F3607608FFE8C450FEC1FEC8CDEC8189" box="[305,387,367,393]" name="Taiwan" pageId="30" pageNumber="71">Taiwan</collectingCountry>
(
<bibRefCitation id="EFE64B69FFE8C450FE61FEC8CE5E8189" author="Aspock, H." box="[401,561,367,393]" pageId="30" pageNumber="71" pagination="33 - 44" refId="ref20619" refString="Aspock, H. (1998) Distribution and biogeography of the order Raphidioptera: updated facts and a new hypothesis. Acta Zoologica Fennica, 209, 33 - 44." type="journal article" year="1998">Aspöck, 1998</bibRefCitation>
;
<bibRefCitation id="EFE64B69FFE8C450FDCCFEC8CF778189" author="Aspock, U. &amp; Liu, X. Y. &amp; Rausch, H. &amp; Aspock, H." box="[572,792,367,393]" pageId="30" pageNumber="71" pagination="259 - 274" refId="ref21624" refString="Aspock, U., Liu, X. Y., Rausch, H. &amp; Aspock, H. (2011) The Inocelliidae of southeast Asia: a review of present knowledge (Raphidioptera). Deutsche Entomologische Zeitschrift, (N. F.), 58, 259 - 274. https: // doi. org / 10.1002 / mmnd. 201100029" type="journal article" year="2011">
Aspöck
<emphasis id="B903EA8AFFE8C450FD68FEC8CEA68189" box="[664,713,367,393]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 2011
</bibRefCitation>
,
<bibRefCitation id="EFE64B69FFE8C450FCD3FEC8CF058189" author="Aspock, U. &amp; Haring, E. &amp; Aspock, H." box="[803,874,367,393]" pageId="30" pageNumber="71" pagination="575 - 582" refId="ref21485" refString="Aspock, U., Haring, E. &amp; Aspock, H. (2012 a) Biogeographical implications of a molecular phylogeny of the Raphidiidae (Raphidioptera). Mitteilungen der Deutschen Gesellschaft fur allgeimeine und angewandte Entomologie, 18, 575 - 582." type="journal article" year="2012">2012a</bibRefCitation>
, b; Aspöck &amp; Aspöck, 2014;
<bibRefCitation id="EFE64B69FFE8C450FB43FEC8C9258189" author="Blades, D. C. A." box="[1203,1354,367,393]" pageId="30" pageNumber="71" pagination="383 - 386" refId="ref21841" refString="Blades, D. C. A. (2019) Raphidioptera of Canada. ZooKeys, 819, 383 - 386. https: // doi. org / 10.3897 / zookeys. 819.26626" type="journal article" year="2019">Blades, 2019</bibRefCitation>
).
</paragraph>
<paragraph id="8BC83698FFE8C450FF37FE34C85582CE" blockId="30.[151,1437,151,2013]" pageId="30" pageNumber="71">
The fossil record of
<taxonomicName id="4C774D1BFFE8C450FE58FE34CE2881AD" authorityName="Navas" authorityYear="1916" box="[424,583,403,429]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Raphidioptera</taxonomicName>
before the Oligocene, especially of the extant families, would then present a biogeographic puzzle. The Eocene/Oligocene boundary represents the end of the globally warm “greenhouse world” climates of the Cretaceous through the Paleocene and Eocene, with higher mean annual temperatures (MAT), a low pole to equator MAT gradient, and lowered temperature seasonality with mild winters. The post-Eocene onset of the modern “icehouse world” regime saw extra-tropical climates with colder MAT, lacking Arctic ice sheets until the later Neogene and Antarctic ice sheets beginning to form at the end of the Eocene, an increased latitudinal MAT gradient, and more severe extra-tropical winters (
<bibRefCitation id="EFE64B69FFE8C450FD6BFDCCCF188285" author="Zachos, J. C. &amp; Dickens, G. R. &amp; Zeebe, R. E." box="[667,887,619,645]" pageId="30" pageNumber="71" pagination="279 - 283" refId="ref26092" refString="Zachos, J. C., Dickens, G. R. &amp; Zeebe, R. E. (2008) An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics. Nature, 451, 279 - 283. https: // doi. org / 10.1038 / nature 06588" type="journal article" year="2008">
Zachos
<emphasis id="B903EA8AFFE8C450FD04FDCCCF498285" box="[756,806,619,645]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 2008
</bibRefCitation>
).
<taxonomicName id="4C774D1BFFE8C450FC7AFDCCC8468285" authorityName="Navas" authorityYear="1916" box="[906,1065,619,645]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Raphidioptera</taxonomicName>
of the Paleocene and Eocene did not then experience cold winters, and in the Mesozoic lived under even warmer conditions, some in low latitudes,
<emphasis id="B903EA8AFFE8C450FF67FD14CCD782CC" box="[151,184,691,716]" italics="true" pageId="30" pageNumber="71">e.g</emphasis>
.,
<collectingCountry id="F3607608FFE8C450FF3EFD14CD7D82CD" box="[206,274,691,717]" name="Brazil" pageId="30" pageNumber="71">Brazil</collectingCountry>
(
<emphasis id="B903EA8AFFE8C450FED2FD14CD2B82CC" box="[290,324,691,716]" italics="true" pageId="30" pageNumber="71">e.g</emphasis>
.,
<bibRefCitation id="EFE64B69FFE8C450FEA9FD14CD9A82CD" author="Oswald, J. D." box="[345,501,691,717]" pageId="30" pageNumber="71" pagination="154 - 163" refId="ref24369" refString="Oswald, J. D. (1990) Raphidioptera. In: Grimaldi, D. A. (Ed.), Insects from the Santana Formation, Lower Cretaceous, of Brazil. Bulletin of the American Museum of Natural History, 195, 154 - 163." type="journal article" year="1990">Oswald, 1990</bibRefCitation>
) and
<collectingCountry id="F3607608FFE8C450FDC4FD14CECE82CD" box="[564,673,691,717]" name="Myanmar" pageId="30" pageNumber="71">Myanmar</collectingCountry>
(
<emphasis id="B903EA8AFFE8C450FD41FD14CEBD82CC" box="[689,722,691,716]" italics="true" pageId="30" pageNumber="71">e.g</emphasis>
.,
<bibRefCitation id="EFE64B69FFE8C450FD18FD14CF1C82CD" author="Engel, M. S." box="[744,883,691,717]" pageId="30" pageNumber="71" pagination="1 - 22" refId="ref22701" refString="Engel, M. S. (2002) The smallest snakefly (Raphidioptera: Mesoraphidiidae): A new species in Cretaceous amber from Myanmar, with a catalog of fossil snakeflies. American Museum Novitates, 3363, 1 - 22." type="journal article" year="2002">Engel, 2002</bibRefCitation>
;
