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20лови
Амурский зоологический журнал, 2022, т. XIV, № 2
Amurian Zoological Journal, 2022, vol. XIV, no. 2
www.azjournal.ru
https://www.doi.org/10.33910/2686-9519-2022-14-2-281-298 http://zoobank.org/References/1498EF7E-BC6B-427A-9CF3-C5F3E272492A
UDC 595.772
Notes on the taxonomy of species of Sciomyzini with a predominantly setulose anepisternum (Diptera, Sciomyzidae)
N. E. Vikhrev1H, W. L. Murphy2
1 Zoological Museum of Moscow University, 2 Bolshaya Nikitskaya Str., 125009, Moscow, Russia 2 Department of Entomology, National Museum of Natural History, Smithsonian, 7835 Tufton Str., Fishers IN 460382257, Washington, The United States of America
Authors Nikita E. Vikhrev
E-mail: nikita6510@yandex.ru
SPIN: 1266-1140
Scopus Author ID: 32467511100
William L. Murphy
E-mail: murph3000@gmail.com
Scopus Author ID: 35797311500
Copyright: © The Authors (2022). Published by Herzen State Pedagogical University of Russia. Open access under CC BY-NC License 4.0.
Abstract. The present paper was conceived originally as a straightforward, simple description of a new African taxon. It grew substantially as more research was conducted. It now consists of four parts. In Part I, we discuss the taxonomy of five species of Sciomyzini with a predominantly setulose anepisternum: Atrichomelina pubera, Ditaeniella grisescens, D. trivittata, Pherbelliapilosa, and Ph. shatalkini. In Parts II and III, we consider the three taxa initially included in the genus Ditaeniella sensu Rozkosny (1987) (D. grisescens, D. parallela, and D. patagonensis) and the taxon described herein, D. milleri sp. nov., analysing this group by use of both the classical morphological approach (Part II) as well as by analysis of molecular data from available DNA barcodes (Part III). In Part IV, we provide a key to the taxa of Sciomyzini worldwide that have a predominantly setulose anepisternum. We also provide a list of specimens examined, the data from which significantly supplement the information previously available regarding the distribution of several Palaearctic species.
Keywords: Diptera, Sciomyzidae, Ditaeniella, identification key, review, new species, synonymy
Таксономические заметки по видам Sciomyzini с большей частью покрытым волосками анэпистернумом (Diptera, Sciomyzidae)
Н. Е. Вихрев1Н, У. Л. Мёрфи2
1 Зоологический музей, МГУ им. М.В. Ломоносова, Большая Никитская ул., д. 2, 125009, г. Москва, Россия 2 Национальный музей естественной истории Смитсоновского института, 7835 Тафтон-стрит, Фишерс, 1Ы 46038-2257, г. Вашингтон, Соединенные Штаты Америки
Сведения об авторах Вихрев Никита Евгеньевич E-mail: nikita6510@yandex.ru SPIN-код: 1266-1140 Scopus Author ID: 32467511100 Уильям Л. Мёрфи E-mail: murph3000@gmail.com Scopus Author ID: 35797311500
Права: © Авторы (2022). Опубликовано Российским государственным педагогическим университетом им. А. И. Герцена. Открытый доступ на условиях лицензии СС БУ-ЫС 4.0.
Аннотация. Настоящая статья изначально задумывалась как простое описание нового африканского таксона. Однако по мере продвижения работы объем статьи существенно вырос, и теперь она состоит из четырех частей. В части I мы обсуждаем систематику пяти видов Sciomyzini с анэпистернумом, большей частью покрытым волосками: Atricomelina pubera, Ditaeniella grisescens, D. trivittata, Pherbellia pilosa и Ph. shatalkini. В частях II и III рассмотрены три таксона, первоначально включенных в род Ditaeniella sensu Rozkosny (1987) (D. grisescens, D. parallela и D. patagonensis), и описанный здесь вид D. milleri sp. nov. Виды Ditaeniella проанализированы с использованием как классического морфологического подхода (Часть II), так и путем анализа молекулярных данных по доступным ДНК-баркодам (Часть III). В части IV дан ключ к таксонам Sciomyzini с анэпистернумом, большей частью покрытым волосками. По рассмотренным таксонам приведен список изученных нами экземпляров, эти данные существенно дополняют ранее известную информацию о распространении нескольких палеарктических видов Sciomyzini.
Ключевые слова: Diptera, Sciomyzidae, Ditaeniella, ключ, обзор, новые виды, синонимы
Introduction
In 1995, Miller reported finding Ditaeni-ella in several localities in southern Africa (Botswana, Namibia, South Africa). He regarded African specimens as representing an undescribed species of Ditaeniella closely related to D. grisescens Meigen, 1830, writing, "A taxonomic study is nearing completion to establish the species' identities and relationships to the similar looking D. grisescens (in prep.)" (Miller 1995: 196). However, until now no description has been published of any species of Ditaeniella from Africa.
In December 2018, N. Vikhrev and M. Yan-bulat collected a series of specimens of an undescribed species of Ditaeniella near Windhoek, Namibia, in southern Africa. We describe the taxon herein as Ditaeniella milleri sp. nov. Our preliminary research included examination of all related taxa to ensure that the African taxon was indeed undescribed. This paper reviews the relevant characters of all taxa related to the new species, i.e., Scio-myzini with a predominantly setulose anepi-sternum.
Sack (1939: 37) described the genus Ditaeniella with D. grisescens (described in the genus Sciomyza) as the type and sole species. However, for many years thereafter, entomologists failed to acknowledge Ditaeniella as a valid taxon, continuing instead to consider D. grise-scens to be a member of the genus Pherbellia Robineau-Desvoidy, 1830 (note: for clarity we herein abbreviate the genera Pherbellia as Ph. and Pteromicra as Pt.). Steyskal (1963: 120) established the Pherbellia grisescens group on the basis of the unusual structure of the male genitalia. He included in the group Ph. grisescens (Palaearctic), Ph. parallela Walker, 1853 (as Ph. humilis Walker, 1853) (Nearctic), and Ph. patagonensis Macquart, 1851 (Neotropical). Rozkosny (1987) resurrected the genus Ditaeniella for species included in Steyskal's Pherbellia grisescens group and redescribed the genus briefly. Knutson et al. (1990: 484, table 1) provided a more detailed comparison of the characters of Ditaeniella and Pherbellia and transferred Pherbellia trivittata Cresson,
1920 (Nearctic) to Ditaeniella. Most recently, Murphy et al. (2018: 54-59) provided a detailed analysis of the history and taxonomy of Ditaeniella as well as extensive data regarding D. parallela and D. trivitatta, the two Nearc-tic species.
Rozkosny (1987: 18) listed 11 morphological characters of Ditaeniella: "Frons mainly matt, mid-frontal stripe elongated and tapered anteriorly, only one orbital seta present. Propleural seta distinct, prosternum haired, also mesopleuron covered with distinct hairs. Inner posterior margin of hind coxa with several hairs. Anal vein on wing reaching posterior wing margin. Male sternum 7 haired, gono-styli partly reduced and postgonites with conspicuous spines." Regarding Ditaeniella, he continued, "grisescens-group of the large genus Pherbellia is considered to be very isolated and regarded as a separate genus...." We cannot fully agree with Rozkosny because the Nearctic Atrichomelina pubera Loew, 1862 and the Palaearctic Ph. pilosa Hendel, 1902 share with Ditaeniella a majority of the morphological characters listed by Rozkosny (1987).
The present paper was conceived originally as a straightforward, simple description of a new African taxon. It grew substantially as more research was conducted. It now consists of four parts. In Part I, we discuss the taxonomy of five taxa of Sciomyzini with a predominantly setulose anepisternum: A. pubera, D. grisescens, D. trivittata, Ph. pilosa, and Ph. shatalkini Rozkosny, 1991. In Parts II and III, we consider the three taxa initially included in the genus Ditaeniella sensu Rozkosny (1987) (D. grisescens, D. parallela, and D. patagonensis) and the taxon described herein, D. milleri sp. nov., analysing this group by use of both the classical morphological approach (Part II) as well as by analysis of molecular data from available DNA barcodes (Part III). In Part IV, we provide a key to the taxa of Sci-omyzini worldwide that have a predominantly setulose anepisternum. We also provide a list of specimens examined, the data from which significantly supplement the information previously available regarding the distribution of several Palaearctic species.
