Two new long-legged in the Santonian amber of France (Diptera: ) André Nel, Romain Garrouste, Christophe Daugeron

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André Nel, Romain Garrouste, Christophe Daugeron. Two new long-legged flies in the Santo- nian amber of France (Diptera: Dolichopodidae). Research, Elsevier, 2017, 69, pp.1-5. ￿10.1016/j.cretres.2016.08.009￿. ￿hal-01369622￿

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1 2 Two new long-legged flies in the Santonian amber of France (Diptera: Dolichopodidae) 3 4 5 6 7 André Nel a, Romain Garrouste a and Christophe Daugeron b 8 9 10 11 a 12 Institut de Systématique, Évolution, Biodiversité, ISYEB - UMR 7205 – CNRS, MNHN, 13 14 UPMC, EPHE, Muséum national d’Histoire naturelle, Sorbonne Universités, 57 rue Cuvier, 15 16 17 CP 50, Entomologie, F-75005, Paris, France ([email protected], [email protected]) 18 19 b Muséum national d'Histoire naturelle, Centre National de la Recherche Scientifique, 20 21 22 Mécanismes adaptatifs et évolution, UMR 7179 MNHN-CNRS MECADEV, CP 50, 45 rue 23 24 Buffon, F-75005 Paris, France. ([email protected]) 25 26 27 28 29 ABSTRACT 30 31 Avenaphora gallica sp. nov., second representative of this dolichopodid genus, and 32 33 34 Cretomicrophorus piolencensis sp. nov., third representative of this genus, are described from 35 36 a new Early Santonian amber outcrop in Southern France. The relationships of Avenaphora in 37 38 39 the Empidoidea are discussed. As for the modern , Avenaphora gallica and 40 41 Cretomicrophorus piolencensis were found in a marine littoral swamp palaeoenvironment. 42 43 44 45 46 Keywords: Diptera; Empidoidea; Cretaceous amber; sp. nov.; palaeoecology 47 48 49 50 51 1. Introduction 52 53 The family Dolichopodidae is one of the five major lineages within the superfamily 54 55 56 Empidoidea. The traditional concept of the family is referred to as Dolichopodidae sensu 57 58 stricto. This concept was expanded to Dolichopodidae sensu lato to include the two small 59 60 61 62 63 64 65 subfamilies and Parathalassiinae (Sinclair and Cumming, 2006). The 1 2 dolichopodid characteristics comprise the Rs vein originating at or near the level of crossvein 3 4 5 h, crossvein r-m situated in the basal fourth of the wing, and the male terminalia rotated 6 7 forward beneath the preceding segments of the abdomen. The Dolichopodidae sensu lato are 8 9 10 not rare in the Cretaceous ambers, the most frequently encountered being Microphorinae. This 11 12 group comprises the modern genera Microphor Macquart, 1827 and Schistostoma Becker, 13 14 1902 and the extinct genera Avenaphora Grimaldi and Cumming, 1999 and Microphorites 15 16 17 Hennig, 1971 (Perrichot and Engel, 2014). Among these taxa, Avenaphora remains quite 18 19 enigmatic. Grimaldi and Cumming (1999) suggested that Avenaphora could have ‘closer 20 21 22 affinities to the Dolichopodidae and/or Parathalassiini’, rather than to the Microphorinae. 23 24 Cumming and Brooks (2002) proposed a sister-group relationship to the higher 25 26 27 Dolichopodidae for this genus, while Ulrich (2004) considered it could be ‘an early 28 29 dolichopodine’. This genus is currently known by a unique male specimen in Early 30 31 Cretaceous Lebanese amber (Grimaldi and Cumming, 1999). Here we describe the second 32 33 34 fossil attributable to this genus together with the third representative of the Cretaceous genus 35 36 Cretomicrophorus Negrobov, 1978, both from the Late Cretaceous amber of Piolenc (South 37 38 39 of France), a new insectiferous amber rediscovered in the beginning of the years 2000 40 41 (Gomez et al., 2003). Very few were recovered in this amber, mainly Diptera 42 43 44 (Choufani et al., 2013). 