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<mods:title id="9B140441AC61A8AB322915ECD0BE0943">Biology and ecology of a deep cave nesting spider wasp, Ageniella evansi Townes, (Hymenoptera: Pompilidae), in Arizona</mods:title>
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<treatment id="1C2A87AAFFA9FF8CFE5E9BDDFDF9FE72" ID-DOI="http://doi.org/10.5281/zenodo.13785664" ID-Zenodo-Dep="13785664" LSID="urn:lsid:plazi:treatment:1C2A87AAFFA9FF8CFE5E9BDDFDF9FE72" httpUri="http://treatment.plazi.org/id/1C2A87AAFFA9FF8CFE5E9BDDFDF9FE72" lastPageId="40" lastPageNumber="1003" pageId="36" pageNumber="999">
<subSubSection id="DC996537FFA9FF80FE5E9BDDFC7BFA37" box="[464,998,1476,1503]" pageId="36" pageNumber="999" type="nomenclature">
<paragraph id="943C36BCFFA9FF80FE5E9BDDFC7BFA37" blockId="36.[317,998,1476,1503]" box="[464,998,1476,1503]" pageId="36" pageNumber="999">
<taxonomicName id="53834D3FFFA9FF80FE5E9BDDFD1AFA37" ID-CoL="5NWP" authority="O.Pickard-Cambridge, 1898" box="[464,647,1476,1503]" class="Arachnida" family="Agelenidae" genus="Melpomene" kingdom="Animalia" order="Araneae" pageId="36" pageNumber="999" phylum="Arthropoda" rank="genus">
<emphasis id="A6F7EAAEFFA9FF80FE5E9BDDFDC1FA37" box="[464,604,1476,1503]" italics="true" pageId="36" pageNumber="999">Melpomene</emphasis>
sp.
</taxonomicName>
O. Pickard-Cambridge, 1898
</paragraph>
</subSubSection>
<subSubSection id="DC996537FFA9FF8CFF2E9BE3FDF9FE72" lastPageId="40" lastPageNumber="1003" pageId="36" pageNumber="999" type="discussion">
<paragraph id="943C36BCFFA9FF81FF2E9BE3FE47FF22" blockId="36.[160,1156,1530,1728]" lastBlockId="37.[160,1157,142,861]" lastPageId="37" lastPageNumber="1000" pageId="36" pageNumber="999">
<taxonomicName id="53834D3FFFA9FF80FF2E9BE3FE82F9FB" ID-CoL="64Q" box="[160,287,1530,1555]" class="Arachnida" family="Agelenidae" kingdom="Animalia" order="Araneae" pageId="36" pageNumber="999" phylum="Arthropoda" rank="family">Agelenidae</taxonomicName>
is a large family with currently in excess of 1300 valid species (WSC 2022). These mostly nocturnal spiders build non-sticky, funnel webs in a variety of habitats, including caves (
<bibRefCitation id="F0124B4DFFA9FF80FED59826FDF1F9B0" author="Bennett RG &amp; Ubick D &amp; Agelenidae. In &amp; Ubick D &amp; Paquin P &amp; Cushing PE &amp; Roth V &amp; Spiders of North America &amp; American Arachnological Society &amp; University of California &amp; Davis" box="[347,620,1599,1624]" pageId="36" pageNumber="999" pagination="56 - 59" refId="ref41791" refString="Bennett RG, Ubick D. Agelenidae. In: Ubick D, Paquin P, Cushing PE, Roth V, editors. Spiders of North America: an identification manual. American Arachnological Society, University of California, Davis; 2005. p. 56 - 59. https: // www. americanarachnology. org / home /." type="book chapter" year="2005">Bennett and Ubick 2005</bibRefCitation>
).
<taxonomicName id="53834D3FFFA9FF80FD0E9826FD62F9B0" ID-CoL="5NWP" authorityName="O.Pickard-Cambridge" authorityYear="1898" box="[640,767,1599,1624]" class="Arachnida" family="Agelenidae" genus="Melpomene" kingdom="Animalia" order="Araneae" pageId="36" pageNumber="999" phylum="Arthropoda" rank="genus">
<emphasis id="A6F7EAAEFFA9FF80FD0E9826FD62F9B0" box="[640,767,1599,1624]" italics="true" pageId="36" pageNumber="999">Melpomene</emphasis>
</taxonomicName>
is a New World group, distributed from southern Arizona and northern Sonora to western Texas and south to
<collectingCountry id="EC94762CFFA9FF80FC69987BFBDCF993" box="[999,1089,1634,1659]" name="Panama" pageId="36" pageNumber="999">Panama</collectingCountry>
(
<bibRefCitation id="F0124B4DFFA9FF80FBC3987BFEC8F976" author="Roth VD &amp; Brame PL" pageId="36" pageNumber="999" pagination="2 - 52" refId="ref47292" refString="Roth VD, Brame PL. 1972. Nearctic genera of the spider family Agelenidae (Arachnida, Araneida). Am Mus Novit. 2505: 2 - 52. http: // digitallibrary. amnh. org / handle / 2246 / 2712." type="journal article" year="1972">Roth and Brame 1972</bibRefCitation>
).
