treatments-xml/data/03/AF/87/03AF87F1F424FFD0FF10FAFCFB427AA4.xml

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<mods:title id="4FFDABA3D4C29A785058630219B8A44E">Drosophila (Sophophora) carrolli n. sp., a new species from Brunei, closely related to Drosophila (Sophophora) rhopaloa Bock &amp; Wheeler, 1972 (Diptera: Drosophilidae)</mods:title>
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<subSubSection id="C31C656CF424FFD7FF10FAFCFC817CB2" pageId="1" pageNumber="503" type="nomenclature">
<paragraph id="8BB936E7F424FFD7FF10FAFCFCB47C93" blockId="1.[151,890,1399,1459]" box="[151,847,1399,1426]" pageId="1" pageNumber="503">
<heading id="D0F1818BF424FFD7FF10FAFCFCB47C93" bold="true" box="[151,847,1399,1426]" fontSize="11" level="1" pageId="1" pageNumber="503" reason="1">
<emphasis id="B972EAF5F424FFD7FF10FAFCFCB47C93" bold="true" box="[151,847,1399,1426]" pageId="1" pageNumber="503">
<taxonomicName id="4C064D64F424FFD7FF10FAFCFCFE7C93" ID-CoL="9HHZY" authority="Gompel &amp; Kopp" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[151,773,1399,1426]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="1" pageNumber="503" phylum="Arthropoda" rank="species" species="carrolli" status="sp. nov." subGenus="Sophophora">
<emphasis id="B972EAF5F424FFD7FF10FAFCFDD57C90" bold="true" box="[151,558,1399,1425]" italics="true" pageId="1" pageNumber="503">Drosophila (Sophophora) carrolli</emphasis>
Gompel &amp; Kopp
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<taxonomicNameLabel id="A241578EF424FFD7FC8AFAF3FCB47C93" box="[781,847,1400,1426]" pageId="1" pageNumber="503" rank="species">n. sp.</taxonomicNameLabel>
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<paragraph id="8BB936E7F424FFD7FF10FA11FC817CB2" blockId="1.[151,890,1399,1459]" box="[151,890,1434,1459]" pageId="1" pageNumber="503">
(
<figureCitation id="133D2A62F424FFD7FF19FA11FE2F7CB3" box="[158,468,1434,1459]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="1" pageNumber="503">Figure 1A, B, E, G, I, K, M</figureCitation>
, 
<figureCitation id="133D2A62F424FFD7FE58FA11FD4F7CB2" box="[479,692,1434,1459]" captionStart="FIGURE 2" captionStartId="4.[151,250,1815,1837]" captionTargetBox="[253,1318,204,1780]" captionTargetId="figure@4.[235,1349,193,1794]" captionTargetPageId="4" captionText="FIGURE 2. Male terminalia. Dissected parts from D. carrolli (A, C, E, G) or D. rhopaloa (B, D, F, H). (A, B) epandrium in dorsal view. Note the differences in sensory bristle length and distribution (blue ovals and arrowheads) as well as genital comb size (blue arrows) between species. The epandrium is always darker overall in D. carrolli. Phallus in ventral view (C, D) and side view (E, F). The phallus conformation in D. carrolli (C, E) is similar to that of D. rhopaloa (D, F), but shows quantitative difference in size and appears sturdier. The looping of the aedeagus base is more open in D. carrolli than in D. rhopaloa (blue double-arrows). (G, H), testes of D. carrolli (G) and D. rhopaloa (H) have similar shapes and coiling, but they appear bigger in D. carrolli, and show a characteristic bright yellow color compared to the dull yellow testes of D. rhopaloa." httpUri="https://zenodo.org/record/1292136/files/figure.png" pageId="1" pageNumber="503">Figure 2A, C, E, G</figureCitation>
, 
<figureCitation id="133D2A62F424FFD7FD38FA11FC897CB2" box="[703,882,1434,1459]" captionStart="FIGURE 3" captionStartId="5.[151,250,1765,1787]" captionTargetBox="[266,1327,198,1733]" captionTargetId="figure@5.[253,1341,193,1748]" captionTargetPageId="5" captionText="FIGURE 3. Female terminalia, eggs and pupae. (A, A’, B) last abdominal segments and spermatheca of D. carrolli (A, A’) or D. rhopaloa (B) females. Note the strongly sclerotized plates (blue arrowheads) at the end of the oviduct in D. carrolli (A, A’), but not D. rhopaloa (B). Also note the teeth (bristles) on the egg-guides are slightly more numerous and stouter in D. carrolli (blue arrows). The rear view of a D. carrolli female laying an egg (A’) shows the relative position of the sclerotized plates, the anal plates and the ovipositor. The spermathecae are marginally bigger and darker in age-matched D. carrolli compared to D. rhopaloa. (C, D) eggs of D. carrolli (C) and D. rhopaloa (D) in ventral view. Both species show characteristic spatula-shape respiratory appendages. These appendages vary in shape within species but show no notable difference between species. (E, F) pupae of D. carrolli (A) or D. rhopaloa (B) males about 24 hours before adult emergence. D. carrolli pupae are more rounded on their sides." httpUri="https://zenodo.org/record/1292138/files/figure.png" pageId="1" pageNumber="503">Figure 3A, C, E</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C31C656CF424FFD7FF10FA69FCDD7F1E" pageId="1" pageNumber="503" type="etymology">
<paragraph id="8BB936E7F424FFD7FF10FA69FCDD7F1E" blockId="1.[151,1437,1506,2038]" pageId="1" pageNumber="503">
<emphasis id="B972EAF5F424FFD7FF10FA69FEE57CFA" bold="true" box="[151,286,1506,1531]" pageId="1" pageNumber="503">Etymology.</emphasis>
The species is named after our former mentor, Sean B. Carroll, in appreciation of his inspiring approach to science in general, and evolution in particular.
</paragraph>
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<subSubSection id="C31C656CF424FFD7FF40F9A1FEE87FD2" pageId="1" pageNumber="503" type="diagnosis">
<paragraph id="8BB936E7F424FFD7FF40F9A1FEE87FD2" blockId="1.[151,1437,1506,2038]" pageId="1" pageNumber="503">
<emphasis id="B972EAF5F424FFD7FF40F9A1FEBA7F42" bold="true" box="[199,321,1578,1603]" pageId="1" pageNumber="503">Diagnosis.</emphasis>
This species resembles 
<taxonomicName id="4C064D64F424FFD7FDD3F9A0FB0E7F42" authority="Bock &amp; Wheeler, 1972" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[596,1269,1578,1603]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="1" pageNumber="503" phylum="Arthropoda" rank="species" species="rhopaloa" subGenus="Sophophora">
<emphasis id="B972EAF5F424FFD7FDD3F9A0FC1F7F42" box="[596,996,1579,1603]" italics="true" pageId="1" pageNumber="503">Drosophila (Sophophora) rhopaloa</emphasis>
<bibRefCitation id="EF974B16F424FFD7FC6BF9A1FB0E7F42" box="[1004,1269,1578,1603]" pageId="1" pageNumber="503" refId="ref3963" refString="Bock, I. R. &amp; Wheeler, M. R. (1972) The Drosophila melanogaster species group. Studies in Genetics VII. University of Texas Publications, 7, 1 - 102." type="journal article">Bock &amp; Wheeler, 1972</bibRefCitation>
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, from which it differs by its darker overall color in male and female adults, a larger size, a region of gray pigmentation at the anterior distal tip of the male wing, the paramere as long as the inner, sclerotized process of the hypandrium, a darker and brighter color of the male testes, and strongly sclerotized plates between the anal plates and the oviscape in females.
</paragraph>
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<subSubSection id="C31C656CF424FFD4FF40F955FE2B7F91" lastPageId="2" lastPageNumber="504" pageId="1" pageNumber="503" type="description">
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<emphasis id="B972EAF5F424FFD7FF40F955FEA27FF6" bold="true" box="[199,345,1758,1783]" pageId="1" pageNumber="503">Description.</emphasis>
Adult dark, total length, from abdominal tip to frons: 2.6± 
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mm (male, n=10), 2.8± 
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(female, n=11).
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Head. Dark brown. Antennae yellowish-dark, 3rd antennal segment ovoid (
<figureCitation id="133D2A62F424FFD7FB81F8A2FB877E43" box="[1030,1148,1833,1858]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="1" pageNumber="503">Figure 1K</figureCitation>
), distinctly longer than in 
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<emphasis id="B972EAF5F424FFD7FF10F8C5FEDB7E64" box="[151,288,1869,1893]" italics="true" pageId="1" pageNumber="503">D. rhopaloa</emphasis>
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(
<figureCitation id="133D2A62F424FFD7FEB7F8C6FE5F7E67" box="[304,420,1869,1894]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="1" pageNumber="503">Figure 1L</figureCitation>
). Arista with 4–5 dorsal and 2 ventral branches, plus a terminal fork. Ocellar triangle and region of orbital setae darker, eyes bright red. Bristle pattern and lengths identical to those of 
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<emphasis id="B972EAF5F424FFD7FB3BF8F9FABE7E88" box="[1212,1349,1905,1929]" italics="true" pageId="1" pageNumber="503">D. rhopaloa</emphasis>
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, with 3 pairs of orbital setae (anterior proclinate, median and posterior reclinate), 3 pairs of vertical setae (anterior and posterior convergent, median reclinate), one pair of ocellar (proclinate, divergent) and one pair of postocellar (reclinate, convergent) setae. Palpus with 2 prominent setae apically and laterally.