<bibRefCitation id="EFE64B69FFE8C450FC8EFD14C84082CE" author="Liu, X. Y. &amp; Lu, X. M. &amp; Zhang, W. W." box="[894,1071,691,718]" pageId="30" pageNumber="71" pagination="301 - 324" refId="ref23579" refString="Liu, X. Y., Lu, X. M. &amp; Zhang, W. W. (2016) New genera and species of the minute snakeflies (Raphidioptera: Mesoraphidiidae: Nanoraphidiini) from the mid Cretaceous of Myanmar. Zootaxa, 4103, 301 - 324. https: // doi. org / 10.11646 / zootaxa. 4103.4.1" type="journal article" year="2016">
Liu
<emphasis id="B903EA8AFFE8C450FC5CFD14CFB282CD" box="[940,989,691,717]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 2016
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C36D6513FFE8C450FF37FD70C8808699" pageId="30" pageNumber="71" type="reference_group">
<paragraph id="8BC83698FFE8C450FF37FD70C82C83C9" blockId="30.[151,1437,151,2013]" pageId="30" pageNumber="71">
<taxonomicName id="4C774D1BFFE8C450FF37FD70CE4A82F1" ID-CoL="BB9K" authority="(Albarda, 1891)" baseAuthorityName="Albarda" baseAuthorityYear="1891" box="[199,549,727,753]" class="Insecta" family="Raphidiidae" genus="Agulla" kingdom="Animalia" order="Raphidioptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="species" species="bicolor">
<emphasis id="B903EA8AFFE8C450FF37FD70CD0582F1" box="[199,362,727,753]" italics="true" pageId="30" pageNumber="71">Agulla bicolor</emphasis>
(
<bibRefCitation id="EFE64B69FFE8C450FE89FD70CE7282F1" author="Albarda, H." box="[377,541,727,753]" pageId="30" pageNumber="71" pagination="65 - 184" refId="ref19852" refString="Albarda, H. (1891) Revision des Rhaphidides. Tijdschrift voor Entomologie, 34, 65 - 184." type="journal article" year="1891">Albarda, 1891</bibRefCitation>
)
</taxonomicName>
from Texas was successfully raised in laboratory conditions at a constant temperature of 23 ± 3°C (
<bibRefCitation id="EFE64B69FFE8C450FE78FD5CCE038315" author="Kovarik, P. W. &amp; Burke, H. R. &amp; Agnew, C. W." box="[392,620,763,789]" pageId="30" pageNumber="71" pagination="353 - 364" refId="ref23307" refString="Kovarik, P. W., Burke, H. R. &amp; Agnew, C. W. (1991) Development and behavior of a snakefly, Raphidia bicolor Albarda (Neuroptera: Raphidiidae). Southwestern Entomologist, 16, 353 - 364." type="journal article" year="1991">
Kovarik
<emphasis id="B903EA8AFFE8C450FE1CFD5CCE738315" box="[492,540,763,789]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 1991
</bibRefCitation>
).
<taxonomicName id="4C774D1BFFE8C450FD8FFD5CCF698315" ID-CoL="FJZ" authorityName="Latreille" authorityYear="1810" box="[639,774,763,789]" class="Insecta" family="Raphidiidae" kingdom="Animalia" order="Raphidioptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Raphidiidae</taxonomicName>
of the North American genus
<taxonomicName id="4C774D1BFFE8C450FBA4FD5CC8FA8315" ID-CoL="RPT" authorityName="Navas" authorityYear="1916" box="[1108,1173,763,789]" class="Insecta" family="Raphidiidae" genus="Alena" kingdom="Animalia" order="Raphidioptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="genus">
<emphasis id="B903EA8AFFE8C450FBA4FD5CC8FA8315" box="[1108,1173,763,789]" italics="true" pageId="30" pageNumber="71">Alena</emphasis>
</taxonomicName>
and Mexican inocelliid species apparently also dont require a cold interval to mature, as these pupate in summer and the imago emerges in late summer (
<bibRefCitation id="EFE64B69FFE8C450FEC2FCE4CDBA835D" author="Aspock, H." box="[306,469,835,861]" pageId="30" pageNumber="71" pagination="35 - 50" refId="ref20676" refString="Aspock, H. (2002) The biology of Raphidioptera: a review of present knowledge. In: G. Sziraki, G. (Ed.), Neuropterology 2000. Proceedings of the Seventh International Symposium on Neuropterology. Acta Zoologica Academiae Scientiarum Hungaricae, 48 (Supplement 2), 35 - 50." type="journal article" year="2002">Aspöck, 2002</bibRefCitation>
). At least two species of
<taxonomicName id="4C774D1BFFE8C450FD02FCE4CF90835D" ID-CoL="4TVY" authority="Steinmann" authorityName="Steinmann" authorityYear="1963" box="[754,1023,835,861]" class="Insecta" family="Raphidiidae" genus="Harraphidia" kingdom="Animalia" order="Raphidioptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="genus">
<emphasis id="B903EA8AFFE8C450FD02FCE4CFEF835D" box="[754,896,835,861]" italics="true" pageId="30" pageNumber="71">Harraphidia</emphasis>
Steinmann
</taxonomicName>
in the Iberian Peninsula inhabit low elevation regions without cold winters, some at locations on the Mediterranean coast such as Tarifa,
<collectingCountry id="F3607608FFE8C450FAF9FCC0C9238381" box="[1289,1356,871,897]" name="Spain" pageId="30" pageNumber="71">Spain</collectingCountry>
, which has a coldest month mean temperature of 13.0°C and minimum temperature of 8°C (
<bibRefCitation id="EFE64B69FFE8C450FB9EFC2CCCBC83C9" author="Monserrat, V. J. &amp; Papenberg, D." pageId="30" pageNumber="71" pagination="203 - 222" refId="ref24008" refString="Monserrat, V. J. &amp; Papenberg, D. (2006) Revision del genero Harraphidia Steinmann, 1963 con la descripcion de dos nuevas especies de la peninsula Iberica y de Marruecos (Insecta, Raphidioptera). Graellsia, 62, 203 - 222. https: // doi. org / 10.3989 / graellsia. 2006. v 62. i 2.67" type="journal article" year="2006">Monserrat and Papenberg, 2006</bibRefCitation>
;
<bibRefCitation id="EFE64B69FFE8C450FF2CFC08CDE983C9" author="AEMET" box="[220,390,943,969]" pageId="30" pageNumber="71" refId="ref19824" refString="AEMET (2020) La Agencia Estatal de Meteorologia. Online. [http: // www. aemet. es / en / serviciosclimaticos / datosclimatologicos]" type="url" year="2020">AEMET, 2020</bibRefCitation>
). A cold interval is then not required for all modern snakeflies.
