Supporting Information

Schmidt et al. 10.1073/pnas.1208464109 SI Text head and is folded around head capsule. Prothoracic leg long and Systematics. slender; lengths of profemur and protibia equal (0.43 mm); width Class: INSECTA of femur up to twice that of tibia. Protarsomeres slender, thinner Order: DIPTERA than apex of tibia; length of probasitarsomere 0.10 mm (segmen- Family: Indet.: tation of more distal tarsomeres is obscure). Protibia and tibia of (Fig. 1 G and H and Fig. S2) unidentified leg (leg C in Fig. 1H) with dorsal and median long- itudinal row of 4–5 slightly longer, thicker, and more erect setu- General description. Specimen is fragmentary, consisting of a par- lae. No apical tibial spurs present, though a thick, short seta tial head with portions of some appendages still attached; an an- occurs at apex of tibia on leg A. No apical comb present on tibiae. tenna (most of it disarticulated from its base); a dorsal portion of Proportions of podomeres on disarticulated legs: leg A femur the thorax; remnants of at least four legs (principally the femora 0.37 mm, tibia 0.28 mm; leg B femur 0.41 mm [tibial apex ambig- and tibiae but some basitarsomeres as well), including a disso- uous]; leg C femur 0.55 mm [tibia incomplete] (Fig. 1H). Leg C is ciated distitarsomere (Fig. 1 G and H and Fig. S2A). The dorsal possibly the hind leg, since these are generally the longest legs in surface of the thorax lies just under or perhaps breaches the sur- many adult . A single distitarsomere, with portion of face of the amber; the head is flexed under the thorax, and some penultimate tarsomere, is preserved, with excellent detail; it is of the head appendages (e.g., maxillary palps) apparently shifted detached and distant from most other parts. Distitarsomere se- during immersion in the resin. There are no remains of wings, tulose, 0.06 mm length; with pair of simple pretarsal claws halteres, or an abdomen. (i.e., not toothed or serrate); length of each claw approximately Head: Slightly pear-shaped, with the presumed dorsal portion equal to width of tarsomere; unguitractor plate visible as light being widest; greatest length 0.25 mm. A cavity on the right side area through dorsal surface. Most significantly: Empodium and (0.20 × 0.15 mm) appears to have been formed by the decay of pair of flanking pulvilli are setiform (definitely not branched, the eye, which exposes a portion of the posterior tentorial arm feathered, or pulvilliform); empodium slightly longer than pulvilli and what is possibly an anterior tentorial arm (Fig. 1H). The only (Fig. 1H and Fig. S2D). eye facets that are preserved appear to be a small group of six (diameter ca. 10 μm) near the posterior or ventral surface of Specimen. MGP 31345, Museum of Geology and Paleontology, the head capsule. Putative eye cavity is pear-shaped (slightly University of Padova, Italy. emarginate), with a smaller cavity anterior and close to the emar- ginate portion (greatest length 0.06 mm). The small cavity is Comments. An ordinal-level attribution of the fossil can be made probably the insertion base of the antenna, such as the scape. Dis- via a process of exclusion, as well as by use of the few specialized tal to the putative scape is a row of four apparent antennomeres; features that are preserved. a roughly triangular basal segment (the pedicel?), then a trape- Ephemeroptera and Odonata can be excluded, based on an- zoidal segment (the largest segment, probably the first flagellar tennal structure (these orders have an aristate flagellum, and segment), and then two smaller segments shaped like parallelo- Odonata are much larger). Polyneoptera (orthopteroids and Dic- grams (Fig. 1H and Fig. S2B). Total length of connected antennal tyoptera) can be excluded based on tarsal structure (i.e., in these section is 0.19 mm. Disarticulated from these segments, and lying orders an arolium and often a pad-like pulvillus are present as some 0.10 mm away, is the remainder of the antenna, composed well as plantular lobes on the more proximal tarsomeres); of seven flagellomeres that diminish in size apicad (Fig. 1H and absence in the fossil of large, spine-like setae on the legs as well Fig. S2C); total length of disarticulated section is 0.27 mm. Basal as the absence of mandibles. mandibles are typically segments of the disarticulated antennal section are barrel- toothed and among the most heavily sclerotized/chitinized struc- shaped, distal two segments narrowed apicad; apical segment tures on the body and so, if originally present, would have been minute, approximately 0.25x the length, 0.3x the width of penul- preserved along with the less durable cephalic appendages. Poly- timate segment. Flagellomeres with sparse setulae, with slightly neoptera also typically have long, flagellate antennae with fili- thicker and longer setae forming a possible subapical whorl. Sec- form antennomeres. Hemiptera