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<mods:title id="FC93D083496B8387BBDA3814574C800D">Taxonomic review and phylogenetic analysis of fifteen North American Entomobrya (Collembola, Entomobryidae), including four new species</mods:title>
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<mods:namePart id="DE2AF72232A68004FC18F1C52CE32664">Katz, Aron D.</mods:namePart>
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<treatment id="6D16AEA78A8C3EA69C87C2822F226F1A" ID-GBIF-Taxon="152062612" LSID="urn:lsid:plazi:treatment:6D16AEA78A8C3EA69C87C2822F226F1A" httpUri="http://treatment.plazi.org/id/6D16AEA78A8C3EA69C87C2822F226F1A" lastPageId="21" lastPageNumber="22" pageId="20" pageNumber="21">
<subSubSection id="09BE078E8991F68A96D2F296A4C073DD" pageId="20" pageNumber="21" type="multiple">
<paragraph id="E5BD686BC0AD45DA1A743EB5D981BB63" pageId="20" pageNumber="21">Taxon classification Animalia Collembola Entomobryidae</paragraph>
</subSubSection>
<subSubSection id="48A693847DC90AAD6EE57B6DE83C72EA" pageId="20" pageNumber="21" type="nomenclature">
<paragraph id="9051E756B4AF2132D4F7EC4F13310E89" pageId="20" pageNumber="21">
<taxonomicName id="4308801210E84B9C93562659FB090484" ID-CoL="6FFFG" authority="Guthrie, 1903" authorityName="Guthrie" authorityYear="1903" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor Guthrie, 1903</taxonomicName>
Figs 2, 11, 12, 13A, 39
</paragraph>
</subSubSection>
<subSubSection id="B1D1900DEB8BCF1FEDC6B0C6433F1779" pageId="20" pageNumber="21" type="description">
<paragraph id="15A21A0B2D8A2D648FF43DC851AF2E62" pageId="20" pageNumber="21">Description.</paragraph>
<paragraph id="FB4B196981A09FB3AF99550360D2EC55" pageId="20" pageNumber="21">Body shape and color pattern. Body elongate and cylindrical. Only one reported color form in adults (Fig. 11A), dark brown or black pigment covering whole body except for a white band across Abd. 1, 2 and medial area of Abd. 3; Abd. 5 sometimes with 1+1 pale spots; legs, furcula, Abd. 6 pale, lacking all dark pigmentation; antennae lightly colored with brown or purple pigment, usually with a white area on distal half of 1st antennal segment. Juvenile pattern distinct from adult, yellow background without dark pigment except for eye patch; legs, furcula, Abd. 6 pale, lacking all dark pigmentation; antennae with light purple pigment. (Fig. 11B)</paragraph>
<caption id="08B7D5C15AE2FBDE164AC1FFC0FB2C19" pageId="20" pageNumber="21">
<paragraph id="CC96A6A718267CB7F90206A6FDECA971" pageId="20" pageNumber="21">
Figure 11. Color patterns of
<taxonomicName id="0C681E968BFA447244600FF9CEA79FEC" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
: A adult B young immature instar. Both photographed specimens were collected from Henderson Co., IL.
</paragraph>
</caption>
<paragraph id="FE9F987A1A6095F919E24FA57B870185" pageId="20" pageNumber="21">
Head. Apical bulb of 4th antennal segment usually bilobed, rarely simple. Long differentiated smooth setae on ventral side of 1st antennal segment ≈ 3
<normalizedToken id="AC9FDD66E626DC01764463A46ED38E86" originalValue="–4×">-4x</normalizedToken>
short seta. Prelabral setae ciliate. Ornamentation of the distal margin of the labral papillae with a single seta or spine-like projection (Fig. 2). Labial papilla E with lateral appendage reaching just above tip of papilla. Labial triangle in one individual with 2 small supplementary ciliate microsetae internal to M1. Dorsal head chaetotaxy as in Figure 12A: macrosetae
<normalizedToken id="85AA0FC253F1823C52BD63765C7B4847" originalValue="An’">An'</normalizedToken>
0, A6,
<normalizedToken id="9A0A7C3D5D26A42D07BC223B7BA92AC0" originalValue="S’">S'</normalizedToken>
0, S6, Ps3, Ps5, Pi1, Pi3, and Pm1i always absent; An3a3 usually absent (present on one side in one specimen). Eyes G and H small and subequal. Eye patch with 5 setae.
