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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.148.1809" ID-GBIF-Dataset="c78f07a8-c6ec-4490-a8a4-d17697b29e6b" ID-PMC="PMC3264415" ID-Pensoft-Pub="1313-2970-148-293" ID-PubMed="22287902" ModsDocAuthor="" ModsDocDate="2011" ModsDocID="1313-2970-148-293" ModsDocOrigin="ZooKeys 148" ModsDocTitle="Brachyceran Diptera (Insecta) in Cretaceous ambers, Part IV, Significant New Orthorrhaphous Taxa" checkinTime="1451249597577" checkinUser="pensoft" docAuthor="Grimaldi, David A., Arillo, Antonio, Cumming, Jeffrey M. &amp; Hauser, Martin" docDate="2011" docId="B8D356C86BDE873F18DFD631BCCC332E" docLanguage="en" docName="ZooKeys 148: 293-332" docOrigin="ZooKeys 148" docSource="http://dx.doi.org/10.3897/zookeys.148.1809" docTitle="Cratotabanus newjerseyensis Grimaldi, sp. n." docType="treatment" docVersion="5" lastPageNumber="303" masterDocId="FF87FFED073F016AFFA1FF97FFD3590E" masterDocTitle="Brachyceran Diptera (Insecta) in Cretaceous ambers, Part IV, Significant New Orthorrhaphous Taxa" masterLastPageNumber="332" masterPageNumber="293" pageNumber="300" updateTime="1668152765975" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Brachyceran Diptera (Insecta) in Cretaceous ambers, Part IV, Significant New Orthorrhaphous Taxa</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Grimaldi, David A.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Arillo, Antonio</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Cumming, Jeffrey M.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Hauser, Martin</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>2011</mods:date>
<mods:detail type="volume">
<mods:number>148</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>293</mods:start>
<mods:end>332</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.148.1809</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.148.1809</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-148-293</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="152032210" LSID="urn:lsid:zoobank.org:act:B8FD8A73-669D-44EB-B923-464378B6AB94" httpUri="http://treatment.plazi.org/id/B8D356C86BDE873F18DFD631BCCC332E" lastPageId="10" lastPageNumber="303" pageId="7" pageNumber="300">
<subSubSection pageId="7" pageNumber="300" type="nomenclature">
<paragraph pageId="7" pageNumber="300">
<taxonomicName LSID="urn:lsid:zoobank.org:act:B8FD8A73-669D-44EB-B923-464378B6AB94" authority="Grimaldi" class="Insecta" family="Tabanidae" genus="Cratotabanus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cratotabanus newjerseyensis" order="Diptera" pageId="7" pageNumber="300" phylum="Arthropoda" rank="species" species="newjerseyensis">Cratotabanus newjerseyensis Grimaldi</taxonomicName>
<taxonomicNameLabel pageId="7" pageNumber="300">sp. n.</taxonomicNameLabel>
Fig. 4
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="300" type="diagnosis">
<paragraph pageId="7" pageNumber="300">Diagnosis.</paragraph>
<paragraph pageId="7" pageNumber="300">
Venation differs from congener by
<taxonomicName class="Insecta" family="Tabanidae" genus="Cratotabanus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cratotabanus stenomyomorphus" order="Diptera" pageId="7" pageNumber="300" phylum="Arthropoda" rank="species" species="stenomyomorphus">Cratotabanus stenomyomorphus</taxonomicName>
having vein R4 not strongly upcurved (vs. strongly upcurved) and R5 slightly downcurved (vs. nearly in line with R4+5).
