The Oldest Species of the Relic Extant Genus Mesochria from Eocene Fushun Amber of China (Diptera: Anisopodidae: Mycetobiinae)

Palaeontologia Electronica palaeo-electronica.org

The oldest species of the relic extant genus Mesochria from Eocene Fushun amber of China (Diptera: Anisopodidae: Mycetobiinae)

Ryszard Szadziewski, Jacek Szwedo, Elżbieta Sontag, and Bo Wang

ABSTRACT

A new species – Mesochria fani Szadziewski and Szwedo sp. nov. – is described from Eocene Fushun amber. This is the oldest record of the genus Mesochria, which includes 10 extant species living in the tropics of the Oriental and Afrotropical regions. Fossil records of the genus from Miocene Dominican amber (20 Ma) and Eocene Fus- hun amber (50 Ma) indicate that this extant group is relictual, and that during its evolution it had a wider, pantropical distribution.

Ryszard Szadziewski. Department of Invertebrate Zoology and Parasitology, University of Gdańsk, 59, Wita Stwosza Street, PL80-308 Gdańsk, Poland. [email protected] Jacek Szwedo (corresponding author). Department of Invertebrate Zoology and Parasitology, University of Gdańsk, 59, Wita Stwosza Street, PL80-308 Gdańsk, Poland. [email protected] Elżbieta Sontag. Department of Invertebrate Zoology and Parasitology, University of Gdańsk, 59, Wita Stwosza Street, PL80-308 Gdańsk, Poland. [email protected] Bo Wang. Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, No.39 East Beijing Road, Nanjing 210008, China; Steinmann Institute, University of Bonn, 53115 Bonn, Germany. [email protected]

Keywords: Diptera; Anisopodidae; amber; Fushun; pantropical; new species

Submission: 21 February 2015 Acceptance: 27 March 2016

INTRODUCTION 4-12 mm), with a small, rounded head and short mouthparts. The compound eyes are dichoptic or Nematoceran dipterans of the family Anisopo- holoptic. Ocelli are present and form an equilateral didae Knab, 1912 (wood gnats, window-gnats) are triangle. The slender antennae with 14 flagel- common flies in forests, feeding on flowers or sap lomeres vary from relatively short to longer than exuding from trees. They are found on all conti- the head and thorax together. The wings are wide, nents except Antarctica; they are less numerous on with two or three median branches, sometimes islands. Adults are small to medium-sized (usually with distinct stigma. The legs are relatively long,

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Szadziewski, Ryszard, Szwedo, Jacek, Sontag, Elżbieta, and Wang, Bo. 2016. The oldest species of the relic extant genus Mesochria from Eocene Fushun amber of China (Diptera: Anisopodidae: Mycetobiinae). Palaeontologia Electronica 19.1.12A: 1-11 palaeo-electronica.org/content/2016/1453-mesochria-from-fushun-amber