</paragraph>
<paragraph id="8BB936E7F427FFD4FF40FF1CFAF27B3D" blockId="2.[151,1437,151,2002]" pageId="2" pageNumber="504">
Thorax. Width of mesonotum at the level of wing hinge: 0.9±0.0 mm (male, n=10), 1.0±0.0 mm (female, n=11). Length of mesonotum: 1.2± 
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(male, n=10), 1.2± 
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(female, n=11). Brown with a pattern of darker markings on mesonotum disc: bilateral patches just anterior to scutellum; bilateral bands extending from wing hinges to intrascutal folds (but not reaching them); prescutal and intrascutal folds as well as scutellum darker. By contrast, 
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has a uniformly light thorax. Mesonotum with eight rows of acrostichal setulae between dorso-central bristles. Legs brown, with apex of femora darker, unlike in 
<taxonomicName id="4C064D64F427FFD4FC5BFEC0FB9D7862" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[988,1126,331,355]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FC5BFEC0FB9D7862" box="[988,1126,331,355]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
where the legs are entirely yellow. Male foretibiae with long, stout, preapical dorsal seta, and foretarsi with 2 rows of sex combs on 1 
<taxonomicName id="4C064D64F427FFD4FABAFEE6FA8C788B" box="[1341,1399,365,394]" pageId="2" pageNumber="504" rank="subSpecies" subSpecies="and">st and</taxonomicName>
2nd tarsomeres, similar to those of 
<taxonomicName id="4C064D64F427FFD4FE7FFE1DFD7978AC" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[504,642,405,429]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FE7FFE1DFD7978AC" box="[504,642,405,429]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
with typically 10–12 teeth per comb (
<figureCitation id="133D2A62F427FFD4FBB0FE1FFB2A78AC" box="[1079,1233,404,429]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="2" pageNumber="504">Figure 1M,N</figureCitation>
). Wing hyaline in both sexes, with a patch of gray pigmentation in males, extending from the anterior margin to the compartment between veins L3 and L4, and from the wing tip to the middle of the wing. This patch becomes distinct 24 hours after emergence and is absent from, or faint in 
<taxonomicName id="4C064D64F427FFD4FD22FD8AFCD57B19" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[677,814,512,536]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FD22FD8AFCD57B19" box="[677,814,512,536]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
males (
<figureCitation id="133D2A62F427FFD4FC02FD8BFBF07B19" box="[901,1035,512,537]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="2" pageNumber="504">
Figure 
<date id="FFB81027F427FFD4FC5FFD8BFC0B7B19" box="[984,1008,512,536]" pageId="2" pageNumber="504">1I</date>
, J
</figureCitation>
; 
<bibRefCitation id="EF974B16F427FFD4FB90FD8BFAE97B19" box="[1047,1298,512,537]" pageId="2" pageNumber="504" refId="ref4665" refString="Setoguchi, S., Takamori, H., Aotsuka, T., Sese, J., Ishikawa, Y. &amp; Matsuo, T. (2014) Sexual dimorphism and courtship behavior in Drosophila prolongata. J Ethology, 32, 91 - 102. https: // doi. org / 10.1007 / s 10164 - 014 - 0399 - z" type="journal article">
Setoguchi 
<emphasis id="B972EAF5F427FFD4FB09FD8AFB447B19" box="[1166,1215,512,536]" italics="true" pageId="2" pageNumber="504">et al</emphasis>
., 2014
</bibRefCitation>
). 
<taxonomicName id="4C064D64F427FFD4FAA2FD8AFA677B19" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[1317,1436,512,536]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F427FFD4FAA2FD8AFA677B19" box="[1317,1436,512,536]" italics="true" pageId="2" pageNumber="504">D. carrolli</emphasis>
</taxonomicName>
females occasionally show a faint trace of darker pigmentation between veins L1 and L2 (
<figureCitation id="133D2A62F427FFD4FB01FDA8FB007B3D" box="[1158,1275,547,572]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="2" pageNumber="504">Figure 1B</figureCitation>
).
</paragraph>
<paragraph id="8BB936E7F427FFD4FF40FDCCFAE97A13" blockId="2.[151,1437,151,2002]" pageId="2" pageNumber="504">
Abdomen. Brown in males, where the darker pigmentation of tergites 5 and 6 typically found in males of many 
<taxonomicName id="4C064D64F427FFD4FF10FDE0FED47B82" box="[151,303,619,643]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="melanogaster">
<emphasis id="B972EAF5F427FFD4FF10FDE0FED47B82" box="[151,303,619,643]" italics="true" pageId="2" pageNumber="504">melanogaster</emphasis>
</taxonomicName>
group species is hardly distinguishable from the pigmentation of anterior segments (see variation in 
<figureCitation id="133D2A62F427FFD4FF10FD05FEF37BA6" box="[151,264,654,679]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="2" pageNumber="504">Figure 1E</figureCitation>
). By contrast, 
<taxonomicName id="4C064D64F427FFD4FE2BFD1BFDCE7BA6" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[428,565,655,679]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FE2BFD1BFDCE7BA6" box="[428,565,655,679]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
males are lightly pigmented on their tergites, except for a broad medial black band on segments 5 and 6, sharply interrupted laterally before sternopleural suture (
<figureCitation id="133D2A62F427FFD4FBDAFD39FB447BCA" box="[1117,1215,690,715]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="2" pageNumber="504">Figure 1</figureCitation>
). Female abdomen dark-brown, with stripes of darker pigmentation of variable width on each segment (
<figureCitation id="133D2A62F427FFD4FBFBFD5DFAE67BEE" box="[1148,1309,726,751]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="2" pageNumber="504">Figure 1B, G</figureCitation>
). As in 
<taxonomicName id="4C064D64F427FFD4FA06FD5CFF067A13" authorityName="Bock &amp; Wheeler" authorityYear="1972" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FA06FD5CFF067A13" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
females, the overall abdominal pigmentation fades on the posterior segments (
<figureCitation id="133D2A62F427FFD4FBE8FD71FAFE7A13" box="[1135,1285,762,787]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="2" pageNumber="504">Figure 1G, H</figureCitation>
).
</paragraph>
<paragraph id="8BB936E7F427FFD4FF40FC96FBE47DE2" blockId="2.[151,1437,151,2002]" pageId="2" pageNumber="504">
Male terminalia. Similar conformation as in 
<taxonomicName id="4C064D64F427FFD4FD5DFC94FC937A37" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[730,872,798,822]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FD5DFC94FC937A37" box="[730,872,798,822]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
, but overall darker and bigger. Genital arch of epandrium with approximately 2 setae near the posterior margin above the insertion of surstylus (primary clasper) (
<figureCitation id="133D2A62F427FFD4FF19FCEEFEEC7A7F" box="[158,279,869,894]" captionStart="FIGURE 2" captionStartId="4.[151,250,1815,1837]" captionTargetBox="[253,1318,204,1780]" captionTargetId="figure@4.[235,1349,193,1794]" captionTargetPageId="4" captionText="FIGURE 2. Male terminalia. Dissected parts from D. carrolli (A, C, E, G) or D. rhopaloa (B, D, F, H). (A, B) epandrium in dorsal view. Note the differences in sensory bristle length and distribution (blue ovals and arrowheads) as well as genital comb size (blue arrows) between species. The epandrium is always darker overall in D. carrolli. Phallus in ventral view (C, D) and side view (E, F). The phallus conformation in D. carrolli (C, E) is similar to that of D. rhopaloa (D, F), but shows quantitative difference in size and appears sturdier. The looping of the aedeagus base is more open in D. carrolli than in D. rhopaloa (blue double-arrows). (G, H), testes of D. carrolli (G) and D. rhopaloa (H) have similar shapes and coiling, but they appear bigger in D. carrolli, and show a characteristic bright yellow color compared to the dull yellow testes of D. rhopaloa." httpUri="https://zenodo.org/record/1292136/files/figure.png" pageId="2" pageNumber="504">Figure 2A</figureCitation>
), in contrast to approximately 5 setae there in 
<taxonomicName id="4C064D64F427FFD4FCA3FCEDFC567A7C" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[804,941,869,893]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FCA3FCEDFC567A7C" box="[804,941,869,893]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
(
<figureCitation id="133D2A62F427FFD4FC3AFCEEFBC87A7F" box="[957,1075,869,894]" captionStart="FIGURE 2" captionStartId="4.[151,250,1815,1837]" captionTargetBox="[253,1318,204,1780]" captionTargetId="figure@4.[235,1349,193,1794]" captionTargetPageId="4" captionText="FIGURE 2. Male terminalia. Dissected parts from D. carrolli (A, C, E, G) or D. rhopaloa (B, D, F, H). (A, B) epandrium in dorsal view. Note the differences in sensory bristle length and distribution (blue ovals and arrowheads) as well as genital comb size (blue arrows) between species. The epandrium is always darker overall in D. carrolli. Phallus in ventral view (C, D) and side view (E, F). The phallus conformation in D. carrolli (C, E) is similar to that of D. rhopaloa (D, F), but shows quantitative difference in size and appears sturdier. The looping of the aedeagus base is more open in D. carrolli than in D. rhopaloa (blue double-arrows). (G, H), testes of D. carrolli (G) and D. rhopaloa (H) have similar shapes and coiling, but they appear bigger in D. carrolli, and show a characteristic bright yellow color compared to the dull yellow testes of D. rhopaloa." httpUri="https://zenodo.org/record/1292136/files/figure.png" pageId="2" pageNumber="504">Figure 2B</figureCitation>
). Anal plates covered with long bristles, especially densely on the ventral portion, strongly elongated below and with one very large, apically somewhat round spine at apex. This spine is a diagnostic character of the 
<taxonomicName id="4C064D64F427FFD4FC74FC25FB857AC4" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[1011,1150,941,965]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FC74FC25FB857AC4" box="[1011,1150,941,965]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
species subgroup (
<bibRefCitation id="EF974B16F427FFD4FADBFC26FF287AE8" pageId="2" pageNumber="504" refId="ref4879" refString="Toda, M. (1991) Drosophilidae (Diptera) in Myanmar (Burma) VII. The Drosophila melanogaster species-group, excepting the D. montium species-subgroup. Oriental Insects, 25, 69 - 94. https: // doi. org / 10.1080 / 00305316.1991.10432216" type="journal article">Toda, 1991</bibRefCitation>
). Primary clasper long (about twice as long as in 
<taxonomicName id="4C064D64F427FFD4FC92FC5AFC5B7AE9" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[789,928,976,1000]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FC92FC5AFC5B7AE9" box="[789,928,976,1000]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
), bearing 2 combs of stout bristles pointing inward. The dorsal comb carries 4–5 teeth and the ventral comb 6–7 longer teeth. 2–3 strong and long bristles are located ventrally to each dorsal comb and 1 long and 1–2 short bristles are located just dorsally to the ventral combs. These combs and bristles exist in 
<taxonomicName id="4C064D64F427FFD4FD02FBB7FCEA7D55" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[645,785,1084,1108]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FD02FBB7FCEA7D55" box="[645,785,1084,1108]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
