Research Article |
Corresponding author: Jindřich Roháček ( rohacek@szm.cz ) Academic editor: Pavel Stoev
© 2020 Jindřich Roháček.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Roháček J (2020) Protanthomyza grimaldii sp. nov., a further member of the extinct subfamily Protanthomyzinae (Diptera, Anthomyzidae) from Baltic amber. ZooKeys 973: 1-15. https://doi.org/10.3897/zookeys.973.51435
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A new fossil species, Protanthomyza grimaldii sp. nov. (Diptera, Anthomyzidae), is described from Baltic amber (Eocene, 48–34 Ma) based on two (male and female) inclusions. It is the ninth species of the †genus Protanthomyza Hennig, 1965 and †subfamily Protanthomyzinae Roháček, 1998. Adult morphology of P. grimaldii sp. nov. revealed that the rich chaetotaxy of the thoracic pleuron, two anal veins and presence of the anteroventral process of the epandrium are plausibly shared by all species of Protanthomyza. Relationships of the new species, which belongs to a group lacking the ctenidial spine on the fore femur, are discussed.
Anthomyzid flies, adult morphology, Eocene, new extinct species, relationships, taxonomy, Tertiary
Fossil taxa of Anthomyzidae (Diptera) were reviewed by
Two amber pieces with 2 anthomyzid inclusions were examined. Abbreviations of amber sources and depositories:
The methods of preparation of amber stones with fly inclusions were described in detail by
The amber inclusions were examined, drawn and measured using two types of binocular stereoscopic microscopes (Reichert, Olympus). Legs were drawn on squared paper using a Reichert binocular microscope with an ocular screen. The specimens were either photographed by a Canon EOS 60D digital camera with Canon MP-E 65 mm 1–5× macro lens or by a Canon EOS 5D Mark III digital camera with a Nikon CFI Plan 10×/0.25NA 10.5mm WD objective attached to a Canon EF 70–200 mm f/4L USM zoom lens. The specimen photographed by means of the latter equipment was repositioned upwards between each exposure using a Cognisys StackShot Macro Rail and the final photograph was compiled from multiple layers (35) using Helicon Focus Pro 7.0.2. The final images were edited in Adobe Photoshop CS6. Some illustrations were drawn from these and some other macrophotographs and details were inked based on direct observation at higher magnification using a binocular microscope. Measurements: Six characteristics were measured – body length (measured from anterior margin of head to end of cercus, thus excluding the antenna), wing length (from wing base to wing tip), wing width (maximum width), index Cs3 : Cs4 (= ratio of length of 3rd costal sector : length of 4th costal sector), index r-m\dm-cu : dm-cu (= ratio of length of section between r-m and dm-cu on cell dm : length of dm-cu) and index r-m\dm-cu : CuA1 (= ratio of length of section between r-m and dm-cu on cell dm: length of apical portion of CuA1).
Morphological terminology follows that used in
A1 first anal vein (= anterior + posterior branch of cubital vein, CuA+ CuP)
A2 second anal vein (= first branch of anal vein, A1)
ac acrostichal (seta) (acr)
ar arista
avp anteroventral process of epandrium;
C costa
ce cercus
CuA1 cubitus (= fourth branch of media, M4)
dc dorsocentral (seta)
dm discal medial cell
dm-cu discal medial-cubital (= discal medial, dm-m) cross-vein
ep epandrium
f1 , f2, f3 fore, mid, hind femur
ha haltere
hu humeral (= postpronotal, pprn) (seta)
lbl labellum
M media (= first branch of media, M1)
mspl mesopleural (= anepisternal, anepst) (seta)
npl notopleural (seta)
oc ocellar (seta)
ors orbital (seta) (orb)
pa postalar (seta) (pal)
pk preapical kink
plp maxillary palpus
poc postocular (setulae)
ppl propleural (= proepisternal + proepimeral, prepst + prepm) (seta)
prs presutural (= presutural intraalar, ial) (seta)
prsc prescutellar acrostichal (seta)
pvt postvertical (= postocellar, poc) (seta)
R1 , R2+3, R4+5 1st, 2nd, 3rd branches of radius
r-m radial-medial cross-vein
S1–S8 abdominal sterna
sa supraalar (seta) (spal)
sc scutellar (seta) (sctl)
Sc subcosta
stpl sternopleural (= katepisternal, kepst) (seta)
T1–T8 abdominal terga
t1 , t2, t3 fore, mid, hind tibia
vi vibrissa (vb)
vte outer vertical (seta) (o vt)
vti inner vertical (seta) (i vt)
Order Diptera Linnaeus, 1758
Superfamily Opomyzoidea Fallén, 1820
Family Anthomyzidae Czerny, 1903
Subfamily Protanthomyzinae Roháček, 1998
Protanthomyza collarti Hennig, 1965; Baltic amber (Eocene).
