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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.755.23939" ID-GBIF-Dataset="7392eb62-5d84-4f6a-8c8a-9f4e351a84a7" ID-PMC="PMC5953965" ID-Pensoft-Pub="1313-2970-755-1" ID-PubMed="29769836" ID-ZBK="AADE14787C914355B776C4AEF28347BF" ModsDocAuthor="" ModsDocDate="2018" ModsDocID="1313-2970-755-1" ModsDocOrigin="ZooKeys 755" ModsDocTitle="A revision of the cleptoparasitic bee genus Epeolus Latreille for Nearctic species, north of Mexico (Hymenoptera, Apidae)" checkinTime="1525801749154" checkinUser="pensoft" docAuthor="Onuferko, Thomas M." docDate="2018" docId="E3CCC42001195F074FB9DD7E527C3059" docLanguage="en" docName="ZooKeys 755: 1-185" docOrigin="ZooKeys 755" docSource="http://dx.doi.org/10.3897/zookeys.755.23939" docTitle="Epeolus nebulosus Onuferko, 2018, sp. n." docType="treatment" docUuid="6C689E06-558F-4C7B-97CE-4BD60D1614E7" docUuidSource="ZooBank" docVersion="5" lastPageNumber="122" masterDocId="D91DD51C1718C109FF8EFFA6FFF3FF9F" masterDocTitle="A revision of the cleptoparasitic bee genus Epeolus Latreille for Nearctic species, north of Mexico (Hymenoptera, Apidae)" masterLastPageNumber="185" masterPageNumber="1" pageNumber="120" updateTime="1668165766199" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>A revision of the cleptoparasitic bee genus Epeolus Latreille for Nearctic species, north of Mexico (Hymenoptera, Apidae)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Onuferko, Thomas M.</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2018</mods:date>
<mods:detail type="volume">
<mods:number>755</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>1</mods:start>
<mods:end>185</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.755.23939</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.755.23939</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-755-1</mods:identifier>
<mods:identifier type="ZBK">AADE14787C914355B776C4AEF28347BF</mods:identifier>
<mods:identifier type="ZooBank">AADE14787C914355B776C4AEF28347BF</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="143842348" LSID="urn:lsid:zoobank.org:act:6C689E06-558F-4C7B-97CE-4BD60D1614E7" httpUri="http://treatment.plazi.org/id/E3CCC42001195F074FB9DD7E527C3059" lastPageId="121" lastPageNumber="122" pageId="119" pageNumber="120">
<subSubSection pageId="119" pageNumber="120" type="nomenclature">
<paragraph pageId="119" pageNumber="120">
34.
<taxonomicName LSID="http://zoobank.org/6C689E06-558F-4C7B-97CE-4BD60D1614E7" class="Insecta" family="Apidae" genus="Epeolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epeolus nebulosus" order="Hymenoptera" pageId="119" pageNumber="120" phylum="Arthropoda" rank="species" species="nebulosus">Epeolus nebulosus</taxonomicName>
<taxonomicNameLabel pageId="119" pageNumber="120">sp. n.</taxonomicNameLabel>
Figs 71, 72, 99A
</paragraph>
</subSubSection>
<subSubSection pageId="119" pageNumber="120" type="diagnosis">
<paragraph pageId="119" pageNumber="120">Diagnosis.</paragraph>
<paragraph pageId="119" pageNumber="120">
The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell
<taxonomicName lsidName="E. nebulosus" pageId="119" pageNumber="120" rank="species" species="nebulosus">E. nebulosus</taxonomicName>
apart from all other North American
<taxonomicName class="Insecta" family="Apidae" genus="Epeolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epeolus" order="Hymenoptera" pageId="119" pageNumber="120" phylum="Arthropoda" rank="genus">Epeolus</taxonomicName>
except
<taxonomicName lsidName="E. basili" pageId="119" pageNumber="120" rank="species" species="basili">E. basili</taxonomicName>
,
<taxonomicName lsidName="E. novomexicanus" pageId="119" pageNumber="120" rank="species" species="novomexicanus">E. novomexicanus</taxonomicName>
, and
<taxonomicName lsidName="E. pusillus" pageId="119" pageNumber="120" rank="species" species="pusillus">E. pusillus</taxonomicName>
: the axilla is large, with the tip extending well beyond the midlength of the mesoscutellum but at most to the band of pale tomentum along its posterior margin, dilated laterally, and ferruginous to some degree whereas the mesoscutellum is typically all black; the
<normalizedToken originalValue="axillas">axilla's</normalizedToken>
free portion is clearly less than 2/5 as long as its entire medial length; the mesopleuron is closely (most i&lt;1d) and evenly punctate, that of the female is obscured by white tomentum only in the upper half (with a large, sparsely hairy circle occupying much of the ventrolateral half) whereas that of the male (excluding the hypoepimeral area) is entirely obscured by white tomentum; T2-T4 have complete and evenly broad fasciae; the T2 fascia has lobe-like anterolateral extensions of tomentum; and the pseudopygidial area of the female is lunate and wider than long (the apex ≤2
<normalizedToken originalValue="×">x</normalizedToken>
the medial length).