Material and methods
Abbreviations of collections in which are deposited specimens that were borrowed for examination during this research are as follows:
USNM—U. S. National Museum of Natural History, Washington, D. C., U. S. A;
ZMUM—Zoological Museum of Moscow University, Russia;
ZIN—Zoological Institute, Saint Petersburg, Russia.
Localities are given as follows: country, region/state/province (in italics), and geographical coordinates in decimal-degree format. The full names of regions of Russian administrative subdivisions are an entangled result of political and historical events of no interest for zoology, so they are listed as name (taken from the English version of Wikipedia) and word "region" (abbreviated in the text as "Reg").
The following generally accepted abbreviations for morphological structures are used: f1, t1, f2, t2, f3, t3 = fore, mid, hind femur or tibia respectively; ac — acrostichal setae; dc — dorsocentral setae; a, p, d, v = anterior, posterior, dorsal, ventral seta(e). The abbreviation for the tarsi as tar followed by a pair of digits separated by a hyphen was proposed by Vikhrev (2011): the first digit (1 to 3) gives the leg number and the second digit (1 to 5) the number of the tarsal segment. For example, tar1-4 = 4th segment of fore tarsus; tar3-1 = hind basitarsus. Instead of using the term gonostyli (Rozkosny and other authors), we use the term surstyli.
Illustrations are original unless otherwise credited. When referring to figures, to avoid confusion we capitalise the first letter (Fig. or Figs.) for those appearing in this paper and use lowercase (fig. or figs.) for those published elsewhere.
COI sequences were taken from the Barcode of Life Data System (BOLD) (http:// boldsystems.org). We also ordered from the Syntol Company (Moscow, Russia) COI sequences of two specimens of D. grisescens and the paratype of D. milleri sp. nov. Genetic distances between the sequences were calculated
in MEGA (https://www.megasoftware.net/) using the Kimura 2-Parameter (Kimura 1980).
Part I. Species of Sciomyzini with a predominantly setulose anepisternum: Classical morphological approach
Atrichomelinapubera Loew, 1862 Sciomyza pubera Loew, 1862 Atrichomelina pubera Loew, 1862 (Cresson 1920)
Atrichomelina pubera Loew, 1862: redescription in Murphy et al. (2018: 48-52) Material examined: see specimens listed in Murphy et al. (2018: 50-51). New specimens examined: USA, Idaho, Moscow (46.7°N 117.0°W), 26.08.1912, J. M. Aldrich, 2$, 2? (with Steyskal's det. label Atrichomelina pubera Loew) (ZIN); Minnesota, Basswood Lake (48.0°N 91.6°W), R. Namba, 16.08.1950, (ZMUM).
Ditaeniella trivittata Cresson, 1920 Melina (Ditaenia) trivittata Cresson, 1920 Ditaeniella trivittata Cresson, 1920: Knutson et al. (1990: table 1)
Ditaeniella trivittata Cresson, 1920: redescription in Murphy et al. (2018: 58-59) Material examined: see specimens listed in Murphy et al. (2018: 59) and Murphy (2020: 9-10).
Remarks. (Sub)genus Ditaenia Hendel, 1902 based on length of midfrontal stripe seems to us quite artificial. (By the way, in D. grisescens the shape and colour of the midfrontal stripe are remarkably variable.)
According to Murphy et al. (2018: 58), D. trivittata often has two pairs of fronto-orbital setae. When present, the anterior pair is short and weak, often appearing as a socketed setula. Distribution. USA (14 states in central and southcentral regions).
Pherbellia czernyi Hendel, 1902 Specimen examined:
SERBIA: Crni Vrh env., 43.407°N 22.587°E, 800 m, N. Vikhrev, 1-7.07.2015, 1$ (ZMUM). Remarks. Described from the Austrian state of Upper Austria (Pfarrkirchen bei Bad Hall, 48.0°N 14.2°E), the type material is in Naturhistorisches Museum, Vienna. We ini-
tially had doubts about the validity of Ph. czernyi as a species separate from Ph. pilosa, but our Serbian specimen differs from all specimens of Ph. pilosa that we examined (listed below) by its much longer aristal hairs, which are longer than the width of the arista at its base; this character was noted in Hendel's (1902) original description of Ph. czernyi. Rozkosny (1991) offered as additional diagnostic characters the chaetotaxy of the fore coxa, the contrasting white tar1-1, and the shape of the surstyli. We examined a large series of Ph. pilosa Hendel, 1902 and found that the chaeto-taxy of the fore coxa and the colour of tar1-1 may be the same as offered diagnostic for Ph. czernyi. We did not examine genitalia of Ph. czernyi.
Examination of more European specimens is required to reach a conclusion regarding the status of this taxon as a species. Distribution. Known only from W Europe.
Pherbellia pilosa Hendel, 1902 Figs. 1, 2
Specimens examined:
BELARUS, Gomel Reg., Mozyr env., 52.02°N, 29.32°E: 20.05.2019, N. Vikhrev, (ZMUM);
RUSSIA: Astrakhan Reg., Baskunchak L., 48.16°N, 46.83°E, 1-6.05.2010, K. Tomko-
Kursk Reg., Selikhovy Dvory, (51.57°N 36.09°E), 12.05.2008, K. Tomkovich, 1? (ZMUM);
Khanty-Mansi Reg., Shapsha, 61.087°N, 69.442°E, 14-16.07.2010, K. Tomkovich, 1$; 61.09°N, 69.46°E, 15.08.2018, K. Tomkovich, 1$ (all ZMUM);
Mordovia Reg., Pushta village env., 54.71°N, 43.22°E: 18-22.05.2020, N. Vikhrev, 1$, 1? (ZMUM); 8-12.05.2020, N. Vikhrev, 5$, 6? (ZMUM and USNM); 22-24.05.2020, M. Yan-bulat, 10$, 10?; 1-5.09.2020, N. Vikhrev, 1$, 1? (all ZMUM);
Moscow Reg., Puschino (54.8°N, 37.6°E), 21.07.1984, M. Krivosheina, 1? (ZMUM);
Omsk Reg: Omsk, Ptichya Gavan, 54.97°N, 73.35°E, 24.06.2008, O. Kosterin, 1$; Cherlak env., 54.1°N, 75.0°E, 10.05.2009, O. Kosterin, 1$, 4? (all ZMUM);
Ryazan Reg., Oksky Natural Reserve (54.7°N, 40.8°E), 15.06.1965, V. Kovalev, 1? (ZMUM);
Voronezh Reg.: Voronezh, Botanical Garden (51.71°N, 39.23°E), Aksenenko, 7.05.2012, 1$, 2? (ZIN); near Khopyor R., 51.36°N, 42.05°E, 7.06.2012, N. Vikhrev, 4$, 3? (ZMUM). Remarks. Surprisingly, in the 120 years during which this species has been known, no one has reported that Ph. pilosa has hairs on the inner posterodorsal margin of the hind
vich, 3$, 4? (ZMUM); coxa.
