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Exceptionally preserved shark fossils from Mexico elucidate the long-standing enigma of the Cretaceous elasmobranch Ptychodus
<p>The fossil fish <em>Ptychodus</em> Agassiz, 1834, characterized by a highly distinctive grinding dentition and an estimated gigantic body size, has remained one of the most enigmatic extinct elasmobranchs (i.e. sharks, skates, and rays) for nearly two centuries. This widespread Cretaceous taxon is common in Albian to Campanian deposits from almost all continents. However, specimens mostly consist of isolated teeth or more or less complete dentitions, whereas cranial and post-cranial skeletal elements are very rare. Here we describe newly discovered material from the early Late Cretaceous of Mexico, including complete articulated specimens with preserved body outline, which reveals crucial information on the anatomy and systematic position of <em>Ptychodus</em>. Our phylogenetic and ecomorphological analyses indicate that ptychodontids were high-speed (tachypelagic) durophagous lamniforms (mackerel sharks), occupying a specialized predatory niche previously unreported among both extant and extinct elasmobranchs. Our results support the view that lamniforms were ecomorphologically highly diverse and represented the dominant group of sharks in Cretaceous marine ecosystems. <em>Ptychodus</em> may have fed predominantly on nektonic hard-shelled prey items such as ammonites and sea turtles rather than on benthic invertebrates, and its extinction during the Campanian, well before the end-Cretaceous crisis, might have been related to competition with emerging blunt-toothed globidensine and prognathodontine mosasaurs.</p>
Fig. 4. Callichimaeraperplexa n. gen. n in Exceptional preservation of mid-Cretaceous marine arthropods and the evolution of novel forms via heterochrony
Fig. 4. Callichimaeraperplexa n. gen. n. sp., reconstruction. (A and B) Line drawing of dorsal (A) and ventral (B) views. Colorsdenoteconvergent traitswith other decapod groups: light gray, dorsal carapace, similar to some lobsters and †Palaeocorystidae (Brachyura: Raninoida); dark gray, large eyes, similar to †Ekalakia (Dromiacea: †Glaessneropsidae) and several homoloids; red, pediform mxp3 bearing acrista dentata, similarto lobsters, most anomurans, and early-branching brachyurans (Homolodromioidea, most Homoloidea); yellow, spanner-like P1, similar to Hippoidea (Anomura), Raninoidea (Brachyura), and †Retrorsichela; green, flattened paddle-like legs P2 and P3, similar to Matutidae (Brachyura: Eubrachyura); blue, reduced legs P4 and P5, as in hermit crabs (Anomura: Paguroidea), podotreme brachyurans (e.g., Homolodromioidea and Cyclodorippoidea) and early-branchingeubrachyurans (e.g., Dorippoidea); orange, sternites S5 and S6 similar to †Retrorsichela, and Heikeopsis (Eubrachyura: Dorippoidea); and purple, symmetrical pleon lacksarticulatedrings anduropods/uropodal plates, likemost brachyurans. (C and D) Digital reconstruction. (C) Dorsal view. (D) Ventralview. A, pleon; Cx, coxa;Is, ischium; Ma, manus. Scalebar,10 mm. See movies S1 and S2 anddata file S2 for three-dimensional (3D) viewableand printable models. Line drawings by J.L. and 3D reconstructions by A.D.
Fig. 3 in Exceptional preservation of mid-Cretaceous marine arthropods and the evolution of novel forms via heterochrony
Fig. 3. Pleon and sexual dimorphism in Callichimaera perplexa n. gen. n. sp., lower Upper Cretaceous, Colombia. (A) Paratype IMG p881217, female, dorsal view showing thelegs 2 to 5 and thepleonites 1 to 4, two of them bearing anaxialtubercle. (B) Paratype IGMp881209b, female,showing the unfoldedpleon in lateralview and pleonites 2 to 5 andmultiple pairs of pleopods (white arrows). (C) Paratype MUN-STRI 27045-06, female, dorsalview, showing pleonites 1 to 6, theroundish telson, and the lack of uropods or uropodal plates. (D and E) Paratype IGM p881202, male. (D) Ventral view showing the chelipeds, pereopods, and pleon with sclerotized gonopods 1 and 2 [white box; see (E)]. (E) Close-upof sclerotized male gonopods 1 and 2 (white arrows), anarrow pleonite 6, anelongate telson,and the lackof uropods oruropodal plates. (F) Linedrawing of (E). (G) Paratype MUN-STRI 27045-09, a very small individual, possibly a megalopa or early postlarvalstage, preserving small and slender legs 2 and 3 compared to adult forms [see (I)]. (H) Paratype IGM p881220, juvenile, with legs 2 and 3, the pleon bearingan axial tubercle, and with a bifidrostrum anda pairof largecompoundeyesbearinghexagonalfacets (Fig.2G). (I) Paratype IGMp881206,adult female, ventralviewpreservingspanner-likechelipeds,longoar-likelegsor P2 and P3, reduced andslender legsor P4 and P5, and abroad pleon. G1 and G2, malegonopods 1 and 2; Pl1 to Pl6, pleonites 1 to 6; T, telson. All specimens were photographed dry; specimens in (A), (C), (D), and (G) to (I) were coated with ammonium chloride; specimens in (B) and (E) were uncoated. Photos by J.L.