<taxonomicName class="Insecta" family="Apidae" genus="Epeolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epeolus basili" order="Hymenoptera" pageId="119" pageNumber="120" phylum="Arthropoda" rank="species" species="basili">Epeolus basili</taxonomicName>
,
<taxonomicName lsidName="E. nebulosus" pageId="119" pageNumber="120" rank="species" species="nebulosus">E. nebulosus</taxonomicName>
,
<taxonomicName lsidName="E. novomexicanus" pageId="119" pageNumber="120" rank="species" species="novomexicanus">E. novomexicanus</taxonomicName>
, and
<taxonomicName lsidName="E. pusillus" pageId="119" pageNumber="120" rank="species" species="pusillus">E. pusillus</taxonomicName>
are all extremely similar to one another.
<taxonomicName class="Insecta" family="Apidae" genus="Epeolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epeolus nebulosus" order="Hymenoptera" pageId="119" pageNumber="120" phylum="Arthropoda" rank="species" species="nebulosus">Epeolus nebulosus</taxonomicName>
is most similar to
<taxonomicName lsidName="E. novomexicanus" pageId="119" pageNumber="120" rank="species" species="novomexicanus">E. novomexicanus</taxonomicName>
, but in
<taxonomicName lsidName="E. novomexicanus" pageId="119" pageNumber="120" rank="species" species="novomexicanus">E. novomexicanus</taxonomicName>
the mesoscutum usually has distinct paramedian bands and at least the integument beneath the T1 apical fascia is ferruginous, as are sometimes the rest of the tergum and other terga, whereas in
<taxonomicName lsidName="E. nebulosus" pageId="119" pageNumber="120" rank="species" species="nebulosus">E. nebulosus</taxonomicName>
the mesoscutum is entirely obscured by pale tomentum and the metasomal terga (excluding the brown translucent apical margins) are entirely black. In
<taxonomicName lsidName="E. basili" pageId="119" pageNumber="120" rank="species" species="basili">E. basili</taxonomicName>
the metasomal terga are also ferruginous to some degree, and the T2 and T3 (for female) or T2-T4 (for male) fasciae are narrowed medially and removed from the apical margin (in
<taxonomicName lsidName="E. nebulosus" pageId="119" pageNumber="120" rank="species" species="nebulosus">E. nebulosus</taxonomicName>
the T2-T4 fasciae are on or very little removed from the apical margin), and the pseudopygidial area of the female is ≥2
<normalizedToken originalValue="×">x</normalizedToken>
the medial length. Whereas in
<taxonomicName lsidName="E. pusillus" pageId="119" pageNumber="120" rank="species" species="pusillus">E. pusillus</taxonomicName>
the flagellum, except sometimes F1, and metasomal sterna are consistently brown or black and clearly not the same reddish-orange color as the legs (tibiae to tarsi), in
<taxonomicName lsidName="E. nebulosus" pageId="119" pageNumber="120" rank="species" species="nebulosus">E. nebulosus</taxonomicName>
the flagellum, at least ventrally, is the same reddish-orange color as the legs (tibiae to tarsi) as are usually the metasomal sterna.