</paragraph>
<paragraph id="8BC83698FFE8C450FF37FC74CF428555" blockId="30.[151,1437,151,2013]" pageId="30" pageNumber="71">
<taxonomicName id="4C774D1BFFE8C450FF37FC74CD2483ED" ID-CoL="624VG" authorityName="Navas" authorityYear="1913" box="[199,331,979,1005]" class="Insecta" family="Inocelliidae" kingdom="Animalia" order="Raphidioptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Inocelliidae</taxonomicName>
and
<taxonomicName id="4C774D1BFFE8C450FE70FC74CE6883ED" ID-CoL="FJZ" authorityName="Latreille" authorityYear="1810" box="[384,519,979,1005]" class="Insecta" family="Raphidiidae" kingdom="Animalia" order="Raphidioptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Raphidiidae</taxonomicName>
of the Okanagan Highlands and Green River Formation are from the latter portion of the Ypresian, which had the hottest sustained global MAT values of the Cenozoic, the Early Eocene Climatic Optimum (
<bibRefCitation id="EFE64B69FFE8C450FF1FFBBCCDA48435" author="Zachos, J. C. &amp; Dickens, G. R. &amp; Zeebe, R. E." box="[239,459,1051,1077]" pageId="30" pageNumber="71" pagination="279 - 283" refId="ref26092" refString="Zachos, J. C., Dickens, G. R. &amp; Zeebe, R. E. (2008) An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics. Nature, 451, 279 - 283. https: // doi. org / 10.1038 / nature 06588" type="journal article" year="2008">
Zachos
<emphasis id="B903EA8AFFE8C450FEB8FBBCCD168435" box="[328,377,1051,1077]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 2008
</bibRefCitation>
). While nearby coastal climates had hot MATs, the interior Okanagan Highlands localities bearing
<taxonomicName id="4C774D1BFFE8C450FED0FB98CDD08459" ID-CoL="6236X" authorityName="Navas" authorityYear="1916" box="[288,447,1087,1113]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Raphidioptera</taxonomicName>
were at some elevation, and had cooler, upper microthermal MAT values (microthermal = MAT ≤13°C), similar to that of south-coastal
<collectingRegion id="49B3F87AFFE8C450FD55FBC4CF0A847D" box="[677,869,1123,1149]" country="Canada" name="British Columbia" pageId="30" pageNumber="71">British Columbia</collectingRegion>
today (
<bibRefCitation id="EFE64B69FFE8C450FC47FBC4C8EA847D" author="Rouse, G. E. &amp; Hopkins, W. S., Jr. &amp; Piel, K. M." box="[951,1157,1123,1149]" pageId="30" pageNumber="71" pagination="213 - 246" refId="ref25089" refString="Rouse, G. E., Hopkins, W. S., Jr. &amp; Piel, K. M. (1970) Palynology of some Late Cretaceous and Early Tertiary deposits in British Columbia and adjacent Alberta. Geological Society of America, Special Paper, 127, 213 - 246. https: // doi. org / 10.1130 / SPE 127 - p 213" type="journal article" year="1970">
Rouse
<emphasis id="B903EA8AFFE8C450FBF5FBC4C859847D" box="[1029,1078,1123,1149]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 1970
</bibRefCitation>
;
<bibRefCitation id="EFE64B69FFE8C450FB7FFBC4C9F8847D" author="Greenwood, D. R. &amp; Archibald, S. B. &amp; Mathewes, R. W. &amp; Moss, P. T." box="[1167,1431,1122,1149]" pageId="30" pageNumber="71" pagination="167 - 185" refId="ref23003" refString="Greenwood, D. R., Archibald, S. B., Mathewes, R. W. &amp; Moss, P. T. (2005) Fossil biotas from the Okanagan Highlands, southern British Columbia and northeastern Washington State: climates and ecosystems across an Eocene landscape. Canadian Journal of Earth Sciences, 42, 167 - 185. https: // doi. org / 10.1139 / e 04 - 100" type="journal article" year="2005">
Greenwood
<emphasis id="B903EA8AFFE8C450FAE8FBC4C926847D" box="[1304,1353,1123,1149]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 2005
</bibRefCitation>
;
<bibRefCitation id="EFE64B69FFE8C450FF67FB20CD7084A1" author="Tribe, S." box="[151,287,1158,1185]" pageId="30" pageNumber="71" pagination="215 - 230" refId="ref25378" refString="Tribe, S. (2005) Eocene paleo-physiography and drainage directions, southern Interior Plateau, British Columbia. Canadian Journal of Earth Sciences, 42, 215 - 230. https: // doi. org / 10.1139 / e 04 - 062" type="journal article" year="2005">Tribe, 2005</bibRefCitation>
;
<bibRefCitation id="EFE64B69FFE8C450FEDBFB20CE4384A1" author="Archibald, S. B. &amp; Greenwood, D. R. &amp; Smith, R. Y. &amp; Mathewes, R. W. &amp; Basinger, J. F." box="[299,556,1159,1185]" pageId="30" pageNumber="71" pagination="145 - 154" refId="ref20229" refString="Archibald, S. B., Greenwood, D. R., Smith, R. Y., Mathewes, R. W. &amp; Basinger, J. F. (2011) Early Eocene lagerstatten of the Okanagan Highlands (British Columbia and Washington State). Geoscience Canada, 38, 145 - 154." type="journal article" year="2011">
Archibald
<emphasis id="B903EA8AFFE8C450FE55FB20CDB684A1" box="[421,473,1159,1185]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 2011
</bibRefCitation>
). These values would usually be associated with cold winters in the modern seasonal Northern Hemisphere, however, winters there appeared to be mild, without frost days; its forests included plants such as palms and their obligate palm feeding bruchids (
<taxonomicName id="4C774D1BFFE8C450FCBEFB68CFA184E9" ID-CoL="C2L" authorityName="Linnaeus" authorityYear="1758" box="[846,974,1231,1257]" class="Insecta" kingdom="Animalia" order="Coleoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
,
<taxonomicName id="4C774D1BFFE8C450FC27FB68C8EC84E9" ID-CoL="87S" authorityName="Latreille" authorityYear="1802" box="[983,1155,1231,1257]" class="Insecta" family="Chrysomelidae" kingdom="Animalia" order="Coleoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Chrysomelidae</taxonomicName>
,
<taxonomicName id="4C774D1BFFE8C450FB7EFB68CD7D850D" authority=": Archibald et al. 2014" authorityName="Archibald" authorityYear="2014" class="Insecta" genus="Pachymerina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="genus">
Pachymerina:
<bibRefCitation id="EFE64B69FFE8C450FADCFB68CD7D850D" author="Archibald, S. B. &amp; Morse, G. E. &amp; Greenwood, D. R. &amp; Mathewes, R. W." pageId="30" pageNumber="71" pagination="8095 - 8100" refId="ref20345" refString="Archibald, S. B., Morse, G. E., Greenwood, D. R. &amp; Mathewes, R. W. (2014) Fossil palm beetles refine upland winter temperatures in the Early Eocene Climatic Optimum. Proceedings of the National Academy of Sciences of the United States of America, 111, 8095 - 8100. https: // doi. org / 10.1073 / pnas. 1323269111" type="journal article" year="2014">
Archibald
<emphasis id="B903EA8AFFE8C450FF67FB54CCA7850D" box="[151,200,1267,1293]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
. 2014
</bibRefCitation>
</taxonomicName>
), as well as other insects restricted to frost-free regions today,
<emphasis id="B903EA8AFFE8C450FC3DFB54CF81850C" box="[973,1006,1267,1292]" italics="true" pageId="30" pageNumber="71">e.g</emphasis>
.,
<taxonomicName id="4C774D1BFFE8C450FBF3FB54C90D850D" ID-CoL="5N7L" authority="Guerin-Meneville" authorityName="Guerin-Meneville" box="[1027,1378,1267,1293]" class="Insecta" family="Dinidoridae" genus="Megymenum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hemiptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="genus">
