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Description of a new species of the genus Rhagoletis Loew (Diptera: Tephritidae) from Nepal, with a key to species from Central and Southern Asia

Authors:
  • I.I.Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine

Abstract and Figures

The genus Rhagoletis is represented in the Oriental Region by four species known to occur in Nepal. A fifth species, Rhagoletis freidbergi n. sp., is described herein. It differs from all known congeners by the colour patterns on the abdominal ter-gites and wing. A key to known Central and southern Asian species of Rhagoletis (from the Caucasus to the Himalayas) is provided. The taxonomic position of "Rhagoletis bezziana" Hendel, 1931 is also discussed.
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DOI: 10.5281/zenodo.3362252; ISSN (online) 2224-6304
urn:lsid:zoobank.org:pub:939743D4-05FB-4911-8A40-CC57D7AE9316
This contribution is published
to honor Dr. Amnon Freidberg,
a scientist, a colleague and a friend,
on the occasion of his 75th birthday.
Description of a new species of the genus Rhagoletis Loew
(Diptera: Tephritidae) from Nepal, with a key to species
from Central and Southern Asia
&
I. I. Schmalhausen Institute of Zoology, NAS of Ukraine, Bogdan Chmielnicki St. 15,
01601 Kiev, Ukraine. E-mail: s.v.korneyev@gmail.com; valery.korneyev@gmail.com
ABSTRACT
The genus Rhagoletis is represented in the Oriental Region by four species known
to occur in Nepal. A fifth species, Rhagoletis freidbergi n. sp., is described herein.
It differs from all known congeners by the colour patterns on the abdominal ter-
gites and wing. A key to known Central and southern Asian species of Rhagoletis
(from the Caucasus to the Himalayas) is provided. The taxonomic position of
Rhago letis bezziana” Hendel, 1931 is also discussed.
Rhagoletis, Oriental Region, Nepal, bio di-
versity, identification key, new species, taxonomy.
INTRODUCTION
The genus Rhagoletis Loew of the tribe Carpomyini includes 73 valid species
from the Holarctic, Oriental and Neotropical Regions (Norrbom et al. 1999, with
ad ditions). Like other Carpomyini, Rhagoletis species breed in fruits, and some of
them are economically important pests of apples, cherries, tomatoes, blueberries
and walnuts (Boller & Prokopy 1976). The systematics and phylogeny of Rhago-
letis
Rohden dorf 1961; Bush 1966; Foote 1981; Norrbom 1989; Hernández-Ortiz 1993;
    
1999; Smith et al.     
been compiled by Foote (1981) for the Americas South of the United States, Bush
(1966) and Foote et al.

et al. (2017) for

known ex cept for two species assigned to this genus by Hendel (1931) and Hardy
(1964). Re cently, Ito (2011) described additional three species and provided a key
to the known Oriental Rhagoletis. He also noted that Rhagoletis bezziana Hendel,
1931 ap pa rently belongs elsewhere.
  

tory, Israel, a previously undescribed and unnamed species of Rhagoletis collected by
Dr Amnon Freidberg in Nepal was found and is described here in his honour. A key
to the known Central and Southern Asian species of Rhagoletis is also provided.
MATERIALSANDMETHODS
The following acronyms refer to collections housing specimens examined in
this study:
        

  
et al.
are given in millimetres. The body length of females includes the oviscape.
The wing and habitus photographs were taken using a Canon PowerShot A640

TAXONOMY
Rhagoletis Loew, 1862
Key to the species of Rhagoletis occurring in the mountains
of Central and Southern Asia

are also included in square brackets)
 
meron, and metatergite with brown or black spots) .........................................2
          
scutellum and parts of pleuron yellow) ..........................................................11
          
complete subapical band connected to anterior apical band (Fig. 7) .................
...................................................... [R. caucasica
 
variable .............................................................................................................3
 

spot-like anterior and posterior apical bands (Fig. 5). Abdominal tergites with
pair of round submedial spots ............ R. rohdendorfi 
 
abdominal tergites and wing pattern variable .................................................. 4
4 Abdominal tergites entirely orange, without black spots.................................5
  ............................................................10
5 Accessory costal band entirely lacking; subapical and apical bands fused (Fig.