</paragraph>
<caption id="433DA68F04EDC6B982A8757779DE9E31" pageId="20" pageNumber="21">
<paragraph id="7F2726869C74D0FE90775FFD0B77CD00" pageId="20" pageNumber="21">
Figure 12. Dorsal chaetotaxy of
<taxonomicName id="84689D4964AE67B0683F896476912D5A" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
: A Head B Mesothorax C Metathorax D 1st abdominal segment E 2nd abdominal segment F 3rd abdominal segment G 4th abdominal segment. Scale bars = 100
<normalizedToken id="D8059735633543A6433BEE081AFC417B" originalValue="µm">µm</normalizedToken>
. See Figure 5 for symbol legend.
</paragraph>
</caption>
<paragraph id="E95B8E58CAA09343E19451977843021A" pageId="20" pageNumber="21">
Thorax. Thoracic chaetotaxy abundantly developed, highly variable with many supplemental macrosetae. Th. 2 macrosetae p6e and p6ep absent and macrosetae m4i2, m4i3 polymorphic (Fig. 12B). Th. 3 macrosetae a5e3, m4, m5p, a6i, a7, and m7 are absent (Fig. 12C). If present, m7 is always a mesoseta. Chaetotaxy of zone Pm of both thoracic segments densely packed with many supplemental macrosetae, forming wing-like patches of posterior setae extending near anterior row, typical for species within the
<taxonomicName id="6674F8BCB006EC464163DFDA77AB4D84" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
complex (Fig. 3D). Position of pseudopores on Th. 3 atypical for
<taxonomicName id="7DD2011CF07FA4F251C45F5898849E9F" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="genus">Entomobrya</taxonomicName>
, displaced anteriorly and closer to macrosetae a1, a2, and a3 than in other species.
</paragraph>
<paragraph id="E8F9EAF5004049349B969878E9C60A15" pageId="20" pageNumber="21">Legs. Trochanteral organ with rectangular setal pattern and up to 37 setae. Unguis with 4 internal teeth; basal teeth enlarged and located approximately middle of inner claw length.</paragraph>
<paragraph id="E9AED3A911F808DB727B820BD467BE14" pageId="20" pageNumber="21">Abdomen. Abdominal chaetotaxy highly developed. Row of microsetae along entire posterior margin present in all segments (not displayed in figure). Abd. 1 with 12 macrosetae (Fig. 12D). Abd. 2 with 10-11 macrosetae: a2, a3, m3, m3e, m3ep, m3ep2, m3ea, m5, p5, and p6 always present. Abd. 2 macroseta m3eai sometimes present, m3ei always absent (Fig. 12E). Abd. 3 macrosetae a2, a3, m3, am6, pm6, p5, p6 present; a1 always absent (Fig. 12F). Abd. 4 elongated, with at least 22 inner macrosetae (Fig. 12G), number of macrosetae extremely variable between individuals and within individuals (Fig. 13A). Position of pseudopores on Abd. 4 unstable, even varying with respect to macroseta and bothriotricha between sides on same individual (Fig. 13A). Mucronal teeth subequal; mucronal spine enlarged.</paragraph>
<caption id="C6AB40E2F7C45973451E70AA3D165461" pageId="20" pageNumber="21">
<paragraph id="A35540E3BA4A6B827FC94C94B91FFB02" pageId="20" pageNumber="21">
Figure 13. Macrosetae variation and asymmetrical polymorphism of Abd. 4 in the
<taxonomicName id="1BCB492746837F0BC1A1226A39D626BF" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
complex: A
<taxonomicName id="CC7CE9D8A543D9533F4DD82CCF91307A" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
B
<taxonomicName id="D8E8E85DDF47E5CED2E683E31357C049" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya decemfasciata" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="decemfasciata">Entomobrya decemfasciata</taxonomicName>
C
<taxonomicName id="13E7634FFC327E5E8191C3D22703C0EE" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya quadrilineata" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="quadrilineata">Entomobrya quadrilineata</taxonomicName>
. Only medial macrosetae (those which occur between bothriotricha T2 and T4) are included in diagram. The dotted line represents the medial division of Abd. 4. Scale bars = 100
<normalizedToken id="4B2EA42CA2E909D2E68810261606F37F" originalValue="µm">µm</normalizedToken>
.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="3073EC85B5BD0520DCA930F322F432FD" pageId="20" pageNumber="21" type="remarks">
<paragraph id="0EF1F6AF9301E43565530176B675FB90" pageId="20" pageNumber="21">Remarks.</paragraph>