Figs. 1-3. 1 — Ph. pilosa, ventral view, postabdomen showing anterior surstyli (at bottom), each with about 15 spinules; 2 — Ph. pilosa, male terminalia, lateral view, anterior surstylus with spinules (from Rozkosny 1991: fig. 49); 3 — Ph. shatalkini, ventral view, postabdomen showing anterior surstyli (at bottom), each with 7-8 spinules stronger than those of Ph. pilosa (from Kurina, Knutson 2019) Рис. 1-3. 1 — Ph. pilosa, постабдомен снизу, видны передние сурстили, каждый с примерно 15 шипиками; 2 — Ph. pilosa, терминалии самца, вид сбоку, передние сурстили с шипиками (по Rozkosny 1991: fig. 49); 3 — Ph. shatalkini, постабдомен снизу, видны передние сурстили, каждый с примерно 7-8 шипиками, которые сильнее, чем таковые у Ph. pilosa (по Kurina, Knutson 2019)
Distribution. Type locality: Danube R. near Vienna, Austria. W Palaearctic species extending to W Siberia to 75°E, north to 61°N, south to 48°N. More southerly records — Kyrgyzstan, Issyk-Kul Lake, and Near East (Rozkosny 1987) — require confirmation. Newly recorded for Belarus, south of European Russia and Western Siberia. Pherbellia shatalkini Rozkosny, 1991 Figs. 3, 4, 6
Type material: Holotype, $: RUSSIA: Amur Reg., Zeya env., (53.7°N, 127.3°E), A. Shatalkin, 13.09.1981. Paratypes: the same locality and collector as the holotype: 21.06.1978, 1$; 10.07.1978, 1$ (all ZMUM). Other specimens examined:
MONGOLIA, Arkhangai aimak (48°N, 101°E), 28.08.1967, V. Zaitsev, 1$ (ZIN).
RUSSIA: Khabarobsk Reg.: Khabarovsk, 48.6°N, 135.1°E, 2-6.06.2014, N. Vikhrev, 1$; Sindinskoe L., 48.92°N, 136.24°E, 7.06.2014, N. Vikhrev, 1$; Komsomolsk-on-Amur, Mylki L., 50.5°N, 136.98°E, 21.06.2022, N. Vikhrev, 1$ (all ZMUM);
Khakassia Reg., Beltirskoe env., 53.038°N, 90.462°E, 10.05.2018, N. Vikhrev, 2$ (ZMUM);
Tuva Reg.: Kyzyl env., 51.7°N, 94.7°E, 17-25.05.2018, N. Vikhrev, 3$, 2?; Uyuk R., 800 m, 52.07°N, 94.04°E, 27.05.2018, N. Vikhrev, 1? (all ZMUM).
USA, Alaska, Matanuska (61.4°N, 150.2°W), 8.10.1944, J. C. Chamberlin, 1$, 1 ? (? with G. C. Steyskal's identification label
Ph. griseola; both $ and ? with presumably K. Elberg's handwritten label "not griseola — hind 1/2 mspl. haired") (all ZIN). Remarks. Specimens from Alaska show that Ph. shatalkini is actually a Holarctic species, while the record of Ph. griseola (Steyskal 1954: 55) is misidentification. Steyskal (1954: fig. 1 and here Fig. 6) wrote, "A figure of the male terminalia of one of the Matanuska specimens is presented to facilitate eventual check against Palaearctic material." This case illustrates how difficult it is to use the male genitalia for identification. Steyskal's drawing of the posterior surstylus shows it to be similar to those of both Ph. shatalkini and Ph. griseola. His drawing of the anterior surstylus shows it to be more similar to that of Ph. shatalkini than to the "dog head profile like" shape of that of Ph. griseola. Illustrations of the genitalia of Ph. shatalkini from the original description by Rozkosny (1991: figs. 13-14) are shown in Fig. 4. Kurina et Knutson (2019) recently redescribed the male genitalia of Ph. shatalkini, noting that the anterior surstylus bears 7-8 rather strong spinules (Fig. 3). The anterior surstylus of Ph. griseola also has been depicted as spinulose (Rozkosny 1991: fig. 37 and our Fig. 5; Vala 1989: fig. 33c; Rivosecchi 1992: fig. 46b and tab. IXe). In fact, the anterior surstylus of Ph. griseola does not bear spinules but bears rather very fine and barely visible hairs on the inner side. To summarise, Steyskal (1954) drew the terminalia of Ph. shatalkini, while the presence of Ph. griseo-
Figs. 4-6. 4 — Ph. shatalkini, male postabdomen, ventral view, male terminalia, lateral view (from Rozkosny 1991: figs. 13-14); 5 — Ph. griseola, male terminalia, lateral view (from Rozkosny 1991: fig. 37); 6 — Ph. shatalkini (as Ph. griseola), male postabdomen, lateral and ventral views (from Steyskal 1954: fig. 1)
Рис. 4-6. 4 — Ph. shatalkini, постабдомен самца, вид снизу и сбоку (по Rozkosny 1991: figs. 1314); 5 — Ph. griseola, терминалии самца сбоку (по Rozkosny 1991: fig. 37); 6 — Ph. shatalkini (as Ph. griseola), постабдомен самца, вид сбоку и снизу (по Steyskal 1954: fig. 1)
la in the Nearctic requires confirmation. These species may be identified as follows:
— Anepisternum with weak setulae on most of it surface. Arista bare. Postpedicel mostly black. Crossveins only slightly darkened. S: anterior surstylus with strong spinules .......................... Ph. shatalkini
— Anepisternum with setulae along posterior margin only, stronger than those on anepisternum of Ph. shatalkini. Aristal hairs almost as long as half width of postpedicel. Postpedicel yellow. Crossveins distinctly darkened. S: anterior surstylus with fine
hardly visible hairs on inner surface.......
............................ Ph. griseola
Distribution. Holarctic species. In eastern Palaearctic known from 90°E to Far East (136°E), also S Korea (Son, Suh 2019); north to 53°N, south to 48°N. Alaskan specimens have been collected much farther north (61.4°N). Other Nearctic records require reexamination as follows from Remarks, above.
Rozkosny (1987: 18) described the genus Ditaeniella as follows: "Frons mainly matt, mid-frontal stripe elongated and tapered anteriorly, only one orbital seta present. Propleural seta distinct, prosternum haired, also mesopleuron covered with distinct hairs. Inner posterior margin of hind coxa with several hairs. Anal vein on wing reaching posterior wing margin. Male sternum 7 haired, gono-styli partly reduced and postgonites with conspicuous spines." He noted, "This combination of characters, the majority of which (especially 1 orbital seta, spines on postgonites, partly reduced gonostyli) belongs obviously to apotypic ones, is not known in any [other] group of the Sciomyzini." As we mentioned in the Introduction, distinguishing between Di-taeniella and other taxa of Sciomyzini with a predominantly setulose anepisternum is not as straightforward as it is with most other taxa of Sciomyzini.
We compare the diagnostic morphological characters of these five taxa in Table 1.
Table 1
Comparison of morphological characters of Ditaeniella with those of other taxa of Sciomyzini
with a predominantly setulose anepisternum*
Таблица 1
Сравнение морфологических признаков Ditaeniella с таковыми у других таксонов Sciomyzini с анэпистернумом, преимущественно покрытым волосками*
Ditaeniella Ditaeniella Àtrichomelina Pherbellia Pherbellia
Characters grisescens trivittata pubera piiosa shatalkini
Rozkosny"s generic diagnosis
midfrontal strip long, blunted tip long, blunted tip short, pointed tip short, pointed tip short, pointed tip
orbital setae one pair one pair one [two) pair two pairs two pairs
prosternum setulose setulose bare bare bare bare
anepisternum entirelly hairy partly hairy entirelly hairy entirelly hairy entirelly hairy
inner margin of hind coxa hairy hairy hairy hairy bare
S genitalia surstyli reduced, postgonites large surstyli small, cerci surstyli well developed, anterior
and with several spines with patch of setulae surstylus with spines
othe characters
minute hairs on meron usually present usually present usually present absent absent
arista bare short pubescent long pubescent short pubescent almost bare
strong setae on anepimeron present present absent present present
v setae on apex of f3 o present present absent absent absent
tarl-1 contarsting whitish ГЮ no yes yes rra
proepistemal seta strong strong hairlike strong strong
*Other taxa of Ditaeniella (D. parallela and D. milleri sp. nov.) are not included because they have the same morphology as D. grisescens. Pherbellia mikiana Hendel, 1900; Ph. czernyi, and Ph. seticoxa Steyskal, 1961 are not included because the authors have not personally examined specimens of those taxa. Comments to Table 1 appear below. *Другие таксоны Ditaeniella (D. parallela and D. milleri sp. nov.) не включены, поскольку они имеют ту же морфологию, что и D. grisescens. Pherbellia mikiana Hendel, 1900; Ph. czernyi и Ph. seticoxa Steyskal, 1961 не включены, поскольку авторы лично не исследовали экземпляры этих видов. Комментарии к таблице 1 приведены ниже.