with distinct tibial spurs. Larvae are saprophagous, taxon ascribed to Anisopodidae from Fushun usually living in decaying or fermenting organic amber – Eoanisopodites fushunensis Hong, 2002 – matter. The family Anisopodidae comprises 25 appears to be an unavailable name, as this species genera with 196 species, including several was proposed after 1999 without any mention of described fossil taxa (Oosterbroek, 2006; Thomp- the type depository (ICZN, 1999; Evenhuis, 2014). son, 2006; Pape et al., 2011; Lukashevich, 2012). The traditional subfamily Mycetobiinae With the transferral of Eopleciidae Rohdendorf, Crampton, 1924 is often treated as a separate fam- 1946 and Sinotendipedidae Hong and Wang, 1990 ily Mycetobiidae of worldwide distribution, with 5 to Anisopodidae (Ansorge and Krzemiński, 1995; genera and about 20 extant species. Members of Pape et al., 2011), as well as the genus Tega Bla- the subfamily are robust, small flies, 4-7 mm long, goderov et al., 1993 from Cramptonomyidae Hen- with a transparent wing membrane, d-cell absent, nig, 1969 (Lukashevich, 2012), the fossil record of and an M vein with 2 branches. The head has three this family is represented by 18 fossil genera, 3 ocelli arranged in a triangle on the frons. The extant genera and 47 species (Evenhuis, 2014). antennae are relatively short, with 14 flagel- The internal subdivisions of Anisopodidae are still lomeres. The proboscis is well-developed, and the a matter of discussion (Amorim and Grimaldi, 4-segmented palpus short. The thorax is convex. 2006; Oosterbroek, 2006; Thompson, 2006; Pape The claws are similar on all the legs; the small, et al., 2011; Lukashevich, 2012). simple, empodium is reduced. Larvae usually are The oldest fossil of the Anisopodidae – Meso- found in moist habitats, in rotting or fermenting rhyphus rhaeticus Rohdendorf, 1962 – comes from organic matter, and are common in exuding tree- the Ak-Bulak-Say locality, Sogyuty, near Lake sap and rot holes (Krivosheina, 1997; Oosterbroek, Issyk-Kul, Dzhil Formation, Kyrgyzstan, which is 2006; Thompson, 2006). aged Hettangian/Sinemurian, Early Jurassic The purpose of the present paper is to (Rohdendorf, 1962; Rasnitsyn and Zherikhin, describe the oldest species of the extant genus 2002; PaleoBioDB, 2016). Various extinct genera Mesochria Enderlein, 1910, from early Eocene are reported from Jurassic and Cretaceous depos- Fushun amber. The only fossil species of its of Asia and Europe (Evenhuis, 1994, 2014). A Mesochria known so far was reported from Mio- few species are known from Eocene Baltic amber, cene Dominican amber (Grimaldi, 1991). but also from sedimentary deposits in England and North America (Evenhuis, 1994, 2014). The Mio- MATERIAL AND METHODS cene record consists of findings in fossil resins of The Fushun amber specimen comes from the the New World, as well as in sedimentary deposits layers of the West Opencast Coalmine (Xilutian from North America and Europe (Evenhuis, 1994, Opencast Coalmine). The West Opencast Coalm- 2014). The general chronological and stratigraphic ine (41°50’ N, 123°54’ E), located to the south of distribution pattern of Anisopodidae illustrates its Fushun City, Liaoning Province, China (Figure 1.1), long and complicated evolutionary history. is the largest opencast coalmine in Asia, but after Some enigmatic taxa placed in Anisopodidae more than 110 years of mining, it is now exhausted (= Rhyphidae Newman, 1834) were described from and being reclaimed (Figure 1.3, 1.4). The coal and Fushun amber by Hong (1981). A new genus and oil shales are found in a relatively small east–west- new species – Caloneura plectilis Hong, 1981 – trending exposure of Mesozoic and Cenozoic sedi- was proposed, with a strange wing venation, mentary and volcanic rocks surrounded by Pre- resulting from its misinterpretation, as one wing cambrian terrain consisting mainly of granitic overlapped the other one (Hong, 1981, p. 91, fig- gneiss (Johnson, 1990; Wu et al., 2002). These ures 96 and 97-IV). Subsequently, Evenhuis (1994, continental sequences consist of swampy to fluvio- 2014) replaced the preoccupied generic name with deltaic and tuffaceous sediments that were depos- Hongocaloneura Evenhuis, 1994: 282 and trans- ited in the basin during the early Palaeogene ferred it to unplaced nematocerous flies. Hong (Hong et al., 1980; Johnson, 1990; Yang and Li, (2002) claimed that his interpretation of the wing 1997; Wu et al., 2002). The mining district in the venation was correct and established the new fam- Fushun basin contains the Palaeocene Laohutai ily Hongocaloneuridae Hong, 2002 in the Anisopo- and Lizigou formations and the Eocene Guchengzi, doidea Knab, 1912. Later, Zhang (2007) Jijuntun, Xilutian, and Gengjiajie formations (Figure commented that the wing venation of Hongocalo- 1.2). The amber-bearing Eocene Guchengzi For- neura plectilis, with four branches of M, had never mation includes thick coal beds intercalated with been reported among known Diptera. Another