<taxonomicName class="Insecta" family="Apidae" genus="Epeolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epeolus nebulosus" order="Hymenoptera" pageId="119" pageNumber="120" phylum="Arthropoda" rank="species" species="nebulosus">Epeolus nebulosus</taxonomicName>
is also similar to
<taxonomicName lsidName="E. scutellaris" pageId="119" pageNumber="120" rank="species" species="scutellaris">E. scutellaris</taxonomicName>
in that the axilla is large, with the lateral margin arcuate, and that the apical fasciae are complete. However, in
<taxonomicName lsidName="E. scutellaris" pageId="119" pageNumber="120" rank="species" species="scutellaris">E. scutellaris</taxonomicName>
the pseudopygidial area of the female is much wider (the apex ~2.5-3
<normalizedToken originalValue="×">x</normalizedToken>
the medial length) than in
<taxonomicName lsidName="E. nebulosus" pageId="119" pageNumber="120" rank="species" species="nebulosus">E. nebulosus</taxonomicName>
, and the mesopleuron of both the female and male is obscured by white tomentum only in the upper half (with a large, sparsely hairy circle occupying much of the ventrolateral half).
</paragraph>
</subSubSection>
<subSubSection lastPageId="120" lastPageNumber="121" pageId="119" pageNumber="120" type="description">
<paragraph pageId="119" pageNumber="120">Description.</paragraph>
<paragraph pageId="119" pageNumber="120">MALE: Length 7.2 mm; head length 2.0 mm; head width 2.7 mm; fore wing length 5.5 mm.</paragraph>
<paragraph pageId="119" pageNumber="120">Integument coloration. Mostly black; notable exceptions as follows: at least partially ferruginous on mandible, antenna, pronotal lobe, tegula, axilla, legs, pygidial plate, and metasomal sterna. Mandible with apex darker than rest of mandible; preapical tooth slightly lighter than mandibular apex. Antenna brown and orange in part. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black. S1-S6 reddish orange.</paragraph>
<paragraph pageId="119" pageNumber="120">Pubescence. Face with tomentum densest on clypeus and around antennal socket, slightly sparser on upper paraocular area and vertexal area. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum largely obscured by pale tomentum. Mesopleuron (excluding hypoepimeral area) entirely obscured by white tomentum. Metanotum with tomentum uninterrupted, uniformly off white. T1 with narrow and short discal patch largely obscured by pale tomentum. T2-T6 each with complete fascia, T2 with fascia with wide basomedially convergent anterolateral extensions of tomentum. S4 and S5 with long coppery to silvery subapical hairs, which individually are often darker apically.</paragraph>
<paragraph lastPageId="120" lastPageNumber="121" pageId="119" pageNumber="120">
Surface sculpture. Punctures dense. Labrum with larger and sparser punctures (i=1-2d) than clypeus (i&lt;1d) (difficult to see in holotype because clypeus entirely obscured
<pageBreakToken pageId="120" pageNumber="121" start="start">by</pageBreakToken>
tomentum; described from paratypes with hair removed). Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate mesally (i≤1d), less so laterally (i=1-2d). Mesopleuron with ventrolateral half densely punctate (i&lt;1d) to rugose; mesopleuron with punctures more or less equally dense throughout (not visible in holotype because mesopleuron entirely obscured by tomentum; described from paratypes). Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.
</paragraph>
<paragraph pageId="120" pageNumber="121">
Structure. Preapical tooth obtuse. Labrum with pair of small subapical denticles not preceded by carinae (difficult to see in holotype; described from paratypes). Frontal keel not strongly raised. Scape with greatest length 2.0
<normalizedToken originalValue="×">x</normalizedToken>
greatest width. F2 noticeably longer than wide (L/W ratio = 1.2). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1.5 MOD at its terminal. Mesoscutellum weakly bigibbous. Axilla large, its lateral margin (L) half as long as mesoscutellar width (W) (L/W ratio = 0.5) and tip extending well beyond midlength of mesoscutellum but not as far back as its posterior margin; axilla with tip clearly visible, but unattached to mesoscutellum for less than 2/5 the medial length of axilla; axilla with lateral margin arcuate. Fore wing with three submarginal cells. Pygidial plate apically rounded, with large deep, well-separated punctures, with the interspaces shining.