88       
 ...........................
....................................................................................R. emiliae Richter, 1976
    
crossband in R. turanica (Fig. 11). Other characters variable ........................6
RHAGOLETIS  89
Figs 1–9: Rhagoletis wings: (1) R. meigenii; (2) R. nakaoi; (3) R. freidbergi n. sp.; (4) R. yasudai; (5) R.
rohdendorfi; (6) R. rumpomaculata; (7) R. caucasica; (8) R. samojlovitshae; (9) R. berberidis.
Scale 1 mm. Arrows point to key characters. Figs 2 and 4 redrawn from Ito (2011); Fig. 6,
et al. (2017), with permission.
1 2
9
8
6
4
7
5
3
discal apical
accessory anterior apical
subapical posterior apical
 
and 6 combined. Accessory costal band fused with anterior apical band; subapical
band widely separated from anterior apical band in cell r4+5 (Fig. 11) ..............
........................................................................R. turanica (Rohdendorf, 1961)
90       
Figs 10 19: Rhagoletis wings: (10) R. emiliae; (11) R. turanica; (12) R. quamplurima; (1315) R.
alternata (variation of pattern); (16) R. almatensis; (17) R. flavicincta; (18) R. flavigenualis;
(19) R. batava. Scale 1 mm. Arrows point to key characters. Fig. 12 redrawn from Ito (2011),
with changes; Figs 17et al. (2017), with permission.
10
19
17
15
13
11
18
16
14
12
 
variable .............................................................................................................7
  ............8
  ..............................................................................9
 
 ................................................................... R. alternata (Fallén, 1817)
  .............
..................................................................................R. quamplurima Ito, 2011
9 Subapical and anterior apical bands broadly connected (Fig. 8). Oviscape mainly

or connected to anterior apical band .....R. samojlovitshae (Rohdendorf, 1961)

 4+5 (Fig. 1).
Oviscape entirely black, conspicuously shorter than tergites 5 and 6 combined.
Accessory costal band always separated from anterior apical band ..................
.................................................................................. R. meigenii (Loew, 1862)

spots separated by yellow interspaces much wider than individual spot (Fig.

of vein R2+3 (Fig. 3) .............................................................. R. freidbergi n. sp.
 
bars medially separated by yellow interspaces much narrower than dark area
..............................................................R. samojlovitshae (Rohdendorf, 1961)

11 Accessory costal band present (Figs 2, 4, 7, 9), although sometimes fused with
subapical and apical bands in R. nigripes and R. berberidis). Other characters
variable ..........................................................................................................12
     
apical bands connected ..................................................................................20
12 Femora yellow. Discal band variable .............................................................13
           
narrower than discal band (or at most as wide as) (Figs 9, 16) .....................18
      
entirely within the dark field; posterior apical band present or absent .......... 14
       
      
markings. Abdominal tergites black, sparsely gray microtrichose, with yellow
bands along posterior margins ........................ R. rumpomaculata Hardy, 1964
14 Subapical and anterior apical bands connected in cell r2+3 (as in Figs 7, 9)
.......................................................................................................................15
        2+3 (Figs 2, 4)
.......................................................................................................................17
RHAGOLETIS  91
15 Discal band conspicuously narrower than hyaline interspace between it and

with Berberis ..................................................................................................16
 