is excluded by the lack of a ond flagellomere in disarticulated portion of antenna with small, sucking beak and by the presence of maxillary palps. Among ring-like sensory organ (doubtfully the Johnston’s Organ, which Holometabola, the mandibulate orders (Coleoptera, Neuropter- occurs on the pedicel). If all antennal segments are preserved, ida, Hymenoptera) can be excluded based on lack of mandibles in flagellum is comprised of 10 flagellomeres; total length 0.59 mm. the fossil. Coleoptera can further be excluded since the fossil has Possible labellum lying between head capsule and thorax, a long, slender legs and does not have a heavily sclerotized, shield- slightly irregular, very slightly sclerotized structure 0.11 mm like pronotum. Hymenoptera have conspicuous, setulose tibial length, with apparently two opposing, setulose lobes; no pseudo- spurs, and the only Triassic Hymenoptera known are a few rare tracheae are visible (Fig. 1H). Distal portions of possible maxil- fossils of the most basal living family, . Amphies- lary palps are deflexed over the head capsule (best seen in view menoptera (Trichoptera and ) can be excluded be- from left side), total preserved length 0.18 mm; distal three pal- cause these two orders also possess conspicuous tibial spurs as pomeres preserved, each palpomere with fine, dense, erect setu- well as a densely setose or scaled body; long, multiarticulate an- lae; palpomeres approximately equal in length (0.05–0.06 mm), tennae; and in Lepidoptera (except a few basal families) a pro- apical one slightly fusiform in shape. boscis composed of long, coiled galeae. While Trichoptera are of Thorax: Remains of at least four legs are preserved (Fig. 1H Triassic age (and some even possess labellum-like mouthparts), and Fig. S2 D–F), though all but one are disarticulated from body Lepidoptera can be further excluded since this appears to be a and thus not identifiable as to pro-, meso- or metathoracic in ori- particularly young order, no older than Jurassic (1). gin. All leg segments with numerous microtrichia, which are den- In lieu of definitive synapomorphies being preserved (e.g., ser on the tibiae than on femora. Left proleg with femur, tibia, wing venation, halteres), attribution to the Diptera is based on and basal two tarsomeres preserved; leg lies over dorsal surface of a combination of features that is most consistent with this order

Schmidt et al. www.pnas.org/cgi/doi/10.1073/pnas.1208464109 1of6 or groups within this order. These features are the following: nating with adhesive structure flanked dorsally by pair of elongate long, slender, setulose legs; antennal flagellum with apparently caudal setae. With only two pairs of anteriorly inserted legs, lack- a reduced number [10] of flagellomeres; antennal flagellum short, ing pairs III–IV. Legs lacking true (paired) claws but with a well- compact, with differentiated flagellomeres (decreased in size api- developed, unpaired empodial featherclaw. Leg setation reduced cad); pretarsal structure, particularly the absence of a lobe-like to maximum of seven setae each, none bothridial; tarsi I-II con- arolium or pulvilliform empodium; and the presence of a putative sistently with a solenidion, usually inserted dorsodistally and of- labellum. ten curved, rod-like in form. Long, slender, setulose legs are widespread in the basal Contains three living families: Eriophyidae, Diptilomiopidae, (“nematocerous”) families of Diptera, and a fleshy labellum (if and Phytopidae. Relationships and classification of the two Trias- correctly identified) is actually a defining feature of the order. sic genera within the Eriophyoidea lineage, below, will be treated Lack of tibial spurs and short coxae preclude most Sciaroidea in a separate paper. (Cecidomyiidae is an exception) and Anisopodidae sensu lato, but otherwise this is a widespread condition among nematocer- Triasacarus Lindquist and Grimaldi, new genus. ans. The monophyletic group Brachycera can be excluded based on flagellomere number, because the groundplan for this subor- Diagnosis. A vermiform eriophyoid with prognathous gnathoso- der is eight flagellomeres. A shortened, compacted antenna with ma; gnathosoma with infracapitulum framed by palpcoxal bases, thick segments occurs sporadically throughout Diptera. The from which other palpal segments extend freely, well separated, pretarsus is unusual, because the empodium and pulvilli are all on either side of a long proboscis flanked dorsally by another pair setiform. Generally, pulvilli are pad-like (pulvilliform) or at least of acuminate, possibly cheliceral structures; prodorsal shield with branched and feathery in most Recent Diptera, although some- anterior margin projected into a frontal lobe; shield with two times quite small. A setiform empodium occurs in the nemato- pairs of setae, one pair inserted anterolaterally in area usually ceran infraorder Culicomorpha (and many Brachycera), and occupied by external vertical setae and one pair inserted poster- pulvilli are lacking altogether in the nematoceran infraorders iorly in area occupied by scapular setae; opisthosoma with ca Tipulomorpha and Psychodomorpha. 