A Diverse Paleobiota in Early Eocene Fushun Amber from China

Current Biology, Volume 24 Supplemental Information

A Diverse Paleobiota in Early Eocene Fushun Amber from China Bo Wang, Jes Rust, Michael S. Engel, Jacek Szwedo, Suryendu Dutta, André Nel, Yong Fan, Fanwei Meng, Gongle Shi, Edmund A. Jarzembowski, Torsten Wappler, Frauke Stebner, Yan Fang, Limi Mao, Daran Zheng, and Haichun Zhang

Supplemental Results and Discussion Geology. Fushun, a city in Liaoning Province, northeastern China, contains six coal-mining pits of which the biggest is the West Opencast Coalmine (41º50’ N, 123º 54’ E) (Figure S1). The coal and oil shale are found in a relatively small east–west-trending exposure of Mesozoic and Cenozoic sedimentary and volcanic rocks surrounded by Precambrian terrain made up mainly of granitic gneiss [S1, S2]. These continental strata consist of swampy to fluvio-deltaic and tuffaceous sediments that were deposited in the basin during the early Paleogene [S2]. In ascending order, the sequence is subdivided into yellow-gray sandstone intercalated with coal seams (Laohutai Formation; 58–65 Ma), overlain by gray-green tuff intercalated with coal seams (Lizigou Formation; 56–58 Ma), followed by a thick coal layer with a roof and floor of dark shale (Guchengzi Formation; 50–56 Ma), oil shale and black shale (Jijuntun Formation; 43–50 Ma), gray-green mudstone and shale (Xilutian Formation; 38–43 Ma), and brown shale and variegated siltstone (Gengjiajie Formation; 30–38 Ma) [S3–S5]. The entire sequence therefore spans the Danian-Bartonian stages [S6]. Amber pieces are usually embedded in the coal or oil shale in the middle and upper coal beds of the Guchengzi Formation in the West Opencast Coalmine (Figure S2) [S7]. Coal seams of the Guchengzi Formation were subjected to abnormally high metamorphism due to the massive intrusion of basic magmas in the Oligocene, which formed diabase sills after coal accumulation [S2]. Consequently, amber pieces were naturally heated and show various colors (Figure S2). Paleoclimate. The floristic composition of the Fushun amber forest is based on available data of macrofossils and fossil palynomorphs. The vegetation of the area was composed of mixed mesophytic forests with woody shrubs and herbs, probably originating from moist and swampy habitats [S4, S5, S8]. The paleoclimate in the Early Eocene in Fushun is reconstructed as moist subtropical, with variability in seasonal precipitation [S8, S9]. The estimated mean annual temperature varied between 15 and 21°C; for the coldest months, the mean temperature varied between 9 and 14°C; and for the warmest months, the mean temperature was between 19 and 25°C. The mean annual precipitation is estimated as varying between 650 and 1500 mm [S4, S5]. Material. Hong published several papers about Fushun amber insects [S10–S13], and later included them and some new taxa in his book [S14]. In total, Hong reported 8 orders and 223 species of insects, most of which were flies, wasps, and wingless aphids, and almost all taxa were based on isolated, single specimens [S14]. Unfortunately, most identifications are systematically suspect and require considerable revision (e.g., refs. [S15–S17]). Most of the taxa are invalid because no depositories were given for the material [S18]. Thus, the taxonomic composition, taxonomic diversity, and morphological disparity of Fushun amber arthropod

inclusions remain unclear. Today authentic Fushun amber is increasingly difficult to obtain on the market and, when available, fetches high prices, some five to ten times the cost of equivalent pieces of Baltic amber. Many treated Baltic, Dominican, and Burmese amber pieces are sold as ‘Chinese Fushun’ amber on the open market (e.g., ref. [S19]). Our research is based mainly on our systematic investigations during the past 20 years. So far we have found abundant arthropod inclusions belonging to 22 orders and more than 80 families (Table S2), which has greatly improved our knowledge of the diversity of the biota. The updated list of orders and families of arthropods is based on collections in Nanjing and Fushun (NIGPAS and Amber Research Institute respectively), several private collections, and revisions of Hong’s descriptions. Nine families named by Hong [S14] are not listed in Table S2 because all these taxa are systematically incorrect, and more importantly, no depositories were given for the material to be reassessed. Paleogene amber insect faunas. There are at least 14 Paleogene insect-bearing ambers from around the world: Arkansas (USA), Baltic (including Bitterfeld), British Columbia (Canada), Cambay (India), Campaolo (Italy), Fushun (China), Gurnigelflysch Formation (Switzerland), Oise (France), Romanian, Rovno (Ukraine), Sakhalin (Russia), Sicilian (Italy), Študlov (Czech), and Washington (USA) [S20–S22]. Most of them remain poorly investigated, and only Baltic, Cambay, Fushun, Oise, and Rovno amber yield abundant insects.

Figure S1. View of the West Opencast Coalmine. Related to Figure 1.