<emphasis id="B903EA8AFFE8C450FBF3FB54C8FD850C" box="[1027,1170,1267,1292]" italics="true" pageId="30" pageNumber="71">Megymenum</emphasis>
Guérin-Méneville
</taxonomicName>
(
<taxonomicName id="4C774D1BFFE8C450FA81FB54CC808531" ID-CoL="HP" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hemiptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Hemiptera</taxonomicName>
,
<taxonomicName id="4C774D1BFFE8C450FF0FFAB0CDE68532" ID-CoL="6248K" box="[255,393,1303,1330]" class="Insecta" family="Dinidoridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hemiptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Dinidoridae</taxonomicName>
),
<taxonomicName id="4C774D1BFFE8C450FE6FFAB0CEBD8531" ID-CoL="8NKN6" authority="(Isoptera)" authorityName="Desneux" authorityYear="1904" baseAuthorityName="Isoptera" box="[415,722,1303,1329]" class="Insecta" family="Mastotermitidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Blattodea" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Mastotermitidae (Isoptera)</taxonomicName>
,
<taxonomicName id="4C774D1BFFE8C450FD11FAB0CF198531" ID-CoL="JXZ" box="[737,886,1303,1329]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="subFamily" subFamily="Myrmeciinae">Myrmeciinae</taxonomicName>
(
<taxonomicName id="4C774D1BFFE8C450FC78FAB0C84A8531" ID-CoL="HYM" box="[904,1061,1303,1329]" class="Insecta" kingdom="Animalia" order="Hymenoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Hymenoptera</taxonomicName>
,
<taxonomicName id="4C774D1BFFE8C450FBC4FAB0C8D78532" ID-CoL="623T5" box="[1076,1208,1303,1330]" class="Insecta" family="Formicidae" kingdom="Animalia" order="Hymenoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Formicidae</taxonomicName>
), and
<taxonomicName id="4C774D1BFFE8C450FAF2FAB0C9F38531" ID-CoL="8MPMF" baseAuthorityName="Saussure" baseAuthorityYear="1864" box="[1282,1436,1303,1329]" class="Insecta" family="Blaberidae" kingdom="Animalia" order="Blattodea" pageId="30" pageNumber="71" phylum="Arthropoda" rank="subFamily" subFamily="Diplopterinae">Diplopterinae</taxonomicName>
(
<taxonomicName id="4C774D1BFFE8C450FF6FFA9CCD608555" ID-CoL="TM" authorityName="Brunner von Wattenwyl" authorityYear="1882" box="[159,271,1339,1365]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Blattodea" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Blattodea</taxonomicName>
,
<taxonomicName id="4C774D1BFFE8C450FEEAFA9CCDFA8556" ID-CoL="8MPBB" box="[282,405,1339,1366]" class="Insecta" family="Blaberidae" kingdom="Animalia" order="Blattodea" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Blaberidae</taxonomicName>
) (
<emphasis id="B903EA8AFFE8C450FE5DFA9CCDBA8554" box="[429,469,1339,1364]" italics="true" pageId="30" pageNumber="71">e.g.</emphasis>
,
<bibRefCitation id="EFE64B69FFE8C450FE12FA9CCEB58555" author="Archibald, S. B. &amp; Morse, G. E. &amp; Greenwood, D. R. &amp; Mathewes, R. W." box="[482,730,1339,1365]" pageId="30" pageNumber="71" pagination="8095 - 8100" refId="ref20345" refString="Archibald, S. B., Morse, G. E., Greenwood, D. R. &amp; Mathewes, R. W. (2014) Fossil palm beetles refine upland winter temperatures in the Early Eocene Climatic Optimum. Proceedings of the National Academy of Sciences of the United States of America, 111, 8095 - 8100. https: // doi. org / 10.1073 / pnas. 1323269111" type="journal article" year="2014">
Archibald
<emphasis id="B903EA8AFFE8C450FDA9FA9CCEE58555" box="[601,650,1339,1365]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 2014
</bibRefCitation>
,
<bibRefCitation id="EFE64B69FFE8C450FD15FA9CCF4D8555" author="Archibald, S. B. &amp; Rasnitsyn, A. P. &amp; Brothers, D. J. &amp; Mathewes, R. W." box="[741,802,1339,1365]" pageId="30" pageNumber="71" pagination="205 - 257" refId="ref20548" refString="Archibald, S. B., Rasnitsyn, A. P., Brothers, D. J. &amp; Mathewes, R. W. (2018) Modernisation of the Hymenoptera: ants, bees, wasps and sawflies of the early Eocene Okanagan Highlands. The Canadian Entomologist, 150, 205 - 257. https: // doi. org / 10.4039 / tce. 2017.59" type="journal article" year="2018">2018</bibRefCitation>
).
</paragraph>
<paragraph id="8BC83698FFE8C450FF37FAF8C8808699" blockId="30.[151,1437,151,2013]" pageId="30" pageNumber="71">
The regional tectonic uplift in the Ypresian that raised the Okanagan Highlands created a landscape of great topographic relief, with high mountains and deeply incised valleys in which the fossil-bearing lacustrine depositional basins formed (
<bibRefCitation id="EFE64B69FFE8C450FEB2FA00CDBA85C1" author="Ewing, T." box="[322,469,1447,1473]" pageId="30" pageNumber="71" pagination="619 - 638" refId="ref22877" refString="Ewing, T. (1980) Paleogene tectonic evolution of the Pacific Northwest. The Journal of Geology, 88, 619 - 638. https: // doi. org / 10.1086 / 628551" type="journal article" year="1980">Ewing, 1980</bibRefCitation>
;
<bibRefCitation id="EFE64B69FFE8C450FE2FFA00CE0A85C1" author="Tribe, S." box="[479,613,1446,1473]" pageId="30" pageNumber="71" pagination="215 - 230" refId="ref25378" refString="Tribe, S. (2005) Eocene paleo-physiography and drainage directions, southern Interior Plateau, British Columbia. Canadian Journal of Earth Sciences, 42, 215 - 230. https: // doi. org / 10.1139 / e 04 - 062" type="journal article" year="2005">Tribe, 2005</bibRefCitation>
). Its snakefly fossils are in good general condition, having not suffered the extensive mechanical damage that would be consistent with post-mortem downslope transport from cooler elevations in streams feeding the depositional lakes. Nor can their presence be easily explained by occasional transport of living adults by winds from local higher elevations. The larvae of modern
<taxonomicName id="4C774D1BFFE8C450FC24F9B4C81C862D" ID-CoL="6236X" authorityName="Navas" authorityYear="1916" box="[980,1139,1555,1581]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Raphidioptera</taxonomicName>
live from one to six years, usually under bark, but some in litter below trees or bushes (
<bibRefCitation id="EFE64B69FFE8C450FCC8F990CFB68651" author="Aspock, H." box="[824,985,1591,1617]" pageId="30" pageNumber="71" pagination="35 - 50" refId="ref20676" refString="Aspock, H. (2002) The biology of Raphidioptera: a review of present knowledge. In: G. Sziraki, G. (Ed.), Neuropterology 2000. Proceedings of the Seventh International Symposium on Neuropterology. Acta Zoologica Academiae Scientiarum Hungaricae, 48 (Supplement 2), 35 - 50." type="journal article" year="2002">Aspöck, 2002</bibRefCitation>
). Most adults live for a very short while, often a few days, and are weak fliers with low vagility, remaining close to their resident tree or bush (
<bibRefCitation id="EFE64B69FFE8C450FB00F9FCC9FF8675" author="Aspock, H." box="[1264,1424,1627,1653]" pageId="30" pageNumber="71" pagination="33 - 44" refId="ref20619" refString="Aspock, H. (1998) Distribution and biogeography of the order Raphidioptera: updated facts and a new hypothesis. Acta Zoologica Fennica, 209, 33 - 44." type="journal article" year="1998">Aspöck, 1998</bibRefCitation>
). We believe that Okanagan Highlands snakeflies lived in close proximity to their depositional lakes.