Lonicera ......................................................... R. almatensis Rohdendorf, 1961

as long as, and surstyli longer than subcostal cell .............................................
................................................................R. magniterebra (Rohdendorf, 1961)
             
shorter, and surstyli hardly as long as subcostal cell .........................................
...................................................................R. chumsanica (Rohdendorf, 1961)

cell r2+3; but separated from subapical band along vein R4+5 (Fig. 2). Thoracic
pleuron anterior and posterior to anterior spiracle widely black. Abdominal

brown at posterior margin, medially wider, in males tergites also yellow on
 ...........
............................................................................................ R. nakaoi Ito, 2011
 4+5,
widely separated from subapical band along costa in cell r2+3 (Fig. 4). Thoracic
pleuron anterior and posterior to anterior spiracle widely with yellowish brown
         
   
female widely brownish yellow, with 3 black dots on anterior margin. Oviscape
shining black, as long as tergite 6 ......................................R. yasudai Ito, 2011
18 Scutellum entirely black .................................... R. nigripes Rohdendorf, 1961
  ...............................................................................19
19 Scutum black with four silvery microtrichose vittae; scutellum entirely yellow,
          
and entering into cell m (as shown on Fig. 16). Larvae in Lonicera and cherry
fruits ........................................................................ R. cerasi (Linnaeus, 1758)
     

entering into cell m (Fig. 9, arrow). Larvae in Berberis seeds ..........................
...............................................................................[R. berberidis
20 Anterior apical band entirely contiguous with apical wing margin (Fig. 17).
Larvae in Lonicera ........................................... R. flavicincta Enderlein, 1934
 
hyaline area (Figs 18, 19). Not associated with Lonicera .............................21
21 Femora yellow. Associated with Juniperus ...................................................22
 Juniperus ...............................................23
92       
22 Posterior surface of head completely yellow, at most with narrow black streaks
 
 .............................................................R. flavigenualis Hering, 1958
 
 ........................R. mongolica

Rhamnus ..........................................................
....................................................... [R. bagheera 
   
Hippophae ........................................
..................................................................................... R. batava Hering, 1958
Rhagoletis freidbergi n. sp.

LSID: urn:lsid:zoobank.org:act:B922E84D-5A2F-4A57-8854-43AC2E740E8F.
Etymology: This species is named in honour of Dr Amnon Freidberg, who collec-
ted the type specimen, in recognition of his contributions to the study of fruit flies
world wide.
Diagnosis: The new species can be easily separated from other Rhagoletis species
by the following combination of characters: thorax yellow with yellow microtrichose
mesonotum bearing black lyrate pattern; yellow setulae and black setae; abdomen

R. alternata, R. caucasica, R. meigenii, R. nakaoi,
R. rum pomaculata, R. rohdendorfi, R. samojlovitshae and R. yasudai in having a
mostly yellow body and wing pattern with five crossbands (Fig. 3), including sub-
basal, discal, accessory costal, subapical, and anterior apical bands (Fig. 1). Differing
from the other species as given in the key above.
This species is believed to belong in the group of species associated with Berberis
spp. (the meigenii group sensu R. samojlovitshae,
sharing the mostly reddish yellow body colour and first flagellomere apically poin-
ted, differing in having a lyrate black pattern on the mesonotum and abdominal
R. samojlovitshae, the mesonotum is at
most diffusely darkened to brown on the posterior half anterior of the scutum, and
the dark spots on the abdominal tergites are subrectangular and submedial, separated
by a yellow medial vitta.
Description: Female. Head
        
longer than wide, matt yellow, sparsely white microsetulose; frontal plates matt,
finely and sparsely yellow microsetulose. Face 1.2× as high as narrowest distance
between eyes; with low facial carina and shallow antennal grooves. Flagellomere 1
dorsoapically pointed, twice as long as wide; arista black except basal 0.2, brownish
RHAGOLETIS  93
yellow, micropubescent, longest rays much shorter than arista width at base. Occiput
entirely yellow, without dark pattern, silvery microtrichose posterior to medial
vertical seta and at margins of genae, sparsely brown setulose at eye level and pale
    
yellow, with short brownish yellow genal seta and several setulae posteroventrally,
0.27× as high as eye and 0.85× as high as flagellomere 1 length; anteroventrally

palp yellow, 0.75× as long as flagellomere 1. Frontal, orbital, ocellar, vertical and
postocular setae black: 3 frontal setae on left and 4 on right side; 1 reclinate orbital
seta; one ocellar seta, half as long as medial vertical seta; lateral vertical seta 0.75×
 