55–60 fine annuli that circumscribe body, annuli subequal dorso- The antennal flagellum has a reduced number [10] of compact, ventrally, not broadened into tergites dorsally; each annulus with differentiated flagellomeres, which occur sporadically throughout single transverse row of minute spicules; opisthosoma with pro- nematocerans [e.g., Simuliidae (blackflies), Axymyiidae, Scatop- minent pair of subcaudal setae f and with ventrolateral setae d, e, sidae], and in basal Brachycera (e.g., Stratiomyiidae, Xylomyii- and caudal setae h1 and h2 seemingly evident, other setae indis- dae, etc.), although Recent Brachycera have only eight or cernible; legs I and II with empodial featherclaws long, not bushy, fewer flagellomeres. Culicomorpha are preserved as compres- main shaft divided, tips of each branch bifurcate; legs I and II with sions (mostly just wings) in Late Triassic sedimentary deposits a prominent seta on each of femur, genu, tibia (two setae), and a from the Anisian to Rhaetian, and stem-group Brachycera (lack- short seta on tarsus (not all these verifiable on leg I); leg II with a ing the antennal specialization of recent brachycerans) are known probable solenidion on each of tibia and tarsus (these not verifi- from the Carnian (2). able on leg I); legs III and IV absent. Five principal Late Triassic sedimentary deposits have preserved diverse Diptera compressions from the Anisian to Type species: T. fedelei, sp. nov. Rhaetian (reviewed in ref. 2). These deposits are Mt. Crosby, Queensland (Australia); Stransham, UK; Vosges, France; Dzhai- Trias loucho, Kyrgyzstan; and the solite quarries of Virginia (the last Etymology. Generic from sic (geological period of origin) and acarus (Latin for mite); species name patronymic for Paolo three deposits have produced 80% of the species). Most recent ’ infraorders of Diptera (and some extinct ones) existed by the Fedele (Cortina d Ampezzo), discoverer of the Triassic amber Carnian, even though Diptera only represented approximately deposit. 1% of all insect compressions during the Triassic (2). Many of Triasacarus fedelei Lindquist and Grimaldi, new species. (Fig. 2) the Triassic Diptera were quite small, which agrees with an esti- mated body length of the specimen reported herein, some 1.5– 2 mm. By the Early Jurassic Diptera comprised 20–30% of all Diagnosis. As for genus (monotypic). fossil insect compressions, and nearly half of all inclusions in am- ber from the Cretaceous and Tertiary. Description. Body shape vermiform, cylindrical, total length 210 μm (6.5x greatest width). Body observable mostly just ven- Class: ARACHNIDA trally. Gnathosoma distinctly prognathous, with infracapitulum Subclass: ACARI framed by fused palpcoxal bases. Among some image focal planes Superorder: ACARIFORMES (“f.p.”), infracapitulum with an anterior, underlying ledge with Order: TROMBIDIFORMES narrowly rounded anterior margin (f.p. 2,346–49, 2,448–50), Infraorder: TETRAPODILI above which a long proboscis, seeming with paired structures, Superfamily: Eriophyoidea projects anteriorly and then somewhat ventrally, between freely standing apical segments of palpi (Fig. 2 A, B, and D). A pair of Gnathosoma prognathous, with stylet-like cheliceral struc- acuminate, possibly cheliceral structures flank proboscis dorsolat- tures. Cheliceral bases not enlarged, not forming a stylophore. erally. Palpi with freely extending segments well separated on Palpi maximally four-segmented, without thumb-claw process either side, slightly convergent apically, each with slightly and without spine-like or elongated setae. Prodorsum covered elbowed, slender palp-femur, longer than wide, bearing a short by a prodorsal shield bearing maximally five simple setae, none seta; palp-genu shorter than wide, indistinctly delineated from bothridial. Opisthosoma more or less vermiform, with numerous more apical segments; palp-tibia and -tarsus partly fused, elon- narrow annuli ventrally and sometimes dorsally, or dorsal annuli gate, with short apical seta, and seeming to end in a sinuous, blunt consolidated into a series of fewer, thicker, less flexible “tergites”; apex (f.p. 2,337–50). Coxisternal region of legs I and II apparently annuli provided at least ventrally with a single transverse row of consolidated, but structural details, including setae, not discern- minute spicules. Opisthosomatic setation reduced to maximally ible. Prodorsal shield details observed with difficulty in ventral eight pairs of setae, of which 3–4 lateral pairs displaced ventro- view; anterior margin of shield projected as narrowly rounded laterally to resemble ventral setae. Caudally, opisthosoma termi- frontal lobe over base of gnathosoma (f.p. 2,351–54); faint indi-