<taxonomicName id="53834D3FFFA9FF80FEE6989CFD91F976" box="[360,524,1669,1694]" class="Arachnida" family="Agelenidae" genus="Melpomene" kingdom="Animalia" order="Araneae" pageId="36" pageNumber="999" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="A6F7EAAEFFA9FF80FEE6989CFE7AF976" box="[360,487,1669,1694]" italics="true" pageId="36" pageNumber="999">Melpomene</emphasis>
sp.
</taxonomicName>
(
<figureCitation id="0CB82A39FFA9FF80FD97989CFD0FF976" box="[537,658,1669,1694]" captionStart="Figure 20" captionStartId="33.[160,225,535,558]" captionTargetBox="[188,1139,174,480]" captionTargetId="figure-331@33.[162,1152,142,503]" captionTargetPageId="33" captionText="Figure 20. The four families of spider hosts taken by Ageniella evansi Townes, 1957 in Colossal Cave Mountain Park, Arizona. A, Selenopidae Selenops sp. Latreille, 1819. B, Gnaphosidae (Herpyllus schwarzi Banks, 1901). C, Lycosidae. D, Agelenidae (Melpomene sp. O. Pickard-Cambridge, 1898)." figureDoi="http://doi.org/10.5281/zenodo.13758318" httpUri="https://zenodo.org/record/13758318/files/figure.png" pageId="36" pageNumber="999">Figure 20D</figureCitation>
) is apparently rarely taken by
<taxonomicName id="53834D3FFFA9FF80FC51989CFBA0F976" ID-CoL="65RTS" authorityName="Townes" authorityYear="1957" box="[991,1085,1669,1694]" class="Insecta" family="Pompilidae" genus="Ageniella" kingdom="Animalia" order="Hymenoptera" pageId="36" pageNumber="999" phylum="Arthropoda" rank="species" species="evansi">
<emphasis id="A6F7EAAEFFA9FF80FC51989CFBA0F976" box="[991,1085,1669,1694]" italics="true" pageId="36" pageNumber="999">A. evansi</emphasis>
</taxonomicName>
in the Park, with only two records, representing one half of one percent of total observed wasptransported host spiders. It seems reasonable that use of this spider represents opportunistic capture by the wasps.
</paragraph>
<paragraph id="943C36BCFFA8FF81FF4E9ECDFEAEFDCD" blockId="37.[160,1157,142,861]" pageId="37" pageNumber="1000">
No field surveys were conducted for any of the spider host species, so no information is available about their phenology or population dynamics. The primary host (by percentage,
<taxonomicName id="53834D3FFFA8FF81FF519F00FEA3FEDA" ID-CoL="7F95" box="[223,318,281,306]" class="Arachnida" family="Selenopidae" genus="Selenops" kingdom="Animalia" order="Araneae" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="genus">
<emphasis id="A6F7EAAEFFA8FF81FF519F00FEA3FEDA" box="[223,318,281,306]" italics="true" pageId="37" pageNumber="1000">Selenops</emphasis>
</taxonomicName>
) is difficult to census due to their cryptic and inaccessible habits in cracks, fissures and caves. The information used in this analysis is based solely on the observed spiders returned to the cave by the wasps. The distribution by month of the total observed spiders taken by the Arkenstone Cave population of
<taxonomicName id="53834D3FFFA8FF81FC049F98FC6DFE72" ID-CoL="65RTS" authorityName="Townes" authorityYear="1957" box="[906,1008,385,410]" class="Insecta" family="Pompilidae" genus="Ageniella" kingdom="Animalia" order="Hymenoptera" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="species" species="evansi">
<emphasis id="A6F7EAAEFFA8FF81FC049F98FC6DFE72" box="[906,1008,385,410]" italics="true" pageId="37" pageNumber="1000">A. evansi</emphasis>
</taxonomicName>
is shown in
<figureCitation id="0CB82A39FFA8FF81FF2E9FBDFE94FE55" box="[160,265,420,445]" captionStart="Figure 23" captionStartId="37.[160,225,1616,1639]" captionTargetBox="[165,1150,944,1582]" captionTargetId="figure-295@37.[163,1153,942,1585]" captionTargetPageId="37" captionText="Figure 23. Monthly total observed spider hosts (N = 404) taken by the Arkenstone Cave population of Ageniella evansi Townes, 1957 between 1992 and 2020. The total includes all identified and 18 unidentified spiders. The average peak of the wasp hunting season over 32 years (1993 to 2024) is 5 March." figureDoi="http://doi.org/10.5281/zenodo.13758324" httpUri="https://zenodo.org/record/13758324/files/figure.png" pageId="37" pageNumber="1000">Figure 23</figureCitation>
. The total (N = 404) includes all identified and 18 unidentified spiders observed between 1992 and 2020. The earliest spider capture record for
<taxonomicName id="53834D3FFFA8FF81FCE19FDFFC4DFE37" ID-CoL="65RTS" authorityName="Townes" authorityYear="1957" box="[879,976,454,479]" class="Insecta" family="Pompilidae" genus="Ageniella" kingdom="Animalia" order="Hymenoptera" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="species" species="evansi">
<emphasis id="A6F7EAAEFFA8FF81FCE19FDFFC4DFE37" box="[879,976,454,479]" italics="true" pageId="37" pageNumber="1000">A. evansi</emphasis>
</taxonomicName>
in the Park is 7 November (1998;
<taxonomicName id="53834D3FFFA8FF81FEF19FF0FD84FDEA" ID-CoL="623K5" box="[383,537,489,514]" class="Arachnida" family="Gnaphosidae" kingdom="Animalia" order="Araneae" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="family">Gnaphosidae</taxonomicName>
) and the latest capture record is 11 May (2003;
<taxonomicName id="53834D3FFFA8FF81FF2E9C15FEB4FDCD" box="[160,297,524,549]" class="Arachnida" family="Selenopidae" genus="Selenops" kingdom="Animalia" order="Araneae" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="A6F7EAAEFFA8FF81FF2E9C15FF62FDCD" box="[160,255,524,549]" italics="true" pageId="37" pageNumber="1000">Selenops</emphasis>
sp.