</paragraph>
</subSubSection>
<subSubSection lastPageId="9" lastPageNumber="302" pageId="7" pageNumber="300" type="description">
<paragraph pageId="7" pageNumber="300">Description.</paragraph>
<paragraph lastPageId="8" lastPageNumber="301" pageId="7" pageNumber="300">
AMNH NJ-1862 (holotype): Body length 1.0 cm, wing length 8.0 mm. Most of left lateral view and some of dorsal, right lateral, and frontal view of face observable. Specimen apparently female. Head: Eyes bare, large, not dichoptic, no differentiation of facets nor apparent color patterns. Details of frons and face not entirely observable (e.g., presence of frontal callus and subcallus unlikely; development of ocelli not discernable). Antenna with scape and pedicel not observable but apparently short (not projected); flagellomere I apically narrowed to 0.5
<normalizedToken originalValue="×">x</normalizedToken>
basal width, with 3 faint annuli; remaining 6 flagellomeres stylate, tapered apicad, articles of approximately equal lengths [best seen in frontal view]. Proboscis robust, palps barely discernable (but apparently short, length 0.4
<normalizedToken originalValue="×">x</normalizedToken>
that of proboscis), labellum well developed; entire proboscis fairly long, length = 0.75
<normalizedToken originalValue="×">x</normalizedToken>
depth of head. Thorax: Standard proportions for
<taxonomicName family="Tabanidae" lsidName="" pageId="7" pageNumber="300" rank="family">Tabanidae</taxonomicName>
; legs without discernable spurs (although apices of hind tibiae not observable). Metathoracic spiracle also not observable [e.g., presence of postspiracular scale]. Wing: Completely hyaline, no patterning. Base of R2-5 nearly perpendicular to R1, not at a sharp, acute angle. Fork of R4-5 widely divergent and encompassing entire wing tip, base of R4 perpendicular to R5, then strongly and concavely curved to meet
<pageBreakToken pageId="8" pageNumber="301" start="start">C</pageBreakToken>
; base of R4 without a small appendix. M1, M2, M3 nearly parallel; M3 and CuA1 convergent (not parallel); CuA1 and A1 meeting just before wing margin. A2 extended nearly to wing margin; alula very large. Abdomen: Details (e.g., segmentation of cerci) not observable.
</paragraph>
<paragraph lastPageId="9" lastPageNumber="302" pageId="8" pageNumber="301">
Specimen. AMNH NJ-1081 (paratype): Thorax + abdomen length 8.2 mm, wing length 8.5 mm (from base of basicosta to wing tip). Wing: Basicosta present as a thick, scale-like lobe at base of vein C. C thickened proximally, circumambient. Short crossvein h present, where costal thickening is narrowed. Sc long, 0.6
<normalizedToken originalValue="×">x</normalizedToken>
length of wing, straight and parallel to vein C. Veins R and base of R1 also straight, parallel, and close to Sc; apices of Sc and R1 diverging apically. Dark, heavily sclerotized pterostigma covers and surrounds R1, vein C, and extends to tip of R2+3. R2+3 straight, turned slightly upward at apex. Stem of R4 and R5 straight, base of R4 nearly perpendicular to this stem, then curved upward and meeting C anterior to tip of wing; R5 nearly in line with stem of R4+R5. Cell d large, length ca. 2.7
<normalizedToken originalValue="×">x</normalizedToken>
the width; with veins M1, M2 and M3 each deriving directly from apical wall of cell. M veins slightly divergent, long; M1 slightly longer than cell d, M3 ca. 0.6
<normalizedToken originalValue="×">x</normalizedToken>
length of cell d. Crossveins r-m and m-cu in line with each other. Veins CuA2 and A1 meet slightly before wing margin, forming long,
<pageBreakToken pageId="9" pageNumber="302" start="start">complete</pageBreakToken>
cua cell with very short vein CuA2+A1. Vein A2 well developed, concave to A1, evanescent apically; anal lobe and anal cell well developed. Alula present but partially obscure. Abdomen: Short, broad, tergites short, typical of tabanids.
</paragraph>
<caption pageId="9" pageNumber="302">
<paragraph pageId="9" pageNumber="302">
Figure 4.
<taxonomicName class="Insecta" family="Tabanidae" genus="Cratotabanus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cratotabanus newjerseyensis" order="Diptera" pageId="9" pageNumber="302" phylum="Arthropoda" rank="species" species="newjerseyensis">Cratotabanus newjerseyensis</taxonomicName>
Grimaldi, sp. n. (
<taxonomicName family="Tabanidae" lsidName="" pageId="9" pageNumber="302" rank="family">Tabanidae</taxonomicName>
) in Turonian amber from New Jersey, USA. Above: Lateral view of holotype, AMNH 1862. (scale 1.0 mm). Below: Wing of paratype, AMNH NJ1081.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="9" pageNumber="302" type="types">
<paragraph pageId="9" pageNumber="302">Types.</paragraph>
<paragraph pageId="9" pageNumber="302">
Holotype (sex unknown), AMNH NJ-1862, New Jersey (USA): Middlesex Co., Sayreville, White Oaks [Old
<normalizedToken originalValue="Crossmans">Crossman's</normalizedToken>
] pits (Turonian), collected by Stephen Swolensky. Observation of the fly was optimized by embedding the amber in epoxy under vacuum and trimming very close to surfaces of the fly, but the specimen is not well preserved, being occluded with a reddish, crazed layer over most of the body and by similar internal fractures in the piece, as well as by a suspension of fine particles in the amber. Piece is irregular in shape, 10
<normalizedToken originalValue="×">x</normalizedToken>
13 mm in largest dimensions. Study of the specimen might benefit from microtomography.