Fig. 6 in Exceptional preservation of mid-Cretaceous marine arthropods and the evolution of novel forms via heterochrony
Fig. 6. Convergent decarcinized body forms in various families of false and true crabs and convergent appendages in swimming and/or fossorial arthropods. (A to I) Decarcinized crabs. (A to D) Anomura. (A to C) Molecrabs,Hippoidea: (A) Hippidae, Hippamarmorata, Taiwan.(B) Blepharipodidae: Blepharipodaoccidentalis. (C) Albuneidae: Albunea occulta, Taiwan. (D) Porcelain crabs, Galatheoidea: Porcellanidae: Euceramus panatelus, Panama. (E to I) Brachyura. (E and F) Frog crabs, Raninoidea: Raninidae: (E) Raninoides benedicti, Panamá. (F) Symethis sp. Panamá. (G and H) Masked burrowing crabs, Eubrachyura: Corystoidea: Corystidae: (G) Corystes cassivelaunus, Belgium. (H) Jonas distinctus, Taiwan. (I) Chimera crab, †Callichimaeroidea: Callichimaeridae: Callichimaeraperplexa n. gen. n. sp., Cenomanian-Turonian (95 to 90 Ma ago) of Colombia. (J to Q) Otheraquatic arthropods with modified appendages for swimming and/or digging. (J) Sea scorpions, Chelicerata: †Eurypterida: Eurypterusremipes, YPM 211521, upper Silurian, New York. (K to M) Insecta. (K) Coleoptera: Dytiscidae: Cybisterfimbriolatus, YPMENT.959234; (L) Hemiptera: Corixidae: Hesperocorixa kennicottii, YPM ENT.452851; (M) Hemiptera: Notonectidae: Notonectaundulate, last instarnymph, Alberta, Canada. (N) Isopoda: Munnopsidae: Munnopsis longiremis, USNM 113282, Baja California, Mexico. (O to Q) Brachyura. (O) Orithyioidea: Orithyiidae: Orithyia sinica, USNM 134243, China. (P) Calappoidea: Matutidae: Matutavictor. (Q) Portunoidea:Portunidae:Arenaeus cribrarius. Photosby T. Y. Chan (A, C, and H), C. Boyko (B), A. Anker (D, E, and N), J.L. (F and O), H. Hillewaert (G), J. Utrup (J), E. Lazo-Wasem (K and L), J. Acorn (M), and O. Radosta (P and Q).
Fig. 2 in Exceptional preservation of mid-Cretaceous marine arthropods and the evolution of novel forms via heterochrony
Fig. 2. Dorsal, frontal, and ocular features in Callichimaera perplexa n. gen. n. sp., from the mid-Cretaceous of Colombia. (A and B) Paratype MUN-STRI 27044-02, showing three specimensin dorsal view (left), ventral view (right), and anisolated dorsal carapace (bottom center). (A) Color image. (B) Inverted color image highlighting details of thecarapace outlines, clawsand legs, andlarge eyes. (C) Paratype IMGp881203, dorsalview, showingdetails of thecarapace groovesandridgesandthelackof true orbits. (D) Paratype IMGp881218, dorsal view, showing details of dorsal carapace, claws, andlarge oar-like legs 2 and 3. (E and F) Paratype IGM p881209a.(E) Scanning electron microscope (SEM) image showing details of large compound eye, optical lobe, rostrum (R), and two pairs of short antennae between the eyes. (F) SEM showing aclose-upof theoptical lobe. (G) Paratype IGMp881220,SEM oflargeeyepreservingmostlyhexagonalfacetsinhexagonalarrangement (left box), althoughtheproximal cornea bears squarishfacets inorthogonal packing (right box). (H to J) Paratype IGMp881208, ventral view,(H) showing thelargeeyes and bifidrostrum,(I) SEMof ventral right eye, and (J) SEMof ventralleft eye.An1, antenna 1 orantennula; An2,antenna 2; Bcg, branchio-cardiac groove; Cg, cervical groove; E, compound eye; Lr,longitudinal ridge; Ol, opticallobe; Pfr, postfrontal longitudinal ridge; Pgr,protogastric longitudinal ridge.All specimenswere photographed dry; specimensin (C) and (D) were coated with ammonium chloride; specimens in (A), (B), and (E) to (J) were uncoated. Photos by J.L.