45 46 47 48 49 2. Material and method 50 51 The specimens are preserved in pieces of relatively clear, yellow amber. The amber 52 53 pieces were cut, shaped, and polished. Then they were prepared between two coverslips with 54 55 56 a Canada balsam medium as described in Azar et al. (2003), before being examined and 57 58 photographed. Fossil was examined and measured under incident light with Olympus SZX9 59 60 61 62 63 64 65 and Nikon SMZ1500 binocular microscopes, equipped with a Nikon D800 camera. We follow 1 2 the terminology of Grimaldi and Cumming (1999) and the empidoid classification of Sinclair 3 4 5 and Cumming (2006). 6 7 8 9 10 3. Systematic palaeontology 11 12 Family: Dolichopodidae Latreille, 1809 13 14 Subfamily: Parathalassinae Chvála, 1981: 15 16 17 Genus Avenaphora Grimaldi and Cumming, 1999 18 19 Type species. Avenaphora hispida Grimaldi and Cumming, 1999. Other species: Avenaphora 20 21 22 gallica sp. n. 23 24 25 26 27 Avenaphora gallica sp. n. 28 29 (Fig. 1) 30 31 Derivation of name. Named after Gallica, the ancient Roman name for France 32 33 34 Material. Holotype MNHN.F.A53892, stored in the Laboratoire de Paléontologie, MNHN, 35 36 Paris, France. 37 38 39 Type locality and horizon. Early Santonian, in amber, ancient sand quarry of Saint Louis, 40 41 Piolenc, Vaucluse, France (Gomez et al., 2003). 42 43 44 Diagnosis. Body and wing very small (wing 0.7 mm long, body 0.7 mm long); veins M1 and 45 46 M2 clearly reaching wing margin; anal lobe narrow. 47 48 49 Description. Body length ca. 0.7 mm; thorax length ca. 0.3 mm; wing length 0.7 mm. Known 50 51 only from female holotype. 52 53 Head relatively broad, slightly broader than thorax. Eye bare. Frons wide, margins of eyes 54 55 56 diverging dramatically posteriad; frons with three pairs of frontal-orbital setae, anterior pair 57 58 inclined and slightly cruciate, middle and posterior pairs reclinate; pair of small ocellar setae; 59 60 61 62 63 64 65 pair of cruciate postocellars, pair of lateroclinate outer verticals. Antenna with scape bearing 1 2 two setae; cuplike pedicel, distal margin with ring of fine setulae; flagellomere 1 dropshaped, 3 4 5 ventral surface with longer setulae; arista with two articles (suture between aristomeres well 6 7 visible), distal article whiplike, with numerous microtrichia slightly longer than width of 8 9 10 aristal trunk; total length of arista slightly more than three times length of flagellomere 1. 11 12 Proboscis not easily discerned, retracted into oral cavity. 13 14 Thorax: notum hardly arched; dorsocentrals and acrostichals differentiated; acrostichals in 15 16 17 two median rows, about six setae per row; dorsocentrals slightly larger, five per row, posterior 18 19 pair of dorsocentrals twice length of anterior three pairs. One supra-alar, one postpronotal, 20 21 22 two pairs of scutellar setae. Scutellar setae longer than posteriormost dorsocentrals, almost 23 24 upright; apical scutellars cruciate. Legs of moderate length, bristly. Forelegs: femur with two 25 26 27 long ventral setae, basitarsomere with ventral comb of fine, short setae; tibial anteroapical 28 29 comb apparently present. Midlegs: trochanter appears unusually long; tibia with three apical 30 31 and one preapical seta. Hind legs: femur slightly stouter than others, basitarsus with pair of 32 33 34 ventro-apical setae; tibial posteroapical comb present; no dorso-medial projection on 35 36 tarsomeres 5. 