vertical seta. Postocellar and postvertical seta brownish yellow, 0.3× and 0.1× as
long as medial vertical seta, respectively.
20 23
2221
94       
Figs 2024: Rhagoletis freidbergi
mesonotum, dorsal; (23) abdomen, dorsal; (24) labels.
24
Thorax (Fig. 22) yellow, with black setae; scutum yellow setulose, densely white
microtrichose, with black lyrate pattern partly hidden by microtrichiae; scutellum

on each side; mediotergite entirely black, without median yellow vitta, sparsely
whitish microtrichose, subshining, with entirely bare posteromedial spot; 2 pairs of
short brown scapular setae; dorsocentral seta aligned with anterior supra-alar seta,
acrostichal prescutellar seta strong, at level of intra-alar seta; 2 pairs of scutellar
setae, apical one 0.8× as long as basal seta; 1 postpronotal, 2 subequal notopleural,
1 intra-alar, 1 postalar, 1 anepisternal, 1 katepisternal, and 1 anepimeral setae; scu-
tellum slightly convex.
Legs yellow, brown setose and setulose, non-thickened; hind femur subapically
 
surface.
Wing pattern as in Fig. 3. Subbasal band pale brown, broken into 3 spots: at hu-
me ral crossband, in base of cell br, and at Cu2 and A1 junction; discal band broad,
entire, reaching from pterostigma to middle of cell cu1; accessory costal band present;
subapical band complete; anterior apical band narrowly separated from subapical
band by hyaline spot at apex of vein R2+3; posterior apical band strongly reduced
to short brown stump connected to subapical band at vein R4+5 and small dark spot

Abdomen 
 
posterior margins; syntergite 1+2 mostly black, with yellow posterior margin (note:
base of abdomen is full of pitchy-black internal tissue obscuring actual colouration
         
wide yellow interspaces from lateral margins (Fig. 23). Oviscape dorsally convex,
pale brown, darkened basally and apically, brown setulose, ventrally 1.5× as long

mem brane with dorsal taeniae fused posteriorly; aculeus (not dissected, almost en-
tirely exposed) moderately narrow, in cross section more cylindrical than flattened.
Spermathecae not examined.
Male unknown.
Measurements: Female. Body length, 5.0 mm; wing length, 4.3 mm, costal cell
length, 0.9 mm; aculeus length, 0.8 mm; aculeus length / costal cell length, 0.9.
Holotype:Nepal:
Host plant: Unknown.
Remarks. This species is morphologically similar to the Central Asian R. sa moj-
lovitshae (Rohdendorf) and apparently belongs in the group of species as sociated
with Berberis spp. This group includes R. berberidis, R. caucasica, R. chumsa-
nica, R. kurentsovi (Rohdendorf), R. magniterebra, R. meigenii, R. rohden dorfi, R.
sa moj lovitshae, and corresponds to the meigenii group of species recognized by

RHAGOLETIS  95
a clearly monophyletic cluster supported by both molecular and larval and adult
mor pho logical data, as well as by the host-plant association. The included species
are mostly specialized for feeding within the seeds rather than fleshy part of the

body (except R. berberidis) and a moderately or (in R. magniterebra) very long,


DISCUSSION
The species of the genus Rhagoletis are most diverse in the Nearctic Region,
with certain groups of species occurring in the Neotropical and Palaearctic regions
et al. 2006;
Hulbert 2018), with local centres of diversity in mountainous areas, especially Cen-

now known, including R. freidbergi and the four species described from Nepal by

asl), where the climate is similar to Central Asian mountains at lower altitudes, and
the flora and fauna are related rather to those of the Palaearctic Region. Nothing
is known about their host plants and biology in general, but due to morphologi-
cal si milarity with Central Asian species, they may be associated with Berberis or
Rosa spp. Furthermore, R. quamplurima Ito shows strong similarity to, and few
rather insufficient differences from the widespread R. alternata and quite possibly
re presents a local population of it.
Rhagoletis bezziana Hendel, 1931 (replacement name for Zonosema dubium
Bezzi, 1913, non Johnson, 1903) from India (Uttar Pradesh, Naini Tal), as has been
al ready noted by Ito (2011), apparently does not belong to Rhagoletis. Indeed, it has
a narrow wing with a dark spot at the apex of cell bcu, which are characters more
com mon for species of the tribe Trypetini rather than for Carpomyini; some species
of these tribes can be classified based only on genital characters. This species is not
in cluded in the key above.
ACKNOWLEDGEMENTS