</taxonomicName>
).
</paragraph>
<paragraph id="943C36BCFFA8FF82FF4E9C37FBDEFAC8" blockId="37.[160,1157,142,861]" lastBlockId="38.[160,1156,975,1728]" lastPageId="38" lastPageNumber="1001" pageId="37" pageNumber="1000">
An analysis of host
<typeStatus id="4B38881EFFA8FF81FE1D9C37FE58FDAF" box="[403,453,558,583]" pageId="37" pageNumber="1000">type</typeStatus>
by month (
<figureCitation id="0CB82A39FFA8FF81FDCA9C37FD31FDAF" box="[580,684,558,583]" captionStart="Figure 24" captionStartId="38.[160,225,852,875]" captionTargetBox="[165,1150,144,817]" captionTargetId="figure-326@38.[163,1153,142,820]" captionTargetPageId="38" captionText="Figure 24. Percent host type by month taken by the Arkenstone Cave population of Ageniella evansi Townes, 1957 between 1992 and 2020." figureDoi="http://doi.org/10.5281/zenodo.13758326" httpUri="https://zenodo.org/record/13758326/files/figure.png" pageId="37" pageNumber="1000">Figure 24</figureCitation>
) reveals exclusive early season (November and December) use of gnaphosid spiders. No
<taxonomicName id="53834D3FFFA8FF81FD129C48FD66FD82" box="[668,763,593,618]" class="Arachnida" family="Selenopidae" genus="Selenops" kingdom="Animalia" order="Araneae" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="genus">
<emphasis id="A6F7EAAEFFA8FF81FD129C48FD66FD82" box="[668,763,593,618]" italics="true" pageId="37" pageNumber="1000">Selenops</emphasis>
</taxonomicName>
were observed taken in November or December, although a total of only 12 spiders were recorded for those 2 months. Use of gnaphosid spiders remains dominant through February. Use of
<taxonomicName id="53834D3FFFA8FF81FCF19C8FFC43FD47" box="[895,990,662,687]" class="Arachnida" family="Selenopidae" genus="Selenops" kingdom="Animalia" order="Araneae" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="genus">
<emphasis id="A6F7EAAEFFA8FF81FCF19C8FFC43FD47" box="[895,990,662,687]" italics="true" pageId="37" pageNumber="1000">Selenops</emphasis>
</taxonomicName>
overlaps gnaphosid use beginning in January, becomes dominant in March and extends through early May, while gnaphosid use drops off significantly by the end of February. Because
<taxonomicName id="53834D3FFFA8FF81FBAA9CC5FB1EFD1D" box="[1060,1155,732,757]" class="Arachnida" family="Selenopidae" genus="Selenops" kingdom="Animalia" order="Araneae" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="genus">
<emphasis id="A6F7EAAEFFA8FF81FBAA9CC5FB1EFD1D" box="[1060,1155,732,757]" italics="true" pageId="37" pageNumber="1000">Selenops</emphasis>
</taxonomicName>
apparently live more than 1 year (
<bibRefCitation id="F0124B4DFFA8FF81FDA49CE7FD6CFCFF" author="Crews SC &amp; Wienskoski E &amp; Gillespie RG" box="[554,753,766,791]" pageId="37" pageNumber="1000" pagination="2747 - 2761" refId="ref42462" refString="Crews SC, Wienskoski E, Gillespie RG. 2008. Life history of the spider Selenops occultus Mello-Leitao (Araneae, Selenopidae) from Brazil with notes on the natural history of the genus. J Natural His. 42 (43 / 44): 2747 - 2761. doi: 10.1080 / 00222930802354183." type="journal article" year="2008">
Crews
<emphasis id="A6F7EAAEFFA8FF81FDF79CE7FD36FCFF" box="[633,683,766,791]" italics="true" pageId="37" pageNumber="1000">et al</emphasis>
. 2008
</bibRefCitation>
), various age cohorts are probably present throughout the year as potential hosts for
<taxonomicName id="53834D3FFFA8FF81FD6B9D38FCDAFCD2" authorityName="Townes" authorityYear="1957" box="[741,839,801,826]" class="Insecta" family="Pompilidae" genus="Ageniella" kingdom="Animalia" order="Hymenoptera" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="species" species="evansi">
<emphasis id="A6F7EAAEFFA8FF81FD6B9D38FCDAFCD2" box="[741,839,801,826]" italics="true" pageId="37" pageNumber="1000">A. evansi</emphasis>
</taxonomicName>
. This study revealed that