Schmidt et al. www.pnas.org/cgi/doi/10.1073/pnas.1208464109 2of6 cations of one pair of long vertical setae discernible and project- At the superfamilial level, the fossil appears to lack an integra- ing anterolaterally from ve position (f.p. 2,179–80, 2,356–58), and tion of gnathosomatic structures into the form that characterizes pair of shorter scapular setae sc, projecting posteriorly, more dis- all known extant members of the superfamily Eriophyoidea. cernibly on right side (f.p. 2,359–63, 2,429). Opisthosoma elon- Although details of form of the cheliceral and associated sty- gate, with ca. 55–60 fine, transverse annuli that encircle the let-like structures are not discernible, the palpi are not formed body circumference and are not differentiated dorsoventrally. as thickened, truncated structures appressed to the infracapitu- Annuli each with single transverse row of minute spicules; these lum and partially enclosing and guiding the feeding structures. longer, spiniform laterally, decreased in size posteriad. Pair of Also, the palpal apices are not truncated distally into adhesive long, fine subcaudal setae f present at level of dorsal opisthoso- lips that flank the feeding structures. This less specialized form matic annuli 55–56, their tips reaching to level of opisthosomatic of the gnathosoma may be viewed as basal (or primitive) to that apex. Other opisthosomatic setae not readily discernible, but in- which characterizes the superfamily and argue for a separate tax- dications of them evident among some focal planes and also on within a higher category that has long been recognized by var- based on positions of faint indications of their insertions, as fol- ious authors as the Tetrapodili. low: d at level of annuli 22–23 (f.p. 2,022–38, 2,156–83), e at level of annuli 37–38 (f.p. 1,926–31, 2,156–83, 2,022–38, 2,482–85), and Ampezzoa Lindquist and Grimaldi, new genus. elongate caudal setae h2 and short accessory setae h1 on terminal – – – annuli before anal lobe (f.p. 2,206 13, 2,412 16, and 2,479 89). Diagnosis. A distinctive fusiform eriophyoid with prognathous Anal lobe small, slender, with possible accessory lobes. Genital gnathosoma; gnathosoma framed by lateral margins of palpal opening and setae not observable. Legs slender, without spine- segments, between which an infracapitulargutterbearsaset like projections or serrations evident on any segments; trochanter of indistinguishable structures (some cheliceral); anterior portion ca. and femur thickest segments on legs I and II; tibia I longer ( of idiosoma broad, posterior end tapered; body dorsoventrally – 1.4x) than tibia II. Femur of legs I II with long, fine, ventral seta flattened; prodorsal shield well developed, with posterior margin – – – (f.p. 2,323 26, 2,447 50); genu of legs I II short, with long, dorsal well defined, posterior region of shield possessing pair of closely – – seta (f.p. 2,335 40); tibia of legs I II slightly longer than tarsus, set dorsal tubercles, each with scapular seta; shield also with pos- with a long ventrobasal seta and a long, fine, dorsoapical seta and sible indications of an unpaired internal vertical seta anteriome- adjacent lateroapical solenidion (f.p. 2,330–37) (solenidion and – dially and a pair of external vertical setae midlaterally; opistho- ventral seta not discernible on tibia I). Tarsus of legs I II with soma with approximately 18 tergites, 1–15 bear pair of digitiform short dorsal seta and moderately long, curved, blunt-tipped sole- lateral lobes; lateral lobes and portions of opisthosomatic tergites nidion subapically (about 1.2x longer than empodial featherclaw) – apparently wax-secreting; telosome with prominent pair of caudal (f.p. 2,330 33; Fig. 2). Empodial featherclaw long, length ap- setae h2; opisthosoma with ventrolateral setae d, e, f, and acces- proximately equal to that of tibia + tarsus on leg II, 8- or 9-rayed, sory caudal setae h1 seemingly evident, other setae indiscernible; rays simply branched, with central shaft divided (f.p. 1,989–92, – – – – A E–G leg I with 5 discernible setae, of which one appears to emanate 2,022 23, 2,068 69, 2,323 26, 2,336 37; Fig. 2 and ). from each of femur, genu, tibia, tarsus, and a second one from tarsus; tarsal solenidion with enlarged tip; tarsus I long and slen- Type. Holotype, no. MGP 31343, in amber from outcrops of the der; empodial featherclaw moderately long, not bushy, main shaft Heiligkreuz Formation [late Julian to early Tuvalian in age (Late possibly divided, tips of each branch bifurcate; leg pair II present, ca. Carnian: 230 Ma)] near the village of Cortina in the Dolomite but structure not discernible; legs III and IV absent. Alps of northeastern Italy. Museum of Geology and Paleontol- ogy, University of Padova, Italy. Type species: A. triassica, n. sp.