, but they are smaller, and the combs have fewer teeth. Aedeagus similar to that of 
<taxonomicName id="4C064D64F427FFD4FE4BFBEBFDA87D76" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[460,595,1119,1143]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FE4BFBEBFDA87D76" box="[460,595,1119,1143]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
, only stronger and darker. Phallus with a strong basal sinuation, wider than in 
<taxonomicName id="4C064D64F427FFD4FF32FB0FFEBB7D9A" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[181,320,1155,1179]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FF32FB0FFEBB7D9A" box="[181,320,1155,1179]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
(blue double-arrows in 
<figureCitation id="133D2A62F427FFD4FDD3FB08FD137D9D" box="[596,744,1155,1180]" captionStart="FIGURE 2" captionStartId="4.[151,250,1815,1837]" captionTargetBox="[253,1318,204,1780]" captionTargetId="figure@4.[235,1349,193,1794]" captionTargetPageId="4" captionText="FIGURE 2. Male terminalia. Dissected parts from D. carrolli (A, C, E, G) or D. rhopaloa (B, D, F, H). (A, B) epandrium in dorsal view. Note the differences in sensory bristle length and distribution (blue ovals and arrowheads) as well as genital comb size (blue arrows) between species. The epandrium is always darker overall in D. carrolli. Phallus in ventral view (C, D) and side view (E, F). The phallus conformation in D. carrolli (C, E) is similar to that of D. rhopaloa (D, F), but shows quantitative difference in size and appears sturdier. The looping of the aedeagus base is more open in D. carrolli than in D. rhopaloa (blue double-arrows). (G, H), testes of D. carrolli (G) and D. rhopaloa (H) have similar shapes and coiling, but they appear bigger in D. carrolli, and show a characteristic bright yellow color compared to the dull yellow testes of D. rhopaloa." httpUri="https://zenodo.org/record/1292136/files/figure.png" pageId="2" pageNumber="504">Figure 2E, F</figureCitation>
). One pair of membranous, palm-like processes is present at laterosubapical portions of the aedeagus and is a diagnostic character for the 
<taxonomicName id="4C064D64F427FFD4FC73FB23FB877DBE" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[1012,1148,1191,1215]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FC73FB23FB877DBE" box="[1012,1148,1191,1215]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
subgroup. Testes orangebrown, darker than in 
<taxonomicName id="4C064D64F427FFD4FE08FB40FDEC7DE2" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[399,535,1227,1251]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FE08FB40FDEC7DE2" box="[399,535,1227,1251]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
, with 2 outer and 3 inner coils (
<figureCitation id="133D2A62F427FFD4FCFBFB41FBE87DE2" box="[892,1043,1226,1251]" captionStart="FIGURE 2" captionStartId="4.[151,250,1815,1837]" captionTargetBox="[253,1318,204,1780]" captionTargetId="figure@4.[235,1349,193,1794]" captionTargetPageId="4" captionText="FIGURE 2. Male terminalia. Dissected parts from D. carrolli (A, C, E, G) or D. rhopaloa (B, D, F, H). (A, B) epandrium in dorsal view. Note the differences in sensory bristle length and distribution (blue ovals and arrowheads) as well as genital comb size (blue arrows) between species. The epandrium is always darker overall in D. carrolli. Phallus in ventral view (C, D) and side view (E, F). The phallus conformation in D. carrolli (C, E) is similar to that of D. rhopaloa (D, F), but shows quantitative difference in size and appears sturdier. The looping of the aedeagus base is more open in D. carrolli than in D. rhopaloa (blue double-arrows). (G, H), testes of D. carrolli (G) and D. rhopaloa (H) have similar shapes and coiling, but they appear bigger in D. carrolli, and show a characteristic bright yellow color compared to the dull yellow testes of D. rhopaloa." httpUri="https://zenodo.org/record/1292136/files/figure.png" pageId="2" pageNumber="504">Figure 2G, H</figureCitation>
).
</paragraph>
<paragraph id="8BB936E7F427FFD4FF40FB65FD907CDF" blockId="2.[151,1437,151,2002]" pageId="2" pageNumber="504">
Female terminalia. Oviscape (egg guides) moderately sclerotized, lined-up with single row of about 15 short and stout setae along its outer edge on each side, and an additional pair of longer and thiner bristles at its posterior tip. This configuration is similar to that of 
<taxonomicName id="4C064D64F427FFD4FDF6FABCFD027C4F" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[625,761,1334,1358]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FDF6FABCFD027C4F" box="[625,761,1334,1358]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
, which shows more irregularity in the alignment of the stout setae (
<figureCitation id="133D2A62F427FFD4FF59FAD2FE8D7C73" box="[222,374,1369,1394]" captionStart="FIGURE 3" captionStartId="5.[151,250,1765,1787]" captionTargetBox="[266,1327,198,1733]" captionTargetId="figure@5.[253,1341,193,1748]" captionTargetPageId="5" captionText="FIGURE 3. Female terminalia, eggs and pupae. (A, A’, B) last abdominal segments and spermatheca of D. carrolli (A, A’) or D. rhopaloa (B) females. Note the strongly sclerotized plates (blue arrowheads) at the end of the oviduct in D. carrolli (A, A’), but not D. rhopaloa (B). Also note the teeth (bristles) on the egg-guides are slightly more numerous and stouter in D. carrolli (blue arrows). The rear view of a D. carrolli female laying an egg (A’) shows the relative position of the sclerotized plates, the anal plates and the ovipositor. The spermathecae are marginally bigger and darker in age-matched D. carrolli compared to D. rhopaloa. (C, D) eggs of D. carrolli (C) and D. rhopaloa (D) in ventral view. Both species show characteristic spatula-shape respiratory appendages. These appendages vary in shape within species but show no notable difference between species. (E, F) pupae of D. carrolli (A) or D. rhopaloa (B) males about 24 hours before adult emergence. D. carrolli pupae are more rounded on their sides." httpUri="https://zenodo.org/record/1292138/files/figure.png" pageId="2" pageNumber="504">Figure 3A, B</figureCitation>
). Anal plates well developed, stronger than in 
<taxonomicName id="4C064D64F427FFD4FC03FAD1FBF67C73" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[900,1037,1370,1394]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FC03FAD1FBF67C73" box="[900,1037,1370,1394]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
. On the dorsal/posterior side of the oviduct, between the oviscape and the anal plates, there are two obvious pairs of strongly sclerotized, pigmented plates (
<figureCitation id="133D2A62F427FFD4FF6FFA2AFE9B7CB8" box="[232,352,1441,1466]" captionStart="FIGURE 3" captionStartId="5.[151,250,1765,1787]" captionTargetBox="[266,1327,198,1733]" captionTargetId="figure@5.[253,1341,193,1748]" captionTargetPageId="5" captionText="FIGURE 3. Female terminalia, eggs and pupae. (A, A’, B) last abdominal segments and spermatheca of D. carrolli (A, A’) or D. rhopaloa (B) females. Note the strongly sclerotized plates (blue arrowheads) at the end of the oviduct in D. carrolli (A, A’), but not D. rhopaloa (B). Also note the teeth (bristles) on the egg-guides are slightly more numerous and stouter in D. carrolli (blue arrows). The rear view of a D. carrolli female laying an egg (A’) shows the relative position of the sclerotized plates, the anal plates and the ovipositor. The spermathecae are marginally bigger and darker in age-matched D. carrolli compared to D. rhopaloa. (C, D) eggs of D. carrolli (C) and D. rhopaloa (D) in ventral view. Both species show characteristic spatula-shape respiratory appendages. These appendages vary in shape within species but show no notable difference between species. (E, F) pupae of D. carrolli (A) or D. rhopaloa (B) males about 24 hours before adult emergence. D. carrolli pupae are more rounded on their sides." httpUri="https://zenodo.org/record/1292138/files/figure.png" pageId="2" pageNumber="504">Figure 3A</figureCitation>
, A’) that are not visible in 
<taxonomicName id="4C064D64F427FFD4FD11FA29FCDB7CB8" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[662,800,1441,1465]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FD11FA29FCDB7CB8" box="[662,800,1441,1465]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
. Spermathecae rounded and hollow, slightly bigger and darker than those of 
<taxonomicName id="4C064D64F427FFD4FEFAFA4DFDFE7CDC" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[381,517,1477,1501]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FEFAFA4DFDFE7CDC" box="[381,517,1477,1501]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
females.
</paragraph>
<paragraph id="8BB936E7F427FFD4FF40FA62FB9D7F48" blockId="2.[151,1437,151,2002]" pageId="2" pageNumber="504">
Pupa. Length: 3.2± 
<quantity id="4CFE9B02F427FFD4FE2FFA62FDF97F01" box="[424,514,1513,1537]" metricMagnitude="-4" metricUnit="m" metricValue="3.0" pageId="2" pageNumber="504" unit="mm" value="0.3">0.3 mm</quantity>
(n=10). Pupal case yellowish, with sides regularly rounded (more so than 
<taxonomicName id="4C064D64F427FFD4FA06FA62FF067F25" authorityName="Bock &amp; Wheeler" authorityYear="1972" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FA06FA62FF067F25" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
pupae, 
<figureCitation id="133D2A62F427FFD4FED1F987FE1A7F24" box="[342,481,1548,1573]" captionStart="FIGURE 3" captionStartId="5.[151,250,1765,1787]" captionTargetBox="[266,1327,198,1733]" captionTargetId="figure@5.[253,1341,193,1748]" captionTargetPageId="5" captionText="FIGURE 3. Female terminalia, eggs and pupae. (A, A’, B) last abdominal segments and spermatheca of D. carrolli (A, A’) or D. rhopaloa (B) females. Note the strongly sclerotized plates (blue arrowheads) at the end of the oviduct in D. carrolli (A, A’), but not D. rhopaloa (B). Also note the teeth (bristles) on the egg-guides are slightly more numerous and stouter in D. carrolli (blue arrows). The rear view of a D. carrolli female laying an egg (A’) shows the relative position of the sclerotized plates, the anal plates and the ovipositor. The spermathecae are marginally bigger and darker in age-matched D. carrolli compared to D. rhopaloa. (C, D) eggs of D. carrolli (C) and D. rhopaloa (D) in ventral view. Both species show characteristic spatula-shape respiratory appendages. These appendages vary in shape within species but show no notable difference between species. (E, F) pupae of D. carrolli (A) or D. rhopaloa (B) males about 24 hours before adult emergence. D. carrolli pupae are more rounded on their sides." httpUri="https://zenodo.org/record/1292138/files/figure.png" pageId="2" pageNumber="504">Figure 3E,F</figureCitation>
). Anterior spiracles similar to those of 
<taxonomicName id="4C064D64F427FFD4FC1CF986FBDF7F25" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[923,1060,1548,1572]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F427FFD4FC1CF986FBDF7F25" box="[923,1060,1548,1572]" italics="true" pageId="2" pageNumber="504">D. rhopaloa</emphasis>
</taxonomicName>
, with 7 (occasionally 8) terminal branches. Horn index (100 x length of the anterior spiracles/total puparium length) = 5.