37 38 39 Wing relatively short, broad, length 2.7 x greatest width, anal lobe of relatively small size, 40 41 alula absent. Costa extended to apex of R4+5; costal seta present, costal vestiture with a 42 43 44 single row of spine-like setae. Sc very difficult to discern; R1 short, no pterostigma; Rs origin 45 46 opposite humeral crossvein, R2+3 and R4+5 nearly straight and simple; rm crossvein very 47 48 49 short, cell dm absent; M1+2 forked, both veins clearly reaching wing margin; CuA1 well 50 51 developed; cells bm and cup slender, of subequal length; bm-cu vein complete, apex of CuA2 52 53 not looped into cell bm; vein A1 extended to midway between cell cup and wing margin. 54 55 56 Abdomen relatively short and stout. Female genitalia hardly visible, except for ca. 3-4 57 58 acanthophorite spines. 59 60 61 62 63 64 65 Discussion. This fossil corresponds to the diagnosis of the genus Avenaphora: eyes bare, 1 2 broadly dichoptic (in male after Grimaldi and Cumming, 1999, but also present in this 3 4 5 female); antenna with two-articled arista; frons with three pairs of frontal-orbital setae, 6 7 anterior pair inclined. Wing with costal vein ending at R4+5; R1 short; cell dm absent (dm-cu 8 9 10 vein absent); M1+2 forked; cell bm slender, bm-cu vein complete; cell cup narrow, subequal 11 12 to length of cell bm, truncate apically; anal vein (A1) present, extended midway to wing 13 14 margin; anal lobe of moderate size. Only the structures of the male genitalia are unknown 15 16 17 because it is a female specimen. Thus we consider that our fossil belongs to this genus. Our 18 19 fossil differs in very few points from the type species Avenaphora hispida, i.e., body and wing 20 21 22 sizes much smaller than in A. hispida (wing 0.7 mm instead of 0.96 mm, body 0.7 mm long 23 24 instead of 1.16 mm); veins M1 and M2 are clearly reaching the wing margin in A. gallica; 25 26 27 anal lobe narrower than in A. hispida. The body chaetotaxy is the same in both species. 28 29 In the key to the families of Sinclair and Cumming (2006), the vein R4+5 unbranched 30 31 would support an attribution to the Dolichopodidae sensu lato, Hybotidae, or Atelestidae. The 32 33 34 base of Rs near the humeral vein and the basal cells shortened so that the crossvein r-m occurs 35 36 in the basal fourth of wing are character states supporting an attribution to the Dolichopodidae 37 38 39 sensu lato (Microphorinae + Dolichopodidae) (Sinclair and Cumming, 2006: 81), rather than 40 41 to the Hybotidae and the Atelestidae. The apex of antenna usually with lengthened, bristle- 42 43 44 like style would also exclude an attribution to the Atelestidae. 45 46 Nevertheless the Microphorinae have a discal cell, M1 and M2 arising independently 47 48 49 on it, and costa running around the wing (Chvála, 1983; Sinclair and Cumming, 2006), unlike 50 51 A. gallica. 52 53 The Dolichopodidae sensu stricto have no M2 and the costa ending on M1, unlike A. 54 55 56 gallica (Chvála, 1983; Sinclair and Cumming, 2006). Among the Parathalassinae (see list of 57 58 modern genera in Sinclair and Cumming (2006), only some species (viz. E. platypalpus 59 60 61 62 63 64 65 Shamshev and Grootaert, 2005) of the genus Eothalassius Shamshev and Grootaert, 2005 1 2 have no discal cell and a vein M2 (Negrobov, 1978; Ulrich, 1991, 2004; Cumming and 3 4 5 Brooks, 2002; Shamshev and Grootaert, 2002; Brooks and Cumming, 2011). E. platypalpus 6 7 has a venation very close to that of Avenaphora. But Eothalassius differs from Avenaphora in 8 9 10 the one-segmented style (Shamshev and Grootaert, 2005). The recent genus Neothalassius 11 12 Brooks and Cumming, 2016 has also no discal cell but differs from Avenaphora in the 13 14 absence of the vein M2 (Brooks and Cumming, 2016). 