</paragraph>
<paragraph pageId="9" pageNumber="302">
Paratype (sex unknown), AMNH NJ-1081, in Late Cretaceous (Turonian) amber from
<normalizedToken originalValue="Crossmans">Crossman's</normalizedToken>
Pits, Sayreville, New Jersey. Fly is partially preserved: besides the entire right wing and a very small portion of left wing, only the dorsal surfaces of the abdomen and thorax remain; the head and legs are entirely lost. The amber piece is triangular and approximately 19
<normalizedToken originalValue="×">x</normalizedToken>
8
<normalizedToken originalValue="×">x</normalizedToken>
5 mm, embedded in epoxy but trimmed and polished so as to expose a dorsal view of the fly. The amber itself is light yellow and turbid, with a thick suspension of organic particles that obscures much of the fly. AMNH NJ-1081 differs from NJ-1862 by the following minor venational details: R1 slightly longer, Rs branches from R1 at a more acute angle, proximal end of cell d slightly more shallow V-shaped; A2 slightly shorter. Both specimens are also very similar in body shape and size.
</paragraph>
</subSubSection>
<subSubSection pageId="9" pageNumber="302" type="etymology">
<paragraph pageId="9" pageNumber="302">Etymology.</paragraph>
<paragraph pageId="9" pageNumber="302">&quot;from New Jersey,&quot; in reference to provenance.</paragraph>
</subSubSection>
<subSubSection lastPageId="10" lastPageNumber="303" pageId="9" pageNumber="302" type="discussion">
<paragraph pageId="9" pageNumber="302">Discussion.</paragraph>
<paragraph pageId="9" pageNumber="302">
These are the only tabanids known to be preserved in Cretaceous amber. Other tabanids in amber are from the Miocene of the Dominican Republic and the Eocene Baltic amber (
<bibRefCitation pageId="9" pageNumber="302">Evenhuis 1994</bibRefCitation>
).
</paragraph>
<paragraph lastPageId="10" lastPageNumber="303" pageId="9" pageNumber="302">
<bibRefCitation author="Ren, D" journalOrPublisher="Acta Zootaxonomica Sinica" pageId="33" pageNumber="326" pagination="65 - 83" title="Late Jurassic Brachycera from northeastern China." volume="23" year="1998">Ren (1998)</bibRefCitation>
described three genera of putative, compression-fossilized tabanids with long proboscides from the Early Cretaceous Yixian Formation of China.
<bibRefCitation author="Grimaldi, D" journalOrPublisher="Annals of the Missouri Botanical Garden" pageId="31" pageNumber="324" pagination="373 - 406" title="The co-radiations of pollinating insects and angiosperms in the Cretaceous." url="doi: 10.2307/2666181" volume="86" year="1999">Grimaldi (1999)</bibRefCitation>
discussed the characters on which his assignment was made, and concluded that these fossils may not be tabanids. For example, features of
<taxonomicName class="Insecta" family="Athericidae" genus="Palaepangonius" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palaepangonius eupterus" order="Diptera" pageId="9" pageNumber="302" phylum="Arthropoda" rank="species" species="eupterus">Palaepangonius eupterus</taxonomicName>
Ren that are inconsistent with
<taxonomicName family="Tabanidae" lsidName="" pageId="9" pageNumber="302" rank="family">Tabanidae</taxonomicName>
are the short, upturned R2+3, very long veins R4 and R5 (half this length and much more divergent in true
<taxonomicName family="Tabanidae" lsidName="" pageId="9" pageNumber="302" rank="family">Tabanidae</taxonomicName>
), and veins A1 and CuA2 that do not fuse but meet the wing margin independently (Fig. 4c). These do appear to be tabanomorphs, but may be stem-group taxa to Recent
<taxonomicName family="Tabanidae" lsidName="" pageId="9" pageNumber="302" rank="family">Tabanidae</taxonomicName>
,
<taxonomicName family="Athericidae" lsidName="" pageId="9" pageNumber="302" rank="family">Athericidae</taxonomicName>
,
<taxonomicName family="Pelecorhynchidae" lsidName="" pageId="9" pageNumber="302" rank="family">Pelecorhynchidae</taxonomicName>
, and possibly even
<taxonomicName family="Rhagionidae" lsidName="" pageId="9" pageNumber="302" rank="family">Rhagionidae</taxonomicName>
(some fossil rhagionids had long, piercing mouthparts). Another early compression fossil,
<taxonomicName class="Insecta" family="Tabanidae" genus="Baissomyia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Baissomyia redita" order="Diptera" pageId="9" pageNumber="302" phylum="Arthropoda" rank="species" species="redita">Baissomyia redita</taxonomicName>
, from the Early Cretaceous Zaza Formation of Russia, was attributed to the
<taxonomicName family="Tabanidae" lsidName="" pageId="9" pageNumber="302" rank="family">Tabanidae</taxonomicName>
essentially on the basis of body shape and styletiform mouthparts (
<bibRefCitation author="Mostovski, MB" journalOrPublisher="Paleontological Journal" pageId="33" pageNumber="326" pagination="162 - 169" title="Horseflies and athericids (Diptera: Tabanidae, Athericidae) from the Lower Cretaceous of England and Transbaikalia." volume="37" year="2003">Mostovski et al. 2003</bibRefCitation>
), since the antennae and most of the wing (and, thus, most features defining the family) were not preserved.