</paragraph>
<paragraph id="8BB936E7F427FFD4FF40F9DEFE2B7F91" blockId="2.[151,1437,151,2002]" pageId="2" pageNumber="504">
Egg. Length: 0.6±0.0 mm (n=10). Two antero-dorsal respiratory appendages with a round basal stem and a flat spatula-like tip (
<figureCitation id="133D2A62F427FFD4FECAF9FCFE397F91" box="[333,450,1655,1680]" captionStart="FIGURE 3" captionStartId="5.[151,250,1765,1787]" captionTargetBox="[266,1327,198,1733]" captionTargetId="figure@5.[253,1341,193,1748]" captionTargetPageId="5" captionText="FIGURE 3. Female terminalia, eggs and pupae. (A, A’, B) last abdominal segments and spermatheca of D. carrolli (A, A’) or D. rhopaloa (B) females. Note the strongly sclerotized plates (blue arrowheads) at the end of the oviduct in D. carrolli (A, A’), but not D. rhopaloa (B). Also note the teeth (bristles) on the egg-guides are slightly more numerous and stouter in D. carrolli (blue arrows). The rear view of a D. carrolli female laying an egg (A’) shows the relative position of the sclerotized plates, the anal plates and the ovipositor. The spermathecae are marginally bigger and darker in age-matched D. carrolli compared to D. rhopaloa. (C, D) eggs of D. carrolli (C) and D. rhopaloa (D) in ventral view. Both species show characteristic spatula-shape respiratory appendages. These appendages vary in shape within species but show no notable difference between species. (E, F) pupae of D. carrolli (A) or D. rhopaloa (B) males about 24 hours before adult emergence. D. carrolli pupae are more rounded on their sides." httpUri="https://zenodo.org/record/1292138/files/figure.png" pageId="2" pageNumber="504">Figure 3C</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C31C656CF427FFD0FF40F910FB54781C" lastPageId="6" lastPageNumber="508" pageId="2" pageNumber="504" type="materials_examined">
<paragraph id="8BB936E7F427FFD4FF40F910FBC77ED0" blockId="2.[151,1437,151,2002]" pageId="2" pageNumber="504">
<materialsCitation id="3B6E3CBAF427FFD4FF40F910FBB37FFA" ID-GBIF-Occurrence="1849336044" collectingDate="2003-10" collectorName="Olga Barmina" country="Brunei" county="Kuala Belalong" latitude="4.975111" location="Eighty-four" longLatPrecision="1" longitude="114.89102" municipality="Artyom Kopp" pageId="2" pageNumber="504" specimenCount="1" stateProvince="Temburong" typeStatus="holotype">
<emphasis id="B972EAF5F427FFD4FF40F910FE837FB5" bold="true" box="[199,376,1691,1716]" pageId="2" pageNumber="504">
<typeStatus id="54BD8845F427FFD4FF40F910FEF97FB5" box="[199,258,1691,1716]" pageId="2" pageNumber="504">Type</typeStatus>
material.
</emphasis>
<location id="8ED9603CF427FFD4FE04F910FDFD7FB5" LSID="urn:lsid:plazi:treatment:03AF87F1F424FFD0FF10FAFCFB427AA4:8ED9603CF427FFD4FE04F910FDFD7FB5" box="[387,518,1691,1716]" country="Brunei" county="Kuala Belalong" latitude="4.975111" longLatPrecision="1" longitude="114.89102" municipality="Artyom Kopp" name="Eighty-four" pageId="2" pageNumber="504" stateProvince="Temburong">Eighty-four</location>
specimens of 
<taxonomicName id="4C064D64F427FFD4FD35F917FCD77FB2" authority="Gompel &amp; Kopp, 2018" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[690,812,1691,1715]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="2" pageNumber="504" phylum="Arthropoda" rank="species" species="carrolli" status="sp. nov.">
<emphasis id="B972EAF5F427FFD4FD35F917FCD77FB2" box="[690,812,1691,1715]" italics="true" pageId="2" pageNumber="504">D. carrolli</emphasis>
</taxonomicName>
<emphasis id="B972EAF5F427FFD4FCBFF910FC807FB5" bold="true" box="[824,891,1691,1716]" pageId="2" pageNumber="504">
<taxonomicNameLabel id="A241578EF427FFD4FCBFF910FC807FB5" box="[824,891,1691,1716]" pageId="2" pageNumber="504" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
, derived from an isofemale line established in 
<date id="FFB81027F427FFD4FF10F934FECF7FD9" box="[151,308,1727,1752]" pageId="2" pageNumber="504" value="2003-10">
<collectingDate id="EFFCE9CFF427FFD4FF10F934FECF7FD9" box="[151,308,1727,1752]" pageId="2" pageNumber="504" value="2003-10">October 2003</collectingDate>
</date>
(
<collectingMunicipality id="6BDDAC9DF427FFD4FEC0F94BFE117FD6" box="[327,490,1728,1752]" pageId="2" pageNumber="504">Artyom Kopp</collectingMunicipality>
and 
<collectorName id="26F35331F427FFD4FDAEF934FD307FD9" box="[553,715,1727,1752]" pageId="2" pageNumber="504">Olga Barmina</collectorName>
<emphasis id="B972EAF5F427FFD4FD50F934FCFB7FD6" box="[727,768,1727,1751]" italics="true" pageId="2" pageNumber="504">leg.</emphasis>
), from a single female collected at 
<collectingCounty id="62D84E6BF427FFD4FB2BF934FA997FD9" box="[1196,1378,1727,1752]" pageId="2" pageNumber="504">Kuala Belalong</collectingCounty>
, Ulu 
<collectingRegion id="49C2F805F427FFD4FF10F969FEE37FFA" box="[151,280,1762,1787]" country="Brunei" name="Temburong" pageId="2" pageNumber="504">Temburong</collectingRegion>
National Park, 
<collectingCountry id="F3117677F427FFD4FE67F969FD3B7FFA" box="[480,704,1762,1787]" name="Brunei" pageId="2" pageNumber="504">Brunei Darussalam</collectingCountry>
[
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<geoCoordinate id="EE325020F427FFD4FC02F969FBBA7FFA" box="[901,1089,1762,1787]" direction="east" orientation="longitude" pageId="2" pageNumber="504" precision="1" value="114.89102">114°53’27.68”E</geoCoordinate>
]
</materialsCitation>
. 
<materialsCitation id="3B6E3CBAF427FFD4FBDCF969FBC77ED0" ID-GBIF-Occurrence="1849335966" collectorName="N. Gompel &amp; UC Davis Bohart Museum &amp; Davis, CA &amp; P. O'Grady &amp; M. J. Toda" country="Germany" county="Sapporo" location="British Museum of National History" municipality="Kita-ku" pageId="2" pageNumber="504" specimenCount="167" specimenCount-female="41" specimenCount-male="42" stateProvince="Hokkaido" typeStatus="holotype">
This material, mounted on cardboard and pinned, will be dispatched as follows: 
<specimenCount id="9D00FD6EF427FFD4FD76F88CFC927E1E" box="[753,873,1798,1823]" pageId="2" pageNumber="504" type="generic" typeStatus="holotype">1 holotype</specimenCount>
(male, dissected) and 
<specimenCount id="9D00FD6EF427FFD4FBE0F88DFB0D7E1E" box="[1127,1270,1798,1823]" pageId="2" pageNumber="504" type="generic" typeStatus="paratypes">26 paratypes</specimenCount>
(
<specimenCount id="9D00FD6EF427FFD4FA81F88CFA977E1E" box="[1286,1388,1798,1823]" pageId="2" pageNumber="504" type="male">12 males</specimenCount>
and 
<specimenCount id="9D00FD6EF427FFD4FF10F8A0FEE77E42" box="[151,284,1834,1859]" pageId="2" pageNumber="504" type="female">14 females</specimenCount>
): coll. 