Figure S2. Amber samples. Related to Figure 1. (A) Amber piece embedded in the coal. (B) Amber pieces with different colours. (C) Amber craftwork (dragon). Red arrow points to the amber piece. Scale bars 20 mm.

Figure S3. FTIR spectrum of Fushun amber. Related to Figure 2.

Figure S4. Some Fushun amber arthropods. Related to Figure 3. (A) True cricket (Orthoptera: Gryllidae), NIGP156974. (B) Dark-winged fungus gnat (Diptera: Sciaridae), NIGP156975. (C) Spider juvenile or female (Araneae), NIGP156976. (D) Plant bug (Hemiptera: Miridae), NIGP156977. (E) Cockroach nymph (Blattodea), NIGP156978. (F) Parasitoid wasp (Hymenoptera: Braconidae), NIGP156979. (G) Froghopper (Hemiptera: Cercopidae), NIGP156980. (H) Fungus gnat (Diptera: Mycetophilidae), NIGP156985. (I) Barklouse (Psocodea: Psyllipsocidae), NIGP156986 (J) Dark-winged fungus gnat (Diptera: Sciaridae), NIGP156987 Scale bars 1 mm.

Table S1. Biomarkers identified in the total extracts of Fushun amber. Related to Figure 2. Peak No

Molecular weight

Compound

1 2 3 4 5 6 7 8 9 10 11 12 13 14

192 192 192 192 188 174 188 170 198 184 274 256 276 274

15 16 17 18 19 20 21

270 252 316 234 284 286 268

Octahydro-tetramethyl naphthalene Octahydro-tetramethyl naphthalene isomer Octahydro-tetramethyl naphthalene isomer Octahydro-tetramethyl naphthalene isomer Unknown compound Ionene Methylionene Trimethyl naphthalene Cadalene Tetramethyl naphthalene C 20 -diterpane 19-Norabieta-8,11,13-triene Abietane C 20 -diterpane Dehydroabietane Simonellite Unknown compound Retene Abieta-8,11,13-trien-7-one Ferruginol 12-hydroxysimonellite

Table S2. Taxa of arthropods in Fushun Amber. Related to Figure 3. MYRIAPODA Diplopoda Family undet. ARACHNIDA Acariformes Families undet. (mites) Parasitiformes Families undet. (ticks) Araneae Dictynoidea

Families undet. Opiliones Families undet. Pseudoscorpiones Cheliferidae HEXAPODA Thysanura Family undet. Collembola Families undet. Ephemeroptera †Philolimniidae1

Dermaptera Family undet. Orthoptera Gryllidae Embiidina Family undet. Blattodea Corydiidae2 Blaberidae †Cainoblattinidae3 Families undet. Psocodea Mesopsocidae Psyllipsocidae Caeciliusidae

Thysanoptera Phlaeothripidae Hemiptera Sternorrhyncha Coccoidea Aphidoidea Aphididae Drepanosiphidae Eriosomatidae ? Hormaphididae Phloeomyzidae †Drepanochaitophoridae4 Cicadomorpha Cercopidae Cicadellidae Heteroptera Aradidae Miridae Pentatomidae Neuroptera Hemerobiidae Lepidoptera Family undet. Coleoptera Staphylinidae Elateridae Throscidae Helodidae Mordellidae Scirtidae Meloidae Cleridae Curculionidae Families undet. Strepsiptera Family undet. Hymenoptera Ichneumonidae

Braconidae Aulacidae Torymidae Perilampidae Eurytomidae Eulophidae Mymarommatidae Scelionidae Bethylidae Diapriidae Formicidae Diptera Nematocera Ptychopteridae Chaoboridae ? Limoniidae Hesperinidae Ceratopogonidae Chironomidae Bibionidae Scatopsidae Sciaridae Mycetophilidae Keroplatidae Macroceridae Psychodidae Anisopodidae Cecidomyiidae Brachycera Empididae Dolichopodidae Phoridae Sciadoceridae Lauxaniidae Rhagionidae † extinct taxa. 1 from Hong [S14]. 2 from Hong [S11] and Jacobus and McCafferty [S23].

3 4

from Ping [S24]. from Zhang and Hong [S25].

