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<mods:title id="176AD8B2D2A33C0CA2D82BC5BC49C63B">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="9F9CF33BE268A68D9E1034DC18A2AC85">Katz, Aron D.</mods:namePart>
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<mods:namePart id="AA8A321415169C6221DCA413B6A1B374">Soto-Adames, Felipe</mods:namePart>
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<treatment id="FB17ACB651FC2651B8B7C51630C8DE11" ID-GBIF-Taxon="152062605" LSID="urn:lsid:plazi:treatment:FB17ACB651FC2651B8B7C51630C8DE11" httpUri="http://treatment.plazi.org/id/FB17ACB651FC2651B8B7C51630C8DE11" lastPageId="45" lastPageNumber="46" pageId="42" pageNumber="43">
<subSubSection id="247BCE13E077A57F8A2C24F77E773700" pageId="42" pageNumber="43" type="multiple">
<paragraph id="826B2E2D84DAD9CE0093955DB914BED8" pageId="42" pageNumber="43">Taxon classification Animalia Collembola Entomobryidae</paragraph>
</subSubSection>
<subSubSection id="5C64FE5510CBD36592C75C08F299FA7A" pageId="42" pageNumber="43" type="nomenclature">
<paragraph id="5845B44E4BF703429ED9903030AF7FF0" pageId="42" pageNumber="43">
<taxonomicName id="78D7A600B8ABD3A1A5F45D83BBB53541" ID-CoL="6FF5L" authority="(Linnaeus), 1758" authorityName="(Linnaeus)" authorityYear="1758" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya nivalis" order="Entomobryomorpha" pageId="42" pageNumber="43" phylum="Arthropoda" rank="species" species="nivalis">Entomobrya nivalis (Linnaeus), 1758</taxonomicName>
Figs 2, 31, 32, 39
</paragraph>
</subSubSection>
<subSubSection id="6A0012A8D07546DCA7562FDBBB157EF2" pageId="42" pageNumber="43" type="description">
<paragraph id="F2F468BF636FE09629180FF27E33C2DC" pageId="42" pageNumber="43">Description.</paragraph>
<paragraph id="310E6724A75514CEEAF6CBE5029FE3EC" pageId="42" pageNumber="43">
Body shape and color pattern. Body cylindrical. One primary, but variable, color form (Fig. 31): yellow or white background with black, dark brown or purple pigment always forming thin transverse bands along the posterior margin of Th. 3, and Abd. 2 through Abd. 6. Additional transverse bands present or absent on Th. 2 and Abd. 1. Abd. 4 usually with U-shaped or
<normalizedToken id="B06186B53043F7608BA89BDB2BDDD70A" originalValue="“11”">&quot;11&quot;</normalizedToken>
-shaped pattern connecting basally with band along posterior margin. Antennae usually lack dark pigmentation, sometimes with light brown or purple pigment, darkening near the apex.
</paragraph>
<caption id="B1CD64C1FE074D44C7856298DE2E10AD" pageId="42" pageNumber="43">
<paragraph id="303DCB42A0B6B2C558CC8BD2CB5A37CB" pageId="42" pageNumber="43">
Figure 31. Color pattern of
<taxonomicName id="644D422AF1EC0674234FC69818A66F47" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya nivalis" order="Entomobryomorpha" pageId="42" pageNumber="43" phylum="Arthropoda" rank="species" species="nivalis">Entomobrya nivalis</taxonomicName>
collected from: A Lamoille Co., VT B Sauk Co., WI C Hancock Co., ME D Lamoille Co., VT E Hancock Co., ME F Sauk Co., WI.
