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<mods:title id="02960914F76F50A3183AE15A0DFEDE7E">Review of Neotropical Chonocephalus Wandolleck (Diptera: Phoridae)</mods:title>
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Genus
<taxonomicName id="8CC14D778727FF8AADEDFD21FD79FD51" ID-CoL="3P2P" ID-ENA="92528" authority="Wandolleck, 1898" authorityName="Wandolleck" authorityYear="1898" box="[238,659,741,767]" class="Insecta" family="Phoridae" genus="Chonocephalus" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">
Chonocephalus
<bibRefCitation id="2F504B058727FF8AACB6FD21FD79FD51" author="Wandolleck, B." box="[437,659,741,767]" pageId="2" pageNumber="3" pagination="412 - 441" refId="ref19679" refString="Wandolleck, B. (1898) Die Stenopathidae, eine neue fluegel- und schwingerlose Familie der Diptera, Zoologische Jahrbucher Abteilung fur Systematik, 11, 412 - 441." type="journal article" year="1898">Wandolleck, 1898</bibRefCitation>
</taxonomicName>
</emphasis>
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</paragraph>
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<paragraph id="4B7E36F48727FF8AAD94FCEEFC91FCEA" blockId="2.[151,1436,810,2036]" box="[151,891,810,836]" pageId="2" pageNumber="3">
<emphasis id="79B5EAE68727FF8AAD94FCEEFED3FCEA" bold="true" box="[151,313,810,836]" pageId="2" pageNumber="3">
<typeStatus id="947A88568727FF8AAD94FCEEFF3EFCEA" box="[151,212,810,836]" pageId="2" pageNumber="3">Type</typeStatus>
species.
</emphasis>
<taxonomicName id="8CC14D778727FF8AAC43FCEEFC9CFCEA" ID-CoL="5YB4J" authority="Wandolleck, 1898: 428" authorityName="Wandolleck" authorityPageNumber="428" authorityYear="1898" box="[320,886,810,836]" class="Insecta" family="Phoridae" genus="Chonocephalus" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="species" species="dorsalis">
<emphasis id="79B5EAE68727FF8AAC43FCEEFDB6FCED" box="[320,604,810,835]" italics="true" pageId="2" pageNumber="3">Chonocephalus dorsalis</emphasis>
<bibRefCitation id="2F504B058727FF8AAF67FCEEFC9CFCEA" author="Wandolleck, B." box="[612,886,810,836]" pageId="2" pageNumber="3" pagination="412 - 441" refId="ref19679" refString="Wandolleck, B. (1898) Die Stenopathidae, eine neue fluegel- und schwingerlose Familie der Diptera, Zoologische Jahrbucher Abteilung fur Systematik, 11, 412 - 441." type="journal article" year="1898">Wandolleck, 1898: 428</bibRefCitation>
</taxonomicName>
.
</paragraph>
<paragraph id="4B7E36F48727FF8AADC5FC96FBBCFC4A" blockId="2.[151,1436,810,2036]" pageId="2" pageNumber="3">
The genus is readily identified in the most recent key to world genera (
<bibRefCitation id="2F504B058727FF8AA90FFC96FB46FCC2" author="Disney, R. H. L." box="[1036,1196,850,876]" pageId="2" pageNumber="3" refId="ref19271" refString="Disney, R. H. L. (1994) Scuttle Flies: The Phoridae (Chapman and Hall, London), xii + 467 pp." type="book" year="1994">Disney, 1994</bibRefCitation>
), with the modification that the genus
<taxonomicName id="8CC14D778727FF8AAC82FCBEFD2EFC3A" ID-CoL="4CYH" authority="Schmitz" authorityName="Schmitz" authorityYear="1927" box="[385,708,890,916]" class="Insecta" family="Phoridae" genus="Epichonocephalus" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="79B5EAE68727FF8AAC82FCBEFDB3FC3D" box="[385,601,890,915]" italics="true" pageId="2" pageNumber="3">Epichonocephalus</emphasis>
Schmitz
</taxonomicName>
has been synonymised with
<taxonomicName id="8CC14D778727FF8AA927FCBEFA72FC3A" ID-CoL="3P2P" authority="(Disney, 2002)" baseAuthorityName="Disney" baseAuthorityYear="2002" box="[1060,1432,890,916]" class="Insecta" family="Phoridae" genus="Chonocephalus" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="79B5EAE68727FF8AA927FCBEFB30FC3D" box="[1060,1242,890,915]" italics="true" pageId="2" pageNumber="3">Chonocephalus</emphasis>