15 16 17 Avenaphora remains a Dolichopodidae (sensu lato) of uncertain affinities, but it shows 18 19 more similarities with the Parathalassinae than with the Microphorinae or the Dolichopodidae 20 21 22 s.str. 23 24 25 26 27 Genus Cretomicrophorus Negrobov, 1978 28 29 Type species. Cretomicrophorus rohdendorfi Negrobov, 1978. Other species: 30 31 Cretomicrophorus novemundus Grimaldi and Cumming, 1999, Cretomicrophorus 32 33 34 piolencensis sp. nov. 35 36 37 38 39 Cretomicrophorus piolencensis sp. nov. 40 41 (Figs. 2-3) 42 43 44 Derivation of name. Named after Piolenc, type locality. 45 46 Material. Holotype MNHN.F.A53893, stored in the Laboratoire de Paléontologie, MNHN, 47 48 49 Paris, France. 50 51 Locality and horizon. Early Santonian, in amber, ancient sand quarry of Saint Louis, Piolenc, 52 53 Vaucluse, France (Gomez et al., 2003). 54 55 56 57 58 59 60 61 62 63 64 65 Diagnosis. Hind tibiae bearing three anterodorsal setae and five stout apicals; middle tibiae 1 2 bearing six stout apicals; six pairs of acrostichals; scutellum with only long, apical pair of 3 4 5 setae; C extending to R4+5; no pterostigma. 6 7 Description. Body ca. 1.8 mm long. Head strongly deformed in its anterior part, 0.3 mm long, 8 9 10 0.4 mm high; ocellar tubercle not displaced forward; frons and face not visible between eyes 11 12 due to the deformation of this part of head; two stout ocellar setae, several setae around 13 14 ocellar tubercle; two stout inner verticals , two stout postverticals; eyes bare, not contacting; 15 16 17 setae along posterior eye margin (postocular bristles) arranged in one row from above to 18 19 below; genae not developed; antennae located on apical third of head; scape very short, 20 21 22 pedicel lacking apical process, but with apical circlet of setae, first flagellomere bulbiform, 23 24 apically pointed; its length exceeding its thickness; arista apical one-segmented; proboscis 25 26 27 hardly visible, short; palpus small. Thorax short, 0.6 mm long with the following distinct 28 29 setae: 1 propleural, 1 postpronotal, 2 notopleurals, 1 supra-alar, 6 pairs of acrostichals, 8 pairs 30 31 of dorsocentrals, 2 stout scutellars; mesopleura bare. Legs comparatively short, with well 32 33 34 differentiated stout setae; femora not thickened; tarsi not broadened. Fore tibia bearing several 35 36 stout antero- and posterodorsal setae, but lacking stout apicals; mid coxa with two stout setae; 37 38 39 mid trochanter with several long apical setae; mid tibia with 2 antero- and posterodorsals, 6 40 41 stout apicals; hind coxa with 1 stout seta; hind tibia with 3 anterodorsals and 5 stout apicals; 42 43 44 no dorso-medial projection on tarsomeres 5. Wings hyaline, 1.26 mm long; venation nearly 45 46 identical to that of the modern praecox (Loew); no pterostigma; C extending to 47 48 49 R4+5, Sc entering C; costal cell large; R2+3 simple; simple R4+5 terminating at wing tip; 50 51 vein bm-cu not complete; anal lobe weakly developed. Abdomen cylindrical, short, 0.9 mm 52 53 long, bearing one stout dorsal seta on the first five segments; segment 7 clearly seen 54 55 56 externally, not reduced; segment 8 located at apex of abdomen; hypopygium displaced onto 57 58 right side, its apex extending onto abdominal tergite 5. 59 60 61 62 63 64 65 Discussion. Following Sinclair and Cumming (2006), Cretomicrophorus piolencensis sp. nov. 1 2 can be attributed to the Dolichopodidae sensu lato for the following diagnostic characters: Rs 3 4 5 originating at or near level of crossvein h; basal cells shortened (less than half length of cell 6 7 dm) so that crossvein r–m occurs in the basal fourth of wing; vein R4+5 unbranched; male 8 9 10 terminalia rotated (lateroflexed) forward beneath the preceding segments with the pregenital 11 12 segments partially twisted or rotated. It shares with the Parathalassiinae the following 13 14 diagnostic characters: antennae inserted high on head; dichoptic males, narrow