1. The midfrontal stripe of Ditaeniella differs slightly in different specimens. It usually appears as a long sharp-angled triangle (Fig. 16). It is not as long and parallel sided as in Ph. cinerella but it is noticeably different from the short, pointed triangle found in more typical Pherbellia taxa.
2. Ditaeniella species have one pair (posterior) of fronto-orbital seta. Similarly, only one pair of fronto-orbital setae is present in Ptero-micra glabricula, in both species of Salticella, and in several taxa of Tetanocerini. In Atri-chomelina, a weak anterior pair of fronto-orbital setae are sometimes present.
3. Among Sciomyzini with a predominantly setulose anepisternum, a setose prosternum is unique in Ditaeniella sensu Rozkosny (bare in D. trivittata). A setose prosternum also is found in Sciomyza species and several taxa of Tetanocerini.
4. The anepisternal setulae are slightly less extensive in Ph. shatalkini and much less extensive in D. trivittata than in D. grisescens. In the Ph. dorsata group, the anepisternum is setulose along the posterior margin only. In the Tetanocerini, the presence of setulae or setae on the anepisternum is a genus-specific character.
5. In the Nearctic Ph. seticoxa, the hind coxa bears a few fine hairs on the inner posterior margin (Murphy et al. 2018: 65, 80). In the Tetanocerini, the presence of hairs on the inner posterior margin of the hind coxa is a genus-specific character.
6. Species of Ditaeniella have very similar genitalia that differ only in fine details (Figs. 7-9, 13, 15). Likewise, the general genitalic structure of Ph. pilosa and Ph. shatalkini is similar to that of each other (Figs. 1, 3). In contrast to these similarities, the genitalic ground plans of Ditaeniella, Atrichomelina, and taxa of Pher-bellia with a setulose anepisternum differ conspicuously from each other. Thus, the genitalic ground plan may be used as generic character, although applicable to male specimens only.
7. Microsetulae are present on the meron of species of Ditaeniella and Atrichomelina (Fig. 10). The diagnostic value of these micro-setulae is limited because they are not distinct
in all specimens and have been overlooked in previous published descriptions of species. The same microsetulae usually are present on meron of Ph. cinerella, and stronger setulae are present on the meron of species in the Ph. dorsata group.
8. As with Atrichomelina, some specimens of Ph. shatalkini lack strong seta(e) on the an-epimeron.
9. Males of Ditaeniella have two rows of strong, short setae on the ventral side of the hind femur, one anteroventral and one pos-teroventral, with no setae between the rows, only setulae (Figs. 11, 12, 17). The rows extend almost the full length of the femur and are denser on the apical half than on the basal half. Neither Atrichomelina nor species of Pherbellia with a predominantly setulose an-episternum have such rows of spinulose av and pv setae, although ventral setulae (longer or shorter, dense or sparse) may be present. Females of Ditaeniella usually have at least 1-2 av seta(e) near the apex of the hind femur, except for D. milleri sp. nov., which has 4-5 av and 2-4pv setae.
Conclusions. 1) Both sexes of the five species considered in Table 1 may be distinguished reliably from each other on the basis of external morphology, without examination of the genitalia. 2) A generic diagnosis of a group consisting of Ditaeniella, Atrichomelina, and species of Pherbellia with a predominantly setulose anepimeron is difficult to formulate; thus, we think it is most reasonable to retain the currently accepted division by genera until further results of molecular phy-logenetic analyses become available.
Part II. Taxa of Ditaeniella sensu Rozkosny, a classical morphological approach
Ditaeniella grisescens Meigen, 1830 Figs. 7, 14, 15
Sciomyza grisescens Meigen, 1830 Melina (Ditaenia) grisescens Meigen, 1830 (Cresson, 1920)
Pherbellia grisescens Meigen, 1830 (Malloch, 1933)
Ditaeniella grisescens Meigen, 1830 (Rozkosny, 1987)
Figs. 7-9. Postabdomen of Ditaeniella species in lateral view (modified from Steyskal 1963: figs. 8-10): 7 — D. grisescens; 8 — D. patagonensis; 9 — D. parallela
Рис. 7-9. Постабдомен таксонов Ditaeniella, вид сбоку (по Steyskal 1963: figs. 8-10): 7 — D. grisescens; 8 — D. patagonensis; 9 — D. parallela
Sciomyza patagonensis Macquart, 1851 = Pherbellia grisescens Meigen, 1830 (Malloch, 1933)
Specimens examined:
BELARUS, Minsk Reg., Borisov, Berezina R., 54.239°N, 28.494°E, D. Gavryushin, 5.07.2013, (ZMUM).
CHINA, Liaoning Prov., 50 km north of Mukden (= Shenyang = Fengtian) (42.3°N, 123.4°E), I. Rubtsov, 20.07.1952, 1? (ZIN).
HUNGARY, Domsod, Apaj-puszta, (47.1°N, 19.1°E), K. Gorodkov, 25.06.1970, 2? (ZIN).
IRAN: Isfahan Prov., Ghomrood R., 1920 m, 33.42°N 50.12°E, O. Kosterin, 21.05.2017, 2?; Markazi Prov., Arak env., 1700 m, 34.13°N, 49.71°E, O. Kosterin, 18-30.05.2017, 1$; Gav Godar env., 1800 m, 34.11°N, 49.36°E, O. Kosterin, 19.05.2017, 1$; Hendoudar R., 2000 m, 33.76°N, 49.22°E, O. Kosterin, 20.05.2017, 1$, 1?; Shazand env., 2000 m, (33.92°N, 49.40°E), I. Grichanov, 20.05.2017, 1$ (all ZMUM).
KAZAKHSTAN: Almaty Reg., Kapcha-gay Lake env., 43.7°N, 77.2°E, N. Vikhrev, 22-28.05.2016, 2$; Kyzylorda Reg., Bazykara env., 45.757°N, 62.311°E, fresh pond, K. Tom-kovich, 15-19.05.2011, 6$, 1?; Perovsk (= Kyzylorda) (« 44.75°N, 65.71°E), A. Zhe-lokhovtsev, 14.05.1925, 1$ (all ZMUM); Turkistan Reg., Saryagash Distr. (« 41.6°N, 68.7°E), P. Ler, 27.04.1957, 1$ (ZIN).
KYRGYZSTAN: Bishkek Reg., Bishkek Park, 42.90°N 74.62°E, N. Vikhrev, 17.09.2013, 2$, 1? (ZMUM); Issyk-Kul Lake, Rybachiy (= Balykchy) (42.46°N,
76.2°E), 1600 m, E. Nartchuk, K. Gorodkov, 16.08.1969, 4$, 2? (ZIN).
MONGOLIA: Dornod aimak, near Choi-balsan, 40 km upstream Kerulen R. (48.0°N, 113.9°E) I. Kerzhner, 27.07.1971, 5$, 1?; Omnogovi aimak, 20 km WSW Bayan-Ovoo (42.9°N, 105.8°E), I. Kerzhner, 8 August 1971, 1$, 1?; Tov aimak, Urga (= Ulaanbaatar), Tola R. (47.88°N, 106.90°E), P. Kozlov, 2930.06.1905, 1$; 5-7.07.1905, 2?; Uvs aimak, 50 km ESE of Ulangom, 49.95°N, 92.65°E, E. Narchuk, 7.08.1975, 1? (all ZIN).