(
<bibRefCitation id="2F504B058727FF8AA9EFFCBEFA65FC3A" author="Disney, R. H. L." box="[1260,1423,890,916]" pageId="2" pageNumber="3" pagination="1 - 36" refId="ref19357" refString="Disney, R. H. L. (2002) Revisionary notes on Chonocephalus Wandolleck (Diptera: Phoridae) with keys to species of the Holarctic Region, Zootaxa, 60, 1 - 36." type="journal article" year="2002">Disney, 2002</bibRefCitation>
)
</taxonomicName>
. The same paper provides a detailed diagnosis of the genus in both sexes. The wings of the males are distinctive (
<figureCitation id="D3FA2A718727FF8AADD1FC0EFEC1FC4A" box="[210,299,970,996]" captionStart="FIGURES 1518" captionStartId="8.[151,269,1544,1568]" captionTargetBox="[161,1421,225,1482]" captionTargetId="figure-146@8.[151,1436,194,1592]" captionTargetPageId="8" captionText="FIGURES 1518. Chonocephalus aripoensis: (15) male, right lobe of hypandrium; (16) male, left face of hypopygium; (17) female, abdominal sternite 8 and spermatheca; (18) male, right wing. Scale bars = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5123995" httpUri="https://zenodo.org/record/5123995/files/figure.png" pageId="2" pageNumber="3">Figs 18</figureCitation>
and
<figureCitation id="D3FA2A718727FF8AAC67FC0EFE69FC4A" box="[356,387,970,996]" captionStart="FIGURES 6465" captionStartId="27.[151,269,1318,1342]" captionTargetBox="[214,1349,195,1290]" captionTargetId="figure-181@27.[151,1436,134,1340]" captionTargetPageId="27" captionText="FIGURES 6465. Chonocephalus heymonsi: (64) male, right wing; (65) female, posterior face of hind leg from femur onwards. Scale bars = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5124047" httpUri="https://zenodo.org/record/5124047/files/figure.png" pageId="2" pageNumber="3">64</figureCitation>
) as is the general appearance of the females (
<figureCitation id="D3FA2A718727FF8AAE95FC0EFC04FC4A" box="[918,1006,970,996]" captionStart="FIGURES 4748" captionStartId="19.[151,269,1258,1282]" captionTargetBox="[390,1229,186,1210]" captionTargetId="figure-159@19.[340,1248,176,1252]" captionTargetPageId="19" captionText="FIGURES 4748. Chonocephalus dominicanus female: (47) whole fly from above; (48) sternite 8, furca and spermatheca. Scale bars = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5124027" httpUri="https://zenodo.org/record/5124027/files/figure.png" pageId="2" pageNumber="3">Figs 47</figureCitation>
and
<figureCitation id="D3FA2A718727FF8AA924FC0EFBA2FC4A" box="[1063,1096,970,996]" captionStart="FIGURES 6668" captionStartId="28.[151,269,1629,1653]" captionTargetBox="[151,1387,401,1570]" captionTargetId="figure-152@28.[151,1436,340,1605]" captionTargetPageId="28" captionText="FIGURES 6668. Chonocephalus heymonsi female: (66) whole fly from above; (67) tergite 8; (68) sternite 8, furca and spermatheca. Scale bars = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5124051" httpUri="https://zenodo.org/record/5124051/files/figure.png" pageId="2" pageNumber="3">66</figureCitation>
).
</paragraph>
<paragraph id="4B7E36F48727FF8AADC5FC36FF34FB9A" blockId="2.[151,1436,810,2036]" pageId="2" pageNumber="3">
The full details of the references for synonyms, etc., covered by
<bibRefCitation id="2F504B058727FF8AAED0FC36FB94FBA2" author="Disney, R. H. L." box="[979,1150,1010,1036]" pageId="2" pageNumber="3" pagination="1 - 36" refId="ref19357" refString="Disney, R. H. L. (2002) Revisionary notes on Chonocephalus Wandolleck (Diptera: Phoridae) with keys to species of the Holarctic Region, Zootaxa, 60, 1 - 36." type="journal article" year="2002">Disney (2002)</bibRefCitation>
are not repeated in this paper.