<taxonomicName class="Insecta" family="Tabanidae" genus="Eotabanoid" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Eotabanoid lordi" order="Diptera" pageId="9" pageNumber="302" phylum="Arthropoda" rank="species" species="lordi">Eotabanoid lordi</taxonomicName>
, from the Early Cretaceous of England, is probably a tabanid, but it too plesiomorphically has long R4 and R5 veins, which are nearly symmetrial (in true tabanids R4 is typically much more curved) (
<bibRefCitation author="Mostovski, MB" journalOrPublisher="Paleontological Journal" pageId="33" pageNumber="326" pagination="162 - 169" title="Horseflies and athericids (Diptera: Tabanidae, Athericidae) from the Lower Cretaceous of England and Transbaikalia." volume="37" year="2003">Mostovski et al. 2003</bibRefCitation>
) (Fig. 4c). Besides the specimens in New Jersey amber described herein, the only
<pageBreakToken pageId="10" pageNumber="303" start="start">other</pageBreakToken>
definitive
<taxonomicName family="Tabanidae" lsidName="" pageId="10" pageNumber="303" rank="family">Tabanidae</taxonomicName>
from the Cretaceous is
<taxonomicName class="Insecta" family="Tabanidae" genus="Cratotabanus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cratotabanus stenomyomorphus" order="Diptera" pageId="10" pageNumber="303" phylum="Arthropoda" rank="species" species="stenomyomorphus">Cratotabanus stenomyomorphus</taxonomicName>
from the Aptian-aged Crato limestone of Brazil (
<bibRefCitation author="Martins-Neto, RG" journalOrPublisher="Acta Geologica Leopoldensia" pageId="32" pageNumber="325" pagination="289 - 297" title="Um novo genero e uma nova especie de Mutuca (Insecta, Diptera, Tabanidae) da Formacao Santana (Cretaceo Inferior). Bacia do Araripe, Nordeste do Brasil." volume="39" year="1994">Martins-Neto and Kucera-Santos 1994</bibRefCitation>
). Venation of
<taxonomicName class="Insecta" family="Tabanidae" genus="Cratotabanus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cratotabanus stenomyomorphus" order="Diptera" pageId="10" pageNumber="303" phylum="Arthropoda" rank="species" species="stenomyomorphus">Cratotabanus stenomyomorphus</taxonomicName>
and
<taxonomicName class="Insecta" family="Tabanidae" genus="Cratotabanus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cratotabanus newjerseyensis" order="Diptera" pageId="10" pageNumber="303" phylum="Arthropoda" rank="species" species="newjerseyensis">Cratotabanus newjerseyensis</taxonomicName>
are extremely similar. Another species of the genus from the Crato Formation is as yet undescribed (
<bibRefCitation author="Martins-Neto, RG" journalOrPublisher="Paleontological Journal" pageId="32" pageNumber="325" title="The fossil tabanids (Diptera Tabanidae): When they began to appreciate warm blood and why they began transmit [sic] diseases? Memorias Instituto Oswaldo Cruz 98 (suppl. 1): 29 - 34." url="doi: 10.1590/S0074-02762003000900006" year="2003">Martins-Neto 2003</bibRefCitation>
). Diverse
<taxonomicName family="Tabanidae" lsidName="" pageId="10" pageNumber="303" rank="family">Tabanidae</taxonomicName>
occur in Tertiary rocks and amber (summarized by Evenhuis, 1994), but generic assignments of those species described prior to 1950 need to be assessed. Cretaceous fossils assigned to the
<taxonomicName family="Tabanidae" lsidName="" pageId="10" pageNumber="303" rank="family">Tabanidae</taxonomicName>
include the following:
</paragraph>
<paragraph pageId="10" pageNumber="303">
<taxonomicName class="Insecta" family="Tabanidae" genus="Baissomyia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Baissomyia redita" order="Diptera" pageId="10" pageNumber="303" phylum="Arthropoda" rank="species" species="redita">Baissomyia redita</taxonomicName>
Mostovski, Jarzembowski &amp; Coram, 2003: Zaza Formation, Baissa, Transbaikalia, Russia.