<taxonomicName id="53834D3FFFA8FF81FBE09D38FF79FCB5" authorityName="Townes" authorityYear="1957" class="Insecta" family="Pompilidae" genus="Ageniella" kingdom="Animalia" order="Hymenoptera" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="species" species="evansi">
<emphasis id="A6F7EAAEFFA8FF81FBE09D38FF79FCB5" italics="true" pageId="37" pageNumber="1000">A. evansi</emphasis>
</taxonomicName>
uses a variety of age cohorts of
<taxonomicName id="53834D3FFFA8FF81FDD79D5DFD25FCB5" box="[601,696,836,861]" class="Arachnida" family="Selenopidae" genus="Selenops" kingdom="Animalia" order="Araneae" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="genus">
<emphasis id="A6F7EAAEFFA8FF81FDD79D5DFD25FCB5" box="[601,696,836,861]" italics="true" pageId="37" pageNumber="1000">Selenops</emphasis>
</taxonomicName>
, and the lack of their use of this species early in the wasp active season suggests a preference for gnaphosid spiders. The drop-off in use of gnaphosid spiders may be the result of a depletion of large individuals of this element of the wasp host base within the wasp foraging range by the end of February. Switching of host
<typeStatus id="4B38881EFFABFF82FEE29A2EFE03FBB8" box="[364,414,1079,1104]" pageId="38" pageNumber="1001">type</typeStatus>
in the
<taxonomicName id="53834D3FFFABFF82FE609A2EFDF1FBB8" baseAuthorityName="Evans" baseAuthorityYear="1966" box="[494,620,1079,1104]" class="Insecta" family="Pompilidae" kingdom="Animalia" order="Hymenoptera" pageId="38" pageNumber="1001" phylum="Arthropoda" rank="family">Pompilidae</taxonomicName>
resulting from depletion of certain elements of the host base has been documented elsewhere (
<bibRefCitation id="F0124B4DFFABFF82FD629A43FBE0FB9B" author="Kurczewski FE &amp; Kurczewski EJ" box="[748,1149,1114,1139]" pageId="38" pageNumber="1001" pagination="1 - 33" refId="ref45001" refString="Kurczewski FE, Kurczewski EJ. 1968 a. Host records for some North American Pompilidae (Hymenoptera) with a discussion of factors in prey selection. J Kansas Entomol Soc. 41 (1): 1 - 33. https: // www. jstor. org / stable / 25083675? seq = 1." type="journal article" year="1968">Kurczewski and Kurczewski 1968a</bibRefCitation>
;
<bibRefCitation id="F0124B4DFFABFF82FF2E9A64FEA9FB7E" author="Martins RP" box="[160,308,1149,1174]" pageId="38" pageNumber="1001" pagination="231 - 236" refId="ref45591" refString="Martins RP. 1991. Nesting behavior and prey of Poecilopompilus algidus fervidus and Tachypompilus xanthopterus (Hymenoptera: Pompilidae). J Kansas Entomol Soc. 91 (2): 231 - 236. https: // www. jstor. org / stable / 25085276? refreqid = excelsior % 3 A 044544337635 e 3201609 db 1 c 7 b 5 a 30 fc." type="journal article" year="1991">Martins 1991</bibRefCitation>
;
<bibRefCitation id="F0124B4DFFABFF82FECD9A64FDABFB7E" author="Endo T &amp; Endo A" box="[323,566,1149,1174]" pageId="38" pageNumber="1001" pagination="225 - 235" refId="ref43036" refString="Endo T, Endo A. 1994. Prey selection by a spider wasp, Batozonellus lacerticida (Hymenoptera: Pompilidae): effects of seasonal variation in prey species, size and density. Ecol Res. 9 (2): 225 - 235. doi: 10.1007 / BF 02347498." type="journal article" year="1994">Endo and Endo 1994</bibRefCitation>
). The wasps then apparently focus on
<taxonomicName id="53834D3FFFABFF82FC759A64FBC7FB7E" box="[1019,1114,1149,1174]" class="Arachnida" family="Selenopidae" genus="Selenops" kingdom="Animalia" order="Araneae" pageId="38" pageNumber="1001" phylum="Arthropoda" rank="genus">
<emphasis id="A6F7EAAEFFABFF82FC759A64FBC7FB7E" box="[1019,1114,1149,1174]" italics="true" pageId="38" pageNumber="1001">Selenops</emphasis>
</taxonomicName>
for the remainder of the active season. The wasps take lycosids beginning in February, peaking in April and continuing into early May. Approximately one-quarter of spiders taken in April are lycosids. The use of lycosids beginning in February may reflect the time at which these spiders reach an acceptable minimum size threshold for the wasps.