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FIGURE 1. 1, Fushun fossil site location and palaeogeographic setting of north-eastern China during the Eocene; Palaeotopographic reconstruction after Wang H. (1995); 2, stratigraphic sequence of the West Opencast Coalmine. The yellow heptagons indicate amber-bearing layers in upper section of the Guchengzi Formation. PETM – Palaeo- cene-Eocene Thermal Maximum (approximately 55 Ma); EECO – Early Eocene Climatic Optimum (51–53 Ma); MECO – Middle Eocene Climatic Optimum (approximately 41.5 Ma); 3, View of the West Opencast Coalmine, from the east, 3 September 2013; arrows indicate amber-bearing strata; 4, View of reclamation works in the West Open- cast Coalmine from the west, 5 September 2014. carbonaceous shale (Wu et al., 2001; Meng et al., al., 2011), palaeomagnetic considerations and iso- 2012, Wang, B. et al., 2014). topic dating (Zhao et al., 1994; Quan et al., 2011). The West Opencast Coalmine was the largest The palaeoclimate in the early Eocene in Fushun is amber deposit in China. The Fushun amber-bear- reconstructed as moist subtropical, with variability ing strata were the middle and upper coal beds of in seasonal precipitation (very probably a mon- the Guchengzi Formation, assigned to an early- soonal system), with dry and wet seasons (Quan et middle Ypresian age (51–55 Ma) and constrained al., 2011, 2012; Wang, D., et al. 2013). The esti- by the palaeofloras (Wang, Q. et al., 2010; Quan et mated mean annual temperature varied between

3 SZADZIEWSKI ET AL.: Mesochria FROM FUSHUN AMBER

15–21°C; the coldest month mean temperature mize light reflections during further investigation. varied between 9–14°C and the warmest month The specimen was examined under Olympus SZ11 mean temperature varied between 19–25°C. The and SZ12 stereomicroscopes and an Olympus mean annual precipitation is estimated as having BX43 microscope. Photographs were taken with a varied between 650–1500 mm (Wang, Q. et al., Nikon DXM1200 digital camera connected to the 2010; Quan et al., 2011, 2012). These estimations above stereomicroscopes and with a Zeiss Discov- concerns Guchengzi Formation and are based on ery V20 system. floral elements (palynofloras) preserved with use of According to Recommendation 40A of the the principle of coexistence approach (CA; Mos- Code (ICZN, 1999) the name of family is correctly brugger and Utescher, 1997) based on the concept cited as ‘Anisopodidae Knab, 1912’. We follow of “nearest living relative (NLR) philosophy”; Quan here the proposition by Thompson (2006) to treat et al. (2012) deals also with leaf physiognomical Mesochria (including Neomesochria Amorim and analyses for the reconstructions and habitat condi- Tozoni, 1994) in the traditional concept as defined tions discussed. The climate was slightly cooler by the fusion of wing veins R2+3 to R1. Morphologi- than that of coeval Europe (Mosbrugger et al., cal terms are adopted from Thompson (2006). 2005; Zachos et al., 2008, Wang, Q. et al., 2010). The colour of Fushun amber could vary from SYSTEMATIC PALAEONTOLOGY honey, light yellow, transparent, through various Order DIPTERA Linnaeus, 1758 varieties of orange, cherry and cognac, translucent Suborder NEMATOCERA Duméril, 1805 to opaque, patchy and non- transparent forms of Infraorder BIBIONOMORPHA Hennig, 1954 dappled earth-amber. This variety is also related to Family ANISOPODIDAE Knab, 1912 the depositional conditions. The amber pieces are Subfamily MYCETOBIINAE Crampton, 1924 usually small, seldom longer than 10 cm. Fushun Genus MESOCHRIA Enderlein, 1910 amber is naturally autoclaved as the amber-bearing coal deposits were affected by a massive intru- Type species. Mesochria scottiana Enderlein, sion of basic magmas in the Oligocene (Wu et al., 1910: 65, by original designation. 2000; Wang, X. et al., 2001) so that the amber was Mesochria fani Szadziewski and Szwedo sp. nov. heated during diagenesis, like Burmese amber (Shi zoobank.org/A59ADD22-AF4B-4033-A0BE-AA0547449AA5 et al., 2012). The Fourier transform infrared (FTIR) spectrum of Fushun amber shows characteristics Etymology. The specific name is dedicated to Mr. of cupressaceous-araucarian resins (Tappert et al., Yong Fan, the director of the Fushun Amber Insti- 2013), which suggest Cupressaceae (especially tute, in recognition of his important contribution to Metasequoia) as the mother plant of the resin. the study of Fushun amber. Fragments of Metasequoia are not only the most Material examined. Holotype male, NIGP156988, common plant inclusions in Fushun amber but also Fushun amber. Deposited in the collections of the very frequent as compressions in the amber-bear- Nanjing Institute of Geology and Palaeontology, ing and succeeding layers, supporting this interpre- Chinese Academy of Sciences, Nanjing, China. tation (Wang, B. et al., 2014). Specimen is slightly distorted and carbonized, due Fushun amber contains a diverse, well-pre- to preservation conditions of amber (Wu et al., served fossil arthropod fauna and abundant botani- 2000; Wang, X. et al., 2001), i.e., natural heating cal and other inclusions such as leaves, fungi and (autoclaving) in the deposit, which is often affecting even mammalian hairs. In addition, some microin- amber inclusions from Fushun. clusions, such as palynomorphs and testate amoe- Diagnosis. The species is characteristic in having bae, are frequently found in Fushun amber (Wang, well-developed anal veins, the median fork much B. et al., 2011, 2014). Arthropod inclusions cover longer than the basal stem, the costal vein extend- an impressive array, including 22 orders and more ing one third of the way between apices R4+5 and than 80 families (Wang, B. et al., 2011, 2014). M1, and the mid leg with two tibial spurs. Fossil M. The piece of amber under scrutiny here neotropica from Dominican amber differs in having comes from the collections of the Nanjing Institute the median fork slightly shorter than the basal of Geology and Palaeontology, Chinese Academy stem, the costal vein ending halfway between R4+5 of Sciences, Nanjing, China. Fresh amber coated and M1, no anal veins, and the mid leg with one tib- in thick coal dust was ground and polished manu- ial spur. Both fossil species are much smaller than ally with a series of wet silicon carbide abrasive extant members of the genus. papers to remove the opaque surface and to mini-