Supplemental References S1. Johnson, E.A. (1990). Geology of the Fushun coalfield, Liaoning Province, People’s Republic of China. Int. J. Coal Geol. 14, 217–236. S2. Wu, C.L., Yang, Q., Zhu, Z.D., Liu, G., and Li, X. (2000). Thermodynamic analysis and simulation of coal metamorphism in the Fushun Basin, China. Int. J. Coal Geol. 44, 149–168. S3. Meng, Q.T., Liu, Z.J., Bruch, A.A., Liu, R., and Hu, F. (2012). Palaeoclimatic evolution during Eocene and its influence on oil shale mineralisation, Fushun basin, China. J. Asian Earth Sci. 45, 95–105. S4. Quan, C., Liu, Y.S., and Utescher, T. (2011). Paleogene evolution of precipitation in northeast China supporting the middle Eocene intensification of the East Asian monsoon. Palaios 26, 743–753. S5. Quan, C., Liu, Y.S., and Utescher, T. (2012). Paleogene temperature gradient, seasonal variation and climate evolution of northeast China. Palaeogeogr. Palaeoclimatol. Palaeoecol. 313–314, 150–161. S6. Gradstein, F.M., Ogg, J.G., Schmitz, M., and Ogg, G. (2012). The Geologic Time Scale 2012 (Amsterdam: Elsevier Science). S7. Xu, S.C., Liu, Z.J., Dong, Q.S., Liu, S.Y., Liu, R., and Meng, Q.T. (2012). Eocene sedimentary evolution and its control over coal & oil shale development in Fushun Coalfield. J. China Univ. Petrol. 36, 45–52. S8.

S9.

S10.

S11. S12. S13.

S14.

Wang, Q., Ferguson, D.K., Feng, G.P., Ablaev, A.G., Wang, Y.F., Yang, J., Li, Y.L., and Li, C.S. (2010). Climatic change during the Palaeocene to Eocene based on fossil plants from Fushun, China. Palaeogeogr. Palaeoclimatol. Palaeoecol. 295, 323–331. Wang, D.H., Lu, S.C., Han, S., Sun, X.Y., and Quan, C. (2013). Eocene prevalence of monsoon-like climate over eastern China reflected by hydrological dynamics. J. Asian Earth Sci. 62, 776–787. Hong, Y.C., Yang, T.C., Wang, S.T., Wang, S.E., Li, Y.K., Sun, M.R., Sun, H.C., and Tu, N.C. (1974). Stratigraphy and Palaeontology of Fushun Coal-field, Liaoning Province. Acta Geol. Sin. 2, 113–149. Hong, Y.C. (1979). On Eocene Philolimnias gen. nov. (Ephemeroptera, Insecta) in amber from Fushun Coalfield, Liaoning Province. Sci. Sin. 12, 331–339. Hong, Y.C. (1981) Eocene Fossil Diptera Insecta in Amber of Fushun Coalfield. (Beijing: Geological Publishing House). Hong, Y.C., Guo, X., and Ren D. (2000). A new genus Eopalomyitis gen. nov. from Eocene Fushun amber and discussion of its taxonomic position. Acta Parasitol. Med. Entomol. Sin. 7, 225–234. Hong, Y.C. (2002). Amber Insect of China (Beijing: Beijing Science and Technology Press).

S15. S16. S17.

S18. S19.

S20.

S21.

S22.

S23.

S24. S25.

Brown, B.V. (1999). Re-evaluation of the fossil Phoridae (Diptera). J. Nat. Hist. 33, 1561–1573. Doitteau, G., and Nel, A. (2007). Chironomid midges from early Eocene amber of France (Diptera: Chironomidae). Zootaxa 1404, 1–66. Liu, Z.W., Engel, M.S., and Grimaldi, D.A. (2007). Phylogeny and geological history of the cynipoid wasps (Hymenoptera: Cynipoidea). Am. Mus. Novit. 3583, 1–48. Heraty, J.M., and Darling, D.C. (2009). Fossil Eucharitidae and Perilampidae (Hymenoptera: Chalcidoidea) from Baltic Amber. Zootaxa 2306, 1–16. Heie, O.E., and Poinar, G.O.Jr. (2011). Using fossils to determine an amber source: aphids and crane flies in Chinese or Baltic amber? Hist. Biol. 23, 431–433. Martínez-Delclòs, X., Briggs, D.E.G., and Peñalver, E. (2004). Taphonomy of insects in carbonates and amber. Palaeogeogr. Palaeoclimatol. Palaeoecol. 203, 19–64. Perkovsky, E.E., Rasnitsyn, A.P., Vlaskin, A.P., and Taraschuk, M.V. (2007). A comparative analysis of the Baltic and Rovno amber arthropod faunas: Representative samples. Afr. Invertebr. 48, 229–245. Szwedo, J., and Sontag, E. (2013). The flies (Diptera) say that amber from the Gulf of Gdańsk, Bitterfeld and Rovno is the same Baltic amber. Pol. J. Entomol. 82, 379–388. Jacobus, L.M., and McCafferty, W.P. (2006). Reevaluation of the phylogeny of the Ephemeroptera infraorder Pannota (Furcatergalia), with adjustments to higher classification. Trans. Am. Entomol. Soc. 132, 81–90. Ping, Ch. (1931). On a blattoid insect in the Fushun amber. Bull. Geol. Soc. China 11, 205–207. Zhang, G.X., and Hong, Y.C. (1999). A new family Drepanochaitophoridae (Homoptera: Aphidoidea) from Eocene Fushun amber of Liaoning Province, China. Entomol. Sin. 6, 127–134.