</paragraph>
<caption id="C0FC6634FFA8FF81FF2E9849FF6AF957" ID-DOI="http://doi.org/10.5281/zenodo.13758324" ID-Zenodo-Dep="13758324" httpUri="https://zenodo.org/record/13758324/files/figure.png" pageId="37" pageNumber="1000" startId="37.[160,225,1616,1639]" targetBox="[165,1150,944,1582]" targetPageId="37" targetType="figure">
<paragraph id="943C36BCFFA8FF81FF2E9849FF6AF957" blockId="37.[160,1156,1616,1727]" pageId="37" pageNumber="1000">
<emphasis id="A6F7EAAEFFA8FF81FF2E9849FE9BF98F" bold="true" box="[160,262,1616,1639]" pageId="37" pageNumber="1000">Figure 23.</emphasis>
Monthly total observed spider hosts (N = 404) taken by the Arkenstone Cave population of
<taxonomicName id="53834D3FFFA8FF81FF2E9874FE48F96C" authority="Townes, 1957" authorityName="Townes" authorityYear="1957" box="[160,469,1645,1668]" class="Insecta" family="Pompilidae" genus="Ageniella" kingdom="Animalia" order="Hymenoptera" pageId="37" pageNumber="1000" phylum="Arthropoda" rank="species" species="evansi">
<emphasis id="A6F7EAAEFFA8FF81FF2E9874FEA2F96C" box="[160,319,1645,1668]" italics="true" pageId="37" pageNumber="1000">Ageniella evansi</emphasis>
<bibRefCitation id="F0124B4DFFA8FF81FEC49874FE48F96C" author="Townes H" box="[330,469,1645,1668]" pageId="37" pageNumber="1000" refId="ref48313" refString="Townes H. 1957. Nearctic wasps of the subfamilies Pepsinae and Ceropalinae. United States National Museum Bull. 209. https: // repository. si. edu / handle / 10088 / 10223." type="journal volume" year="1957">Townes, 1957</bibRefCitation>
</taxonomicName>
between 1992 and 2020. The total includes all identified and 18 unidentified spiders. The average peak of the wasp hunting season over 32 years (1993 to 2024) is 5 March.
</paragraph>
</caption>
<caption id="C0FC6634FFABFF82FF2E9D4DFDBBFC60" ID-DOI="http://doi.org/10.5281/zenodo.13758326" ID-Zenodo-Dep="13758326" httpUri="https://zenodo.org/record/13758326/files/figure.png" pageId="38" pageNumber="1001" startId="38.[160,225,852,875]" targetBox="[165,1150,144,817]" targetPageId="38" targetType="figure">
<paragraph id="943C36BCFFABFF82FF2E9D4DFDBBFC60" blockId="38.[160,1155,852,904]" pageId="38" pageNumber="1001">
<emphasis id="A6F7EAAEFFABFF82FF2E9D4DFE9AFC83" bold="true" box="[160,263,852,875]" pageId="38" pageNumber="1001">Figure 24.</emphasis>
Percent host type by month taken by the Arkenstone Cave population of
<taxonomicName id="53834D3FFFABFF82FC679D4DFEBAFC60" authority="Townes, 1957" authorityName="Townes" authorityYear="1957" class="Insecta" family="Pompilidae" genus="Ageniella" kingdom="Animalia" order="Hymenoptera" pageId="38" pageNumber="1001" phylum="Arthropoda" rank="species" species="evansi">
<emphasis id="A6F7EAAEFFABFF82FC679D4DFB1EFC83" box="[1001,1155,852,875]" italics="true" pageId="38" pageNumber="1001">Ageniella evansi</emphasis>
<bibRefCitation id="F0124B4DFFABFF82FF2E9D68FEBAFC60" author="Townes H" box="[160,295,881,904]" pageId="38" pageNumber="1001" refId="ref48313" refString="Townes H. 1957. Nearctic wasps of the subfamilies Pepsinae and Ceropalinae. United States National Museum Bull. 209. https: // repository. si. edu / handle / 10088 / 10223." type="journal volume" year="1957">Townes, 1957</bibRefCitation>
</taxonomicName>
between 1992 and 2020.
</paragraph>
</caption>
<paragraph id="943C36BCFFABFF82FF4E9B33FC08F9B0" blockId="38.[160,1156,975,1728]" pageId="38" pageNumber="1001">
An analysis of host
<typeStatus id="4B38881EFFABFF82FE189B33FE55FAAB" box="[406,456,1322,1347]" pageId="38" pageNumber="1001">type</typeStatus>
by year (
<figureCitation id="0CB82A39FFABFF82FDA09B33FD0BFAAB" box="[558,662,1322,1347]" captionStart="Figure 25" captionStartId="39.[160,225,892,915]" captionTargetBox="[165,1150,144,857]" captionTargetId="figure-301@39.[163,1153,142,860]" captionTargetPageId="39" captionText="Figure 25. Percent of observed host type by year taken by the Arkenstone Cave population of Ageniella evansi Townes, 1957 between 1993 and 2020." figureDoi="http://doi.org/10.5281/zenodo.13758330" httpUri="https://zenodo.org/record/13758330/files/figure.png" pageId="38" pageNumber="1001">Figure 25</figureCitation>
) shows a greater diversity of spiders used in the early years of the study (1996–1997), where both lycosid and agelenid spiders show up in the data. This may be due to the greater number of spiders returned to the cave in years when the female hunting population was large. Lycosid and agelenid spiders are probably regular components of
<taxonomicName id="53834D3FFFABFF82FD9A9BACFDE9FA26" authorityName="Townes" authorityYear="1957" box="[532,628,1461,1486]" class="Insecta" family="Pompilidae" genus="Ageniella" kingdom="Animalia" order="Hymenoptera" pageId="38" pageNumber="1001" phylum="Arthropoda" rank="species" species="evansi">
<emphasis id="A6F7EAAEFFABFF82FD9A9BACFDE9FA26" box="[532,628,1461,1486]" italics="true" pageId="38" pageNumber="1001">A. evansi</emphasis>
</taxonomicName>
hosts taken in a typical year, but are less often observed as they are apparently minor elements of total spiders taken. The broader implication is that when larger data sets are obtained for other pompilid species a greater diversity of host use may be found. Thus the
<taxonomicName id="53834D3FFFABFF82FD1E9804FC8CF9DE" baseAuthorityName="Evans" baseAuthorityYear="1966" box="[656,785,1565,1590]" class="Insecta" family="Pompilidae" kingdom="Animalia" order="Hymenoptera" pageId="38" pageNumber="1001" phylum="Arthropoda" rank="family">Pompilidae</taxonomicName>
, in general, will likely be found to be more flexible in their host selection that is currently recognised.