</paragraph>
<paragraph pageId="10" pageNumber="303">
<taxonomicName class="Insecta" family="Tabanidae" genus="Eotabanoid" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Eotabanoid lordi" order="Diptera" pageId="10" pageNumber="303" phylum="Arthropoda" rank="species" species="lordi">Eotabanoid lordi</taxonomicName>
Mostovski, Jarzembowski &amp; Coram, 2003: Durlston Formation (Berriasian), Purbeck Group, Dorset UK.
</paragraph>
<paragraph pageId="10" pageNumber="303">
&quot;
<taxonomicName class="Insecta" family="Apataniidae" genus="Allomyia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Allomyia" order="Trichoptera" pageId="10" pageNumber="303" phylum="Arthropoda" rank="genus">Allomyia</taxonomicName>
&quot; [sensu Ren] ruderalis Ren, 1998: Yixian Formation, China.
</paragraph>
<paragraph pageId="10" pageNumber="303">
<taxonomicName class="Insecta" family="Tabanidae" genus="Eopangonius" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Eopangonius pletus" order="Diptera" pageId="10" pageNumber="303" phylum="Arthropoda" rank="species" species="pletus">Eopangonius pletus</taxonomicName>
Ren, 1998: Yixian Formation, China.
</paragraph>
<paragraph pageId="10" pageNumber="303">
<taxonomicName class="Insecta" family="Athericidae" genus="Palaepangonius" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palaepangonius eupterus" order="Diptera" pageId="10" pageNumber="303" phylum="Arthropoda" rank="species" species="eupterus">Palaepangonius eupterus</taxonomicName>
Ren, 1998: Yixian Formation, China.
</paragraph>
<paragraph pageId="10" pageNumber="303">
<taxonomicName class="Insecta" family="Tabanidae" genus="Cratotabanus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cratotabanus stenomyomorphus" order="Diptera" pageId="10" pageNumber="303" phylum="Arthropoda" rank="species" species="stenomyomorphus">Cratotabanus stenomyomorphus</taxonomicName>
Martins-Neto &amp; Santos, 1994: Crato Formation (Aptian), Ceara, Brazil.
</paragraph>
<paragraph pageId="10" pageNumber="303">
&quot;
<taxonomicName class="Insecta" family="Tabanidae" genus="Cratotabanus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cratotabanus" order="Diptera" pageId="10" pageNumber="303" phylum="Arthropoda" rank="genus">Cratotabanus</taxonomicName>
sp. n.&quot;: Crato Formation (Aptian), Ceara, Brazil (in
<bibRefCitation author="Martins-Neto, RG" journalOrPublisher="Paleontological Journal" pageId="32" pageNumber="325" title="The fossil tabanids (Diptera Tabanidae): When they began to appreciate warm blood and why they began transmit [sic] diseases? Memorias Instituto Oswaldo Cruz 98 (suppl. 1): 29 - 34." url="doi: 10.1590/S0074-02762003000900006" year="2003">Martins-Neto 2003</bibRefCitation>
: pg. 31, ex: Grimaldi 1990).
</paragraph>
<paragraph pageId="10" pageNumber="303">
<taxonomicName class="Insecta" family="Tabanidae" genus="Cratotabanus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cratotabanus newjerseyensis" order="Diptera" pageId="10" pageNumber="303" phylum="Arthropoda" rank="species" species="newjerseyensis">Cratotabanus newjerseyensis</taxonomicName>
sp.n.: Raritan Formation amber (Turonian), New Jersey, USA (herein).
</paragraph>
</subSubSection>
</treatment>
</document>