</paragraph>
</caption>
<paragraph id="F012077AB349B0C7BE433189FE1F7DE8" pageId="42" pageNumber="43">
Head. Apical bulb of 4th antennal segment usually bi- or trilobed. Long differentiated smooth setae on ventral side of 1st antennal segment ≈3
<normalizedToken id="005AD59D879B65D4A7C5E1EB33FAF254" originalValue="×">x</normalizedToken>
short setae. Ornamentation of the distal margin of the labral papillae with 3-4 small seta or spine-like projections per papilla (Fig. 2). Labial lateral appendage slightly curved, relatively thick, nearly reaching tip of papilla. Labial triangle chaetotaxy normal, one specimen with a supplemental ciliate seta internal to M1 (one side only). Dorsal head chaetotaxy (Fig. 32A) with macrosetae
<normalizedToken id="2E6DC3D8C619295340F8FAA13372275C" originalValue="An">An'</normalizedToken>
0, A3a2, A3a3, M3i,
<normalizedToken id="C37B45CE23B699EBC9806ED429B9443F" originalValue="S">S'</normalizedToken>
0, Ps3, Pi2, Pi3, Pm1i, and Pm2 absent; S1 usually present, A6, S6, Ps3, and Pa2 sometimes present. Eyes G and H small and subequal. Eye patch with 5 or 6 setae.
</paragraph>
<caption id="16F17571B2E01D8CAE6B8E331CC8E6C3" pageId="42" pageNumber="43">
<paragraph id="B5836C21CD3D9882FE57D174C62A673A" pageId="42" pageNumber="43">
Figure 32. Dorsal chaetotaxy of
<taxonomicName id="B7F5C2CC52C562A3CAB2B3912B197F4D" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya nivalis" order="Entomobryomorpha" pageId="42" pageNumber="43" phylum="Arthropoda" rank="species" species="nivalis">Entomobrya nivalis</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="844671BCA79800FAC40637C3BD848714" originalValue="µm">µm</normalizedToken>
. See Figure 5 for symbol legend.
</paragraph>
</caption>
<paragraph id="EB23A08884CFA362FA17D9AFD8B48CA4" pageId="42" pageNumber="43">Legs. Trochanteral organ with triangular setal pattern and up to 34 setae. Unguis with 4 internal teeth; basal teeth located approximately 60% of inner claw length. Unguiculus acuminate with small serrations on internal edge.</paragraph>
<paragraph id="117326B6AFE4E5965E33D5D94727485B" pageId="42" pageNumber="43">Thorax. Th. 2 macrosetae a5, m1, m2, m4, m4i, m4pi, and m5 present (Fig. 32B); m2i and m2i2 usually present; m1i and m4p sometimes present; all posterior macrosetae (series P) are present. Th. 3 macrosetae a1, a2, a3, a4, a4i, a5, a6, a7, m5, m5i, m6, and m7 present (Fig. 32C); most posterior macrosetae (series P) present, p3 present or absent. Both thoracic segments with a moderate number of supplemental macrosetae in zone Pm (Fig. 32B,C).</paragraph>
<paragraph id="304F71A326F7AD852B4A3E6870CBED06" pageId="42" pageNumber="43">Abdomen. Abd. 1 with 7-10 macrosetae (Fig. 32D). Abd. 2 macroseta a2, a3, m3, m3e, m3ep, and m5 present; m3ei and m3ea usually absent (Fig. 32E). Abd. 3 macroseta a1, m3, am6, pm6, and p6 present (Fig. 32F); a3 usually absent. Abd. 4 with 7-8 inner macrosetae (Fig. 32G). Mucronal teeth subequal.</paragraph>
</subSubSection>
<subSubSection id="119FDC6026393B9DBBD35559FD5F3F2C" lastPageId="45" lastPageNumber="46" pageId="43" pageNumber="44" type="remarks">
<paragraph id="5B3B1E0A993569BA5FBFBC6167BCC81E" pageId="43" pageNumber="44">
<pageBreakToken id="3717D573FB2A5682A3645EDA6E893A45" pageId="43" pageNumber="44" start="start">Remarks</pageBreakToken>
.