</paragraph>
<paragraph id="4B7E36F48727FF8AADC5FB86FBC0F91A" blockId="2.[151,1436,810,2036]" pageId="2" pageNumber="3">
Males. The recognition of species depends primarily on the details of the hypopygium, which is at first sight very complex. However, if one ignores the, often elaborate, penis complex (X), the details of the epandrium (E) and hypandrium (H) that are referred to in the key below are indicated in
<figureCitation id="D3FA2A718727FF8AA97DFB56FB2FFB02" box="[1150,1221,1170,1196]" captionStart="FIGURE 1" captionStartId="3.[151,255,1295,1319]" captionTargetBox="[294,1247,275,1259]" captionTargetId="figure-256@3.[151,1436,256,1290]" captionTargetPageId="3" captionText="FIGURE 1. Chonocephalus male hypopygium, diagrammatic ventral view of a generalised species. A = anal tube (with median hypoproct and paired cerci); E = epandrium; H = hypandrium; X = penis complex (simplified). AP, PP = anterior and posterior processes (probably = subepandrial sclerites) of epandrium; L = lobe(s) of hypandrium; G = gonopods; B = bridge (probably = sternite 10); C = left surstylus (clasper); S = spine (modified bristle) of epandrium." figureDoi="http://doi.org/10.5281/zenodo.5123979" httpUri="https://zenodo.org/record/5123979/files/figure.png" pageId="2" pageNumber="3">Fig. 1</figureCitation>
. This represents a generalised, stylized, hypopygium. While the anal tube (A) is seldom utilised in the recognition of species it is a useful point of reference. Any one of the labelled structures may be modified and may sometimes be very elaborate in form. In the inferred groundplan state the rear margin of the hypandrium bears four processes. These are the gonopods (G) and a pair of median lobes (L). In the plesiomorphic state the gonopods articulate with the dorsal face of the hypandrium, but more frequently they arise from it. One (usually the left) or both may be reduced or absent. The median lobes (L) may form a single bilobed structure or be reduced to a single lobe (which may be greatly elongated) or be lost altogether. Ventrally each side of the epandrium (E) typically bears an anterior process (AP) and a posterior process (PP). In some species the anterior processes (AP) may appear to be appendages (attaching to the inner face of the epandrium) rather than simple projections of the epandrium. A presumed true left surstylus or clasper (C) is frequently present (e.g.
<figureCitation id="D3FA2A718727FF8AA971F9E6FB20F992" box="[1138,1226,1570,1596]" captionStart="FIGURES 1518" captionStartId="8.[151,269,1544,1568]" captionTargetBox="[161,1421,225,1482]" captionTargetId="figure-146@8.[151,1436,194,1592]" captionTargetPageId="8" captionText="FIGURES 1518. Chonocephalus aripoensis: (15) male, right lobe of hypandrium; (16) male, left face of hypopygium; (17) female, abdominal sternite 8 and spermatheca; (18) male, right wing. Scale bars = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5123995" httpUri="https://zenodo.org/record/5123995/files/figure.png" pageId="2" pageNumber="3">Figs 17</figureCitation>
and
<figureCitation id="D3FA2A718727FF8AA806F9E6FACCF992" box="[1285,1318,1570,1596]" captionStart="FIGURES 24 26" captionStartId="10.[151,269,1702,1726]" captionTargetBox="[161,1349,195,1638]" captionTargetId="figure-71@10.[151,1436,164,1712]" captionTargetPageId="10" captionText="FIGURES 24 26. Chonocephalus browni male, details of hypopygium: (24) left side of epandrium from below; (25) right side of epandrium and right gonopod from below; (26) hypandrium and gonopods from left side. Scale bar = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5124001" httpUri="https://zenodo.org/record/5124001/files/figure.png" pageId="2" pageNumber="3">24</figureCitation>
), variably fused to the epandrium, and is sometimes elaborate. Occasionally it is fused to the bridge (B), which is sometimes present and probably represents sternite 10. The most anterior (lower) bristle of the left side of the epandrium is sometimes modified into a spine (S), which may be further modified.