</paragraph>
<paragraph pageId="120" pageNumber="121">FEMALE: Description as for male except for usual secondary sexual characters and as follows: F2 even longer than wide (L/W ratio = 1.5); mesopleuron densely hairy, except for two almost entirely bare patches (one beneath base of fore wing (hypoepimeral area), a larger circular patch occupying much of ventrolateral half of mesopleuron); T5 with large, continuous patch of pale tomentum bordering and contacting pseudopygidial area present only in female; T5 with pseudopygidial area lunate, its apex twice as wide as medial length, indicated by silvery setae on disc of apicomedial region elevated from rest of tergum; S4 and S5 with much shorter hairs (S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by much more than 1/4 MOD); pygidial plate apically truncate, with small, denser punctures.</paragraph>
<caption pageId="120" pageNumber="121">
<paragraph pageId="120" pageNumber="121">
Figure 71.
<taxonomicName class="Insecta" family="Apidae" genus="Epeolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epeolus nebulosus" order="Hymenoptera" pageId="120" pageNumber="121" phylum="Arthropoda" rank="species" species="nebulosus">Epeolus nebulosus</taxonomicName>
A female allotype, lateral habitus (scale bar 3 mm) B female allotype, dorsal habitus (scale bar 3 mm) C male holotype, lateral habitus (scale bar 3 mm), and D female allotype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="120" pageNumber="121" type="etymology">
<paragraph pageId="120" pageNumber="121">Etymology.</paragraph>
<paragraph pageId="120" pageNumber="121">
The name is in reference to the pale tomentum obscuring much of the integument of this species. From the Latin,
<normalizedToken originalValue="“nebulosus”">&quot;nebulosus&quot;</normalizedToken>
(hazy).
</paragraph>
</subSubSection>
<subSubSection pageId="120" pageNumber="121" type="distribution">
<paragraph pageId="120" pageNumber="121">Distribution.</paragraph>
<paragraph pageId="120" pageNumber="121">California and probably western Nevada (Fig. 72).</paragraph>
<caption pageId="120" pageNumber="121">
<paragraph pageId="120" pageNumber="121">
Figure 72. Occurrence records of
<taxonomicName lsidName="E. nebulosus" pageId="120" pageNumber="121" rank="species" species="nebulosus">E. nebulosus</taxonomicName>
known to the author (yellow circles).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="120" pageNumber="121" type="ecology">
<paragraph pageId="120" pageNumber="121">Ecology.</paragraph>
<paragraph pageId="120" pageNumber="121">
HOST RECORDS: The host species of
<taxonomicName lsidName="E. nebulosus" pageId="120" pageNumber="121" rank="species" species="nebulosus">E. nebulosus</taxonomicName>
is/are presently unknown.