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      Rhagoletis Loew (Diptera, Trypetidae)
und verwandte Bohrffiegengattungen. Entomologicheskoe obozrenie 40

 &     
Rhagoletis. InFruit flies (Diptera: Tephritidae): Phy-
logeny and evolution of behavior
 &  
morphological characters in the subtribe Carpomyina (Diptera: Tephritidae). Israel Journal
of Entomology 35– 36
http://doi.org/10.5281/zenodo.3362080
 &    In  
Norr bom, A.L. (Eds), Fruit flies (Tephritidae): Phylogeny and evolution of behavior. CRC
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RHAGOLETIS  97
... Th e genus Rhagoletis Loew, 1862 belongs in the tribe Carpomyini and includes more than 75 described species of the true fruit fl ies occurring mostly in the Holarctic and Neotropical Regions and, to the lesser degree, in the Oriental Region (Smith & Bush, 1999;Korneyev & Korneyev, 2019). Many Rhagoletis species are economically important, including pests of apples, cherries, blueberries, and walnuts (Boller & Prokopy, 1976). ...
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Rhagoletis merzi sp. n., is described and illustrated based on specimens swept and reared from Juniperus sabina L. in Switzerland. A comparative review of Palaearctic species and a key to Palearctic and Nearctic species similar to R. merzi is provided. Based on DNA sequences from the COI, CAD, 28S, period, and AATS genes (4270 bp) of 92 isolates from two outgroup species (Anastrepha ludens, Euphranta canadensis), one species of Carpomya and 35 species representing most of species groups of Rhagoletis, a MrBayes analysis recovered a monophyletic lineage of Juniper-infesting species within a monophyletic cluster of R. fausta, R. batava, as well as the suavis, cingulata, pomonella, tabellaria and juniperina groups. Th e juniperina group includes both Nearctic (R. juniperina and undescribed forms) and Palaearctic species (R. fl avigenualis and R. merzi). Rhagoletis merzi is more similar to the Nearctic R. juniperina in both morphological characters (wing pattern, occiput, mesonotum and legs coloration, shape of male surstyli) and molecular sequences than to the Palearctic R. fl avigenualis.
... Th e genus Rhagoletis Loew, 1862 belongs in the tribe Carpomyini and includes more than 75 described species of the true fruit fl ies occurring mostly in the Holarctic and Neotropical Regions and, to the lesser degree, in the Oriental Region (Smith & Bush, 1999;Korneyev & Korneyev, 2019). Many Rhagoletis species are economically important, including pests of apples, cherries, blueberries, and walnuts (Boller & Prokopy, 1976). ...
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S. V. Korneyev (https://orcid.org/0000-0001-8599-7695) J. J. Smith (https://orcid.org/0000-0003-1492-3095) J. E. Frey (https://orcid.org/0000-0001-6628-8834) V. A. Korneyev (https://orcid.org/0000-0001-9631-1038) A new species of Rhagoletis (Diptera, Tephritidae) from Switzerland, with discussion of its relationships within the genus. Korneyev, S. V., Smith, J. J., Hulbert, D. L., Frey, J. E., Korneyev, V. A.-Rhagoletis merzi sp. n., is described and illustrated based on specimens swept and reared from Juniperus sabina L. in Switzerland. A comparative review of Palaearctic species and a key to Palearctic and Nearctic species similar to R. merzi is provided. Based on DNA sequences from the COI, CAD, 28S, period, and AATS genes (4270 bp) of 92 isolates from two outgroup species (Anastrepha ludens, Euphranta canadensis), one species of Carpomya and 35 species representing most of species groups of Rhagoletis, a MrBayes analysis recovered a monophyletic lineage of Juniper-infesting species within a monophyletic cluster of R. fausta, R. batava, as well as the suavis, cingulata, pomonella, tabellaria and juniperina groups. Th e juniperina group includes both Nearctic (R. juniperina and undescribed forms) and Palaearctic species (R. fl avigenualis and R. merzi). Rhagoletis merzi is more similar to the Nearctic R. juniperina in both morphological characters (wing pattern, occiput, mesonotum and legs coloration, shape of male surstyli) and molecular sequences than to the Palearctic R. flavigenualis.