wing (anal 15 16 17 lobe not developed); crossvein bm-cu incomplete; single segmented arista. The 18 19 Microphorinae are excluded because they have holoptic male, broad wings (i.e., anal lobe 20 21 22 developed), two-segmented arista, and costa circumambient. The modern Parathalassiinae 23 24 includes the following genera: Amphithalassius Ulrich, 19991, Chimerothalassius Shamshev 25 26 27 and Grootaert, 2003, Eothalassius Shamshev and Grootaert, 2005, Microphorella Becker, 28 29 1909, Mik, 1891, Plesiothalassius Ulrich, 1991, Thalassophorus Saigusa, 30 31 1986, and Neothalassius Brooks and Cumming, 2010. Ulrich (2004) listed the Cretaceous 32 33 34 genus Archichrysotus Negrobov, 1978 in this group and maybe also Avenaphora (see above), 35 36 while Sinclair and Cumming (2006) listed the genera Cretomicrophorus Negrobov, 1978, 37 38 39 Archichrysotus, and Negrobov, 1978 in this group on the basis of the single articled 40 41 stylus. 42 43 44 C. piolencensis differs from Microphorella, Thalassophorus, and Parathalassius in the 45 46 bare eye, and the presence of several strong setae on the legs (Smith, 1972; Chvála, 1983; 47 48 49 Saigusa, 1986). Chimerothalassius, Eothalassius, Plesiothalassius, Amphithalassius, and 50 51 Neothalassius have also no strong and long setae on legs (Ulrich, 1991, Shamshev and 52 53 Grootaert, 2002, 2005; Brooks and Cumming, 2016). Chimerothalassius has also distinctly 54 55 56 shorter setae on mesoscutum. Eothalassius has distinctly longer arista than in C. piolencensis. 57 58 59 60 61 62 63 64 65 Archichrysotus and Retinitus differ from C. piolencensis in the presence of very few setae on 1 2 tibiae (Negrobov, 1978). 3 4 5 C. piolencensis fits quite well with the diagnosis of Cretomicrophorus (Negrobov, 6 7 1978). Differences with C. rohdendorfi (Taymyr amber, Siberia, Santonian) are very few: 8 9 10 hind tibiae bearing three anterodorsal setae (instead of two) and five stout apicals (instead of 11 12 three); mid tibia bearing six stout apicals instead of four; scutellum with only long, apical pair 13 14 of setae, no pair of smaller, anterior ones; C extending to R4+5 instead of M1+2. C. 15 16 17 piolencensis differs from C. novemundus (New Jersey amber, Turonian) in the presence of six 18 19 pairs of acrostichals (vs. four); no pterostigma. 20 21 22 23 24 4. Palaeoecology 25 26 27 After Ulrich (2004), the Microphorinae (recent and fossil) are supposed to be terrestrial, while 28 29 the modern Parathalassiinae and the most ‘basal’ representatives of the ‘higher’ 30 31 Dolichopodidae are living in the marine littoral. Interestingly the outcrop of lignites and sands 32 33 34 of Piolenc is considered as resulting from marine littoral swamps and sediments (Gomez et 35 36 al., 2003). Thus the presence of two Parathalassiinae would fit well with Ulrich’s hypothesis. 37 38 39 40 41 Acknowledgements. We sincerely thank xxxxx referees ….. 42 43 44 45 46 References 47 48 49 Azar, D., Perrichot, V., Néraudeau, D., Nel, A., 2003. New psychodids from the Cretaceous 50 51 ambers of Lebanon and France, with a discussion of Eophlebotomus connectens Cockerell, 52 53 1920 (Diptera, Psychodidae). Annals of the Entomological Society of America 96, 117-126. 54 55 56 57 58 59 60 61 62 63 64 65 Brooks, S.E., Cumming, J.M., 2011. The New World genera of Parathalassiinae (Diptera: 1 2 Empidoidea: Dolichopodidae s.l.), with new species of Thalassophorus and Eothalassius. The 3 4 5 Canadian Entomologist 143, 423-446. 6 7 Brooks, S.E., Cumming, J.M., 2016. Neothalassius, a new genus of Parathalassiinae (Diptera: 8 9 10 Dolichopodidae s.lat.) from the Pacific coast of South America. Zootaxa 4066, 311-322. 