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   |-Hupehsuchus nanchengensis
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Ichthyosauria


, , (c) Stanton F. Fink


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The Ichthyosaurs ("fish lizards") were a successful group of air-breathing that bore an uncanny resemblance to modern day (late Cenozoic) dolphins. Along with the , the ichthyosaurs were among the earliest marine reptiles to be discovered, and the two groups had a strong effect on the 19th century Victorian imagination.

Ichthyosaurs were very diverse early in their history (during the period), where they included some of the largest marine reptiles that ever lived (up to 15 to 23 meters - the size of whales). During the late Triassic they developed their familiar streamlined fish-like form. The late Triassic and early Jurassic were actually the high points of the Ichthyosaur reign. The Toarcian turnover affected them heavily and only a single family made it through to the middle Jurassic. Although the Ichthyosaurs straggled on to the mid Cretaceous (Cenomanian age) they never regained their previous dominance, there place as top marine predator being taken by large pliosaurs.

Despite their dolphin-like form ichthyosaurs lacked the stamina of modern marine mammals. They were sprinters, not endurance swimmers, with a typically reptilian metabolism. The reason for their decline was probably tied up a lot with their inability to keep up with the new fast swimming Teleost fish that appeared during the Jurassic.

Being so fish-like in form, ichthyosaurs could not crawl up onto the land to lay their eggs. Instead, the eggs remained in the mother until they hatched. She then gave birth to live young. Skeletons of baby ichthyosaurs have been found inside the skeletons of adults. MAK991007.


(The following menu and phylogeny refers to subtopics of this page)


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Taxonomy Phylogeny

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Ichthyosaur mounted skeleton (Charmouth Heritage Coast Centre).The genus had first been described in 1699 from fossil fragments discovered in Wales.
The first fossil vertebrae were published twice in 1708 as tangible mementos of the Universal Deluge. The first complete ichthyosaur fossil was found in 1811 by in , along what is now called the Jurassic Coast. She subsequently discovered three separate species.

In 1905, the led by of the University of California and financed by Annie Alexander, found 25 specimens in central Nevada, which during the Triassic was under a shallow ocean. Several of the specimens are now in the collection of the University of California Museum of Paleontology. Other specimens are embedded in the rock and visible at Berlin-Ichthyosaur State Park in Nye County. In 1977 the Triassic ichthyosaur became the State Fossil of Nevada. Nevada is the only state to possess a complete skeleton, 55 ft (17 m) of this extinct marine reptile. In 1992, Canadian ichthyologist (Curator of Marine Reptiles at the Royal Tyrrell {"tur ell"} Museum) uncovered the largest fossil specimen ever, a 23 meter long example.

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The , looking more like finned lizards than the familiar fish or dolphin forms, are known from the Early and Early-Middle ( and ) strata of , , , and in . These primitive forms included the genera Chaohusaurus, Grippia, and Utatsusaurus. These very early proto-ichthyosaurs, which are now classified as rather than as ichthyosaurs proper (Motani 1997, Motani et al. 1998), quickly gave rise to true ichthyosaurs sometime in the latest Early Triassic or earliest Middle Triassic. These latter diversified into a variety of forms, including the sea-serpent like , which reached 10 meters, and smaller more typical forms like .

Ichthyosauria, strictly speaking, are + . Cymbospondylus is traditionally included in the . However, in the phylogenetic arrangement of (adopted here) this genus is shown to be much more primitive, even more primitive than the . Probably Cymbospondylus belongs in its own family ("Cymbospondylidae").

Cymbospondylus had a very elongate body, and the head, although large taken in isolation, was unusually small relative to the overall body. As befits its primitive status, Cymbospondylus lacked the familiar on its back and had only a very slight tail kink. It did however have the typical ichthyosaur pointed teeth in the long, beak-like jaws. MAK990620.