Comments. A superfamily-level attribution of the fossil can be Etymology. Generic from Valle d’Ampezzo (Anpezo in local dia- made via a process of exclusion as well as by use of the specialized lect) in northeastern Italy, source of Triassic amber outcrops; spe- features that are preserved. Among acariform mites, no other tax- cies name for period of geological origin. on than Eriophyoidea is characterized by the apomorphic attri- butes of an elongated, vermiform body that terminates caudally Ampezzoa triassica Lindquist and Grimaldi, new species (Fig. 3) with an adhesive anal structure, loss of legs III–IV, and the re- maining legs equipped with empodial featherclaws (3). Within the superfamily, the immediately most notable attributes are Diagnosis. As for genus (monotypic). the presence of a ventral seta and a solenidion on tibia II (the presence of these structures on leg I could not be confirmed). Description. Body fusiform, dorsoventrally flattened, total length Among extant taxa of Eriophyoidea, the presence (retention) 124 μm, greatest width ca. 50 μm (Fig. 3A and B). Body as pre- of a tibial seta has been noted as an atavism in only one species served is largely just visible from dorsal surface. Gnathosoma of the relatively derived eriophyid genus Abacarus (4), while a prognathous, clearly demarcated from idiosoma. Greatest width tibial solenidion is found only among some of the genera of of gnathosoma 0.4x that of prodorsal shield; length of sclerotized the putatively early-derived family Phytoptidae and, even among portion of gnathosoma slightly less than that of prodorsal shield. these taxa, only on leg I. Therefore, the presence of these struc- Infracapitulum with an anterior, underlying ledge with broadly tures on the tibia of leg II is thought to be even more ancestral rounded anterior margin (f.p. 2,606–09; 3,006–09). Lateral within the lineage. The presence of a pair of prodorsal setae with margins of gnathosoma outlined by palpcoxal bases and outer insertions in the area occupied by the pair of external vertical margins of apparently three segments (palpal trochanter-femor- setae, ve, is also notable, because only a handful of genera in ogenu, palp-tibia, palp-tarsus). Within these margins is a pair of the same family Phytoptidae retain these setae. Other attributes undulate dorsal ridges that may represent walls of a stylet sheath characteristic of Phytoptidae are not discernible in the fossil of or infracapitular gutter (Fig. 3 A–C), within which lie a complex Triasacarus, in particular the form of the female genitalia, with of structures; each ridge bears a bluntly rounded dorsal projec- spermathecal tubes longer than the diameter of the spermathecal tion, alongside the palp-tibial base, that may represent a chelic- sacs. Not only are these structures internal (and therefore not eral retainer. Chelicerae not discernible amidst adjacent visible externally in the fossil), but what surfaces are visible do gnathosomatic structures. Prodorsal shield large, well sclerotized, not allow determining whether the specimen is a female, or somewhat trapezoidal in shape, with anterior margin as shortest indeed, even an adult. side, lateral margins divergent posteriad, posterior margin long-

Schmidt et al. www.pnas.org/cgi/doi/10.1073/pnas.1208464109 3of6 est and slightly convex (Fig. 3 A and B). Pair of low, paramedian the presence of strongly differentiated tergites is generally ex- dorsal tubercles near posterior margin of prodorsal shield, each pressed only in the adult stage of eriophyoid mites. tubercle with short internal scapular seta sc directed posterome- Unlike Triasacarus, the specimen of Ampezzoa presents an in- dially; length of seta ca. 2x the diameter of tubercle; possible in- tegration of gnathosomatic structures into much of the form that dications of other prodorsal setae evident among a few focal is apotypical of all known extant members of the superfamily plane images—i.e., unpaired internal vertical seta vi anterome- Eriophyoidea. Therefore, these two fossil taxa may be regarded dially (f.p. 2,607–09, 3,006–08)—paired external vertical setae as only distantly related to one another in the eriophyoid lineage ve midlaterally (f.p. 2,597–2,601, 2,663–68, 2,999–3,002). of the Tetrapodili, with no possibility of their being dimorphic