</paragraph>
</subSubSection>
<subSubSection id="C36D6513FFE8C451FF37F904CD958141" lastPageId="31" lastPageNumber="72" pageId="30" pageNumber="71" type="description">
<paragraph id="8BC83698FFE8C450FF37F904CDF2874D" blockId="30.[151,1437,151,2013]" pageId="30" pageNumber="71">
The Okanagan Highlands communities also included a suite of other insects that today inhabit cooler boreal regions, higher elevations in lower latitudes, or mostly so. These include aphids (
<taxonomicName id="4C774D1BFFE8C450FBB1F960C92086E2" authority=", Aphidoidea" authorityName="Aphidoidea" box="[1089,1359,1735,1762]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hemiptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Hemiptera, Aphidoidea</taxonomicName>
), scorpionflies (
<taxonomicName id="4C774D1BFFE8C450FEFAF94CCDE98705" box="[266,390,1771,1797]" class="Insecta" kingdom="Animalia" order="Mecoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Mecoptera</taxonomicName>
,
<taxonomicName id="4C774D1BFFE8C450FE63F94CCE7B8705" box="[403,532,1771,1797]" class="Insecta" family="Panorpidae" kingdom="Animalia" order="Mecoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Panorpidae</taxonomicName>
, species of
<taxonomicName id="4C774D1BFFE8C450FD6BF94CCF1E8705" authority="Linnaeus" authorityName="Linnaeus" authorityYear="1758" box="[667,881,1771,1797]" class="Insecta" family="Panorpidae" genus="Panorpa" kingdom="Animalia" order="Mecoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="genus">
<emphasis id="B903EA8AFFE8C450FD6BF94CCE928704" box="[667,765,1771,1796]" italics="true" pageId="30" pageNumber="71">Panorpa</emphasis>
Linnaeus
</taxonomicName>
:
<bibRefCitation id="EFE64B69FFE8C450FC8CF94CC8E98706" author="Archibald, S. B. &amp; Mathewes, R. W. &amp; Greenwood, D. R." box="[892,1158,1771,1798]" pageId="30" pageNumber="71" pagination="677 - 695" refId="ref20290" refString="Archibald, S. B., Mathewes, R. W. &amp; Greenwood, D. R. (2013 b) The Eocene apex of panorpoid scorpionfly family diversity. Journal of Paleontology, 87, 677 - 695. https: // doi. org / 10.1666 / 12 - 129" type="journal article" year="2013">
Archibald
<emphasis id="B903EA8AFFE8C450FC05F94CC8478705" box="[1013,1064,1771,1797]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 2013b
</bibRefCitation>
) and sawflies and wood wasps (
<taxonomicName id="4C774D1BFFE8C450FF1AF8A8CE688729" authority=", Symphyta" authorityName="Symphyta" box="[234,519,1807,1833]" class="Insecta" kingdom="Animalia" order="Hymenoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Hymenoptera, Symphyta</taxonomicName>
:
<taxonomicName id="4C774D1BFFE8C450FDE0F8A8CED48729" box="[528,699,1807,1833]" class="Insecta" family="Tenthredinidae" kingdom="Animalia" order="Hymenoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Tenthredinidae</taxonomicName>
,
<taxonomicName id="4C774D1BFFE8C450FD35F8A8CF448729" authorityName="Billberg" authorityYear="1820" box="[709,811,1807,1833]" class="Insecta" family="Siricidae" kingdom="Animalia" order="Hymenoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Siricidae</taxonomicName>
,
<taxonomicName id="4C774D1BFFE8C450FCC6F8A8CFA28729" authorityName="Cameron" authorityYear="1890" box="[822,973,1807,1833]" class="Insecta" family="Pamphiliidae" kingdom="Animalia" order="Hymenoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Pamphiliidae</taxonomicName>
,
<taxonomicName id="4C774D1BFFE8C450FC28F8A8C8298729" box="[984,1094,1807,1833]" class="Insecta" family="Cephidae" kingdom="Animalia" order="Hymenoptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Cephidae</taxonomicName>
: Archibald &amp; Rasnitsyn, 2015;
<bibRefCitation id="EFE64B69FFE8C450FF67F894CDFE874D" author="Archibald, S. B. &amp; Rasnitsyn, A. P. &amp; Brothers, D. J. &amp; Mathewes, R. W." box="[151,401,1843,1869]" pageId="30" pageNumber="71" pagination="205 - 257" refId="ref20548" refString="Archibald, S. B., Rasnitsyn, A. P., Brothers, D. J. &amp; Mathewes, R. W. (2018) Modernisation of the Hymenoptera: ants, bees, wasps and sawflies of the early Eocene Okanagan Highlands. The Canadian Entomologist, 150, 205 - 257. https: // doi. org / 10.4039 / tce. 2017.59" type="journal article" year="2018">
Archibald
<emphasis id="B903EA8AFFE8C450FEFEF894CD50874D" box="[270,319,1843,1869]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 2018
</bibRefCitation>
).
</paragraph>
<paragraph id="8BC83698FFE8C451FF37F8F0CD958141" blockId="30.[151,1437,151,2013]" lastBlockId="31.[151,1437,151,1510]" lastPageId="31" lastPageNumber="72" pageId="30" pageNumber="71">
The Okanagan Highlands green lacewings (
<taxonomicName id="4C774D1BFFE8C450FD43F8F0CF598771" box="[691,822,1879,1905]" class="Insecta" kingdom="Animalia" order="Neuroptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="order">Neuroptera</taxonomicName>
,
<taxonomicName id="4C774D1BFFE8C450FCB2F8F0CFBE8772" box="[834,977,1879,1906]" class="Insecta" family="Chrysopidae" kingdom="Animalia" order="Neuroptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="family">Chrysopidae</taxonomicName>
) consist of at least nine species of
<taxonomicName id="4C774D1BFFE8C450FAABF8F0CD7F8795" authorityName="Navas" authorityYear="1910" class="Insecta" family="Chrysopidae" kingdom="Animalia" order="Neuroptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="subFamily" subFamily="Nothochrysinae">Nothochrysinae</taxonomicName>
and one of the predominantly Mesozoic
<taxonomicName id="4C774D1BFFE8C450FD1FF8DCCF0C8795" baseAuthorityName="Martins-Neto &amp; Vulcano" baseAuthorityYear="1988" box="[751,867,1915,1941]" class="Insecta" family="Chrysopidae" kingdom="Animalia" order="Neuroptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="subFamily" subFamily="Limaiinae">Limaiinae</taxonomicName>
, its youngest occurrence. Today,
<taxonomicName id="4C774D1BFFE8C450FB1BF8DCC9F38795" authorityName="Navas" authorityYear="1910" box="[1259,1436,1915,1941]" class="Insecta" family="Chrysopidae" kingdom="Animalia" order="Neuroptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="subFamily" subFamily="Nothochrysinae">Nothochrysinae</taxonomicName>
mostly prefer Mediterranean climates of milder winters (
<bibRefCitation id="EFE64B69FFE8C450FCE3F838C86087B9" author="Archibald, S. B. &amp; Morse, G. E. &amp; Greenwood, D. R. &amp; Mathewes, R. W." box="[787,1039,1951,1977]" pageId="30" pageNumber="71" pagination="8095 - 8100" refId="ref20345" refString="Archibald, S. B., Morse, G. E., Greenwood, D. R. &amp; Mathewes, R. W. (2014) Fossil palm beetles refine upland winter temperatures in the Early Eocene Climatic Optimum. Proceedings of the National Academy of Sciences of the United States of America, 111, 8095 - 8100. https: // doi. org / 10.1073 / pnas. 1323269111" type="journal article" year="2014">
Archibald
<emphasis id="B903EA8AFFE8C450FC7CF838CFD287B9" box="[908,957,1951,1977]" italics="true" pageId="30" pageNumber="71">et al</emphasis>
., 2014
</bibRefCitation>
). Green lacewings of the
<taxonomicName id="4C774D1BFFE8C450FADCF838CCD187DD" class="Insecta" family="Chrysopidae" kingdom="Animalia" order="Neuroptera" pageId="30" pageNumber="71" phylum="Arthropoda" rank="subFamily" subFamily="Chrysopinae">Chrysopinae</taxonomicName>
thrive in a wide range of climates, including regions of very cold winters. They appear in the fossil record after Okanagan Highlands time at the end of the Eocene, subsequently diversifying to vastly dominate the family today across most of the globe from the equator to northern Siberia in our post-Eocene icehouse world as the
<taxonomicName id="4C774D1BFFE9C451FAABFF1CCD7F80F9" authorityName="Navas" authorityYear="1910" class="Insecta" family="Chrysopidae" kingdom="Animalia" order="Neuroptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="subFamily" subFamily="Nothochrysinae">Nothochrysinae</taxonomicName>
became relictual.