<collectorName id="26F35331F427FFD4FEF7F8A0FE0E7E42" box="[368,501,1834,1859]" pageId="2" pageNumber="504">N. Gompel</collectorName>
, Munich, 
<collectingCountry id="F3117677F427FFD4FDE8F8A0FD207E42" box="[623,731,1835,1859]" name="Germany" pageId="2" pageNumber="504">Germany</collectingCountry>
; 
<specimenCount id="9D00FD6EF427FFD4FD6CF8A0FC867E42" box="[747,893,1835,1859]" pageId="2" pageNumber="504" type="generic" typeStatus="paratypes">11 paratypes</specimenCount>
(
<specimenCount id="9D00FD6EF427FFD4FC15F8A1FC157E42" box="[914,1006,1834,1859]" pageId="2" pageNumber="504" type="male">6 males</specimenCount>
and 
<specimenCount id="9D00FD6EF427FFD4FBB7F8A1FB5C7E42" box="[1072,1191,1834,1859]" pageId="2" pageNumber="504" type="female">5 females</specimenCount>
): 
<location id="8ED9603CF427FFD4FB39F8A1FEA87E67" LSID="urn:lsid:plazi:treatment:03AF87F1F424FFD0FF10FAFCFB427AA4:8ED9603CF427FFD4FB39F8A1FEA87E67" country="Germany" county="Sapporo" municipality="Kita-ku" name="British Museum of National History" pageId="2" pageNumber="504" stateProvince="Hokkaido">British Museum of National History</location>
, London; 
<specimenCount id="9D00FD6EF427FFD4FE44F8C5FDAB7E66" box="[451,592,1870,1895]" pageId="2" pageNumber="504" type="generic" typeStatus="paratypes">11 paratypes</specimenCount>
(
<specimenCount id="9D00FD6EF427FFD4FDD8F8C6FD4E7E67" box="[607,693,1869,1894]" pageId="2" pageNumber="504" type="male">6 males</specimenCount>
and 
<specimenCount id="9D00FD6EF427FFD4FD6CF8C6FCA67E67" box="[747,861,1869,1894]" pageId="2" pageNumber="504" type="female">5 females</specimenCount>
): 
<collectorName id="26F35331F427FFD4FCE8F8C5FB627E67" box="[879,1177,1869,1894]" pageId="2" pageNumber="504">
UC 
<location id="8ED9603CF427FFD4FC1BF8C6FB627E67" LSID="urn:lsid:plazi:treatment:03AF87F1F424FFD0FF10FAFCFB427AA4:8ED9603CF427FFD4FC1BF8C6FB627E67" box="[924,1177,1869,1894]" country="Germany" county="Sapporo" municipality="Kita-ku" name="Davis Bohart Museum of Entomology" pageId="2" pageNumber="504" stateProvince="Hokkaido">Davis Bohart Museum</location>
</collectorName>
of Entomology, 
<collectorName id="26F35331F427FFD4FAD3F8C6FF3A7E8B" pageId="2" pageNumber="504">Davis, CA</collectorName>
, USA; 
<specimenCount id="9D00FD6EF427FFD4FE94F8F9FE5B7E8B" box="[275,416,1906,1930]" pageId="2" pageNumber="504" type="generic" typeStatus="paratypes">11 paratypes</specimenCount>
(
<specimenCount id="9D00FD6EF427FFD4FE37F8FAFDFC7E8B" box="[432,519,1905,1930]" pageId="2" pageNumber="504" type="male">6 males</specimenCount>
and 
<specimenCount id="9D00FD6EF427FFD4FDB9F8FAFD4A7E8B" box="[574,689,1905,1930]" pageId="2" pageNumber="504" type="female">5 females</specimenCount>
): Musée National d’Histoire Naturelle, Paris, 
<collectingCountry id="F3117677F427FFD4FB35F8F9FAF97E88" box="[1202,1282,1906,1929]" name="France" pageId="2" pageNumber="504">France</collectingCountry>
; 
<specimenCount id="9D00FD6EF427FFD4FA89F8F9FA677E8B" box="[1294,1436,1906,1930]" pageId="2" pageNumber="504" type="generic" typeStatus="paratypes">12 paratypes</specimenCount>
(
<specimenCount id="9D00FD6EF427FFD4FF18F81EFF017EAF" box="[159,250,1941,1966]" pageId="2" pageNumber="504" type="male">6 males</specimenCount>
and 
<specimenCount id="9D00FD6EF427FFD4FEBEF81EFE557EAF" box="[313,430,1941,1966]" pageId="2" pageNumber="504" type="female">6 females</specimenCount>
): coll. 
<collectorName id="26F35331F427FFD4FD87F81DFD787EAF" box="[512,643,1941,1966]" pageId="2" pageNumber="504">P. O’Grady</collectorName>
, Cornell University, Ithaca, NY, USA; 
<specimenCount id="9D00FD6EF427FFD4FBC8F81DFB1A7EAF" box="[1103,1249,1942,1966]" pageId="2" pageNumber="504" type="generic" typeStatus="paratypes">12 paratypes</specimenCount>
(
<specimenCount id="9D00FD6EF427FFD4FB72F81EFAAB7EAF" box="[1269,1360,1941,1966]" pageId="2" pageNumber="504" type="male">6 males</specimenCount>
and 
<specimenCount id="9D00FD6EF427FFD4FA09F81EFF0F7ED3" pageId="2" pageNumber="504" type="female">6 females</specimenCount>
): coll. 
<collectorName id="26F35331F427FFD4FEB9F831FE437ED3" box="[318,440,1977,2002]" pageId="2" pageNumber="504">M. J. Toda</collectorName>
, 
<collectingRegion id="49C2F805F427FFD4FE44F832FDCA7ED3" box="[451,561,1977,2002]" country="Japan" name="Hokkaido" pageId="2" pageNumber="504">Hokkaido</collectingRegion>
University Museum, 
<collectingMunicipality id="6BDDAC9DF427FFD4FCA1F832FC847ED3" box="[806,895,1977,2002]" pageId="2" pageNumber="504">Kita-ku</collectingMunicipality>
, 
<collectingCounty id="62D84E6BF427FFD4FC0EF832FC127ED0" box="[905,1001,1977,2001]" pageId="2" pageNumber="504">Sapporo</collectingCounty>
, 
<collectingCountry id="F3117677F427FFD4FC72F831FBC37ED0" box="[1013,1080,1978,2001]" name="Japan" pageId="2" pageNumber="504">Japan</collectingCountry>
.
</materialsCitation>
</paragraph>
<caption id="DF79666FF426FFD5FF10F9D6FDEB7EE9" httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="3" pageNumber="505" startId="3.[151,250,1629,1651]" targetBox="[227,1287,212,1577]" targetPageId="3">
<paragraph id="8BB936E7F426FFD5FF10F9D6FDEB7EE9" blockId="3.[151,1437,1629,2024]" pageId="3" pageNumber="505">
<emphasis id="B972EAF5F426FFD5FF10F9D6FD0D7F72" bold="true" box="[151,758,1629,1651]" pageId="3" pageNumber="505">FIGURE 1. Adult morphology and geographical range.</emphasis>
(A–D) dorsal views of 5–7 day old adult 
<taxonomicName id="4C064D64F426FFD5FB2BF9D6FA887F72" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[1196,1395,1629,1651]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F426FFD5FB2BF9D6FA887F72" box="[1196,1395,1629,1651]" italics="true" pageId="3" pageNumber="505">Drosophila carrolli</emphasis>
</taxonomicName>
(A, male; B, female) and 
<taxonomicName id="4C064D64F426FFD5FEF4F9F6FE147F93" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[371,495,1660,1682]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F426FFD5FEF4F9F6FE147F93" box="[371,495,1660,1682]" italics="true" pageId="3" pageNumber="505">D. rhopaloa</emphasis>
</taxonomicName>
(C, male; D, female). (E, F) male abdomens of 
<taxonomicName id="4C064D64F426FFD5FC51F9F6FBB97F93" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[982,1090,1660,1682]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F426FFD5FC51F9F6FBB97F93" box="[982,1090,1660,1682]" italics="true" pageId="3" pageNumber="505">D. carrolli</emphasis>
</taxonomicName>
(E) and 
<taxonomicName id="4C064D64F426FFD5FB1CF9F6FAEC7F93" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[1179,1303,1660,1682]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F426FFD5FB1CF9F6FAEC7F93" box="[1179,1303,1660,1682]" italics="true" pageId="3" pageNumber="505">D. rhopaloa</emphasis>
</taxonomicName>
(F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in 
<taxonomicName id="4C064D64F426FFD5FC3FF931FBC87FCE" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[952,1075,1721,1743]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F426FFD5FC3FF931FBC87FCE" box="[952,1075,1721,1743]" italics="true" pageId="3" pageNumber="505">D. rhopaloa</emphasis>
</taxonomicName>
, but not 
<taxonomicName id="4C064D64F426FFD5FB09F931FB007FCE" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[1166,1275,1721,1743]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F426FFD5FB09F931FB007FCE" box="[1166,1275,1721,1743]" italics="true" pageId="3" pageNumber="505">D. carrolli</emphasis>
</taxonomicName>
(green arrows). Also note the bright yellow testes of 
<taxonomicName id="4C064D64F426FFD5FD9EF952FD7D7FEF" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[537,646,1752,1774]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F426FFD5FD9EF952FD7D7FEF" box="[537,646,1752,1774]" italics="true" pageId="3" pageNumber="505">D. carrolli</emphasis>
</taxonomicName>
, contrasting with the paler testes of 
<taxonomicName id="4C064D64F426FFD5FB86F952FB847FEF" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[1025,1151,1752,1774]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F426FFD5FB86F952FB847FEF" box="[1025,1151,1752,1774]" italics="true" pageId="3" pageNumber="505">D. rhopaloa</emphasis>
</taxonomicName>
(green arrowheads). (G, H) female abdomens of 
<taxonomicName id="4C064D64F426FFD5FEEEF973FE2E7E0D" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[361,469,1782,1805]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F426FFD5FEEEF973FE2E7E0D" box="[361,469,1782,1805]" italics="true" pageId="3" pageNumber="505">D. carrolli</emphasis>
</taxonomicName>
(G) and 
<taxonomicName id="4C064D64F426FFD5FDA9F973FD527E0D" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[558,681,1782,1805]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F426FFD5FDA9F973FD527E0D" box="[558,681,1782,1805]" italics="true" pageId="3" pageNumber="505">D. rhopaloa</emphasis>
</taxonomicName>
(H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of 
<taxonomicName id="4C064D64F426FFD5FE5BF89DFDB37E2A" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[476,584,1813,1835]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F426FFD5FE5BF89DFDB37E2A" box="[476,584,1813,1835]" italics="true" pageId="3" pageNumber="505">D. carrolli</emphasis>
</taxonomicName>
(I) and 
<taxonomicName id="4C064D64F426FFD5FD1DF89DFCEC7E2A" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[666,791,1813,1835]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F426FFD5FD1DF89DFCEC7E2A" box="[666,791,1813,1835]" italics="true" pageId="3" pageNumber="505">D. rhopaloa</emphasis>
</taxonomicName>
(J). Note the darker pigmentation at the anterior distal tip of of 
<taxonomicName id="4C064D64F426FFD5FF10F8BEFEFF7E4B" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[151,260,1844,1866]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F426FFD5FF10F8BEFEFF7E4B" box="[151,260,1844,1866]" italics="true" pageId="3" pageNumber="505">D. carrolli</emphasis>
</taxonomicName>
but not 
<taxonomicName id="4C064D64F426FFD5FEDBF8BEFE217E4B" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[348,474,1844,1866]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F426FFD5FEDBF8BEFE217E4B" box="[348,474,1844,1866]" italics="true" pageId="3" pageNumber="505">D. rhopaloa</emphasis>
</taxonomicName>
wings. (K, L) right antennae of males, showing no notable difference in the arista branching between 
<taxonomicName id="4C064D64F426FFD5FF75F8DDFEA57E6A" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[242,350,1877,1899]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F426FFD5FF75F8DDFEA57E6A" box="[242,350,1877,1899]" italics="true" pageId="3" pageNumber="505">D. carrolli</emphasis>
</taxonomicName>
(K) and 
<taxonomicName id="4C064D64F426FFD5FE3EF8DDFDCD7E6A" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[441,566,1877,1899]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F426FFD5FE3EF8DDFDCD7E6A" box="[441,566,1877,1899]" italics="true" pageId="3" pageNumber="505">D. rhopaloa</emphasis>
</taxonomicName>
(L), but a relatively shorter 3rd antennal segment in 
<taxonomicName id="4C064D64F426FFD5FBC1F8DDFB397E6A" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[1094,1218,1877,1899]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F426FFD5FBC1F8DDFB397E6A" box="[1094,1218,1877,1899]" italics="true" pageId="3" pageNumber="505">D. rhopaloa</emphasis>
</taxonomicName>
(double arrows). (M, N) sex combs on the 1 
<taxonomicName id="4C064D64F426FFD5FE08F8FFFE3C7E8D" box="[399,455,1908,1932]" pageId="3" pageNumber="505" rank="subSpecies" subSpecies="and">st and</taxonomicName>
2nd tarsal segments of forelegs in 
<taxonomicName id="4C064D64F426FFD5FCBCF8F3FC507E8D" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[827,939,1910,1933]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F426FFD5FCBCF8F3FC507E8D" box="[827,939,1910,1933]" italics="true" pageId="3" pageNumber="505">D. carrolli</emphasis>
</taxonomicName>
(M) and 
<taxonomicName id="4C064D64F426FFD5FB92F8F3FB6E7E8D" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[1045,1173,1910,1933]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F426FFD5FB92F8F3FB6E7E8D" box="[1045,1173,1910,1933]" italics="true" pageId="3" pageNumber="505">D. rhopaloa</emphasis>
</taxonomicName>
(N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where 
<taxonomicName id="4C064D64F426FFD5FD83F83EFD7A7ECB" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[516,641,1972,1994]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F426FFD5FD83F83EFD7A7ECB" box="[516,641,1972,1994]" italics="true" pageId="3" pageNumber="505">D. rhopaloa</emphasis>
</taxonomicName>
(white circles) and 
<taxonomicName id="4C064D64F426FFD5FCCAF83EFC427ECB" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[845,953,1972,1994]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="3" pageNumber="505" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F426FFD5FCCAF83EFC427ECB" box="[845,953,1972,1994]" italics="true" pageId="3" pageNumber="505">D. carrolli</emphasis>
</taxonomicName>
(red circle) were found. The circle with dotted outline denotes an imprecise locality.