</paragraph>
<paragraph id="961A909F4961F5E5538FFE10616CC904" pageId="43" pageNumber="44">
<taxonomicName id="22B4ED136C6EF3A7F3CD9CD94FA38137" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya nivalis" order="Entomobryomorpha" pageId="43" pageNumber="44" phylum="Arthropoda" rank="species" species="nivalis">Entomobrya nivalis</taxonomicName>
can be diagnosed by the presence of a U-shaped or
<normalizedToken id="B84AD7F065DACDDE52A4FB17CC72E901" originalValue="“11”">&quot;11&quot;</normalizedToken>
shaped pattern on Abd. 4 combined with the presence of macrosetae head S4i, Th. 2 m5, Abd. 2 m3ep, and Abd. 3 a1, and the absence of Abd. 3 a2 (see Table 2 for additional diagnostic characters). This species has a highly variable color pattern with many intermediate forms that intergrade with
<taxonomicName id="B2C88BD1CB3AC22C33BA7EF4408146C5" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya atrocincta" order="Entomobryomorpha" pageId="43" pageNumber="44" phylum="Arthropoda" rank="species" species="atrocincta">Entomobrya atrocincta</taxonomicName>
females,
<taxonomicName id="58FF804AE8E4CDD964939B4522CA285C" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya intermedia" order="Entomobryomorpha" pageId="43" pageNumber="44" phylum="Arthropoda" rank="species" species="intermedia">Entomobrya intermedia</taxonomicName>
, and
<taxonomicName id="92F0B733DC93D08C8C18391D6A6AD4AF" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya multifasciata" order="Entomobryomorpha" pageId="43" pageNumber="44" phylum="Arthropoda" rank="species" species="multifasciata">Entomobrya multifasciata</taxonomicName>
. However,
<taxonomicName id="213E4E4A1E5925EF89C280A52CC37FA9" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya nivalis" order="Entomobryomorpha" pageId="43" pageNumber="44" phylum="Arthropoda" rank="species" species="nivalis">Entomobrya nivalis</taxonomicName>
can be separated from these species by chaetotaxy (Table 3) and, with careful consideration, color pattern; the presence of a U-shaped or
<normalizedToken id="0E4091CB024E9AC0E62E842F6A1CA7F0" originalValue="“11”">&quot;11&quot;</normalizedToken>
shaped pattern on Abd. 4 is unique to this species. Therefore, it is critical to evaluate chaetotaxy in addition to color pattern when making a species diagnosis.
</paragraph>
<paragraph id="672B8F5F0613B3EEFAEA602B1EFB5B07" lastPageId="45" lastPageNumber="46" pageId="44" pageNumber="45">
<pageBreakToken id="A29D2585D7D46F67CCE4FD375625B0B8" pageId="44" pageNumber="45" start="start">It</pageBreakToken>
is important to note that the large genetic distances between presumably conspecific individuals (
<bibRefCitation id="353236DCE13F384647C8F45C19AA6821" 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>
; Feng Zhang, personal communication), differences in chaetotaxy between populations in North America and Europe (
<bibRefCitation id="BDC19B1C63274A2070FCE39F73A5F05A" author="Jordana, R" journalOrPublisher="Soil Organisms" pageId="60" pageNumber="61" pagination="1 - 390" title="Synopses of Palaearctic Collembola: Capbryinae &amp; Entomobryini." volume="84" year="2012">Jordana 2012</bibRefCitation>
), and
<pageBreakToken id="D6F476D91CE85DBBA8C41166B1B4DAAF" pageId="45" pageNumber="46" start="start">variable</pageBreakToken>
color pattern among populations across its world-wide distribution, suggests that
<taxonomicName id="FDEB1662DABC5F31869B54A14AABE8A0" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya nivalis" order="Entomobryomorpha" pageId="45" pageNumber="46" phylum="Arthropoda" rank="species" species="nivalis">Entomobrya nivalis</taxonomicName>
likely represents a cryptic species complex.