Opisthosoma pyriform, with caudal end tapered narrowly (telo- forms of the same species. Although details of structure of the some width ca. 0.3x that of anteriormost segment); with 18 fully cheliceral and associated stylet-like structures are not discernible, transverse, delineated tergites. Tergites 1–14eachwithsingle in Ampezzoa the palpi are formed as thickened, truncated struc- transverse row of minute spicules along posterior margin, and lat- tures appressed to the infracapitulum and partially enclosing and erally projecting into thin digitiform lobes (some perhaps bifur- guiding the feeding structures. In contrast, in Triasacarus, the pal- cate at apex, e.g., t10–13; Fig. 3 A and B). Very fine, transparent, pi appear to be well-separated, free-standing structures some- striated “webbing” appears to connect between lobes and extends what reminiscent of the form seen in some cunaxid mites. dorsally over opisthosoma in places; striations oblique, but fine striations over t8–12 transverse. Tergite 15 without denticles or Amber from the Late Triassic (Carnian) of Europe and Other Sites in striations, but with pair of sharp, posterolateral, spine-like lobes. the World Tergites 16 and 17 simple. Pair of long caudal setae h2 present Two types of Triassic amber have been found in the Dolomite (having ventral insertion) (f.p. 2,533–35; Fig. 3 B and E), lengths outcrops. The first occurs as fractured brittle pieces of up to approximately equal to lengths of segments 15 þ 16 þ 17. Other 30-mm size embedded in sandstone and hybrid-sandstone, mixed opisthosomatic setae not readily discernible but possible indica- with shells of large bivalves. These fossils indicate a marginal mar- tions evident among some focal plane images—i.e., ventrolateral ine environment. Abundant plant fragments, including horsetails, seta d on both sides at level under tergite 6 or 7 (f.p. 2,597–98, conifer twigs, wood, and leaflets, suggest a rich coastal vegetation 2,952–53), ventrolateral seta e on right side at level under tergite (9). This resin was transported prior to deposition. The second 10 (f.p. 2,596–99, 2,892–94, 2,952–54, 2,999–3,000), ventrolateral amber type is represented by abundant, isolated, and well-pre- seta f on both sides at level under tergite 13 (f.p. 2,589–92, 2,945– served autochthonous drops, which occur in large number within 48, 2,994–95), on right side (f.p. 2,632–34, 2,873–74) and on left a lignitic paleosol (2–5% by volume). These amber droplets are side (f.p. 2,780–81, 2,832–34, 2,886–88), and short accessory cau- ovoid to elongated (Fig. 1F and Fig. S1) and mostly 2–6mmin dal seta h1 (f.p. 2,529–30, 2,765–66). A pair of long coxisternal size. Their color ranges from yellow to russet and they possess a setae, either 1a or 2a, seem evident in a few images (f.p. resinous luster. The surface of the amber droplets often shows 2,561–66). Leg pairs I and II present (right legs obscured beneath reticulate desiccation marks, suggesting exposure to air or sun prodorsal shield), left leg I best revealed, some segmentation pre- that caused a fast evaporation of volatile components in the resin. served but basal segments difficult to interpret (Fig. 3B and D). The small size of the amber droplets and the presence of a small Leg I with 5 long, fine setae discernible, of which one each inter- peduncle suggest a narrow point of exudation from the plant. preted to insert on femur (ventrally), genu (dorsally) and tibia Coeval findings of Triassic amber (see Fig. S3) suggest that small (dorsally), and two on tarsus (dorsobasally) (f.p. 2,690–94, droplets are typical for Triassic ambers. Although strong com- 2,752–57, 3,015–19). Leg I tarsus long, slender, length 1.75x that pression due to the weight of superimposed sediments occurred of empodial featherclaw; featherclaw simply branched (central during the geological history of the Dolomites (the paleosol is shaft possibly divided) (f.p. 2,611–12, 2,904–06), not bushy; tarsal interposed between compact white dolomite levels), the softness solenidion with enlarged tip (f.p. 2,755–57, 2,916–19, 3,020–21). of the sediment constituting the paleosol (mainly clay and cuticle Minute portion of left leg II preserved with attached setae. remains) permitted optimal preservation of the amber. Consider- ing the sometimes larger