<taxonomicName id="4C774D1BFFE9C451FE15FF78CE0780F9" authorityName="Rambur" authorityYear="1842" box="[485,616,223,249]" class="Insecta" family="Nymphidae" kingdom="Animalia" order="Neuroptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="subFamily" subFamily="Nymphinae">Nymphinae</taxonomicName>
(
<taxonomicName id="4C774D1BFFE9C451FD88FF78CE9480F9" box="[632,763,223,249]" class="Insecta" kingdom="Animalia" order="Neuroptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="order">Neuroptera</taxonomicName>
,
<taxonomicName id="4C774D1BFFE9C451FCF8FF78CFE080FA" authorityName="Rambur" authorityYear="1842" box="[776,911,223,250]" class="Insecta" family="Nymphidae" kingdom="Animalia" order="Neuroptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Nymphidae</taxonomicName>
), found at two Okanagan Highlands localities (
<bibRefCitation id="EFE64B69FFE9C451FF6FFEA4CDF6811D" author="Archibald, S. B. &amp; Makarkin, V. N. &amp; Ansorge, J." box="[159,409,259,285]" pageId="31" pageNumber="72" pagination="59 - 68" refId="ref20487" refString="Archibald, S. B., Makarkin, V. N. &amp; Ansorge, J. (2009) New fossil species of Nymphidae (Neuroptera) from the Eocene of North America and Europe. Zootaxa, 2157 (1), 59 - 68. https: // doi. org / 10.11646 / zootaxa. 2157.1.4" type="journal article" year="2009">
Archibald
<emphasis id="B903EA8AFFE9C451FEE8FEA4CD26811D" box="[280,329,259,285]" italics="true" pageId="31" pageNumber="72">et al</emphasis>
., 2009
</bibRefCitation>
;
<bibRefCitation id="EFE64B69FFE9C451FE53FEA4CE83811D" author="Archibald, S. B. &amp; Makarkin, V. N." box="[419,748,259,285]" pageId="31" pageNumber="72" pagination="269 - 287" refId="ref20421" refString="Archibald, S. B. &amp; Makarkin, V. N. (2020) A new genus and species of split-footed lacewings (Neuroptera) from the early Eocene of western Canada and revision of the subfamily affinities of Mesozoic Nymphidae. The Canadian Entomologist, 152, 269 - 287. https: // doi. org / 10.4039 / tce. 2020.10" type="journal article" year="2020">Archibald &amp; Makarkin, 2020</bibRefCitation>
) is today restricted to
<collectingCountry id="F3607608FFE9C451FC14FEA4C821811D" box="[996,1102,259,285]" name="Australia" pageId="31" pageNumber="72">Australia</collectingCountry>
, where it is less seasonal than at equivalent northern latitudes.
</paragraph>
</subSubSection>
<subSubSection id="C36D6513FFE9C451FF37FEECCF8B82F1" pageId="31" pageNumber="72" type="discussion">
<paragraph id="8BC83698FFE9C451FF37FEECCF8B82F1" blockId="31.[151,1437,151,1510]" pageId="31" pageNumber="72">
At the same time, snakeflies to the east of the Okanagan Highlands from the Green River Formation of Colorado inhabited much warmer lowland climates, with high mesothermal (mesothermal = MAT&gt;13°, &lt;20° C) to megathermal (MAT ≥20° C) MAT value estimates ranging from 19.6°C to 23.0°C (Wilf, 2000;
<bibRefCitation id="EFE64B69FFE9C451FB12FE34CCA181D1" author="Archibald, S. B. &amp; Greenwood, D. R. &amp; Smith, R. Y. &amp; Mathewes, R. W. &amp; Basinger, J. F." pageId="31" pageNumber="72" pagination="145 - 154" refId="ref20229" refString="Archibald, S. B., Greenwood, D. R., Smith, R. Y., Mathewes, R. W. &amp; Basinger, J. F. (2011) Early Eocene lagerstatten of the Okanagan Highlands (British Columbia and Washington State). Geoscience Canada, 38, 145 - 154." type="journal article" year="2011">
Archibald
<emphasis id="B903EA8AFFE9C451FAABFE34C9E181AD" box="[1371,1422,403,429]" italics="true" pageId="31" pageNumber="72">et al</emphasis>
., 2011
</bibRefCitation>
and references therein). Its forests also included frost-intolerant palms. Green River
<taxonomicName id="4C774D1BFFE9C451FB62FE10C95E81D1" authorityName="Navas" authorityYear="1916" box="[1170,1329,439,465]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="order">Raphidioptera</taxonomicName>
probably inhabited similar or warmer temperatures than
<taxonomicName id="4C774D1BFFE9C451FD5CFE7CCF5581F5" authorityName="Steinmann" authorityYear="1963" box="[684,826,475,501]" class="Insecta" family="Raphidiidae" genus="Harraphidia" kingdom="Animalia" order="Raphidioptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="genus">
<emphasis id="B903EA8AFFE9C451FD5CFE7CCF5581F5" box="[684,826,475,501]" italics="true" pageId="31" pageNumber="72">Harraphidia</emphasis>
</taxonomicName>
at Tarifa,
<collectingCountry id="F3607608FFE9C451FC5FFE7CCF9D81F5" box="[943,1010,475,501]" name="Spain" pageId="31" pageNumber="72">Spain</collectingCountry>
, where MAT is 17.2° C and had mild coldest months as above (
<bibRefCitation id="EFE64B69FFE9C451FE44FE58CF648219" author="Monserrat, V. J. &amp; Papenberg, D." box="[436,779,511,537]" pageId="31" pageNumber="72" pagination="203 - 222" refId="ref24008" refString="Monserrat, V. J. &amp; Papenberg, D. (2006) Revision del genero Harraphidia Steinmann, 1963 con la descripcion de dos nuevas especies de la peninsula Iberica y de Marruecos (Insecta, Raphidioptera). Graellsia, 62, 203 - 222. https: // doi. org / 10.3989 / graellsia. 2006. v 62. i 2.67" type="journal article" year="2006">Monserrat &amp; Papenberg, 2006</bibRefCitation>
;
<bibRefCitation id="EFE64B69FFE9C451FCE4FE58CFD28219" author="AEMET" box="[788,957,511,537]" pageId="31" pageNumber="72" refId="ref19824" refString="AEMET (2020) La Agencia Estatal de Meteorologia. Online. [http: // www. aemet. es / en / serviciosclimaticos / datosclimatologicos]" type="url" year="2020">AEMET, 2020</bibRefCitation>
). Although the Green River Formation was deposited in a lowland intermontane basin, the abundance of snakefly fossils indicates that they were part of the local community, as river transport from a distant mountain source would have been a rare event. There are numerous rocks with more than one specimen, in one case, four (
<figureCitation id="134C2A1DFFE9C451FCCDFDCCCFC88285" box="[829,935,619,645]" captionStart="FIGURE 14" captionStartId="20.[151,250,1455,1479]" captionTargetBox="[151,1436,181,1431]" captionTargetId="figure-26@20.[151,1436,181,1431]" captionTargetPageId="20" captionText="FIGURE 14. Archiinocellia protomaculata (Engel, 2011).A, UCM 61231, rock with four specimens of A. protomaculata (ac) and one undeterminable Raphidioptera (d); B, UCM 62523. Scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.4655680" httpUri="https://zenodo.org/record/4655680/files/figure.png" pageId="31" pageNumber="72">Fig. 14A</figureCitation>
). Further, almost all specimens in the UCM collection examined have a body with wings,
<emphasis id="B903EA8AFFE9C451FD50FD28CED382A8" box="[672,700,655,680]" italics="true" pageId="31" pageNumber="72">i.e</emphasis>
., are complete or at least relatively so. The wings may be folded in some, but in general, these do not appear to have suffered mechanical degradation characteristic of long-distance river transport. We believe that these also lived near their depositional lake.