</paragraph>
<paragraph pageId="120" pageNumber="121">
FLORAL RECORDS: Labels of examined voucher specimens indicate a floral association with
<taxonomicName class="Magnoliopsida" family="Asteraceae" genus="Ericameria" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ericameria nauseosa" order="Asterales" pageId="120" pageNumber="121" phylum="Tracheophyta" rank="species" species="nauseosa">Ericameria nauseosa</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection lastPageId="121" lastPageNumber="122" pageId="120" pageNumber="121" type="discussion">
<paragraph pageId="120" pageNumber="121">Discussion.</paragraph>
<paragraph lastPageId="121" lastPageNumber="122" pageId="120" pageNumber="121">
<taxonomicName class="Insecta" family="Apidae" genus="Epeolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epeolus nebulosus" order="Hymenoptera" pageId="120" pageNumber="121" phylum="Arthropoda" rank="species" species="nebulosus">Epeolus nebulosus</taxonomicName>
is a cryptic species within the &quot;pusillus group&quot; that closely resembles some specimens of
<taxonomicName lsidName="E. novomexicanus" pageId="120" pageNumber="121" rank="species" species="novomexicanus">E. novomexicanus</taxonomicName>
, and the ranges of the two species overlap to some extent. The morphological differences (in integument coloration and patterns of pubescence) among the four members of the &quot;pusillus group&quot; are subtle. The status of
<taxonomicName lsidName="E. nebulosus" pageId="120" pageNumber="121" rank="species" species="nebulosus">E. nebulosus</taxonomicName>
as a separate species is further supported by a separate BIN and large barcode sequence divergence (&gt;3.2%) from its nearest neighbor,
<taxonomicName lsidName="E. novomexicanus" pageId="120" pageNumber="121" rank="species" species="novomexicanus">E. novomexicanus</taxonomicName>
. Although most species of
<taxonomicName class="Insecta" family="Apidae" genus="Epeolus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Epeolus" order="Hymenoptera" pageId="120" pageNumber="121" phylum="Arthropoda" rank="genus">Epeolus</taxonomicName>
were described from a female name-bearing type, a male specimen is designated as the holotype of
<taxonomicName lsidName="E. nebulosus" pageId="120" pageNumber="121" rank="species" species="nebulosus">E. nebulosus</taxonomicName>
because a barcode-compliant sequence is associated with it and because much of the pubescence
<pageBreakToken pageId="121" pageNumber="122" start="start">is</pageBreakToken>
discolored or rubbed off in the available female specimen, which is herein designated as the allotype. Since this species is described from very few specimens, efforts should be made to collect additional representatives of
<taxonomicName lsidName="E. nebulosus" pageId="121" pageNumber="122" rank="species" species="nebulosus">E. nebulosus</taxonomicName>
for DNA barcoding to determine if the morphological differences between it and
<taxonomicName lsidName="E. novomexicanus" pageId="121" pageNumber="122" rank="species" species="novomexicanus">E. novomexicanus</taxonomicName>
reported here are consistent.
</paragraph>
</subSubSection>
<subSubSection pageId="121" pageNumber="122" type="materials_examined">
<paragraph pageId="121" pageNumber="122">Material studied.</paragraph>
<paragraph pageId="121" pageNumber="122">
Type material. Primary: USA: California: Gilbert Pass on Hwy 168 (
<geoCoordinate direction="north" orientation="latitude" precision="5" value="37.4305">37.4305°N</geoCoordinate>
;
<geoCoordinate direction="west" orientation="longitude" precision="5" value="-117.9388">117.9388°W</geoCoordinate>
) (N Deep Springs Valley, Inyo County), 14.ix.2013, M.C. Orr (holotype ♂ [CCDB-28239 F01], BBSL).
</paragraph>
<paragraph pageId="121" pageNumber="122">
Secondary: USA: California: 3.2 km S Pearblossom (Los Angeles County), 07.xi.1977, R.R. Snelling (allotype ♀, LACM); Gilbert Pass on Hwy 168 (
<geoCoordinate direction="north" orientation="latitude" precision="5" value="37.4305">37.4305°N</geoCoordinate>
;
<geoCoordinate direction="west" orientation="longitude" precision="5" value="-117.9388">117.9388°W</geoCoordinate>
) (N Deep Springs Valley, Inyo County), 14.ix.2013, M.C. Orr (paratypes 2♂, BBSL).
</paragraph>
</subSubSection>
<subSubSection pageId="121" pageNumber="122" type="materials_examined">
<paragraph pageId="121" pageNumber="122">DNA barcoded material with BIN-compliant sequences.</paragraph>
<paragraph pageId="121" pageNumber="122">Available. BOLD:ACZ0767. See Type material for specimens examined and sequenced (indicated by unique CCDB-plate and well number).</paragraph>
</subSubSection>
</treatment>
</document>