11 12 Choufani, J., Perrichot, V., Girard, V., Garrouste, R., Azar, D., Nel, A., 2013. Two new biting 13 14 midges of the modern type from Santonian amber of France (Diptera: Ceratopogonidae). pp. 15 16 17 75-95. In: Azar, D., Engel, M.S., Jarzembowski, E., Krogmann, L., Nel, A., Santiago-Blay, J. 18 19 (eds). Evolution in an Amberiferous and Stone Alphabet. Proceedings of the 6th 20 21 22 International Congress on Fossil Insects, and Amber. 23 24 Chvála, M., 1983. The Empidoidea (Diptera) of Fennoscandia and Denmark. 2. General part. 25 26 27 The families Hybotidae, Atelestidae and Microphoridae. Fauna Entomologica Scandinavica 28 29 12, 1-279. 30 31 Cumming, J.M., Brooks, S.E., 2002. Electrophorella, a new genus of parathalassiine flies 32 33 34 from Baltic amber, with a cladistic analysis of the Microphorinae + Dolichopodidae lineage 35 36 (Diptera: Empidoidea). Studia Dipterologica 9, 41-54. 37 38 39 Gomez, B., Barale, G., Saad, D., 2003. Santonian angiosperm-dominated leaf-assemblage 40 41 from Piolenc (Vaucluse, Sud-Est de la France). Comptes-Rendus Palevol 2, 197-204. 42 43 44 Grimaldi, D.A., Cumming, J., 1999. Brachyceran Diptera in Cretaceous ambers and Mesozoic 45 46 diversification of the Eremoneura. Bulletin of the American Museum of Natural History 239, 47 48 49 1-124. 50 51 Melander, A.L., 1927. Diptera fam. Empididae. Genera Insectorum 185, 1-434. 52 53 Negrobov, O.P., 1978. Flies of the superfamily Empidoidea (Diptera) from Cretaceous retinite 54 55 56 in Northern Siberia. Paleontological Journal 12, 221-228. 57 58 59 60 61 62 63 64 65 Perrichot, V., Engel, M.S., 2014. Youngest occurrence of the genus Microphorites (Diptera: 1 2 Dolichopodidae): A new species in Late Cretaceous Vendean amber. Paleontological 3 4 5 Contributions 10G: 30-33. 6 7 Saigusa, T., 1986. New genera of Empididae (Diptera) from Eastern Asia. Sieblodia 5, 97- 8 9 10 118. 11 12 Shamshev, I.V., Grootaert, P., 2002. A new genus of Microphorinae (Diptera: Empidoidea) 13 14 from New Zealand. Belgian Journal of Entomology 4, 129-144. 15 16 17 Shamshev, I.V., Grootaert, P., 2005. Eothalassius, a new genus of parathalassiine flies 18 19 (Diptera: Empidoidea: Dolichopodidae) from Southeast Asia and Pupua New Guinea. 20 21 22 European Journal of Entomology 102, 107-118. 23 24 Sinclair, B.J., 1997. Icasma Collin and an allied new genus Glyphidopeza, from New Zealand 25 26 27 (Diptera: Empidoidea; Ceratomerinae). Records of the Australian Museum, 49 (2): 195-211. 28 29 Sinclair, B.J. and Cumming, J.M. 2006. The morphology, higher-level phylogeny and 30 31 classification of the Empidoidea (Diptera). Zootaxa 1180, 1-172. 32 33 34 Smith, K.G.V., 1972. A remarkable new Parathalassius (Diptera: Empididae) from South 35 36 Africa, with a key to world species. Annals of the Natal Museum 21, 455-458. 37 38 39 Ulrich, H., 1991. Two new genera of parathalassiine-like flies from South Africa (Diptera: 40 41 Empidoidea). Bonner Zoologische Beiträge 42, 187-216. 42 43 44 Ulrich, H., 2004. Phylogenetic considerations about an early colonization of the sea coasts by 45 46 Dolichopodidae (Diptera). Studia Dipterologica 11, 233-243. 47 48 49 50 51 Fig. 1. Avenaphora gallica sp. n., holotype MNHN.F.A53892. A. Habitus, ventral view; B. 52 53 Wing. Scale bar = 0.1 mm (A,B). 54 55 56 Fig. 2. Cretomicrophorus piolencensis sp. nov., holotype MNHN.F.A53893. Habitus. Scale 57 58 bar = 0.5 mm. 59 60 61 62 63 64 65 Fig. 3. Cretomicrophorus piolencensis sp. nov., holotype MNHN.F.A53893. A. Head; B. 1 2 wing. Scale bar = 0.1 mm (A), 0.5 mm (B). 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 Figure 1 Click here to download high resolution image Figure 2 Click here to download high resolution image Figure 3 Click here to download high resolution image