RUSSIA: Altai Krai Reg., Zmeinogorsk Distr., Kolyvanskoe L. (« 51.37°N, 82.17°E), O. Kosterin, 8.09.2007, 1$, 2? (ZMUM);
Amur Reg., Zeya env. (53.7°N, 127.2°E), A. Shatalkin, 6.06.1978, 1? (ZMUM);
Astrakhan Reg., Baskunchak Salt Lake, 48.167°N, 46.830°E, near lake, K. Tomko-vich, 3-6.05.2010, 1$; 48.193°N, 48.813°E, fresh pond silt, K. Tomkovich, 1-2.05.2010, 1? (all ZMUM);
Buryatia Reg.: Ust-Kiran env., R. Chikoy (50.42°N, 106.83°E), Khomze, 29.07.1908, 1? (ZIN); Mondy env., 51.62°N, 100.91°E, 1960 m, 22-24.06.2021, N. Vikhrev, 1$ (ZMUM).
Crimea Reg., Koktebel env. (44.95°N, 35.25°E), Dyakonov, 27.08.1923, 1? (ZIN);
Irkutsk Reg.: Bratsk (« 56.29°N, 101.72°E), A. Monchadsky, 29.06.1956, 1?; (Olkhon L.) Peschanka (« 53.285°N, 107.58°E), P. Mikhno, 25.07.1925, 1? (all ZIN);
Kaliningrad Reg., Khrabrovo env., 54.88°N, 20.60°E, K. Tomkovich, 23.08.2013, 1?; 54.90°N, 20.65°E, fresh ponds, K. Tomkovich, 13-15.08.2013, 1$, 1? (ZMUM);
Karachay-Cherkess Reg., Bol. Zelenchuk R., 44.22°N, 41.72°E, 580 m, O. Kosterin, 6.07.2019, 1$, 1? (ZMUM);
Khanty-Mansi Reg., Khanty-Mansiysk Airport, 61.04°N, 69.11°E, fir forest, K. Gorod-kov, 28.08.1976, 4$, 4? (ZIN).
Krasnodar Reg.: Adler env., Imereti Lowlands (43.41°N, 39.95°E), N. Vikhrev, 2.10.2008, 2?; Gelendzhik Distr., Krinitsa env., Pshada R., 44.396°N, 38.342°E, K. Tom-kovich, 6-13.09.2009, 1$ (all ZMUM);
Krasnoyarsk Reg., Ergaki National Park, 1450 m, 52.84°N, 93.25°E, N. Vikhrev, 2729.06.2017, 1$ (ZMUM);
North Ossetia-Alania Reg., Mozdok Distr., Sukhotskoe (« 43.67°N, 44.44°E), A. Ozerov, 3.08.1988, 1? (ZMUM);
Omsk Reg.: Irtysh R. floodplain, oxbow of Zamarayka R. (54.964°N, 73.353°E), O. Kosterin: 14-18.07.2007, 1?; Krugloe L., 54.89°N, 73.36°E, O. Kosterin, 24.07.2012, 1$, 1?; Ptichya Gavan oxbow, 54.97°N, 73.35°E, O. Kosterin, 23.07.2008, 1? (all ZMUM);
Primorsky Reg.: Vladivostok, Sedanka (43.2°N, 132.0°E), A. Stackelberg, 20.06.1927, 1? (ZIN); Khanka Lake, 45.06°N, 131.99°E, N. Vikhrev, 15-19.07.2014, 2$, 1? (ZMUM);
Rostov Reg., Tsimlyansk env. (47.65°N, 42.10°E), O. Negrobov, 27.09.1965, 1$ (ZIN);
Saint Petersburg Reg.: Yukki (60.11°N, 30.28°E), A. Stackelberg, 27.08.1953, 6$, 9?; Luga Distr., Yaschera (59.15°N, 29.91°E), A. Stackelberg, 17.07.1963, 1? (all ZIN);
Tuva Reg., Dus-Khol Salt Lake, 700 m, 51.36°N, 94.45°E, N. Vikhrev, 2-5.07.2017, 1$ (ZMUM);
Tver Reg., Volgo L., 56.877°N, 33.244°E, A. Ozerov, 18-19.08.2011, 1? (ZMUM);
Ulyanovsk Reg., Radischevo Distr., Vjazov-ka env., 52.82°N, 48.35°E, K. Tomkovich, 2-5.05.2011, 1$ (ZMUM);
Yamalo-Nenetsk Reg.: Salekhard env., 66.6°N, 66.8°E, N. Vikhrev, M. Yanbulat, 16-19.07.2019, 4$, 4? (ZMUM); Muzhi vill. (65.4°N, 64.7°E), bank of Ob R., K. Gorodkov, 23.08.1976, 2$, 2? (ZIN).
Yaroslavl Reg., Berditsino (57.45°N 40.11°E), A. Yakovlev, 14.07.1907, 1$ (ZIN).
TAJIKISTAN: Dushanbe env. (38.5°N, 68.7°E), E. Gurjeva, 2.04.1962, 1?; Gorno-Ba-
dakhshan Reg., Shakhdara R. near Roshtqala (« 37.20N, 72.00E), E. Nartchuk, 15.07.1965, 1$ (all ZIN).
TURKEY: Ankara Prov., Karagol L. env., 1540 m, 40.3460N, 31.9290E, N. Vikhrev, 2.09.2009, 3$, 2?; Antalya Prov., Titreyen L., 36.7570N, 31.4550E, N. Vikhrev, 1-4.04.2006, 3$, 2?; 27-30.03.2007, 4$, 1?; 27.09.2007, 2?; 4.10.2007, 1$; A. Ozerov, 5.10.2007, 1? (all ZMUM); bank of Manavgat R., 36.87PN, 31.5270E, N. Vikhrev, 29 March 2007, 1?; A. Ozerov, 3.10.2007, 1$; Mersin Prov., near Silifke, 36.310N, 34.010E, N. Vikhrev, 22.04.2010, 2$, 1? (all ZMUM).
TURKMENISTAN, Lebap Prov., Charjou (= Turkmenabat) env. (« 38.760N, 64.200E), A. Ozerov, 25.04.1990, 1? (ZMUM); Da-soguz Prov., Tashauz (= Dasoguz, 41.80N, 60.00E), lower Amu-Daria River, Ushinsky, 21-24.04.1931, 3$, 3? (ZIN).
UKRAINE, Kherson Reg, Aleshky, lower Dnepr River (46.640N, 32.720E), L. Zimin, 26.07.1926, 1?; 18.08.1926, 1? (ZIN).
UZBEKISTAN: Tashkent Reg.: Karjantau Ridge (« 41.830N, 69.840E), K. Obukhova, 22.07.1938, 1?; 30.07.1938, 1$ (ZMUM); Chinaz, Syr-Darya R. (40.90N, 68.70E), F. Do-brzhansky, 28.04.1925, 1$ (ZIN); Xorazm Reg., Khiva (41.40N, 60.40E), L. Zimin, 2127.04.1927, 2? (ZIN).
VIETNAM, Lao Cai Prov., Sin Chai, 1400 m, 22.3380N, 103.8080E, A. Ozerov, 15.04.2012, 1? (ZMUM). Distribution. Remarkably widely distributed in the Palaearctic region from west to east, south to the Near East and Egypt, north to the Polar Circle; also recorded from North Vietnam. Has been recorded in the mountains up to 2000 m asl (Buryatia). Notes on synonymy of Ditaeniella patago-nensis. Malloch (1933) synonymised D. pa-tagonensis Macquart, 1851 with D. grisescens Meigen, 1830 after examination of material from Argentina, Chile and Uruguay. Steyskal (1963) refuted this synonymy and proposed D. patagonensis as a valid species. Steyskal's (1963: 121, figs. 6-10) drawings of male genitalia of species of Ditaeniella (Figs. 7-9) were provided with no explanation as to which dif-
ferences were diagnostically important. In the figures, the postabdomens of D. patagonensis and D. grisescens appear quite similar, while that of D. parallela differs slightly, for example, in D. patagonensis and D. grisescens the postgonites bear several spines, rather than a single spine as in D. parallela. Thus, Steyskal's drawings did not convince us that the differences between D. patagonensis and D. grisescens are significant. We agree with Malloch's synonymy that D. grisescens = D. patagonensis. If this synonymy is confirmed, it will turn out that D. parallela occurs in North America, whereas D. grisescens occurs in Eurasia and South America. The only reasonable explanation for this disjunct distribution would be that D. grisescens was introduced relatively recently from Eurasia to South America.