Protanthomyza
sp. nov.:
The species is dedicated to Prof. Dr. David Grimaldi (New York, U. S. A.), the distinguished American dipterist and palaeontologist, in recognition of his valuable contribution to the knowledge of amber fossil flies, including acalyptrates (largely from Dominican amber).
Holotype
♂ labelled “Faszination Bernstein, Christel Hoffeins, Hans Werner Hoffeins” (framed on obverse), ”1040-5a Diptera: Acalyptratae, Anthomyzidae ♂” (handwritten by C. Hoffeins, on reverse), “Baltic amber, Russia: Kaliningrad region, Yantarny”, “obtained in May 2010 from Dr. Andrey Krylov, Kaliningrad, Russia”, and “Holotypus ♂, Protanthomyza grimaldii sp.n., J. Roháček det. 2020” (red label) [amber piece embedded in polyester resin, size 8.6 × 7.9 × 4.8 mm], deposited in
Russia: Kaliningrad region, Yantarny mine. Middle to Late Eocene, 48–34 Ma (cf.
Ocellar triangle delimited by a groove; frontal triangle not delimited; 1st antennal flagellomere normal, not enlarged; arista bare; 3 or 4 dc macrosetae; f1 without a ctenidial spine; male epandrium elongate and posteriorly tapered, with a robust, hook-like, curved anteroventral process; female T7 short; female S6 and S7 broadly transverse.
Male (Figs
Head
(Figs
Thorax
hardly narrower than head, brown to blackish brown, with shining mesonotum and duller pleural part. Mesonotum relatively convex, separated from scutellum by deep suture. Scutellum rounded subtriangular, wider than long, convex dorsally; postscutellum not visible. Thoracic chaetotaxy (see Figs
Protanthomyza grimaldii sp. nov., holotype male (Baltic amber) 5 head, left sublateral view 6 base of left wing 7 right mesopleuron, sternopleuron and pteropleuron, lateral view 8 abdomen and haltere, right lateral view 9 terminalia, right lateral view. Scale bars: 0.5 mm (5); 0.3 mm (6–8); 0.1 mm (9). For abbreviations see p. 4.
Legs
brown to ochreous, femora darkest, fore coxa pale ochreous. f1 lacking ctenidial spine; 5 (2 longer) distinct widely-spaced setae in posterodorsal row; setae in posteroventral row more numerous but short and weak. f2 with 2 or 3 anterior setae near middle (cf. Fig.
Wing
(Figs
Abdomen
(Figs
Genitalia. Epandrium (Figs
Female (Figs
Abdomen
(Figs
Postabdomen. T6 distinctly shorter than T5, transverse; T7 yet shorter and also narrower than T6. S6 strikingly broad and transverse, wider but shorter than S5 and apparently larger than adjacent T6 and laterally almost reaching the latter; S7 only half length of S6, strongly transverse and laterally meeting with sides of T7 (Fig.
Protanthomyza grimaldii sp. nov. was previously recognized as a new species by
Protanthomyza grimaldii belongs to a group of species lacking a ctenidial spine on the fore femur and both sexes are correctly keyed by
Based on the data presented above, it can be concluded that:
(1) P. grimaldii is the ninth named species of the genus Protanthomyza, making this genus the most species-rich fossil genus of Anthomyzidae.
(2) The description of a new Protanthomyza species extends the morphological diversity evident in the male and female terminalia of this genus. This adds to the previously documented wide morphological diversity among species seen in the head structures, chaetotaxies, and formation of the sclerites of the abdomen besides the male and female terminalia.
(3) Two anal veins (A1, A2) can continue to be treated as a subfamily/generic character for Protanthomyzinae and Protanthomyza, respectively.
(4) The chaetotaxy of the mesopleuron (usually with 3 posterior mspl macrosetae + numerous setulae more anteriorly) and sternopleuron (with 1 posterior stpl macroseta + setulae) seems to be rather uniform within the genus, although in a few species some setulae on the mesopleuron can be enlarged and/or one (usually the dorsal) mspl macroseta is reduced (cf. Roháček 1913a).
(5) The anteroventral process of the epandrium is a synapomorphic character of Protanthomyza. It is expected that this process will be found in all other species where the male is unknown, viz. in P. loewi, P. meunieri and P. presli.
I am very grateful to Mrs Christel Hoffeins (Hamburg, Germany) and Dr Michael von Tschirnhaus (Bielefeld, Germany) for making interesting amber specimens available for examination. Dr Andrey Krylov (Kaliningrad, Russia) is thanked for his continuing support with Baltic amber Diptera, especially acalyptrates, and Mr Peter Chandler (Melksham, England, UK) and both reviewers for their comments, improvements and language corrections of the manuscript. This research on fossil Anthomyzidae was supported by the Ministry of Culture of the Czech Republic, with institutional financing of the long-term conceptual development of the research institution (the Silesian Museum, MK000100595).