<paragraph id="1E4DD57E60233E4A521274F37E626F58" pageId="20" pageNumber="21">
<taxonomicName id="2CAD951149037CCCEF57833144D2E359" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
adults can be easily distinguished by the unique color pattern described above combined with the absence of head macroseta Ps5 (see Table 2 for additional diagnostic characters). Adults have a single, highly stable color form that is easily recognizable and perfectly acceptable for diagnosis (Fig. 11A). Molecular data from
<bibRefCitation id="E46FFCF5D4B2AC113236FBDBF19F172F" author="Katz, AD" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="60" pageNumber="61" pagination="810 - 840" title="Operational criteria for cryptic species delimitation when evidence is limited, as exemplified by North American Entomobrya (Collembola: Entomobryidae)." url="10.1111/zoj.12220" volume="173" year="2015">Katz et al. (2015)</bibRefCitation>
revealed that a small specimen found with
<taxonomicName id="3EB74115C3DD87792E23EF212D184852" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
, which was completely yellow and lacking all dark pigment, is the juvenile form of
<taxonomicName id="568A398B3E083EB085BC0CA048C4C769" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
(Fig. 11B). The
<taxonomicName id="3857A1A01F734BDA30754424BF6D8E2A" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
juvenile color form has never been reported in the literature. Unfortunately, it may be difficult to diagnose juvenile members of this species if collected without adults present in the sample.
</paragraph>
<paragraph id="B1C31C1E33DE55D0C75145C5CCB2F306" pageId="20" pageNumber="21">
<bibRefCitation id="F4E3F5809F89B6285C750F7F1FC3D80F" author="Christiansen, K" journalOrPublisher="Bulletin of the Museum of Comparative Zoology" pageId="59" pageNumber="60" pagination="439 - 594" title="The Nearctic members of the genus Entomobrya (Collembola)." volume="118" year="1958 b">Christiansen (1958b)</bibRefCitation>
placed this species in what he termed &quot;the
<taxonomicName id="2ABF8E62C959809D54F15AC12C4052AA" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
group&quot;; a complex of three closely related species;
<taxonomicName id="CD6168A73E7A50DD6B7B45A28FA4B9E0" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya quadrilineata" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="quadrilineata">Entomobrya quadrilineata</taxonomicName>
,
<taxonomicName id="F3514133F184775B2E40FDCE328ED28B" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya decemfasciata" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="decemfasciata">Entomobrya decemfasciata</taxonomicName>
, and
<taxonomicName id="0BC93D000BD61A35F6ACA1823D61D2D8" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
. This group is characterized by a highly elongate, cylindrical body, a relatively long furcula, antennae and legs, and by their abundant and highly variable chaetotaxy; many duplicate, supplementary macrosetae, distinctive and augmented setal patterns in thoracic zone Pm, expanded chaetotaxy of Abd. 4, and high levels of asymmetry (Fig. 13A). The extreme setal variation obscures potentially informative characters and results in a lack of discrete, useful diagnostic chaetotaxy between species in this complex. The absence of head macroseta Ps5 is the only character (other than color pattern) that differentiates
<taxonomicName id="514B1AD19F615096CB98132D1056D53B" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
from
<taxonomicName id="5B75F73743496B9B86AACEA95E814882" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya decemfasciata" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="decemfasciata">Entomobrya decemfasciata</taxonomicName>
and
<taxonomicName id="1E5DCA4CE34F603705C17ABE82312B50" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya quadrilineata" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="quadrilineata">Entomobrya quadrilineata</taxonomicName>
. Color pattern is critical for species diagnosis within this complex. See Table 5 for a summary of the important diagnostic characters to separate species in this complex.