Ditaeniella milleri sp. nov.
https://zoobank.org/NomenclaturalActs/89FE532F-0687-49FD-B5B5-C5C49A51E5F0 Figs. 10, 11, 12, 13
Type material. Holotype $: NAMIBIA, Windhoek env., 22.545°S, 17.255°E, 1870 m, 11.12.2018, N. Vikhrev. Paratypes, 16$, 15$:
NAMIBIA, 15$, 14$, the same data as holotype;
ETHIOPIA: Amhara Reg., 10.87°N, 39.81°E, 1450 m, 8.08.2012, I. Gomyranov, 1$; Oromya Reg., Bale Mt, 3370 m, 7.088°N, 39.671°E, N. Vikhrev, 17.03.2012, 1$.
Type material is deposited in ZMUM apart from 1$, 1$ paratypes from Namibia in USNM.
Distribution. S Africa, Namibia, Botswana, Ethiopia.
Description. Male, body size: 3.8-5.2 mm.
Head. Frons matt yellow; fronto-orbital plates greyish-white; face and gena white; occiput yellowish white. Midfrontal stripe elongate, usually nearly reaching anterior margin of frons, tapered anteriorly; in dorsolateral view midfrontal stripe appears whitish matt, more pointed at apex, in dorsal view appears whitish brown with a blunted apex. Only one strong fronto-orbital seta (posterior). Gena half as wide as eye width. Antenna yellow ba-sally, postpedicel partly blackish. Arista bare even under high magnification. Palpus yellow.
Thorax. Entirely light-yellowish-grey or scutum dark grey with dirty-yellow pleura, latter colouration probably present in aged specimens. A pair of brownish vittae mesad of dorsocentral present on scutum. Prosternum with a pair of setulae (sometimes bare, rarely with 2 pairs setulae). Anepisternum
Figs. 10-11. Ditaeniella milleri sp. nov.: 10 — microsetulae on meron; 11 — male holotype Рис. 10-11. Ditaeniella milleri sp. nov.: 10 — волоски на мероне; 11 — голотип, самец
Figs. 12-15. Ditaeniella milleri sp. nov., male (12,13): 12 — ventral spines at apex of anterior femora; 13 — postgonites; Ditaeniella grisescens, male (14,15): 14 — ventral setae at apex of anterior femora; 15 — postgonites
Рис. 12-15. Ditaeniella milleri sp. nov., самец (12,13): 12 — вентральные шипы на переднем бедре; 13 — постгониты; Ditaeniella grisescens, самец (14, 15): 14 — вентральные шипы на переднем бедре; 15 — постгониты
evenly setulose. Thoracic setae: 1 strong pro-episternal, 1 postpronotal, 1 presutural supra-alar, 1 postsutural supra-alar, 2 notopleural, 2 postalar, 2 postsutural dorsocentral, 1 pres-cutellar acrostichal, and 2 pairs of scutellar setae. Subalar setae absent; anepimeron with 2 strong setae and 4-7 weak setulae; katepi-sternum setulose, without longer setae on upper margin; meron with 2-4 microsetulae (Fig. 10). Wing hyaline, without pattern.
Legs. Yellow, only apical half of t1 and fore tarsus dark. Fore coxa with 1 d seta at middle. f1: in basal 2/3 covered with dense fine ventral setulae; in apical third with dense rows of 7-8 pv and 4-6 weaker av spines, both pv and av spines much stronger than ventral spinulose setae present at apex of f1 in D. grisescens (see Figs. 12, 14). t1 with 1 dorsal preapical seta. f2 with typical "sciomyzid" a seta at middle. t2 and t3 without submedi-an setae. Inner posterior margin of hind coxa with 3 (2-4) setulae. f3: 2 pd in apical half; apical half with rows of 9-10 av and 8-9 pv spines similar to those of other species of
Ditaeniella (these spines in D. parallela are shown in Fig. 17); ventral area between rows of spines bare in apical third and covered with dense fine setulae in basal 2/3.
Abdomen. Yellow, with indistinct median vitta. Tergites densely setulose. Surstyli reduced to small protrusions; postgonites large and visible even on intact abdomen. Postgonites (Fig. 13) with external lobe with 2 longer and 2 shorter spines, although these spines are relatively shorter than those in D. grises-cens; inner spineless lobe tridentate and wider that in D. grisescens.
FEMALE differs from male as follows:
— body size 4.1-4.9 mm;
—f3 at apical third with 4-5 av and 4-5 pv spines;
— abdominal tergites less setulose than in male but setulae stronger.
Diagnosis. Similar to D. grisescens. f1: in basal 2/3 covered with dense fine ventral setulae; in apical third with dense rows of 7-8 pv and 4-6 weaker av spines, both pv and av spines much stronger than ventral spinulose setu-
lae present at apex of f1 in D. grisescens. Postgonites (Fig. 13) with external lobe with 2 longer and 2 shorter spines, although these spines are relatively shorter than those in D. grisescens; inner spineless lobe tridentate and wider that in D. grisescens. f3 at apical third with 4-5 av and 4-5 pv spines. Etymology. Named in honor of Ray M. Miller, who first found this genus in southern Africa (Miller 1995) and correctly identified his specimens as belonging to an undescribed species of Ditaeniella but never published a description of it.
Habitat. The Namibian series was collected near a riverbed that dries up completely during the dry season and overflows during the rainy season. In 2018, on November 26, heavy rain fell on this riverbed. When we visited this place on December 3, we did not find any Ditaeniella. We collected our series during our second visit on December 11. In January-February 2021, we returned to this place twice during the height of the rainy season. The riverbed was overflowing, but again we found no Ditaeniella.
Ditaeniella parallela Walker, 1853 Figs. 9, 16, 17, 18, 19 Sciomyza parallela Walker 1853 Sciomyza humilis Loew 1876 Sciomyza serena Wulp 1897 Melina (Ditaenia) grisescens Meigen, 1830 (Cresson, 1920: 49-50) Pherbellia parallela (Johnson 1925: 250) Ditaeniella parallela (Rozkosny 1987: 18) Ditaeniella parallela Walker, 1853: redescription in Murphy et al. (2018: 56-58) Material examined: see specimens listed in Murphy et al. (2018: 57-58) and Murphy (2020: 9). New specimens examined: USA: Texas: Laguna Madre, 25 km SE Harlingen (26.0°N, 97.6°W), D. E. Hardy, 25.01.1945, 2$ (with L. V. Knutson's determination label "Ph. humilis Loew" and G.C. Steyskal's determination label "Ph. grisescens") (ZIN); Brazos Co., College Station, 30.60°N, 96.35°W, 10.10.2015, V. Belov, 1$ (ZMUM).
Distribution. Widespread in USA and Canada, also known from Mexico and Dominican Republic.
Figs. 16-19. Ditaeniella parallela, male: 16 — dorsal view; 17 — ventral view with arrow indicating ventral spines on f3; 18 — postabdomen, ventral view, postgonites are visible, each bearing only a single spine; 19 — postabdomen, lateral view with arrow pointing to apically curved sclerotised portion of cerci
Рис. 16-19. Ditaeniella parallela, самец: 16 — вид сверху; 17 — вид снизу, стрелка указывает на вентральные шипики на f3; 18 — постабдомен снизу, видны постгониты, каждый из которых несет по одному шипику; 19 — постабдомен сбоку, стрелка указывает на склеротизированные выросты на церках
Remarks. Cresson (1920) and Sack (1939) regarded D. parallela as a synonym of D. grisescens on the basis of non-genitalic characters; we also found no non-genitalic differences. Steys-kal (1963) examined the genitalia of D. grisescens and D. parallela and published drawings of them, having found them to differ (Figs. 7, 9), but he failed to explain which differences in his drawings were diagnostically important. Males of D. parallela differ from those of D. grisescens by the structure of their genitalia: in D. parallela, the sclerotised portion of the cercus forms a flat paired plate, broadened and upcurved api-cally (Figs. 9, 19); and the postgonites each bear only a single strong spine (Fig. 18). Females of D. parallela are undistinguishable from those of D. grisescens.