</paragraph>
</caption>
<caption id="DF79666FF421FFD2FF10F89CFB217EE4" httpUri="https://zenodo.org/record/1292136/files/figure.png" pageId="4" pageNumber="506" startId="4.[151,250,1815,1837]" targetBox="[253,1318,204,1780]" targetPageId="4">
<paragraph id="8BB936E7F421FFD2FF10F89CFB217EE4" blockId="4.[151,1436,1815,2021]" pageId="4" pageNumber="506">
<emphasis id="B972EAF5F421FFD2FF10F89CFE357E2C" bold="true" box="[151,462,1815,1837]" pageId="4" pageNumber="506">FIGURE 2. Male terminalia.</emphasis>
Dissected parts from 
<taxonomicName id="4C064D64F421FFD2FD28F893FCE77E2C" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[687,796,1815,1837]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="506" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F421FFD2FD28F893FCE77E2C" box="[687,796,1815,1837]" italics="true" pageId="4" pageNumber="506">D. carrolli</emphasis>
</taxonomicName>
(A, C, E, G) or 
<taxonomicName id="4C064D64F421FFD2FC44F893FBC47E2C" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[963,1087,1815,1837]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="506" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F421FFD2FC44F893FBC47E2C" box="[963,1087,1815,1837]" italics="true" pageId="4" pageNumber="506">D. rhopaloa</emphasis>
</taxonomicName>
(B, D, F, H). (A, B) epandrium in dorsal view. Note the differences in sensory bristle length and distribution (blue ovals and arrowheads) as well as genital comb size (blue arrows) between species. The epandrium is always darker overall in 
<taxonomicName id="4C064D64F421FFD2FC46F8DEFBD57E6B" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[961,1070,1876,1898]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="506" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F421FFD2FC46F8DEFBD57E6B" box="[961,1070,1876,1898]" italics="true" pageId="4" pageNumber="506">D. carrolli</emphasis>
</taxonomicName>
. Phallus in ventral view (C, D) and side view (E, F). The phallus conformation in 
<taxonomicName id="4C064D64F421FFD2FDEAF8FFFD227E88" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[621,729,1907,1929]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="506" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F421FFD2FDEAF8FFFD227E88" box="[621,729,1907,1929]" italics="true" pageId="4" pageNumber="506">D. carrolli</emphasis>
</taxonomicName>
(C, E) is similar to that of 
<taxonomicName id="4C064D64F421FFD2FC6CF8FFFB9C7E88" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[1003,1127,1907,1929]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="506" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F421FFD2FC6CF8FFFB9C7E88" box="[1003,1127,1907,1929]" italics="true" pageId="4" pageNumber="506">D. rhopaloa</emphasis>
</taxonomicName>
(D, F), but shows quantitative difference in size and appears sturdier. The looping of the aedeagus base is more open in 
<taxonomicName id="4C064D64F421FFD2FBA2F819FB697EA6" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[1061,1170,1937,1959]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="506" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F421FFD2FBA2F819FB697EA6" box="[1061,1170,1937,1959]" italics="true" pageId="4" pageNumber="506">D. carrolli</emphasis>
</taxonomicName>
than in 
<taxonomicName id="4C064D64F421FFD2FB62F819FA997EA6" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[1253,1378,1937,1959]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="506" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F421FFD2FB62F819FA997EA6" box="[1253,1378,1937,1959]" italics="true" pageId="4" pageNumber="506">D. rhopaloa</emphasis>
</taxonomicName>
(blue double-arrows). (G, H), testes of 
<taxonomicName id="4C064D64F421FFD2FE63F83AFDAB7EC7" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[484,592,1968,1990]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="506" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F421FFD2FE63F83AFDAB7EC7" box="[484,592,1968,1990]" italics="true" pageId="4" pageNumber="506">D. carrolli</emphasis>
</taxonomicName>
(G) and 
<taxonomicName id="4C064D64F421FFD2FD2DF83AFCDE7EC7" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[682,805,1968,1990]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="506" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F421FFD2FD2DF83AFCDE7EC7" box="[682,805,1968,1990]" italics="true" pageId="4" pageNumber="506">D. rhopaloa</emphasis>
</taxonomicName>
(H) have similar shapes and coiling, but they appear bigger in 
<taxonomicName id="4C064D64F421FFD2FF10F85BFEF97EE4" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[151,258,1999,2021]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="506" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F421FFD2FF10F85BFEF97EE4" box="[151,258,1999,2021]" italics="true" pageId="4" pageNumber="506">D. carrolli</emphasis>
</taxonomicName>
, and show a characteristic bright yellow color compared to the dull yellow testes of 
<taxonomicName id="4C064D64F421FFD2FBDEF85BFB2E7EE4" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[1113,1237,1999,2021]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="4" pageNumber="506" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F421FFD2FBDEF85BFB2E7EE4" box="[1113,1237,1999,2021]" italics="true" pageId="4" pageNumber="506">D. rhopaloa</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<caption id="DF79666FF420FFD3FF10F96EFEDE7EF1" httpUri="https://zenodo.org/record/1292138/files/figure.png" pageId="5" pageNumber="507" startId="5.[151,250,1765,1787]" targetBox="[266,1327,198,1733]" targetPageId="5">
<paragraph id="8BB936E7F420FFD3FF10F96EFEDE7EF1" blockId="5.[151,1437,1765,2032]" pageId="5" pageNumber="507">
<emphasis id="B972EAF5F420FFD3FF10F96EFD687FFA" bold="true" box="[151,659,1765,1787]" pageId="5" pageNumber="507">FIGURE 3. Female terminalia, eggs and pupae.</emphasis>
(A, A’, B) last abdominal segments and spermatheca of 
<taxonomicName id="4C064D64F420FFD3FB4DF96DFACF7FFA" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[1226,1332,1765,1787]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F420FFD3FB4DF96DFACF7FFA" box="[1226,1332,1765,1787]" italics="true" pageId="5" pageNumber="507">D. carrolli</emphasis>
</taxonomicName>
(A, A’) or 
<taxonomicName id="4C064D64F420FFD3FF10F88EFEE87E1B" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[151,275,1796,1818]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F420FFD3FF10F88EFEE87E1B" box="[151,275,1796,1818]" italics="true" pageId="5" pageNumber="507">D. rhopaloa</emphasis>
</taxonomicName>
(B) females. Note the strongly sclerotized plates (blue arrowheads) at the end of the oviduct in 
<taxonomicName id="4C064D64F420FFD3FB5BF88EFAB37E1B" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[1244,1352,1796,1818]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F420FFD3FB5BF88EFAB37E1B" box="[1244,1352,1796,1818]" italics="true" pageId="5" pageNumber="507">D. carrolli</emphasis>
</taxonomicName>
(A, A’), but not 
<taxonomicName id="4C064D64F420FFD3FF61F8A8FE997E39" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[230,354,1826,1848]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F420FFD3FF61F8A8FE997E39" box="[230,354,1826,1848]" italics="true" pageId="5" pageNumber="507">D. rhopaloa</emphasis>
</taxonomicName>
(B). Also note the teeth (bristles) on the egg-guides are slightly more numerous and stouter in 
<taxonomicName id="4C064D64F420FFD3FAB7F8A8FA677E39" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[1328,1436,1826,1848]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F420FFD3FAB7F8A8FA677E39" box="[1328,1436,1826,1848]" italics="true" pageId="5" pageNumber="507">D. carrolli</emphasis>
</taxonomicName>
(blue arrows). The rear view of a 
<taxonomicName id="4C064D64F420FFD3FE6AF8C9FDA37E56" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[493,600,1857,1879]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F420FFD3FE6AF8C9FDA37E56" box="[493,600,1857,1879]" italics="true" pageId="5" pageNumber="507">D. carrolli</emphasis>
</taxonomicName>
female laying an egg (A’) shows the relative position of the sclerotized plates, the anal plates and the ovipositor. The spermathecae are marginally bigger and darker in age-matched 
<taxonomicName id="4C064D64F420FFD3FB0DF8EAFB0D7E74" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[1162,1270,1887,1910]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F420FFD3FB0DF8EAFB0D7E74" box="[1162,1270,1887,1910]" italics="true" pageId="5" pageNumber="507">D. carrolli</emphasis>
</taxonomicName>
compared to 
<taxonomicName id="4C064D64F420FFD3FA04F8EAFF087E95" authorityName="Bock &amp; Wheeler" authorityYear="1972" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F420FFD3FA04F8EAFF087E95" italics="true" pageId="5" pageNumber="507">D. rhopaloa</emphasis>
</taxonomicName>
. (C, D) eggs of 
<taxonomicName id="4C064D64F420FFD3FE1AF8F4FDF17E95" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[413,522,1918,1940]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F420FFD3FE1AF8F4FDF17E95" box="[413,522,1918,1940]" italics="true" pageId="5" pageNumber="507">D. carrolli</emphasis>
</taxonomicName>
(C) and 
<taxonomicName id="4C064D64F420FFD3FDE0F8F4FD1F7E95" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[615,740,1918,1940]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F420FFD3FDE0F8F4FD1F7E95" box="[615,740,1918,1940]" italics="true" pageId="5" pageNumber="507">D. rhopaloa</emphasis>
</taxonomicName>
(D) in ventral view. Both species show characteristic spatula-shape respiratory appendages. These appendages vary in shape within species but show no notable difference between species. (E, F) pupae of 
<taxonomicName id="4C064D64F420FFD3FF71F836FE9A7ED3" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[246,353,1980,2002]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F420FFD3FF71F836FE9A7ED3" box="[246,353,1980,2002]" italics="true" pageId="5" pageNumber="507">D. carrolli</emphasis>
</taxonomicName>
(A) or 
<taxonomicName id="4C064D64F420FFD3FE29F836FDD17ED3" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[430,554,1980,2002]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F420FFD3FE29F836FDD17ED3" box="[430,554,1980,2002]" italics="true" pageId="5" pageNumber="507">D. rhopaloa</emphasis>
</taxonomicName>
(B) males about 24 hours before adult emergence. 