</paragraph>
<paragraph id="943C36BCFFABFF83FF4E987BFE61FB1F" blockId="38.[160,1156,975,1728]" lastBlockId="39.[160,1156,1003,1721]" lastPageId="39" lastPageNumber="1002" pageId="38" pageNumber="1001">
Spider size is apparently the crucial element in host selection, even among pompilid species that take spiders of more than one kind (
<bibRefCitation id="F0124B4DFFABFF82FD5A989CFC5BF976" author="Endo T &amp; Endo A" box="[724,966,1669,1694]" pageId="38" pageNumber="1001" pagination="225 - 235" refId="ref43036" refString="Endo T, Endo A. 1994. Prey selection by a spider wasp, Batozonellus lacerticida (Hymenoptera: Pompilidae): effects of seasonal variation in prey species, size and density. Ecol Res. 9 (2): 225 - 235. doi: 10.1007 / BF 02347498." type="journal article" year="1994">Endo and Endo 1994</bibRefCitation>
;
<bibRefCitation id="F0124B4DFFABFF82FC5A989CFEDAF928" author="Kurczewski FE &amp; Edwards GB" pageId="38" pageNumber="1001" pagination="1 - 71" refId="ref44842" refString="Kurczewski FE, Edwards GB. 2012. Hosts, nesting behavior, and ecology of some North American spider wasps (Hymenoptera: Pompilidae). Southeast Nat. 11 (m 4): 1 - 71. doi: 10.1656 / 058.011. m 401." type="journal article" year="2012">Kurczewski and Edwards 2012</bibRefCitation>
).
<bibRefCitation id="F0124B4DFFABFF82FEEC98BEFD62F928" author="Kurczewski FE &amp; Kurczewski EJ" box="[354,767,1703,1728]" pageId="38" pageNumber="1001" pagination="1 - 33" refId="ref45001" refString="Kurczewski FE, Kurczewski EJ. 1968 a. Host records for some North American Pompilidae (Hymenoptera) with a discussion of factors in prey selection. J Kansas Entomol Soc. 41 (1): 1 - 33. https: // www. jstor. org / stable / 25083675? seq = 1." type="journal article" year="1968">Kurczewski and Kurczewski (1968a</bibRefCitation>
,
<bibRefCitation id="F0124B4DFFABFF82FC9C98BEFCC4F928" author="Kurczewski FE &amp; Kurczewski EJ" box="[786,857,1703,1728]" pageId="38" pageNumber="1001" pagination="367 - 382" refId="ref45060" refString="Kurczewski FE, Kurczewski EJ. 1968 b. Host records for some North American Pompilidae (Hymenoptera), first supplement. J Kansas Entomol Soc. 41 (3): 367 - 382. https: // www. jstor. org / stable / 25083722? seq = 1." type="journal article" year="1968">1968b</bibRefCitation>
) provide abundant data supporting the use of large and/or adult spiders by larger pompilid species and smaller and/or immature spiders by smaller pompilid species.
<taxonomicName id="53834D3FFFAAFF83FC949A17FC51FBCF" authorityName="Townes" authorityYear="1957" box="[794,972,1038,1063]" class="Insecta" family="Pompilidae" genus="Ageniella" kingdom="Animalia" order="Hymenoptera" pageId="39" pageNumber="1002" phylum="Arthropoda" rank="species" species="evansi">
<emphasis id="A6F7EAAEFFAAFF83FC949A17FC51FBCF" box="[794,972,1038,1063]" italics="true" pageId="39" pageNumber="1002">Ageniella evansi</emphasis>
</taxonomicName>
generally takes spiders that are subequal to or slightly shorter than their body length. Spiders much longer or larger than the wasps quickly become cumbersome and inefficient to haul long distances to the nesting site. This is particularly important for these cave nesting populations since once they enter the cave they still have a considerable distance to go before reaching their nest site. Whether or how this may affect the extent of their epigean hunting territory is not known.