</paragraph>
</subSubSection>
<subSubSection id="C36D6513FFE9C451FF37FD5CCECA83A6" pageId="31" pageNumber="72" type="description">
<paragraph id="8BC83698FFE9C451FF37FD5CCECA83A6" blockId="31.[151,1437,151,1510]" pageId="31" pageNumber="72">
A generalised climatic profile has been estimated for the Kishenehn formation by a nearest living relative analysis of mollusks (
<bibRefCitation id="EFE64B69FFE9C451FEBFFCB8CEED8339" author="Pierce, H. &amp; Constenius, K. N." box="[335,642,799,825]" pageId="31" pageNumber="72" pagination="1 - 112" refId="ref24710" refString="Pierce, H. &amp; Constenius, K. N. (2001) Late Eocene-Oligocene nonmarine mollusks of the northern Kishenehn Basin, Montana and British Columbia. Annals of the Carnegie Museum, 70, 1 - 112." type="journal article" year="2001">Pierce &amp; Constenius, 2001</bibRefCitation>
). These include taxa associated with both megathermal and microthermal climates. Pierce and Constenius rejected the notion that these coexisted, assuming the mixing of populations from different elevations. Such a mixed community is, however, consistent with a temperate climate with mild winters as in the Okanagan Highlands (above).
</paragraph>
</subSubSection>
<subSubSection id="C36D6513FFE9C451FF37FC08CF8F8459" pageId="31" pageNumber="72" type="reference_group">
<paragraph id="8BC83698FFE9C451FF37FC08CF8F8459" blockId="31.[151,1437,151,1510]" pageId="31" pageNumber="72">
Global MAT declined through the remainder of the Eocene (
<bibRefCitation id="EFE64B69FFE9C451FC89FC08C83583C9" author="Zachos, J. C. &amp; Dickens, G. R. &amp; Zeebe, R. E." box="[889,1114,943,969]" pageId="31" pageNumber="72" pagination="279 - 283" refId="ref26092" refString="Zachos, J. C., Dickens, G. R. &amp; Zeebe, R. E. (2008) An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics. Nature, 451, 279 - 283. https: // doi. org / 10.1038 / nature 06588" type="journal article" year="2008">
Zachos
<emphasis id="B903EA8AFFE9C451FC24FC08C86883C9" box="[980,1031,943,969]" italics="true" pageId="31" pageNumber="72">et al</emphasis>
., 2008
</bibRefCitation>
), but by the late Priabonian, close to the end of the greenhouse world, MAT estimates for upland Florissant, indicate an upper microthermal climate (
<bibRefCitation id="EFE64B69FFE9C451FF09FC50CDAE8411" author="Allen, S. E. &amp; Lowe, A. J. &amp; Peppe, D. J. &amp; Meyer, H. W." box="[249,449,1015,1041]" pageId="31" pageNumber="72" pagination="109678" refId="ref19873" refString="Allen, S. E., Lowe, A. J., Peppe, D. J. &amp; Meyer, H. W. (2020) Paleoclimate and paleoecology of the latest Eocene Florissant flora (Central Colorado, USA). Palaeogeography, Palaeoclimatology, Palaeoecology, 551, 109678. https: // doi. org / 10.1016 / j. palaeo. 2020.109678." type="journal article" year="2020">
Allen
<emphasis id="B903EA8AFFE9C451FEB0FC50CD1D8411" box="[320,370,1015,1041]" italics="true" pageId="31" pageNumber="72">et al</emphasis>
., 2020
</bibRefCitation>
) like those for the Okanagan Highlands, apparently also with no frost days by the presence of
<emphasis id="B903EA8AFFE9C451FF03FBBCCD7B8434" box="[243,276,1051,1076]" italics="true" pageId="31" pageNumber="72">e.g</emphasis>
., palms and Pachymerina (
<bibRefCitation id="EFE64B69FFE9C451FDB9FBBCCF788435" author="Manchester, S. R." box="[585,791,1051,1077]" pageId="31" pageNumber="72" pagination="137 - 162" refId="ref23896" refString="Manchester, S. R. (2001) Update on the Megafossil Flora of Florissant, Colorado. In: Evanoff, E., Gregory-Wodzicki, K. &amp; Johnson, K. (Eds.), Fossil flora and stratigraphy of the Florissant Formation, Colorado. Proceedings of the Denver Museum of Nature and Science, Series 4, 1, 137 - 162." type="journal article" year="2001">
<collectingRegion id="49B3F87AFFE9C451FDB9FBBCCEA08435" box="[585,719,1051,1077]" country="United Kingdom" name="Manchester" pageId="31" pageNumber="72">Manchester</collectingRegion>
, 2001
</bibRefCitation>
;
<bibRefCitation id="EFE64B69FFE9C451FCD3FBBCC84D8435" author="Archibald, S. B. &amp; Morse, G. E. &amp; Greenwood, D. R. &amp; Mathewes, R. W." box="[803,1058,1051,1077]" pageId="31" pageNumber="72" pagination="8095 - 8100" refId="ref20345" refString="Archibald, S. B., Morse, G. E., Greenwood, D. R. &amp; Mathewes, R. W. (2014) Fossil palm beetles refine upland winter temperatures in the Early Eocene Climatic Optimum. Proceedings of the National Academy of Sciences of the United States of America, 111, 8095 - 8100. https: // doi. org / 10.1073 / pnas. 1323269111" type="journal article" year="2014">
Archibald
<emphasis id="B903EA8AFFE9C451FC6CFBBCCFA18435" box="[924,974,1051,1077]" italics="true" pageId="31" pageNumber="72">et al</emphasis>
., 2014
</bibRefCitation>
). The youngest occurrence of the otherwise Cretaceous raphidiopteran family
<taxonomicName id="4C774D1BFFE9C451FD79FB98CF418459" ID-CoL="6ZR" authorityName="Martynova" authorityYear="1961" box="[649,814,1087,1113]" class="Insecta" family="Baissopteridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Baissopteridae</taxonomicName>
is at Florissant.