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Description. Male. Body dark, altered in amber, time. Later, the genus Neomesochria Amorim and length 4.1 mm. Natural colours not preserved. Left Tozoni, 1994 was proposed combining number of mid leg separated from thorax more robust than New World and New Caledonia species of the unnaturally slender right leg (Figure 2.1, 2.5). Dis- genus Mycetobia Meigen, 1818, comprising also tal flagellomeres missing, lost during preparation. the fossil Mesochria neotropica Grimaldi, 1991. Head barely visible (Figure 2.3). Eyes bare, very This statement was argued by Thompson (2006), probably touching or narrowly separated. Ocelli on as according to his opinion the genus Neome- raised tubercle present. Antenna apparently with sochria lacks any synapomorphy, and the only 14 more or less cylindrical flagellomeres. Probos- known to that date Mesochria neotropica venation cis (Figure 2.3.) short. Palpus 4-segmented, with characters, i.e., terminal R2+3 fused to Rl,, places it distal three segments well visible. Fourth palpal clearly in the genus Mesochria. Number of species segment cylindrical with evenly pointed apex, 4 are known exclusively from type specimens with times longer than third one (Figure 2.3, 2.4). very brief and/or incomplete descriptions. The con- Scutum barely preserved. Scutellum bearing 4 stituting features of the genus Mesochria are: ter-