<taxonomicName id="4C064D64F420FFD3FBB4F836FB647ED3" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[1075,1183,1980,2002]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="5" pageNumber="507" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F420FFD3FBB4F836FB647ED3" box="[1075,1183,1980,2002]" italics="true" pageId="5" pageNumber="507">D. carrolli</emphasis>
</taxonomicName>
pupae are more rounded on their sides.
</paragraph>
</caption>
<paragraph id="8BB936E7F423FFD0FF40FF1CFB54781C" blockId="6.[151,1436,151,933]" pageId="6" pageNumber="508">
<emphasis id="B972EAF5F423FFD0FF40FF1CFE2679B1" bold="true" box="[199,477,151,176]" pageId="6" pageNumber="508">Reproductive isolation.</emphasis>
Multiple attempts to cross 
<taxonomicName id="4C064D64F423FFD0FC9FFF12FC6B79B1" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[792,912,152,176]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F423FFD0FC9FFF12FC6B79B1" box="[792,912,152,176]" italics="true" pageId="6" pageNumber="508">D. carrolli</emphasis>
</taxonomicName>
(KB866) to 
<taxonomicName id="4C064D64F423FFD0FBA3FF12FB5579B1" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[1060,1198,152,176]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F423FFD0FBA3FF12FB5579B1" box="[1060,1198,152,176]" italics="true" pageId="6" pageNumber="508">D. rhopaloa</emphasis>
</taxonomicName>
(BaVi0067), in both directions, failed. Multiple replicate cultures consisting of 10–20 virgin females of one species, and 10– 
<specimenCount id="9D00FD6EF423FFD0FA9DFF37FA8579D4" box="[1306,1406,188,213]" pageId="6" pageNumber="508" type="male">20 males</specimenCount>
of the other species, were maintained for several weeks. Since no larvae were ever observed, several females per replicate were dissected, and the absence of sperm in their reproductive tracts was confirmed.
</paragraph>
</subSubSection>
<subSubSection id="C31C656CF423FFD0FF40FEACFEE278D1" pageId="6" pageNumber="508" type="description">
<paragraph id="8BB936E7F423FFD0FF40FEACFEE278D1" blockId="6.[151,1436,151,933]" pageId="6" pageNumber="508">
<emphasis id="B972EAF5F423FFD0FF40FEACFE3F7841" bold="true" box="[199,452,295,320]" pageId="6" pageNumber="508">Geographical range.</emphasis>
<taxonomicName id="4C064D64F423FFD0FE53FEA3FD437841" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[468,696,296,320]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F423FFD0FE53FEA3FD437841" box="[468,696,296,320]" italics="true" pageId="6" pageNumber="508">Drosophila carrolli</emphasis>
</taxonomicName>
is known from 
<collectingCountry id="F3117677F423FFD0FC0AFEACFB957841" box="[909,1134,295,320]" name="Brunei" pageId="6" pageNumber="508">Brunei Darussalam</collectingCountry>
at Kuala Belalong, Ulu 
<collectingRegion id="49C2F805F423FFD0FF10FEC7FEE37864" box="[151,280,332,357]" country="Brunei" name="Temburong" pageId="6" pageNumber="508">Temburong</collectingRegion>
National Park [
<geoCoordinate id="EE325020F423FFD0FE64FEC7FD797864" box="[483,642,332,357]" direction="north" orientation="latitude" pageId="6" pageNumber="508" precision="1" value="4.975111">4°58’30.40&quot;N</geoCoordinate>
<geoCoordinate id="EE325020F423FFD0FD14FEC7FCB77864" box="[659,844,332,357]" direction="east" orientation="longitude" pageId="6" pageNumber="508" precision="1" value="114.89102">114°53’27.68”E</geoCoordinate>
] (
<figureCitation id="133D2A62F423FFD0FCEDFEC7FC127865" box="[874,1001,332,357]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="6" pageNumber="508">Figure 1O</figureCitation>
), a locality that falls in the broad geographic range of 
<taxonomicName id="4C064D64F423FFD0FE02FEFAFDEB7889" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[389,528,368,392]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F423FFD0FE02FEFAFDEB7889" box="[389,528,368,392]" italics="true" pageId="6" pageNumber="508">D. rhopaloa</emphasis>
</taxonomicName>
. It was also collected from 
<collectingCountry id="F3117677F423FFD0FCD2FEE4FC447889" box="[853,959,367,392]" name="Malaysia" pageId="6" pageNumber="508">Malaysia</collectingCountry>
, 
<collectingRegion id="49C2F805F423FFD0FC4BFEE4FBE87889" box="[972,1043,367,392]" country="Malaysia" name="Sabah" pageId="6" pageNumber="508">Sabah</collectingRegion>
, Mt. Kinabalu, Poring [
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<geoCoordinate id="EE325020F423FFD0FF10FE1FFED278AC" box="[151,297,404,429]" direction="east" orientation="longitude" pageId="6" pageNumber="508" precision="15" value="116.69861">116°41'55&quot;E</geoCoordinate>
], approximately 
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.
</specimenCount>
above sea level (M.J. Toda 
<emphasis id="B972EAF5F423FFD0FCEEFE1FFC6878AD" box="[873,915,404,428]" italics="true" pageId="6" pageNumber="508">leg.</emphasis>
; Dr. Masanori Toda, personal communication; 
<figureCitation id="133D2A62F423FFD0FF10FE3CFEF078D1" box="[151,267,439,464]" captionStart="FIGURE 1" captionStartId="3.[151,250,1629,1651]" captionTargetBox="[227,1287,212,1577]" captionTargetId="figure@3.[170,1422,189,1603]" captionTargetPageId="3" captionText="FIGURE 1. Adult morphology and geographical range. (A–D) dorsal views of 5–7 day old adult Drosophila carrolli (A, male; B, female) and D. rhopaloa (C, male; D, female). (E, F) male abdomens of D. carrolli (E) and D. rhopaloa (F) in lateral (left) and ventral (right) views, showing variation in pigmentation intensity and testes color in mature specimens. Note the sharp lateral boundary of the darkly pigmented area on segment A5 and A6 in D. rhopaloa, but not D. carrolli (green arrows). Also note the bright yellow testes of D. carrolli, contrasting with the paler testes of D. rhopaloa (green arrowheads). (G, H) female abdomens of D. carrolli (G) and D. rhopaloa (H) in lateral views, showing variation in pigmentation intensity in mature specimens. (I, J) male wings of D. carrolli (I) and D. rhopaloa (J). Note the darker pigmentation at the anterior distal tip of of D. carrolli but not D. rhopaloa wings. (K, L) right antennae of males, showing no notable difference in the arista branching between D. carrolli (K) and D. rhopaloa (L), but a relatively shorter 3 rd antennal segment in D. rhopaloa (double arrows). (M, N) sex combs on the 1 st and 2 nd tarsal segments of forelegs in D. carrolli (M) and D. rhopaloa (N) showing no notable differences in teeth shape or pattern. Note that image on panel M is modified from Barmina &amp; Kopp (2007). (O) Map of South East Asia showing localities where D. rhopaloa (white circles) and D. carrolli (red circle) were found. The circle with dotted outline denotes an imprecise locality." httpUri="https://zenodo.org/record/1292134/files/figure.png" pageId="6" pageNumber="508">Figure 1O</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C31C656CF423FFD0FF40FE57FB427AA4" pageId="6" pageNumber="508" type="biology_ecology">
<paragraph id="8BB936E7F423FFD0FF40FE57FC1F7B85" blockId="6.[151,1436,151,933]" pageId="6" pageNumber="508">
<emphasis id="B972EAF5F423FFD0FF40FE57FE6778F4" bold="true" box="[199,412,476,501]" pageId="6" pageNumber="508">Ecology, ethology.</emphasis>
<taxonomicName id="4C064D64F423FFD0FE24FE56FDE378F5" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[419,536,476,500]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F423FFD0FE24FE56FDE378F5" box="[419,536,476,500]" italics="true" pageId="6" pageNumber="508">D. carrolli</emphasis>
</taxonomicName>
was collected in primary rainforest, sweeping over a banana bait. It must be rare locally, as of over 1000 isofemale strains established over two weeks of field work, only two were of this species, one of which (KB866) is the origin of the specimens used here to describe the species. No 
<taxonomicName id="4C064D64F423FFD0FB01FDAEFAF57B3D" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[1158,1294,548,572]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F423FFD0FB01FDAEFAF57B3D" box="[1158,1294,548,572]" italics="true" pageId="6" pageNumber="508">D. rhopaloa</emphasis>
</taxonomicName>
were caught at that location. At 20°C, it takes 8 days after egg deposition for 
<taxonomicName id="4C064D64F423FFD0FCEFFDC2FC247B61" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[872,991,584,608]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F423FFD0FCEFFDC2FC247B61" box="[872,991,584,608]" italics="true" pageId="6" pageNumber="508">D. carrolli</emphasis>
</taxonomicName>
to enter metamorphosis and the total of 15–16 days for the adult fly to emerge from pupa, exactly like 
<taxonomicName id="4C064D64F423FFD0FCD3FDE6FC267B85" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[852,989,620,644]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F423FFD0FCD3FDE6FC267B85" box="[852,989,620,644]" italics="true" pageId="6" pageNumber="508">D. rhopaloa</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BB936E7F423FFD0FF40FD04FE057BCD" blockId="6.[151,1436,151,933]" pageId="6" pageNumber="508">
The male courtship behavior of 
<taxonomicName id="4C064D64F423FFD0FDAAFD1AFD5F7BA9" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[557,676,656,680]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="carrolli">
<emphasis id="B972EAF5F423FFD0FDAAFD1AFD5F7BA9" box="[557,676,656,680]" italics="true" pageId="6" pageNumber="508">D. carrolli</emphasis>
</taxonomicName>
has been described in detail and compared to that of closely related species (
<bibRefCitation id="EF974B16F423FFD0FF71FD3FFE097BCD" box="[246,498,692,717]" pageId="6" pageNumber="508" refId="ref4665" refString="Setoguchi, S., Takamori, H., Aotsuka, T., Sese, J., Ishikawa, Y. &amp; Matsuo, T. (2014) Sexual dimorphism and courtship behavior in Drosophila prolongata. J Ethology, 32, 91 - 102. https: // doi. org / 10.1007 / s 10164 - 014 - 0399 - z" type="journal article">
Setoguchi 
<emphasis id="B972EAF5F423FFD0FEE8FD3EFE647BCD" box="[367,415,692,716]" italics="true" pageId="6" pageNumber="508">et al</emphasis>
., 2014
</bibRefCitation>
).