</paragraph>
<paragraph id="4B7E36F48727FF8AADC5F906FE3CF8FA" blockId="2.[151,1436,810,2036]" pageId="2" pageNumber="3">
Females. The critical details of the terminal segments of the abdomen are indicated in
<figureCitation id="D3FA2A718727FF8AA9F2F906FA7DF972" box="[1265,1431,1730,1756]" captionStart="FIGURES 109 110" captionStartId="43.[151,269,1344,1368]" captionTargetBox="[151,1337,444,1303]" captionTargetId="figure-295@43.[151,1436,434,1320]" captionTargetPageId="43" captionText="FIGURES 109 110. Details of the terminal segments of the abdomen of the basic (presumed groundplan) state of a female Chonocephalus (the Oriental C. blackithorum Disney, 1986) from above (109) and from below (110). C = cercus, H = hypoproct, Sp = spermatheca, S8 = sternite 8, S9 = the internalised sternite 9 (= furca), T7T10 = tergites 7 to 10. Scale bar = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5124099" httpUri="https://zenodo.org/record/5124099/files/figure.png" pageId="2" pageNumber="3">Figs 109110</figureCitation>
, which depicts a basic (close to the groundplan) state of a female
<taxonomicName id="8CC14D778727FF8AAE93F92FFB15F8AA" ID-CoL="3P2P" authority="(Disney, 1986)" baseAuthorityName="Disney" baseAuthorityYear="1986" box="[912,1279,1770,1796]" class="Insecta" family="Phoridae" genus="Chonocephalus" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="79B5EAE68727FF8AAE93F92FFBACF8AA" box="[912,1094,1771,1796]" italics="true" pageId="2" pageNumber="3">Chonocephalus</emphasis>
(
<bibRefCitation id="2F504B058727FF8AA955F92EFB1CF8AA" author="Disney, R. H. L." box="[1110,1270,1770,1796]" pageId="2" pageNumber="3" pagination="77 - 87" refId="ref19226" refString="Disney, R. H. L. (1986) Morphological and other observations on Chonocephalus (Phoridae) and phylogenetic implications for the Cyclorrhapha (Diptera), Journal of Zoology, London (A), 210, 77 - 87." type="journal article" year="1986">Disney, 1986</bibRefCitation>
)
</taxonomicName>
. In most species the T9 is lost apart from its anterolateral apodemes, which are fused to those of T10 so that the latter appear to be two-branched.
</paragraph>
<paragraph id="4B7E36F48727FF8AADC5F8A6FC27F85A" blockId="2.[151,1436,810,2036]" pageId="2" pageNumber="3">
<emphasis id="79B5EAE68727FF8AADC5F8A6FE43F8D2" bold="true" box="[198,425,1890,1916]" pageId="2" pageNumber="3">Review of species.</emphasis>
The descriptions of the new species are brief as the significant features of the males are largely restricted to the details of the hypopygia. Likewise in the females there are relatively few features that distinguish the species apart from the details of the abdominal terminalia. The formal diagnoses, therefore, are the features used to distinguish the species in the keys above.
</paragraph>
<paragraph id="4B7E36F48726FF8BADC5FF5CFBC7FF74" blockId="3.[151,1436,152,218]" pageId="3" pageNumber="4">
The brief notes on the natural histories of the species, when known, are taken from
<bibRefCitation id="2F504B058726FF8BA99CFF5CFAADFF1C" author="Disney, R. H. L." box="[1183,1351,152,178]" pageId="3" pageNumber="4" pagination="1 - 36" refId="ref19357" refString="Disney, R. H. L. (2002) Revisionary notes on Chonocephalus Wandolleck (Diptera: Phoridae) with keys to species of the Holarctic Region, Zootaxa, 60, 1 - 36." type="journal article" year="2002">Disney (2002)</bibRefCitation>
except where indicated otherwise. Typical larvae and pupae are shown in
<figureCitation id="D3FA2A718726FF8BAEA1FF04FBC3FF74" box="[930,1065,192,218]" captionStart-0="FIGURES 7071" captionStart-1="FIGURES 7274" captionStartId-0="30.[151,269,784,808]" captionStartId-1="30.[151,269,1725,1749]" captionTargetBox-0="[182,1400,178,688]" captionTargetBox-1="[223,1380,866,1713]" captionTargetId-0="figure-68@30.[151,1436,126,760]" captionTargetId-1="figure-71@30.[151,1436,812,1740]" captionTargetPageId-0="30" captionTargetPageId-1="30" captionText-0="FIGURES 7071. Chonocephalus heymonsi larvae: (70) first instar; (71) third instar. (After Borgmeier, 1935)." captionText-1="FIGURES 7274. Chonocephalus heymonsi pupae: (72) eclosion plate of puparium; (73) anterior cap of puparium; (74) left face of pupa. (After Borgmeier, 1935)." figureDoi-0="http://doi.org/10.5281/zenodo.5124055" figureDoi-1="http://doi.org/10.5281/zenodo.5124059" httpUri-0="https://zenodo.org/record/5124055/files/figure.png" httpUri-1="https://zenodo.org/record/5124059/files/figure.png" pageId="3" pageNumber="4">Figs 7074</figureCitation>
.
</paragraph>
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