size of the first amber type embedded in Type. Holotype, no. MGP 31344, in amber from outcrops of the the sandstone (9), we might hypothesize that future field work Heiligkreuz Formation [late Julian to early Tuvalian in age (Late within the paleosol will give access to hydrodynamically selected Carnian: ca. 230 Ma)] near the village of Cortina in the Dolomite larger amber pieces that might consequently be a source of larger Alps of northeastern Italy. Museum of Geology and Paleontol- Triassic arthropods. ogy, University of Padova, Italy. Triassic amber from the Dolomite Alps of Italy is preserved in the Heiligkreuz Formation, which corresponds to an interval be- Comments. A superfamily-level attribution of the fossil can be tween the Early and Late Carnian. In Italy, Triassic amber has made with the same rationale given above for Triasacarus. The also been found in a second distinct stratigraphic unit in the most notable attribute of this specimen is the form of the opistho- Southern Alps [Rio del Lago Formation in the Julian Alps; Fig. S3 soma, with its thickened tergites individually extended into lateral (9)]. Both sites are located in the westernmost Tethyan margin of projections that appear to be capable of forming waxy secretions. the Eurasian plate during the Mesozoic. In the same chronostra- Among extant taxa of Eriophyoidea, the body form of Cymeda tigraphic interval, amber outcrops have also been found in several zealandica Manson & Gerson (5) is remarkably similar to this fos- other areas of Europe, including Austria [Raibler Schichten (10), sil. A monotypic genus in the tribe Acaricalini (subfamily Phyllo- Lunzerschichten (11)], Hungary [Sándorhegy Formation (12)], coptinae, family Eriophyidae), this taxon is known only from Spain [Keuper facies, Alicante (13)], and Switzerland [Schilfsand- fernsinNewZealand(6).AsinthefossilAmpezzoa,derived stein Formation (14–16)]. A comparative stratigraphic section but possibly homoplasious attributes characteristic of extant shows that the occurrence of Triassic amber in Europe as well Acaricalini are the empodial featherclaw of legs I–II moderately as in other parts of the world (Arizona, South America, and South to deeply divided, and the dorsal opisthosomatic annuli coalesced Africa) was roughly contemporaneous in the Carnian (Fig. S3; into fewer, enlarged tergite-like thickenings, sometimes extended refs. 17–21; stratigraphy based on ref. 22). laterally as lobes (7, 8). However, none of the extant acaricalines These findings suggest three possible scenarios for the Carnian retains either of the vertical pairs of prodorsal setae. Although amber anomaly: (i) changes in sedimentation that facilitated the details of the ventral surface of the fossil are not clear enough preservation of resins; (ii) increased production of resins; and (iii) to determine whether it is an adult (with external genital struc- both factors acting together. After the Permian mass extinction, tures) or immature (without them), it is likely an adult, because and the following slow recovery of environments and biota in the

Schmidt et al. www.pnas.org/cgi/doi/10.1073/pnas.1208464109 4of6 Early Triassic, the Middle and Late Triassic was a relatively stable demonstrated (25), major climatic changes and mass extinctions period, interrupted only by a global episode of wet conditions appear to be associated with C-isotope anomalies in the atmo- (23), with a consequent transition to more mesic floras (24) and sphere-ocean system. In this context, amber is potentially a pa- an increase in the production of conifer resins (9). As recently leoclimatic indicator (9, 26).

1. Grimaldi DA, Engel MS (2005) Evolution of the Insects (Cambridge Univ. Press, Cam- 14. Soom M (1984) Amber from the northern rim of the Swiss Alps. Stutt Beitr Natkd Ser C bridge, UK). 18:15–20, German. 2. Blagoderov V, Grimaldi DA, Fraser NC (2007) How time for flies: Diverse Diptera 15. Kelber KP (1990) The sunken flora of the Main swamps and of Main-Franconia 203 from the Triassic of Virginia and early radiation of the order. Amer Mus Novit million years ago. Beringeria 1:1–67, German. 