</paragraph>
<caption id="C0FC6634FFAAFF83FF2E9D65FD55FC58" ID-DOI="http://doi.org/10.5281/zenodo.13758330" ID-Zenodo-Dep="13758330" httpUri="https://zenodo.org/record/13758330/files/figure.png" pageId="39" pageNumber="1002" startId="39.[160,225,892,915]" targetBox="[165,1150,144,857]" targetPageId="39" targetType="figure">
<paragraph id="943C36BCFFAAFF83FF2E9D65FD55FC58" blockId="39.[160,1156,892,944]" pageId="39" pageNumber="1002">
<emphasis id="A6F7EAAEFFAAFF83FF2E9D65FE91FC7B" bold="true" box="[160,268,892,915]" pageId="39" pageNumber="1002">Figure 25.</emphasis>
Percent of observed host type by year taken by the Arkenstone Cave population of
<taxonomicName id="53834D3FFFAAFF83FF2E9D80FE54FC58" authority="Townes, 1957" authorityName="Townes" authorityYear="1957" box="[160,457,921,944]" class="Insecta" family="Pompilidae" genus="Ageniella" kingdom="Animalia" order="Hymenoptera" pageId="39" pageNumber="1002" phylum="Arthropoda" rank="species" species="evansi">
<emphasis id="A6F7EAAEFFAAFF83FF2E9D80FEA6FC58" box="[160,315,921,944]" italics="true" pageId="39" pageNumber="1002">Ageniella evansi</emphasis>
<bibRefCitation id="F0124B4DFFAAFF83FECC9D80FE54FC58" author="Townes H" box="[322,457,921,944]" pageId="39" pageNumber="1002" refId="ref48313" refString="Townes H. 1957. Nearctic wasps of the subfamilies Pepsinae and Ceropalinae. United States National Museum Bull. 209. https: // repository. si. edu / handle / 10088 / 10223." type="journal volume" year="1957">Townes, 1957</bibRefCitation>
</taxonomicName>
between 1993 and 2020.
</paragraph>
</caption>
<paragraph id="943C36BCFFAAFF8CFF4E9B19FBDEFF05" blockId="39.[160,1156,1003,1721]" lastBlockId="40.[160,1156,142,410]" lastPageId="40" lastPageNumber="1003" pageId="39" pageNumber="1002">
The length of incoming spiders was estimated using a combination of the visual length of the wasp and the proportional length of the spider. Only the small handful of spiders that were separated from female wasps were physically measured, and are included in the data. The combined average length of all spider hosts observed was
<quantity id="537B9B59FFAAFF83FC1C9B71FC78FA69" box="[914,997,1384,1409]" metricMagnitude="-3" metricUnit="m" metricValue="9.2" pageId="39" pageNumber="1002" unit="mm" value="9.2">9.2 mm</quantity>
(N = 311). The average length of spiders by
<typeStatus id="4B38881EFFAAFF83FE519B92FD8CFA4C" box="[479,529,1419,1444]" pageId="39" pageNumber="1002">type</typeStatus>
was:
<taxonomicName id="53834D3FFFAAFF83FDC39B92FD7CFA4C" box="[589,737,1419,1444]" class="Arachnida" family="Gnaphosidae" kingdom="Animalia" order="Araneae" pageId="39" pageNumber="1002" phylum="Arthropoda" rank="family">Gnaphosidae</taxonomicName>
<quantity id="537B9B59FFAAFF83FD689B92FCA5FA4C" box="[742,824,1419,1444]" metricMagnitude="-3" metricUnit="m" metricValue="9.7" pageId="39" pageNumber="1002" unit="mm" value="9.7">9.7 mm</quantity>
(N = 76),
<taxonomicName id="53834D3FFFAAFF83FC2E9B92FBB7FA4C" authorityName="Simon" authorityYear="1897" box="[928,1066,1419,1444]" class="Arachnida" family="Selenopidae" kingdom="Animalia" order="Araneae" pageId="39" pageNumber="1002" phylum="Arthropoda" rank="family">Selenopidae</taxonomicName>
<quantity id="537B9B59FFAAFF83FBBE9B92FB1EFA4C" box="[1072,1155,1419,1444]" metricMagnitude="-3" metricUnit="m" metricValue="9.3" pageId="39" pageNumber="1002" unit="mm" value="9.3">9.3 mm</quantity>
(N = 208),
<taxonomicName id="53834D3FFFAAFF83FE9B9BB7FE1EFA2F" box="[277,387,1454,1479]" class="Arachnida" family="Lycosidae" kingdom="Animalia" order="Araneae" pageId="39" pageNumber="1002" phylum="Arthropoda" rank="family">Lycosidae</taxonomicName>
<quantity id="537B9B59FFAAFF83FE079BB7FE40FA2F" box="[393,477,1454,1479]" metricMagnitude="-3" metricUnit="m" metricValue="7.6" pageId="39" pageNumber="1002" unit="mm" value="7.6">7.6 mm</quantity>
(N = 25) and
<taxonomicName id="53834D3FFFAAFF83FDF69BB7FD6AFA2F" box="[632,759,1454,1479]" class="Arachnida" family="Agelenidae" kingdom="Animalia" order="Araneae" pageId="39" pageNumber="1002" phylum="Arthropoda" rank="family">Agelenidae</taxonomicName>
<quantity id="537B9B59FFAAFF83FD709BB7FCCCFA2F" box="[766,849,1454,1479]" metricMagnitude="-3" metricUnit="m" metricValue="8.5" pageId="39" pageNumber="1002" unit="mm" value="8.5">8.5 mm</quantity>
(N = 2). The slightly higher average length in gnaphosid over
<taxonomicName id="53834D3FFFAAFF83FDA59BC9FD17FA01" box="[555,650,1488,1513]" class="Arachnida" family="Selenopidae" genus="Selenops" kingdom="Animalia" order="Araneae" pageId="39" pageNumber="1002" phylum="Arthropoda" rank="genus">