Fig. 1 in Exceptional preservation of mid-Cretaceous marine arthropods and the evolution of novel forms via heterochrony
Fig. 1. Ventraland appendicularfeatures in Callichimaera perplexa n. gen. n. sp., fromthemid-Cretaceous of Colombia. (A to C) Holotype IGM p881215.(A) Ventral view. (B) Close-up of sternal plates, sutures, and linea media (lm); the white arrow shows a spine in the third leg coxae. (C) Close-up of sternal crown, mandibles, and maxillipeds 2 and 3, thelatterbearingarowof spinesorcristadentata (cd) intheischium. (D and E) Paratype IGMp881196. (D) Ventralview. (E) Close-upof sternumand sutures. (F) Paratype IGM p881185, showingdetails of thespanner-like claw. (G to I) Paratype IGM p881214. (G) Close-up of the large oar-like legs 2 and 3. (H) Ventralview showing the sternites, claws, legs, and pleon. (I) Close-up of the reduced and slender legs 4 and 5. Ba, basis; Ca, carpus; Da, dactylus; Pr, propodus; Ma, mandibula; Me, merus; Mxp2 and Mxp3, second and third maxillipeds, respectively; P1, claw or cheliped; P2 to P5, pereopods or legs 2 to 5; Po, pollex or fixed finger tip; Pl, pleon or "abdomen"; S1 to S7, sternites 1 to 7; 4/5–6/7, sternal sutures. All specimens were photographed dry; specimens in (A), (D), (E), (G), and (H) were coated with ammonium chloride; specimensin (B), (C), (F), and (I) were uncoated. Photos by J.L.
Fig. 5 in Exceptional preservation of mid-Cretaceous marine arthropods and the evolution of novel forms via heterochrony
Fig. 5. Phylogenetic relationships among the main families, superfamilies, and sections of "true" crabs or Brachyura. Tree topology after figs. S5 and S6. Each color and letter (encircled) represents one of the nine major brachyuran evolutionary branches. Dromiacea (red) (A) and Homoloida (purple) (B) are first known from the Jurassic, while †Callichimaeroida section nov. (dark blue) (C), †Torynommoida (light blue) (D), †Etyoida (dark green) (E), Raninoida (light green) (F), †Dakoticancroida (yellow) (G), Cyclodorippoida (orange) (H), and Eubrachyura or "higher" brachyurans (brown) (I) are all first known from the Cretaceous.Thick solid lines represent the ages of the first and last occurrences of each family within the main lineages. Dotted lines and daggers indicate extinct taxa; solid lines indicate living taxa. Black triangles indicate that in Anomura,decarcinization has occurred twice (anomuran clades not illustrated).White triangles indicate the three Brachyura lineages where decarcinization has occurred. Figure by J.L.