As will be shown in Part III, the distance between COI sequences among Palaearctic specimens of Ditaeniella grisescens is much greater than that among Nearctic specimens. This suggests an Old-World origin of Ditaeniella. If so, D. parallela must have colonised North America via the Bering Land Bridge, which existed 30,000-11,000 years BPE.
Ditaeniella patagonensis Macquart, 1851
See: Ditaeniella grisescens: Notes on synonymy of Ditaeniella patagonensis.
Part III. Taxa of Ditaeniella sensu Rozkosny, analysis of molecular data
As follows from the Parts I and II (and the identification key in Part IV, below), the majority of Sciomyzini with a predominantly setulose anepisternum may be distinguished reliably by a set of morphological characters. On the other hand, taxa of Ditaeniella sensu Rozkosny differ only slightly from each other. Females of the Nearctic D. parallela are indistinguishable from those of D. grisescens, while males differ only minutely in the structure of the postabdomen. Historically, authors regarded D. paral-lela as a synonym of D. grisescens (Cresson 1920; Sack 1939), whereas present authors regard it as a valid species (Murphy et al. 2018; Steyskal 1963).
Males of the Afrotropical D. milleri sp. nov. have very similar genitalia to those of D. grisescens; both sexes differ only by stron-
ger ventral setae on fore- and hind femora. As follows from Part II, we see no reason to consider D. patagonensis as a valid species, and we have adopted Malloch's synonymy until results of further study elucidates the matter.
All species of Ditaeniella are allopatric, so we do not know if small differences (non-genitalic or genitalic) lead to reproductive isolation. We tried to clarify the status of taxa of Ditaeniella sensu Rozkosny using COI sequences of D. grisescens and D. parallela available in the Barcode of Life Data System (BOLD) (http://boldsystems.org). We also ordered from the Syntol Company (Russia, Moscow) COI sequences of two specimens of D. grisescens and the paratype of D. milleri sp. nov. To estimate the genetic variability of taxa of Ditaeniella, we compared those data with the COI variability of several other species of Scio-myzidae with a Holarctic distribution.
These Holarctic species were selected on the basis of two criteria: (1) easily recognisable, so erroneous identification was unlikely; (2) although most of the data in the BOLD database represent the North American fauna, at least one good COI sequence was available for the Eurasian specimen. We also included in our study Atrichomelina pubera with only a Nearctic distribution, because this taxon was considered in the present work.
The following COI sequences were analysed:
Atrichomelina pubera Nearctic (NA): TTDFW043-08, Canada, Saskatchewan; TTMDI1107-10, Canada, Ontario; DI PUR272-10, USA, Arizona; BBDIV183-12, USA, California;
Ditaeniella (as Pherbellia) parallela NA: BBDIP018-09, Canada, Alberta; DIPUS644-10, USA, Texas;
Ditaeniella grisescens Palaearctic (PA): GBMIN60863-17, France; MAMTJ157-12, Pakistan.
To evaluate the genetic differences between Nearctic and Palaearctic specimens of Ditaeniella, we calculated the same pairwise intraspecific distances in COI sequences of the following additional Holarctic species of Sciomyzidae:
Pherbellia albocostata NA: BBDEC763-09, Canada, Newfoundland and Labrador;
BBDCN381-10, Canada, British Columbia; AMCAF1462-19, Canada, Yukon; SSEIC6898-13, Canada, Alberta; KNWR008-11, USA, Alaska;
Ph. albocostata PA: GMFIF642-12, Finland; AMTPD2876-15, Germany; GMNWJ1970-14, Norway.
Ph. nana NA: UAMIC3305-15, USA, Alaska; UAMIC3307-15, USA, Alaska;
Ph. nana PA: GBMIN18482-13, Ireland.
Pteromicra angustipennis NA:
UAMIC3270-15, USA, Alaska;
Pt. angustipennis PA: AMTPD2668-15, Germany;
Tetanocera robusta NA: JWDCF297-10, Canada, Manitoba; CNNHA178-14, Canada, Northwest Territories;
T. robusta PA: VMDIP001-16, Norway.
Three more COI sequences were newly obtained:
Ditaeniella grisescens PA: Russia: Primor-sky Reg., Khanka Lake, 45.06°N, 131.99°E and Yamalo-Nenetsk Reg, Salekhard env., 66.6°N, 66.8°E;
D. milleri sp. nov.: Namibia, Windhoek env., 22.545°S, 17.255°E.
The COI secuences obtained in the course of this study were submitted to European Nucleotide Archive / Gene Bank and got the following accession numbers:
OW537104 (D. grisescens, Salekhard), OW537105 (D. grisescens, Lake Khanka) and ON331721 (D. milleri sp. n., as Ditaeniella sp.).
The Neighbour-Joining tree for studied taxa is shown in Fig. 20.
Genetical distances between the sequences were calculated in MEGA by use of the following parameters: Kimura 2-Parameter (K2P) distance model with pairwise deletion of gaps/missing data and inclusion of all substitutions (transitions and transversions).
As a first step, we calculated K2P distances for each species within groups of specimens collected in Nearctic or Palaearctic regions except when such groups consisted of a single specimen. These results are shown in Table 2. As a second step, we calculated K2P distances between five groups of specimens collected in Nearctic or Palaearctic biogeographic realms, the results are shown in Table 3. Of course,
Table 2
K2P distances of COI sequences within groups of specimens in the same biogeographic realm
Таблица 2
K2P расстояния последовательностей COI внутри особей из одного и того же биогеографического
региона
Table 3
K2P distances COI between groups of specimens collected in different biogeographic realms: Palaearctic
(PA), Nearctic (NA) and Afrotropical (AF)
Таблица 3
K2P расстояния последовательностей COI между особями из разных биогеографических регионов: Палеарктика (PA), Неарктика (NA) и Афротропика (AF)
Tabl 2 Tabl 3
Distances within groups specimen K2P Distances between groups
GROUPS in group distance from different biogeographic realms
Ditaeniella grisescens PA 4 1.4% D. grisescens PA D. milleri sp.n. AF
Ditaeniella milleri sp.n. AF 1 n/c Ditaeniella grisescens PA XXX XXX
Ditaeniella para!lela NA 2 0.3% Ditaeniella milleri sp.n. AF 3.0% XXX
Ditaeniella parallels NA 3.5% 3.2%
medium
Atrichomelina púbera NA 4 1.0% T. robusta NA 0.8%
Pherbellia albocostata NA 5 1.3% T. robusta PA low
Pherbellia albocostata PA 3 0.5% Pt. angustipennis NA 0.9%
Pherbellia nana NA 2 0.0% Pt. angustipennis PA low
Pherbellia nana PA 1 n/c Ph. nana NA 1.2%
Pteromicra angustipennis NA 1 n/c Ph. nana PA low
Pteromicra angustipennis PA 1 n/c
Tetanocera robusta NA 2 0.3% Ph. a/bocosfafa NA 10.5%
Tetanocera robusta PA 1 n/c Ph. albocostata PA high
54
83
100
100
92
100
100
50
98
60
60
100
66
lOO
57
60 100
100
70
Atrichomelina púbera US-Ariz Atrichomelina púbera US-Cal Atrichomelina púbera CAN-Sask Atrichomelina púbera CAN-Ont Pherbellia nana IRL Pherbellia nana USAALASKA1 Pherbellia nana USAALASKA2 Ditaeniella milleri NAMIBIA Ditaeniella parallela CAN-AB Ditaeniella parallela USA-Texas Ditaeniella grisescens PAK Ditaeniella grisescens FRA Ditaeniella grisescens RUS-Yamal Ditaeniella grisescens RUS-FarEast Pherbellia albocostata FIN Pherbellia albocostata GERM Pherbellia albocostata NORW Pherbellia albocostata CAN-YT Pherbellia albocostata CAN-NL Pherbellia albocostata USA-ALASKA Pherbellia albocostata CAN-BC Pherbellia albocostata CAN-AB Pteromicra angustipennis GER Reromicra angustipennis USA-ALASKA Tetanocera robusta CAN-NWT Tetanocera robusta NORW Tetanocera robusta CAN MB
Fig. 20. Evolutionary relationships of taxa. The tree was inferred using the Neighbour-Joining method. The percentage of replicate trees in which the associated taxa clustered together in the bootstrap test (1000 replicates) are shown next to the branches. The evolutionary distances were computed by use of the K2P method. The rate variation among sites was modelled with a gamma distribution. This analysis involved 27 nucleotide sequences. Codon positions included were 1st + 2nd + 3rd + Noncoding. All ambiguous positions were removed for each sequence pair (pairwise deletion option). The final dataset included 657 positions. Evolutionary analyses were conducted in MEGA X14 Рис. 20. Эволюционные взаимоотношения таксонов. Дерево сделано с использованием метода Neighbor-Joining. Процент повторяющихся деревьев, в которых связанные таксоны сгруппированы вместе в тесте bootstrap (1000 повторов), показан рядом с ветвями. Эволюционные расстояния были вычислены с использованием метода K2P. Вариабельность участков была смоделирована с использованием гамма-распределения. Этот анализ включал 27 нуклеотидных последовательностей. Включены все позиции кодонов (1-й + 2-й + 3-й + некодирующий). Все неоднозначные позиции были удалены для каждой пары последовательностей (опция попарного удаления). Окончательный набор данных включал 657 позиций. Эволюционный анализ был проведен в MEGA X14
these results are quite preliminary, as the comparison is made only for the COI gene fragment and because only a few sequences were examined. Nevertheless, some suggestions can be made:
1. The distance among four sequences from D. grisescens (Palaearctic) is much greater than that between sequences from two specimens of D. parallela (Nearctic). This suggests an Old-World origin of Ditaeniella.