</paragraph>
<paragraph id="8BB936E7F423FFD0FF40FD5CFB427AA4" blockId="6.[151,1436,151,933]" pageId="6" pageNumber="508">
<emphasis id="B972EAF5F423FFD0FF40FD5CFE307BF1" bold="true" box="[199,459,727,752]" pageId="6" pageNumber="508">Molecular phylogeny.</emphasis>
Based on phylogenetic analysis of 12 nuclear and 2 mitochondrial loci, 
<taxonomicName id="4C064D64F423FFD0FA82FD52FA877BF1" authority="Gompel &amp; Kopp, 2018" authorityName="Gompel &amp; Kopp" authorityYear="2018" box="[1285,1404,728,752]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="carrolli" status="sp. nov.">
<emphasis id="B972EAF5F423FFD0FA82FD52FA877BF1" box="[1285,1404,728,752]" italics="true" pageId="6" pageNumber="508">D. carrolli</emphasis>
</taxonomicName>
<emphasis id="B972EAF5F423FFD0FA02FD5CFF437A14" bold="true" pageId="6" pageNumber="508">
<taxonomicNameLabel id="A241578EF423FFD0FA02FD5CFF437A14" pageId="6" pageNumber="508" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
belongs to the 
<taxonomicName id="4C064D64F423FFD0FEF3FD77FE217A15" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[372,474,764,788]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F423FFD0FEF3FD77FE217A15" box="[372,474,764,788]" italics="true" pageId="6" pageNumber="508">rhopaloa</emphasis>
</taxonomicName>
subgroup of the 
<taxonomicName id="4C064D64F423FFD0FD2FFD77FCBB7A15" box="[680,832,764,788]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="melanogaster">
<emphasis id="B972EAF5F423FFD0FD2FFD77FCBB7A15" box="[680,832,764,788]" italics="true" pageId="6" pageNumber="508">melanogaster</emphasis>
</taxonomicName>
species group, and may be the sister species to 
<taxonomicName id="4C064D64F423FFD0FA06FD76FDEA7A39" authority="Bock &amp; Wheeler, 1972" authorityName="Bock &amp; Wheeler" authorityYear="1972" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F423FFD0FA06FD76FF067A39" italics="true" pageId="6" pageNumber="508">D. rhopaloa</emphasis>
<bibRefCitation id="EF974B16F423FFD0FE80FC94FDEA7A39" box="[263,529,799,824]" pageId="6" pageNumber="508" refId="ref3963" refString="Bock, I. R. &amp; Wheeler, M. R. (1972) The Drosophila melanogaster species group. Studies in Genetics VII. University of Texas Publications, 7, 1 - 102." type="journal article">Bock &amp; Wheeler, 1972</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="EF974B16F423FFD0FDA5FC94FCC27A39" box="[546,825,799,824]" pageId="6" pageNumber="508" refId="ref3916" refString="Barmina, O. &amp; Kopp, A. (2007) Sex-specific expression of a HOX gene associated with rapid morphological evolution. Dev Biol, 311, 277 - 286. https: // doi. org / 10.1016 / j. ydbio. 2007.07.030" type="journal article">Barmina &amp; Kopp, 2007</bibRefCitation>
), within a clade that also contains the more distantly related and morphologically different species 
<taxonomicName id="4C064D64F423FFD0FD1AFCCEFBCC7A5D" authority="Gupta &amp; Singh, 1978" authorityName="Gupta &amp; Singh" authorityYear="1978" box="[669,1079,836,861]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="prolongata">
<emphasis id="B972EAF5F423FFD0FD1AFCCEFCC67A5D" box="[669,829,836,860]" italics="true" pageId="6" pageNumber="508">D. prolongata</emphasis>
Gupta &amp; Singh, 1978
</taxonomicName>
and 
<taxonomicName id="4C064D64F423FFD0FBF7FCCEFEE47A81" authority="Gupta &amp; Singh, 1977" authorityName="Gupta &amp; Singh" authorityYear="1977" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="kurseongensis">
<emphasis id="B972EAF5F423FFD0FBF7FCCEFAC97A5D" box="[1136,1330,836,860]" italics="true" pageId="6" pageNumber="508">D. kurseongensis</emphasis>
<bibRefCitation id="EF974B16F423FFD0FABDFCCFFEE47A81" pageId="6" pageNumber="508" refId="ref4732" refString="Singh, B. K. &amp; Gupta, J. P. (1977) Two new and two unrecorded species of the genus Drosophila Fallen (Diptera: Drosophilidae) from Shillong, Meghalaya, India. Proceedings of the Zoological Society, 30, 31 - 38." type="journal article">Gupta &amp; Singh, 1977</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="EF974B16F423FFD0FEA9FCECFDC67A81" box="[302,573,871,896]" pageId="6" pageNumber="508" refId="ref3916" refString="Barmina, O. &amp; Kopp, A. (2007) Sex-specific expression of a HOX gene associated with rapid morphological evolution. Dev Biol, 311, 277 - 286. https: // doi. org / 10.1016 / j. ydbio. 2007.07.030" type="journal article">Barmina &amp; Kopp, 2007</bibRefCitation>
; 
<bibRefCitation id="EF974B16F423FFD0FDCEFCECFCBA7A81" box="[585,833,871,896]" pageId="6" pageNumber="508" refId="ref4665" refString="Setoguchi, S., Takamori, H., Aotsuka, T., Sese, J., Ishikawa, Y. &amp; Matsuo, T. (2014) Sexual dimorphism and courtship behavior in Drosophila prolongata. J Ethology, 32, 91 - 102. https: // doi. org / 10.1007 / s 10164 - 014 - 0399 - z" type="journal article">
Setoguchi 
<emphasis id="B972EAF5F423FFD0FD47FCE2FD0A7A81" box="[704,753,872,896]" italics="true" pageId="6" pageNumber="508">et al</emphasis>
., 2014
</bibRefCitation>
) (see 
<tableCitation id="C684035CF423FFD0FC03FCECFC2C7A81" box="[900,983,871,896]" captionStart="TABLE 1" captionStartId="6.[151,239,1016,1039]" captionText="TABLE 1. List of species of the D. rhOPalOa subgroup and their geographical range." pageId="6" pageNumber="508">Table 1</tableCitation>
). No molecular information is available for 
<taxonomicName id="4C064D64F423FFD0FF39FC06FD4E7AA5" authority="Takada, Momma &amp; Shima, 1973" authorityName="Takada, Momma &amp; Shima" authorityYear="1973" box="[190,693,908,933]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="palmata">
<emphasis id="B972EAF5F423FFD0FF39FC06FEC67AA5" box="[190,317,908,932]" italics="true" pageId="6" pageNumber="508">D. palmata</emphasis>
<bibRefCitation id="EF974B16F423FFD0FEC3FC07FD4E7AA5" box="[324,693,908,933]" pageId="6" pageNumber="508" refId="ref4818" refString="Takada, H., Momma, E. &amp; Shima, T. (1973) Distribution and Population Constitution of Drosophila in South East Asia and Oceania: I. Drosophilidae at Mt. Kinabalu, East Malaysia (With 52 Text-figures). Journal of the Faculty of Science Hokkaido University, 19, 73 - 94." type="journal article">Takada, Momma &amp; Shima, 1973</bibRefCitation>
</taxonomicName>
, another species of the 
<taxonomicName id="4C064D64F423FFD0FC3DFC06FBB97AA5" authorityName="Bock &amp; Wheeler" authorityYear="1972" box="[954,1090,908,932]" class="Insecta" family="Drosophilidae" genus="Drosophila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diptera" pageId="6" pageNumber="508" phylum="Arthropoda" rank="species" species="rhopaloa">
<emphasis id="B972EAF5F423FFD0FC3DFC06FBB97AA5" box="[954,1090,908,932]" italics="true" pageId="6" pageNumber="508">D. rhopaloa</emphasis>
</taxonomicName>
subgroup.
</paragraph>
</subSubSection>
</treatment>
</document>