Data from: Crypsis in the pelagic realm: evidence from exceptionally preserved fossil fish larvae from the Eocene Stolleklint Clay of Denmark
<p>Marine deposits of earliest Eocene age in northern Jutland, Denmark, are renowned for yielding diverse teleost assemblages that have proved central for enhancing our understanding of the early evolution of many extant actinopterygian clades. In this study, we investigate diminutive larval fish fossils from the Stolleklint Clay<b>,</b> Ølst Formation, that retain multiple soft-tissue features preserved as distinct dark-coloured stains. In order to examine the elemental and molecular composition of these soft parts, we employed a combination of time-of-flight secondary ion mass spectrometry (ToF-SIMS), scanning electron microscopy (SEM) and energy-dispersive X-ray spectroscopy (EDS). Our analyses revealed that the preserved structures contain chemically identifiable eumelanin intimately associated with densely aggregated microbodies that are morphologically consistent with melanosome organelles. Thus, we conclude that the carbonaceous structures represent traces of originally melanized body parts, including the eyes and peritoneum. Comparable pigmentation patterns are seen in many extant teleost larvae that use semi-transparency as a means of camouflage in pelagic environments, to suggest a similar visual appearance of the Stolleklint Clay fish fossils. This in turn suggests that adaptations for concealment and UV-protection had evolved already by the beginning of the Eocene, notably during a time interval characterized by an extreme greenhouse climate, when the global fish fauna become increasingly modern in composition.</p>
Exceptionally preserved shark fossils from Mexico elucidate the long-standing enigma of the Cretaceous elasmobranch Ptychodus
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Data from: Crypsis in the pelagic realm: evidence from exceptionally preserved fossil fish larvae from the Eocene Stolleklint Clay of Denmark
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Data from: An exceptionally preserved Jurassic skink suggests lizard diversification preceded fragmentation of Pangaea
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Data from: Exceptionally preserved stomach contents of a young tyrannosaurid reveal an ontogenetic dietary shift in an iconic extinct predator (TMP 2009.12.14 Citipes tail segment)
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Data from: Exceptional preservation requires fast biodegradation: the case study of thylacocephalan specimens from La Voulte-sur-Rhône (Callovian, Jurassic, France)
<p>Konservat-Lagerstätten are seen as snapshots of past biodiversity for a given location and time. Yet, processes leading to the exceptional morphological preservation of fossils in these deposits remain incompletely understood. This results in the deficient assessment of taphonomic biases and limits the robustness/relevance of palaeobiological reconstructions. Here, we report the mineralogical characterization of crustacean fossils preserved within carbonate-rich concretions from the Jurassic Konservat-Lagerstätte of La Voulte-sur-Rhône (Ardèche, France). The combination of SEM-EDS, TEM, synchrotron-based XRF, XRD and XANES allows identifying the mineralogical phases composing these fossils (i.e. fluorapatite, Fe-sulfides (pyrite, pyrrhotite) and Mg-calcite) and the surrounding matrix (i.e. Mg-calcite, clays and detrital silicates). Fluorapatite and pyrite (and pyrrhotite) precipitated during decay under anoxic conditions, replacing delicate organic structures and preserving anatomical details. These mineral structures were subsequently consolidated by a Mg-calcite cement. Of note, histologically similar tissues were replaced by the same mineral phases, confirming that 'fossilization [in La Voulte] occurred rapidly enough to be influenced by tissue composition'. Altogether, the present study shows that exceptional preservation requires fast biodegradation, thereby confirming recent experimental evidence.</p>
Data from: A new ankylosaurine dinosaur from the Judith River Formation of Montana, USA, based on an exceptional skeleton with soft tissue preservation
The terrestrial Judith River Formation of northern Montana was deposited over an approximately 4 Myr interval during the Campanian (Late Cretaceous). Despite having been prospected and collected continuously by palaeontologists for over a century, few relatively complete dinosaur skeletons have been recovered from this unit to date. Here we describe a new genus and species of ankylosaurine dinosaur, Zuul crurivastator, from the Coal Ridge Member of the Judith River Formation, based on an exceptionally complete and well-preserved skeleton (ROM 75860). This is the first ankylosaurin skeleton known with a complete skull and tail club, and it is the most complete ankylosaurid ever found in North America. The presence of abundant soft tissue preservation across the skeleton, including in situ osteoderms, skin impressions and dark films that probably represent preserved keratin, make this exceptional skeleton an important reference for understanding the evolution of dermal and epidermal structures in this clade. Phylogenetic analysis recovers Zuul as an ankylosaurin ankylosaurid within a clade of Dyoplosaurus and Scolosaurus, with Euoplocephalus being more distantly related within Ankylosaurini. The occurrence of Z. crurivastator from the upper Judith River Formation fills a gap in the ankylosaurine stratigraphic and geographical record in North America, and further highlights that Campanian ankylosaurines were undergoing rapid evolution and stratigraphic succession of taxa as observed for Laramidian ceratopsids, hadrosaurids, pachycephalosaurids and tyrannosaurids.