2. The Nearctic and Palaearctic populations of Pherbellia nana, Pteromicra angustipennis and Tetanocera robusta are very similar.
3. In contrast, the Nearctic and Palaearctic populations of Pherbellia albocostata have very divergent COI sequences. Therefore, Nearctic and Palaearctic specimens of this species should be re-examined for morphological differences suggesting two species rather than one.
4. The sequences of taxa of Ditaeniella from the Nearctic (D. parallela), the Palaearctic (D. grisescens) and the newly described taxon from the Afrotropical region have a medium distance between them. This variability may correspond to either subspecies or species level. Taking into account that our results are preliminary, with results obtained only for the COI gene, we decided on a conservative approach so as to reduce to a minimum any changes in accepted taxonomy. Thus, if D. parallela is generally accepted as a valid species, then the new African taxon also should be regarded as a valid species: D. milleri sp. nov.
Part IV. Key to species of Sciomyzini with a predominantly setulose anepisternum, SS,
1. One (posterior) fronto-orbital seta (rarely very weak second (anterior) seta present). Midfrontal stripe long, with blunt tip; if stripe short with pointed tip, then proepi-sternal seta very weak. S: surstyli reduced or small, not spinulose.................2
— Two strong fronto-orbital setae. Midfron-tal stripe short, with pointed tip. S: anterior surstylus large, with strong spinules (Figs. 1, 3).................6 (Pherbellia)
2. Proepisternal seta very weak, hardly distinct from adjacent setulae. Aristal hairs longer than basal width of arista. tar1-1 whitish,
contrasting with colour of t1 and other segments of fore tarsus. Anepimeron densely covered with 20-30 setulae but without strong setae. Midfrontal stripe short, tip pointed. S: f3 ventrally with fine setulae Postgonites without spines ............Atrichomelinapubera Loew
— Proepisternal seta strong. Arista bare or with hairs much shorter than basal width of arista. tar1-1 not whitish, con-colorous with t1 and other segments of fore tarsus. Anepimeron with 4-7 set-ulae and 2 strong setae. Midfrontal stripe long, tip blunt. S: f3 ventrally with av and pv rows of strong, short setae, these rows extended almost the full length of the femur, denser in apical part. Postgonites with strong spines .......................... 3 (Ditaeniella)
3. Prosternum bare. Anepisternum sparsely setulose with longer setulae in a row along posterior margin. Arista with distinct although short hairs. Postpedicel entirely yellow. Crossveins clouded. Most abdominal tergites with dark spots (1 median, 2 lateral) appearing as 3 stripes. Body length 4.1-7.6 mm........D. trivittata Cresson
— Prosternum with several setulae on either side. Anepisternum evenly setulose, without row of longer setulae along posterior margin. Arista bare. Postpedicel mostly blackish, distinctly darker than scape. Crossveins not clouded. Abdominal ter-gites without stripes or spots but with pale posterior margins. Body length 3.5-5.2 mm ...........4 (Ditaeniella sensu Rozkosny)
4. f1 in basal 2/3 covered with fine, dense ven-
tral setulae; apical third with dense rows of 7-8 pv and 4-6 weaker av spines, both pv and av spines much stronger than ventral setae, which are present at apex of f1 in D. grisescens (see Fig. 12). S: Postgonites (Fig. 13) with external lobe with 2 longer and 2 shorter spines, although these spines are relatively shorter than those in D. grises-cens; inner spineless lobe tridentate and wider than in D. grisescens. f3 at apical third with 4-5 av and 4-5 pv short spinulose setae. Afrotropical.......D. milleri sp. nov.
— Apical third off1 with 3-5 pv and 2-4 av short and weak setae (Fig. 14). $: f3 at apical third with 0-3 av and without short
spinulose pv setae. Not Afrotropical ......
....................................... 5
5. $: sclerotised portion of cerci forming a flat, paired plate, broadened and upcurved apically (Figs. 9, 19). Postgonites (Figs. 9, 18) each bearing a single strong spine. Ne-arctic...............D. parallela Walker
— $: Cerci not matching above description (Fig. 7). Postgonites (Fig. 13) each bearing 2 strong and 1-2 weaker spines. Palaearctic south to border with Indomalayan realm, also South America (probably introduced; currently regarded as full species, D. pa-tagonensis).........D. grisescens Meigen
6. Inner posterior margin of hind coxa with setulae. Postpedicel yellow. Arista with short but distinct hairs. tar1-1 whitish to yellowish, lighter than colour of t1 and other segments of foretarsus. Anepisternum
densely setulose. $: anterior surstylus with about 15 strong spinules; posterior surstylus
small (Figs. 1, 2).........................
.......................Ph. pilosa Hendel
— Inner posterior margin of hind coxa bare. Postpedicel mostly dark. Arista virtually bare. tar1-1 concolorous with t1 and other segments of foretarsus. Anepisternum less densely covered with setulae than in Ph. pilosa, anterior half mostly bare. $: anterior surstylus with 7-8 strong spinules; posterior surstylus large (Fig. 3) ..............
.................Ph. shatalkini Rozkosny
Acknowledgements
We are especially grateful to Maria Yanbu-lat (Russia, Moscow) for her help in collecting and sorting the material used in this research. We thank Dr. Oleg Kosterin (Russia, Novosibirsk) and Dr. Eric Chapman (Kentucky, USA) for reviewing and correcting a preliminary version of this paper.
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For citation: Vikhrev, N. E. , Murphy, W. L. (2022) Notes on the taxonomy of species of Sciomyzini with a predominantly setulose anepisternum (Diptera, Sciomyzidae). Amurian Zoological Journal, vol. XIV, no. 2, pp. 281-298. https://www.doi.org/10.33910/2686-9519-2022-14-2-281-298
Received 6 March 2022; reviewed 12 April 2022; accepted 16 April 2022.
Для цитирования: Вихрев, Н. Е., Мёрфи, У. Л. (2022) Таксономические заметки по видам Sciomyzini с большей частью покрытым волосками анэпистернумом (Diptera, Sciomyzidae). Амурский зоологический журнал, т. XIV, № 2, с. 281-298. https://www.doi.org/10.33910/2686-9519-2022-14-2-281-298 Получена 6 марта 2022; прошла рецензирование 12 апреля 2022; принята 16 апреля 2022.