</paragraph>
<caption id="D1DF9588705571265671818ABB51B0A1" pageId="20" pageNumber="21">
<paragraph id="BE83DE2099198109AF7555C4C1A0220C" pageId="20" pageNumber="21">
Table 5. Important diagnostic characters for the separation of species within the bicolor complex:
<taxonomicName id="AD716D08303A304B93F8B08B67FC43D3" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
,
<taxonomicName id="19D3C2C8FBA4B10AB25B8291B3B760F7" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya decemfasciata" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="decemfasciata">Entomobrya decemfasciata</taxonomicName>
, and
<taxonomicName id="274A9574343A5D6389B7FD22D1E57E81" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya quadrilineata" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="quadrilineata">Entomobrya quadrilineata</taxonomicName>
. Character states: absent (0), present (1).
</paragraph>
</caption>
<paragraph id="74E67AF90134CBE15CAE6889AC8302B3" pageId="20" pageNumber="21">
<table id="0283FDB458C12B917CDBAA227C9A09D2" pageId="20" pageNumber="21">
<tr id="BE02C932670CCCA75BE717E4ED08638B" pageId="20" pageNumber="21">
<th id="4AE7D9465E7F07258D7C58DA54E803F7" colspan="1" pageId="20" pageNumber="21" rowspan="1">Species</th>
<th id="3CA80A53B2D79171F63758AAA6B0EF00" colspan="1" pageId="20" pageNumber="21" rowspan="1">
Head macrosetae Ps
<sub id="98D45E4D992718FC4CAAEB4DE8F3ADF7" pageId="20" pageNumber="21">5</sub>
</th>
<th id="26B33FA3585D722AABE5C8DB3C981BF5" colspan="1" pageId="20" pageNumber="21" rowspan="1">Parallel, longitudinal bands on thorax</th>
</tr>
<tr id="18DF03D7D20FB9A799CB5C6794030E1D" pageId="20" pageNumber="21">
<td id="8D9185E9193FB6D08695513AC2ADCB72" colspan="1" pageId="20" pageNumber="21" rowspan="1">
<taxonomicName id="FA9D759A7C10EC317A901B5CEA39613A" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya bicolor" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bicolor">Entomobrya bicolor</taxonomicName>
</td>
</tr>
<tr id="A83CC6AAA116E76ABF9049D3399978E1" pageId="20" pageNumber="21">
<td id="A0D5D198BB74D06E447649EBAE688149" colspan="1" pageId="20" pageNumber="21" rowspan="1">
<taxonomicName id="0E7D3DFCE78576DADBC010B2C54D034A" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya decemfasciata" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="decemfasciata">Entomobrya decemfasciata</taxonomicName>
</td>
</tr>
<tr id="D96CD4CDA984E7477C3D3BED4410087B" pageId="20" pageNumber="21">
<td id="12B366CD121F17A7533F25E58D57BAA7" colspan="1" pageId="20" pageNumber="21" rowspan="1">
<taxonomicName id="4BA82B02D8F0D8E2418B23AC002FE7A2" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya quadrilineata" order="Entomobryomorpha" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="quadrilineata">Entomobrya quadrilineata</taxonomicName>
</td>
</tr>
</table>
</paragraph>
</subSubSection>
<subSubSection id="C828915C403631C2729B5A0E6455FD5D" pageId="21" pageNumber="22" type="distribution">
<paragraph id="38086BB48C9F486F8168A4147FFA8EF0" pageId="21" pageNumber="22">
<pageBreakToken id="BA4CF02EB1FE8D0274953EB40A2FF9AD" pageId="21" pageNumber="22" start="start">Distribution</pageBreakToken>
.
</paragraph>
<paragraph id="E094ACBC768C2B94DA4463758CDF721D" pageId="21" pageNumber="22">Endemic to North America (Suppl. material 2: C).</paragraph>
</subSubSection>
<subSubSection id="5BBBDF0EC18102BC5C1B800A2572D76A" pageId="21" pageNumber="22" type="materials_examined">
<paragraph id="017A59C030F2C07593A829C3DB08392B" pageId="21" pageNumber="22">Material examined.</paragraph>
<paragraph id="CCC54557219F4417FBF813573122B0C9" pageId="21" pageNumber="22">USA: 9 on slides, 6 in vial, Illinois, Henderson Co., Big River State Forest (41.03435,-90.91620), vacuum sand prairie, 8.vi.2011 (C. H. Dietrich).</paragraph>
</subSubSection>
</treatment>
</document>
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