Data from: An exceptionally preserved Late Devonian actinopterygian provides a new model for primitive cranial anatomy in ray-finned fishes
Actinopterygians (ray-finned fishes) are the most diverse living osteichthyan (bony vertebrate) group, with a rich fossil record. However, details of their earliest history during the middle Palaeozoic (Devonian) 'Age of Fishes' remains sketchy. This stems from an uneven understanding of anatomy in early actinopterygians, with a few well-known species dominating perceptions of primitive conditions. Here we present an exceptionally preserved ray-finned fish from the Late Devonian (Middle Frasnian, ca 373 Ma) of Pas-de-Calais, northern France. This new genus is represented by a single, three-dimensionally preserved skull. CT scanning reveals the presence of an almost complete braincase along with near-fully articulated mandibular, hyoid and gill arches. The neurocranium differs from the coeval Mimipiscis in displaying a short aortic canal with a distinct posterior notch, long grooves for the lateral dorsal aortae, large vestibular fontanelles and a broad postorbital process. Identification of similar but previously unrecognized features in other Devonian actinopterygians suggests that aspects of braincase anatomy in Mimipiscis are apomorphic, questioning its ubiquity as stand-in for generalized actinopterygian conditions. However, the gill skeleton of the new form broadly corresponds to that of Mimipiscis, and adds to an emerging picture of primitive branchial architecture in crown gnathostomes. The new genus is recovered in a polytomy with Mimiidae and a subset of Devonian and stratigraphically younger actinopterygians, with no support found for a monophyletic grouping of Moythomasia with Mimiidae.
Data from: Exceptional preservation and the fossil record of tetrapod integument
The fossil record of exceptionally preserved soft tissues in Konservat-Lagerstätten provides rare yet significant insight into past behaviours and ecologies. Such deposits are known to occur in bursts rather than evenly through time, but reasons for this pattern and implications for the origins of novel structures remain unclear. Previous assessments of these records focused on marine environments preserving chemically heterogeneous tissues from across animals. Here, we investigate the preservation of skin and keratinous integumentary structures in land-dwelling vertebrates (tetrapods) through time, and in distinct terrestrial and marine depositional environments. We also evaluate previously proposed biotic and abiotic controls on the distribution of 143 tetrapod Konservat-Lagerstätten from the Permian to the Pleistocene in a multivariate framework. Gap analyses taking into account sampling intensity and distribution indicate that feathers probably evolved close to their first appearance in the fossil record. By contrast, hair and archosaur filaments are weakly sampled (five times less common than feathers), and their origins may significantly pre-date earliest known occurrences in the fossil record. This work suggests that among-integument variation in preservation can bias the reconstructed first origins of integumentary novelties and has implications for predicting where, and in what depositional environments, to expect further discoveries of exquisitely preserved tetrapod integument.
FIGURE 1 in A exceptionally-preserved new species of Barchaboilus (Orthoptera: Prophalangopsidae) from the Middle Jurassic of Daohugou, China
FIGURE 1. CNU-ORT-NN2011001P, A. photograph of habitus (dry, negative imprint), scale bar represents 10 mm. B. photograph of mandibles (under ethanol, negative imprint), scale bar represent 1 mm.
FIGURE 2 in A exceptionally-preserved new species of Barchaboilus (Orthoptera: Prophalangopsidae) from the Middle Jurassic of Daohugou, China
FIGURE 2. CNU-ORT-NN2011001PC, A. Photograph of left forewing (dry, negative imprint), B. line drawing of left forewing, C. photograph of right forewing (dry, negative imprint, flipped horizontally), D. line drawing of right forewing, E. photograph of hind wings (dry, negative imprint) and F. line drawing of hind wings. Scale bar represent 10 mm.
FIGURE 3 in Exceptionally preserved Leptolepidae (Actinopterygii, Teleostei) from the late Early Jurassic Fossil-Lagerstätten of Grimmen and Dobbertin (Mecklenburg-Western Pomerania, Germany)
FIGURE 3. Proleptolepis-like (MV 202612) from Grimmen. Line drawing of cranial bones in medial view. Dashed lines indicate moulds. Abbreviations: iop, interopercle; io3, infraorbital 3; io.c, infraorbital sensory canal; op, opercle; pop, preopercle; pop.c, preopercular sensory canal; sop, subopercle; suo, suborbital.
FIGURE 11 in Exceptionally preserved Leptolepidae (Actinopterygii, Teleostei) from the late Early Jurassic Fossil-Lagerstätten of Grimmen and Dobbertin (Mecklenburg-Western Pomerania, Germany)
FIGURE 11. Leptolepis jaegeri (GG 431/10) from Grimmen. Line drawing of caudal skeleton. Abbreviations: d.sc, dorsal caudal scute; ebf, epaxial basal fulcra; E1-3, epurals 1 to 3; ff, fringing fulcrum; H1, hypural 1; H3, hypural 3; H9, hypural 9; nsPU3, neural spine of preural vertebra 3; nsU1+2, neural spines of ural vertebra 1+2; PU1, preural vertebra 1; UN1, uroneural 1; UN7, uroneural 7; U1+2, ural vertebra 1+2; U3, ural vertebra 3; v.sc, ventral caudal scute; 1stPR, first principal caudal ray; 19thPR, 19th principal caudal ray.
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