</paragraph>
</subSubSection>
<subSubSection id="C36D6513FFE9C451FF37FBC4C8B685E5" pageId="31" pageNumber="72" type="discussion">
<paragraph id="8BC83698FFE9C451FF37FBC4CDD9850D" blockId="31.[151,1437,151,1510]" pageId="31" pageNumber="72">
Therefore, all known
<taxonomicName id="4C774D1BFFE9C451FE47FBC4CE39847D" authorityName="Navas" authorityYear="1916" box="[439,598,1123,1149]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="order">Raphidioptera</taxonomicName>
before the Oligocene, including
<taxonomicName id="4C774D1BFFE9C451FC32FBC4C829847D" authorityName="Navas" authorityYear="1913" box="[962,1094,1123,1149]" class="Insecta" family="Inocelliidae" kingdom="Animalia" order="Raphidioptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Inocelliidae</taxonomicName>
and
<taxonomicName id="4C774D1BFFE9C451FB8AFBC4C96B847D" authorityName="Latreille" authorityYear="1810" box="[1146,1284,1123,1149]" class="Insecta" family="Raphidiidae" kingdom="Animalia" order="Raphidioptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Raphidiidae</taxonomicName>
, inhabited regions of mild winters without cold intervals whether in hot or temperate climates. An adaptation to winter cold must have evolved independently in both extant families sometime after the Eocene/Oligocene boundary in the modern icehouse world climatic regime, with some members retaining or subsequently reverting to the plesiomorphic adaptation to warm winters.
</paragraph>
<paragraph id="8BC83698FFE9C451FF37FAB0C8B685E5" blockId="31.[151,1437,151,1510]" pageId="31" pageNumber="72">
After the onset of cold extra-tropical icehouse world winters, biota of latitudes outside of the tropics either went extinct (perhaps accounting for the extinctions of,
<emphasis id="B903EA8AFFE9C451FCF3FA9CCF4B8554" box="[771,804,1339,1364]" italics="true" pageId="31" pageNumber="72">e.g</emphasis>
., the
<taxonomicName id="4C774D1BFFE9C451FC91FA9CCFBC8555" baseAuthorityName="Martins-Neto &amp; Vulcano" baseAuthorityYear="1988" box="[865,979,1339,1365]" class="Insecta" family="Chrysopidae" kingdom="Animalia" order="Neuroptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="subFamily" subFamily="Limaiinae">Limaiinae</taxonomicName>
and
<taxonomicName id="4C774D1BFFE9C451FBF7FA9CC8DE8556" authorityName="Martynova" authorityYear="1961" box="[1031,1201,1339,1366]" class="Insecta" family="Baissopteridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Baissopteridae</taxonomicName>
), moved to or restrict their ranges to lower latitudes and Southern Hemisphere regions of mild winters (
<emphasis id="B903EA8AFFE9C451FBDEFAF8C8208578" box="[1070,1103,1375,1400]" italics="true" pageId="31" pageNumber="72">e.g</emphasis>
.,
<taxonomicName id="4C774D1BFFE9C451FB94FAF8C8858579" authorityName="Rambur" authorityYear="1842" box="[1124,1258,1375,1401]" class="Insecta" family="Nymphidae" kingdom="Animalia" order="Neuroptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Nymphidae</taxonomicName>
, palm bruchids,
<taxonomicName id="4C774D1BFFE9C451FF67FA24CD4F859D" box="[151,288,1411,1437]" class="Insecta" family="Dinidoridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hemiptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Dinidoridae</taxonomicName>
,
<taxonomicName id="4C774D1BFFE9C451FEDDFA24CD87859D" authorityName="Desneux" authorityYear="1904" box="[301,488,1411,1437]" class="Insecta" family="Mastotermitidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Blattodea" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Mastotermitidae</taxonomicName>
, Mymeciinae, and
<taxonomicName id="4C774D1BFFE9C451FD32FA24CF0E859E" baseAuthorityName="Saussure" baseAuthorityYear="1864" box="[706,865,1411,1438]" class="Insecta" family="Blaberidae" kingdom="Animalia" order="Blattodea" pageId="31" pageNumber="72" phylum="Arthropoda" rank="subFamily" subFamily="Diplopterinae">Diplopterinae</taxonomicName>
), or remained and adapted to more severe winters (
<emphasis id="B903EA8AFFE9C451FF51FA00CCAD85C0" box="[161,194,1447,1472]" italics="true" pageId="31" pageNumber="72">e.g</emphasis>
., aphids,
<taxonomicName id="4C774D1BFFE9C451FEC2FA00CDFB85C0" authorityName="Linnaeus" authorityYear="1758" box="[306,404,1447,1472]" class="Insecta" family="Panorpidae" genus="Panorpa" kingdom="Animalia" order="Mecoptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="genus">
<emphasis id="B903EA8AFFE9C451FEC2FA00CDFB85C0" box="[306,404,1447,1472]" italics="true" pageId="31" pageNumber="72">Panorpa</emphasis>
</taxonomicName>
,
<taxonomicName id="4C774D1BFFE9C451FE54FA00CE2085C1" box="[420,591,1447,1473]" class="Insecta" family="Tenthredinidae" kingdom="Animalia" order="Hymenoptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Tenthredinidae</taxonomicName>
,
<taxonomicName id="4C774D1BFFE9C451FDACFA00CEAD85C1" authorityName="Billberg" authorityYear="1820" box="[604,706,1447,1473]" class="Insecta" family="Siricidae" kingdom="Animalia" order="Hymenoptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Siricidae</taxonomicName>
,
<taxonomicName id="4C774D1BFFE9C451FD3FFA00CF0985C1" authorityName="Cameron" authorityYear="1890" box="[719,870,1447,1473]" class="Insecta" family="Pamphiliidae" kingdom="Animalia" order="Hymenoptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Pamphiliidae</taxonomicName>
,
<taxonomicName id="4C774D1BFFE9C451FC83FA00CF8F85C1" box="[883,992,1447,1473]" class="Insecta" family="Cephidae" kingdom="Animalia" order="Hymenoptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Cephidae</taxonomicName>
,
<taxonomicName id="4C774D1BFFE9C451FC1DFA00C80185C1" box="[1005,1134,1447,1473]" class="Insecta" family="Panorpidae" kingdom="Animalia" order="Mecoptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Panorpidae</taxonomicName>
, shift to dominance of the
<taxonomicName id="4C774D1BFFE9C451FF67FA6CCD4A85E5" box="[151,293,1483,1509]" class="Insecta" family="Chrysopidae" kingdom="Animalia" order="Neuroptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="subFamily" subFamily="Chrysopinae">Chrysopinae</taxonomicName>
within
<taxonomicName id="4C774D1BFFE9C451FE8AFA6CCE6285E6" box="[378,525,1483,1510]" class="Insecta" family="Chrysopidae" kingdom="Animalia" order="Neuroptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="family">Chrysopidae</taxonomicName>
).
<taxonomicName id="4C774D1BFFE9C451FDEFFA6CCED185E5" authorityName="Navas" authorityYear="1916" box="[543,702,1483,1509]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Raphidioptera" pageId="31" pageNumber="72" phylum="Arthropoda" rank="order">Raphidioptera</taxonomicName>
appears to have mostly followed the latter path.
</paragraph>
</subSubSection>
</treatment>
</document>