</paragraph>
</subSubSection>
<subSubSection id="344D4A5FFBB3202660BB3938A97B0552" pageId="45" pageNumber="46" type="distribution">
<paragraph id="E1F5CDD3449192E271C7FEACC6DA24A0" pageId="45" pageNumber="46">Distribution.</paragraph>
<paragraph id="94F5FB3BDA162751232D7134561213EA" pageId="45" pageNumber="46">
North America and Europe. Records of
<taxonomicName id="A9B096161E17684A5C8196DF43F54308" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya nivalis" order="Entomobryomorpha" pageId="45" pageNumber="46" phylum="Arthropoda" rank="species" species="nivalis">Entomobrya nivalis</taxonomicName>
in North America are suspect if diagnosed without considering chaetotaxy given the superficial similarities in color form expressed by
<taxonomicName id="7C2CDF180D4A573169AFEBFE0588A296" class="Collembola" family="Entomobryidae" genus="Entomobrya" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Entomobrya atrocincta" order="Entomobryomorpha" pageId="45" pageNumber="46" phylum="Arthropoda" rank="species" species="atrocincta">Entomobrya atrocincta</taxonomicName>
females (See Fig. 9G,H). See Suppl. material 2: L for a distribution map and below for a list of material examined with collection and locality information.
</paragraph>
</subSubSection>
<subSubSection id="08C46245A56085088FAA23648569F2A8" pageId="45" pageNumber="46" type="materials_examined">
<paragraph id="8CBD8E2B8D7659A07164C1A9850C69BF" pageId="45" pageNumber="46">Material examined.</paragraph>
<paragraph id="5C245C5B3344CD34310865E2BFDBB9A9" pageId="45" pageNumber="46">
USA: 2 on slides, 50+ in vial, Maine, Hancock Co., Acadia National Park (44.353823,-68.224754), moss, veg. sweep (blueberry, juniper, populus), 17.viii.2011 (E. C. Bernard), #2009-37; 3 on slides, 5 in vial, Pennsylvania, Allegheny Co., Allegheny National Forest, Dewdrop campground (41.83092,-7895937), leaf litter, 8.vii.2008 (S. M. Shreve); 1 on slide, Vermont, Chittenden Co., Red Rock, Locality I (44.44493,-73.23040), leaf litter, 6.vi.2011 (J. Fisher); 2 on slides, Vermont, Chittenden Co., Farrell Park, Locality II (44.44454,-73.20178), leaf litter, 13.vi.2001 (J. Fisher); 1 on slide, Vermont, Chittenden Co., Farrell Park, Locality II (44.44454,-73.20178), leaf litter, 4.vii.2001 (J. Fisher); 2 on slides, Vermont, Chittenden Co., Farrell Park, Locality II (44.44454,-73.20178), leaf litter, 26.vii.2001 (J. Fisher); 1 on slide, Vermont, Lamoille Co., Stowe (44.48377,-72.69859), leaf litter, 24.vii.2001 (J. Fisher); 1 on slide, Vermont, Lamoille Co. (44.54858,-72.79393), DNA ID#: 12-FSVTlam-ni-1; 1 on slide, Vermont, Washington Co., Locality II, Barre (44.19968,-72.50135), leaf litter, 8.vi.2001 (J. Fisher); 2 on slides, Vermont, Washington Co., Locality II, Barre (44.19968,-72.50135), leaf litter, 24.vii.2001 (J. Fisher); 2 on slides, 6 in vial, Vermont, Rutland Co., Green Mountain National Forest, Greendale Recreation Area (43.35112,-72.82225), leaf litter, 10.vii.2008 (S. M. Shreve); 1 on slide, 6 in vial, Wisconsin, Dodge Co., Horicon Marsh National Wildlife Refuge, end of Dike Rd (43.52736,-88.64381), 12.vi.2011, AK11-47; 2 on slides, 7 in vial, Wisconsin, Sauk Co.,
<normalizedToken id="F35E64A39D262E6782A377CD13A6DBD0" originalValue="Devils">Devil's</normalizedToken>
Lake State Park, 0.5mi down Steinke Basin Loop trail (43.4255,-89.71039), 12.vi.2011, AK11-50.
</paragraph>
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</treatment>
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