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The cherry-infesting fruit fly Rhagoletis cerasi Loew is a significant commercial pest in Europe that has recently invaded North America. To date, it has been trapped only in Canada and northwestern counties of New York. It has the potential to spread further and threaten production and movement of cherry commodities. Timely diagnosis of the pest will facilitate surveys and quick response to new detections. Adult morphology of the pest is distinct from other flies in North America. However, when flies are significantly damaged on traps or the immature life stages are found in fruits, molecular methods of identification are important to confirm presence and host-use records. Other than DNA sequencing of genes from flies which takes over a day to complete, there are no timely methods of molecular identification for this pest. In this study, we report the first sequence record of the internal transcribed spacer 1 (ITS1) from R. cerasi and develop two diagnostic tests for the pest based on ITS1 differences among species in North America. The tests use loop-mediated isothermal amplification (LAMP) and multiplex, conventional polymerase chain reaction (mcPCR) technologies that target the same region of the R. cerasi ITS1 sequence. Both tests performed well when tested against collections of R. cerasi from North America and Europe, generating Diagnostic Sensitivity estimates of 98.4-99.5%. Likewise, the tests had relatively high estimates of Diagnostic Specificity (97.8-100%) when tested against Rhagoletis Loew species present in North America that also use cherry as a developmental host.
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Records of Pest Species, with Improved Keys. Korneyev, V. A., Mishustin, R. I., Korneyev, S. V.-Based on previously unpublished specimens from the collection of I.I.Schmalhausen Institute of Zoology (Kyiv), all the species of the tribe Carpomyini known to occur in Europe and Asia westwards of the Caspian Sea and Afghanistan, are listed. The melon fruit fly, Carpomya (Myiopardalis) pardalina (Bigot) is recorded for the first time from Ukraine and Europe, and Carpomya (s. str.) vesuviana (A. Costa, 1854) for Continental Ukraine, based on collection material. Carpomya (Goniglossum) liat (Freidberg, 2016) comb. n. is established as generic status of Goniglossum has not been accepted. New records and illustrated keys to species are given.
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Data on type specimens and new findings, and a key to the Middle Asian species of Rhagoletis are provided. Taxonomic statuses of Rh. cerasi fasciata Rohd, Rh. nigripes Rohd, stat. n. (= Rh. cerasi nigripes Rohd.), Rh. flavicincta End., Rh. obsoleta Hering and Rh. reducta Hering are discussed. Males of Rh. chumsanica and Rh. emiliae V. Richter are found; their morphological features are discussed. Rh. rohdendorfi sp. n. (type locality: Kyrghyzstan, Tash Kumyr) is described. The new species is similar to Rh. samoffovitshae Rohd. and Rh. berberidis Jermy, differing by small size, combination of entirely orange (without dark spots) scutum and black spotted abdomen a.nd presence of second apical crossband on its wing.
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Illustrated ley to genera and species of Tephritidae occurring in Far East Russia and bordering parts of China, Japan and Korean Peninsula. In Russian.
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Genus ACANTHIOPHILUS Acanthiophilus Becker 1908[374]: 136, Tetanocera walkeri Wol-laston (OD). [6600182] REFS—Bezzi 1924[472]: 139 (key to 3 spp. (obsolete) [AF]); Bezzi 1926[476]: 295 (key to 7 spp. (obsolete) [AF: South Africa]); Hendel 1927[2108]: 203 (key to 2 spp. [PA]); Kapoor 1993[2600]: 56 (key to 2 spp. [OR: India]).