marginal bristles. minal R2+3 fused to Rl, M two-branched; discal cell Wing typical of the genus (Figure 2.2, 2.7): length absent, metatibia with apical comb, and addition- measured from base 2.9 mm. Wing membrane ally, contiguous state of the male eyes (Michelsen, hyaline. Costal vein distinctly extending one third of 1999; Thompson, 2006). The other distinguishing the way between apices R4+5 and M1. Subcostal features are based on coloration of particular ele- vein bare, ending in C slightly proximal to base of ments of body, but these data are lacking for some radius. Radial sector Rs setose. Veins R1 and R2+3 modern species (Thompson, 2006). Very little is convergent, contiguous apically. Median fork dis- known on the biology of Mesochria. The stout tinctly longer than basal stem (1.8 times). Cross hooks on the 8th abdominal segment are present veins complete. Two distinct anal veins present. as five pairs in Mesochria (Thompson, 2006); pupal Fore and hind tibiae with single apical spur. Mid skin of Mesochria medicorum Edwards, 1928b, tibia with two apical spurs. Hind tibia with distinct was obtained from rotting banana fibre in Ghana tibial comb composed of dense row of setae. (Edwards, 1928b; Keilin and Tate, 1940). Its sister- Claws small, equal, apices bifid; empodium absent. group, Mycetobia is saprophagous and live in fer- Measurements are given in Table 1. menting saps runs or slime fluxes from wounds in Male genitalia barely visible (Figure 2.5, 2.6). trees or in tree holes (Keilin, 1919; Keilin and Tate, Gonocoxites large, fused. Cerci well developed, 1940; Krivosheina, 1997). long. Ten extant species of Mesochria are known Female unknown. from Fiji (M. schlingeri Thompson, 2006; M. vul- Age and occurrence. Early Eocene, Guchengzi garis Thompson, 2006), Samoa (M. buxtoniana Formation, Fushun amber. Xilutian coal mine, Fus- Edwards, 1928a), Java (M. cinctipes de Meijere, hun, Liaoning, China (41.8° N, 123.9° E; palaeoco- 1913), Borneo (M. intermedia Edwards, 1931), ordinates 43.5° N, 116.0° E). Madagascar (M. sylvatica Stuckenberg, 1961; M. griveaudi Stuckenberg, 1961), the Seychelles (M. DISCUSSION scottiana Enderlein, 1910), Ghana (M. medicorum The male of the new species is very similar to Edwards, 1928b) and the Democratic Republic of the female of fossil Mesochria neotropica Grimaldi, Congo (Zaire) (M. congoensis Tollet, 1956) – Fig- 1991 from Miocene Dominican amber. The ure 3. During the Palaeogene and Neogene the colouration cannot be used for identification as it is genus was more widely distributed, in the lower not preserved in inclusions from Fushun amber. Eocene it was present in North China (M. fani, The only differences are in the wing venation and present report) and in the Miocene in the Carib- in the numbers of tibial spurs on the mid leg (see bean Region (M. neotropica Grimaldi, 1991). The diagnosis). Extant species are distinctly larger with genus Mesochria was exemplified as illustrating wing lengths over 4.0 mm. Both fossil species are the tropical Gondwanan track (Matile, 1990) with smaller with wing lengths from 2.9 to 3.1 mm. sporadic “recent extensions” into the Holarctic The clear definition and unambiguous set of (Amorim and Pires, 1996; Cranston, 2005). The character of the Mesochria Enderlein and related tropical Gondwanan track (Matile, 1990) was pro- genera are still far from resolution. The generic key posed for explanation of Gondwanan distributions of Thompson and Rogers (1992) gave the features of groups, which phylogenies are concordant with of all modern genera placed in the family at that subsequent breakups of Gondwanan landmasses

5 SZADZIEWSKI ET AL.: Mesochria FROM FUSHUN AMBER

FIGURE 2. Holotype male of Mesochria fani Szadziewski and Szwedo, sp. nov. 1, habitus, lateral aspect; 2, wings; 3, mouthparts; 4, palpus; 5, legs; 6, wing reconstructed; 7, genitalia, lateral aspect.

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TABLE 1. Length measurements of legs (in mm). TR (ta1/ta2) – tarsal ratio; fe – femur; ti – tibia; ta1-ta5 – tarsomeres 1-5.

TR fe ti ta1 ta2 ta3 ta4 ta5

Fore leg 2.8 0.90 0.95 0.61 0.22 0.15 0.09 0.09 Mid leg 2.7 1.00 0.88 0.60 0.22 0.16 0.09 0.09 Hind leg 3.1 1.18 1.37 0.81 0.26 0.16 0.09 0.09