<emphasis id="A6F7EAAEFFAAFF83FDA59BC9FD17FA01" box="[555,650,1488,1513]" italics="true" pageId="39" pageNumber="1002">Selenops</emphasis>
</taxonomicName>
spiders does not seem of a magnitude that alone would explain the apparent preference for gnaphosids. Proportionally, however, gnaphosids have abdomens of greater mass and this could conceivably be a driving factor in preference for those spiders. Since the wasps probably select for larger spiders, the noticeably smaller average length of lycosids may suggest these are smaller species than the other hosts used. Body length measurements for female wasps averaged
<quantity id="537B9B59FFAAFF83FC779867FBC7F97F" box="[1017,1114,1662,1687]" metricMagnitude="-2" metricUnit="m" metricValue="1.1199999999999999" pageId="39" pageNumber="1002" unit="mm" value="11.2">11.2 mm</quantity>
(
<quantity id="537B9B59FFAAFF83FBE99867FF6FF951" metricMagnitude="-2" metricUnit="m" metricValue="1.1" metricValueMax="1.4" metricValueMin="0.8" pageId="39" pageNumber="1002" unit="mm" value="11.0" valueMax="14.0" valueMin="8.0">8– 14 mm</quantity>
; N = 1548) and males 9.0 mm (
<quantity id="537B9B59FFAAFF83FDDA98B9FD5FF951" box="[596,706,1696,1721]" metricMagnitude="-3" metricUnit="m" metricValue="8.0" metricValueMax="10.0" metricValueMin="6.0" pageId="39" pageNumber="1002" unit="mm" value="8.0" valueMax="10.0" valueMin="6.0">6–10 mm</quantity>
; N = 347). Thus, on average, the female wasps are approximately 22% longer than the spiders they take. The group of spider host
<typeStatus id="4B38881EFFA5FF8CFF2E9EA8FF41FF22" box="[160,220,177,202]" pageId="40" pageNumber="1003">types</typeStatus>
selected by
<taxonomicName id="53834D3FFFA5FF8CFEFC9EA8FE48FF22" authorityName="Townes" authorityYear="1957" box="[370,469,177,202]" class="Insecta" family="Pompilidae" genus="Ageniella" kingdom="Animalia" order="Hymenoptera" pageId="40" pageNumber="1003" phylum="Arthropoda" rank="species" species="evansi">
<emphasis id="A6F7EAAEFFA5FF8CFEFC9EA8FE48FF22" box="[370,469,177,202]" italics="true" pageId="40" pageNumber="1003">A. evansi</emphasis>
</taxonomicName>
suggests hunting of spiders is based on guild and/or hunt niche specificity rather than host species fealty (Evans 1953;
<bibRefCitation id="F0124B4DFFA5FF8CFCC79ECDFBABFF05" author="Rodriguez J &amp; Pitts JP &amp; Florez JA &amp; Bond JE &amp; von Dohlen CD" box="[841,1078,212,237]" pageId="40" pageNumber="1003" pagination="55 - 64" refId="ref47074" refString="Rodriguez J, Pitts JP, Florez JA, Bond JE, von Dohlen CD. 2016. Molecular phylogeny of Pompilinae (Hymenoptera: Pompilidae): evidence for rapid diversification and host shifts in spider wasps. Mol Phylogen Evol. 94: 55 - 64. doi: 10.1016 / j. ympev. 2015.08.014." type="journal article" year="2016">
Rodriguez
<emphasis id="A6F7EAAEFFA5FF8CFC4C9ECDFC6FFF05" box="[962,1010,212,237]" italics="true" pageId="40" pageNumber="1003">et al</emphasis>
. 2016
</bibRefCitation>
).
</paragraph>
<paragraph id="943C36BCFFA5FF8CFF4E9EEFFDF9FE72" blockId="40.[160,1156,142,410]" pageId="40" pageNumber="1003">
Plotting annual percentage of hosts by
<typeStatus id="4B38881EFFA5FF8CFD199EEFFD54FEE7" box="[663,713,246,271]" pageId="40" pageNumber="1003">type</typeStatus>
against the size of the yearly female hunting population shows the proportion of host
<typeStatus id="4B38881EFFA5FF8CFD6C9F00FC83FEDA" box="[738,798,281,306]" pageId="40" pageNumber="1003">types</typeStatus>
used remains consistent when the annual female population is approximately 19 or greater (
<figureCitation id="0CB82A39FFA5FF8CFCD39F25FC5AFEBD" box="[861,967,316,341]" captionStart="Figure 26" captionStartId="40.[160,226,1675,1698]" captionTargetBox="[165,1150,987,1641]" captionTargetId="figure-252@40.[163,1153,985,1644]" captionTargetPageId="40" captionText="Figure 26. Total annual host utilisation, percent by type, relative to maximum annual female population of Ageniella evansi Townes, 1957 at Arkenstone Cave (composite data from 1993–2020)." figureDoi="http://doi.org/10.5281/zenodo.13758333" httpUri="https://zenodo.org/record/13758333/files/figure.png" pageId="40" pageNumber="1003">Figure 26</figureCitation>
). Smaller annual wasp populations do not provide enough spider return data to show whether host
<typeStatus id="4B38881EFFA5FF8CFBDF9F47FB1EFE9F" box="[1105,1155,350,375]" pageId="40" pageNumber="1003">type</typeStatus>
distribution is consistent at those levels.
</paragraph>
</subSubSection>
</treatment>
</document>
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