3572:1–39 16. Kelber KP, Hansch W (1995) Keuperpflanzen. Die Enträtselung einer über 200 Million- 3. Lindquist EE (1996), Eriophyoid Mites: Their Biology, Natural Enemies, and Control, eds en Jahre alten Flora. [Keuper plants. The unraveling of a flora more than 200 million – Lindquist EE, Sabelis MW, Bruin J (Elsevier, Amsterdam), pp 301–327. years old] Museo 11:1 157, German. 4. Navia D, et al. (2011) A new species of Abacarus (Acari: Prostigmata: Eriophyidae) 17. Koken E (1913) Contributions to the knowledge about the strata of Heiligenkreuz (Ab- – damaging sugarcane, Sacharrum officinarum L., from Costa Rica—The first eriophyoid tei Valley, Southern Tirol). Abh K K Geol Reichsanstalt 16:1 43, German. mite described with a tibial seta on leg II. Zootaxa 3025:51–58. 18. Csillag C, Földvári M (2007) Upper Triassic amber fragments from the Balaton High- lands, Hungary. Annual Report of the Geological Institute of Hungary, 2005. (Geolo- 5. Manson DCM, Gerson U (1986) Eriophyoid mites associated with New Zealand ferns. gical Institute of Hungary, Budapest), pp 37, 46, Hungarian. NZ J Zool 13:117–129. 19. Litwin RJ, Ash SS (1991) First early Mesozoic amber in the western hemisphere. Geol- 6. Gerson U (1996) Eriophyoid Mites: Their Biology, Natural Enemies, and Control, eds ogy 19:273–276. Lindquist EE, Sabelis MW, Bruin J (Elsevier, Amsterdam), pp 227–258. 20. Ansorge J (2007) FossilsX3: Insects, Arthropods, Amber (Abstracts), ed. J. Alonso (Alava 7. Lindquist EE, Amrine JW (1996) Eriophyoid Mites: Their Biology, Natural Enemies, and Museum of Natural Science, Vitoria-Gasteiz, Spain), pp 52–54. Control, eds Lindquist EE, Sabelis MW, Bruin J (Elsevier, Amsterdam), pp 33–87. 21. Colombi C, Parrish JT (2008) Taphonomy of the paleofloral assemblages in the Ischi- 8. Amrine JW, Stasny TA, Flechtmann CHW (2003) Revised Keys to World Genera of Erio- gualasto Formation, Upper Triassic (Carnian), Argentina. Palaios 23:778–795. phyoidea (Acari: Prostigmata) (Indira Publishing, West Bloomfield, MI). 22. Ogg JG, Ogg G, Gradstein FM (2008) The Concise Geologic Time Scale (Cambridge Univ. 9. Roghi G, Ragazzi E, Gianolla P (2006) Triassic amber of the southern Alps (Italy). Palaios Press, Cambridge, UK), p. 102, Alternate Time Scale for the Triassic. – 21:143 154. 23. Simms MJ, Ruffell AH (1989) Synchroneity of climatic change and extinctions in the 10. Pichler A (1868) Contributions to the geognosis of Tyrol. XI. Fossil Resin. Jb K K Geol Late Triassic. Geology 17:265–268. Reichsanstalt 18:45–52, German. 24. Roghi G, et al. (2010) Palynological correlation of Carnian humid sub-events through- 11. Sigmund A (1937) Minerals of Lower Austria (Deuticke, Wien-Leipzig), German. out western Tethys. Palaeogeogr Palaeoclimatol Palaeoecol 290:89–106. 12. Budai T, et al. (1999) Geology of the Balaton Uplands (Geological Institute of Hungary, 25. Dal Corso J, et al. (2012) Discovery of a major negative δ13C spike in the Carnian (Late Budapest), Hungarian. Triassic) linked to the eruption of Wrangellia flood basalts. Geology 40:79–82. 13. Peñalver E, Delclòs X (2010) Biodiversity of Fossils in Amber from the Major World 26. Gianolla P, Ragazzi E, Roghi G (1998) Upper Triassic amber in the Dolomites (northern Deposits, ed. Penney D. (Siri Scientific Press, Manchester, UK), pp 236–270. Italy). A paleoclimatic indicator? Riv Ital Paleontol Stratigr 102:381–390.

Fig. S1. Typical appearance of numerous amber droplets found in the paleosol of the Dolomite Alps Triassic outcrop. Scale bar∶5 mm.

Schmidt et al. www.pnas.org/cgi/doi/10.1073/pnas.1208464109 5of6 Fig. S2. Photomicrographs of Diptera specimen in Dolomite Triassic amber, MGP (Museum of Geology and Paleontology, University of Padova, Italy) 31345. (A) Entire specimen with dissociated appendages. (B) Head, detail. (C) Detail of antenna and adjacent leg, showing oval sensory area, possibly Johnston’s organ. (D) Detail of dissociated distitarsus. (E) Dissociated legs. (F) Same dissociated legs as in E, from opposite side and taken before embedding preparation. All photos using bright field (BF) illumination; all images are stacked composites of individual focal planes. Scale bars: A, 200 μm; B, E, and F, 100 μm; C and D, 50 μm.

Fig. S3. Stratigraphic chart of the main Triassic amber deposits of the world. NCA, Northern Calcareous Alps; FM., Formation; MB., Member. References are indicated in parentheses. Geochronological scale according to Ogg et al. (22).

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