(dated minimally to the early Cretaceous). In fact, 1992; Sanmartín and Ronquist, 2004; McCarthy et this track, was recognized earlier by Harrison al., 2007), with various explanations given. Vicari- (1928), a perceptive early adopter of the Wegener ance and long distance dispersion events are allo- paradigm. Vast majority of the extinct Anisopodi- patry based, geographical explanations for the dae is known exclusively from the Northern Hemi- process of speciation and, although both had a role sphere, since the early Jurassic to the Pleistocene in the diversification of lineages, often are treated (Evenhuis 1994, 2014). Within the Anisopodidae, as exclusive alternative models (Crisp et al., 2011; as Matile (1990) and Amorim and Pires (1996) Gillespie et al., 2012). The present finding of the observed, extant members have fossil material oldest species of Mesochria from the early Eocene from the northern hemisphere, but tropical Gond- of the Northern Hemisphere challenges tropical wanan track is seen in the pairs of sister genera Gondwanan track hypothesis (Matile, 1990). We Eogaster (Afro-Oriental)/Olbiogaster (Neotropical), believe, the present distribution of Mesochria could and Mesochria (Afro-Oriental)/Neomesochria result from its long history, with both vicariance and (Neotropical/Australian). The problem of so-called long dispersal events. More mechanisms could be Gondwanan taxa was widely discussed (Eskov, involved in explanation of their distribution and

FIGURE 3. Geographical distribution of fossil and extant species of the genus Mesochria.

7 SZADZIEWSKI ET AL.: Mesochria FROM FUSHUN AMBER presence of the known fossils. The most rates of decreased and better correspond to the conditions taxa formation are observed in such locations in reconstructed for the Early Eocene Fushun area. which the struggle for existence against abiotic fac- Most of the amber specimens were found in the tors is reduced, as proposed by concepts of zonal upper layer of the Guchengzi Formation, according stratification (Zherikhin, 1978) and phytospreading to Quan et al. (2012) these strata were deposited (Meyen, 1987a, 1987b), which are the extensions during the Early Eocene warming period. In our of Darlington’s “equatorial pump” concept (Darling- opinion, presence of fossil Mesochria in Fushun ton, 1957, 1959). The model of the “tropical pump” amber corroborates the hypothesis on subtropical assumes that formation of the new taxa is climate, or at least fluctuation of climate during the restricted mainly to the tropical equatorial zone, terminal early Eocene as already suggested by and proliferated descendants oust the ancestors to Quan et al. (2012). the extratropical regions. The concept of zonal Despite of scarcity of biological data on stratification assumes that due to deterioration of Mesochria, its biological and environmental more uniform climatic and ecological conditions of requirements, limited recent distribution and fossil the late Cretaceous-early Palaeogene times and record some explanations on its evolutionary his- re-organisation of biosphere, with separation and tory can be presented. In our opinion the distribu- differentiation of bioclimatic zones numerous late tional pattern of the modern Mesochria resulted Mesozoic and early Cenozoic taxa gone extinct from concentration of relictual descendants of late (Zherikhin, 1978; Eskov, 1996). The phenomenon Mesozoic-Palaeogene forms in the tropical and of phytospreading (Meyen, 1987a, 1987b; Eskov, subtropical zones. Both vicariance and long dis- 1996) is not so much geographic as ecological in tance dispersal events, as well as other evolution- nature. It assumes that the newly formed taxa fol- ary mechanisms probably shaped the long history low by anisotropic propagation (“spreading”) from of this relictual genus, which, with present finding regions of the world where the climatic conditions could be traced back to early Eocene times. are most into regions with harsher abiotic conditions. The late Mesozoic and early Cenozoic were ACKNOWLEDGEMENTS characterised by weak climatic zonality (Gornitz, We greatly appreciate the efforts of Mr. M. 2009), which resulted in intermittent and ceasing Gąsior, MSc (Museum and Institute of Zoology, operation of “equatorial pump” mechanism (Eskov, Polish Academy of Sciences, Warsaw) for his help 1995, 1996), and increasing role of second-rank with the georeferencing software and map drawing. gradients (humid-arid, lowland-highland, etc.), We wish also to thank Mr. P. Senn (Gdynia) for lan- enabling the phytospreading mechanism operant guage advice. B.W. was supported by the National in the regime of Dobruskina’s (1982) diffuse cen- Natural Science Foundation of China (41572010) tres of speciation. This could trigger the diversifica- and a Research Fellowship from the Alexander von tion of Mesochria. 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