Mixosaurus includes several species of small creatures about a meter in length. They are among the earliest and most primitive of the Ichthyosaurs and were the sister group of the more derived merriamosaurs. Mixosaurus remained more reptilian than fishlike in appearance, though with flippers and a small fin on the tail. It also had a small dorsal fin on the back. Mixosaurus was morphologically intermediate between Cymbospondylus and more advanced Triassic Ichthyosaurs, such as the shastasaurs. The two lineages, the mixosaurs and Cymbospondylus, actually coexisted for some ten million or so years, along with other, even more basal early ichthyosaurs such as Hupehsuchus and Parvenator. MAK990620.

The make up the clade uniting the shastasaurs with the more fish-like, Jurassic ichthyosaurs.

In the shastasaurs we see an example of convergent or parallel evolution with the toothed whales of the Cenozoic oceans; although in the rapid evolutionary tendency to huge size there is a similarity with the baleen whales, which likewise grew from medium-sized to enormous creatures in the space of some five or ten-odd million years (Miocene-Pliocene).

By the , ichthyosaurs consisted of both classic Shastasauria and more advanced, "dolphin"-like Euichthyosauria (Californosaurus, Toretocnemus) and Parvipelvia (Hudsonelpidia, Macgowania). Experts disagree over whether these represent an evolutionary continum, with the less specialised shastosaurs a grade that was evolving into the more advanced forms (Maisch and Matzke 2000), or whether the two were separate that evolved from a common ancestor earlier on (Nicholls and Manabe 2001).

Reaching lengths of 10 to 15 meters, and with bodies that were quite deep and sturdy, the shastasaurs were not only the largest ichthyosaurs, but also among the largest of the marine animals of the Mesozoic. These giant ichthyosaurs, which were far larger than any of the contemporary Triassic land-living archosauromorphs and early dinosaurs, would have lived a lifestyle much like that of the present day toothed whales, feeding primarily on fish and other marine reptiles. They might, however, have been slow swimmers, rowing themselves slowly along with their long paddle-like limbs.

During the and , shastosaurs reached huge sizes. , known from a number of specimens from the Carnian of Nevada, was 15 meters long. Norian shonisaurs are known from both sides of the Pacific. Himalayasaurus tibetensis and Tibetosaurus (probably a synonym) have been found in . These large (10 to 15 meters long) ichthyosaurs probably belong to the same genus as Shonisaurus (Motani et al, 1999; Lucas, 2001, pp.117-119). While the gigantic Shonisaurus sikanniensis, whose remains were found in the Pardonet formation of British Columbia by Elizabeth Nicholls, reached as much as 21 meters in length - the largest marine reptile known to date.

These giants (along with their smaller cousins) seemed to have disappeared at the end of the Norian. (latest Triassic) ichthyosaurs are known from England, and these are very similar to those of the . Like the dinosaurs, the ichthyosaurs and their contemporaries the plesiosaurs survived the end-Triassic extinction event, and immediately diversified to fill the vacant ecological niches of the earliest Jurassic.

Painting of the Ichthyosaurus by . It is not known if these animals really could leap like dolphins
The Early Jurassic, like the Late Triassic, was the heyday of the ichthyosaurs, which are represented by four families and a variety of species, ranging from one to ten meters in length. Genera include Eurhinosaurus, , Leptonectes, Stenopterygius, and the large predator , along with the persistently primitive Suevoleviathan, which was little changed from its Norian ancestors. All these animals were streamlined, dolphin-like forms, although the more primitive animals were perhaps more elongated than the advanced and compact and .

Ichthyosaurs were still common in the , but had now decreased in diversity. All belonged to the single clade Ophthalmosauria. Represented by the 4 meter long and related genera, they were very similar to Ichthyosaurus, and had attained a perfect "tear-drop" streamlined form. The eyes of Ophthalmosaurus were huge, and it is likely that these animals hunted in dim and deep water (Motani 2000).

Ichthyosaurs seemed to decrease in diversity even further with the . Only a single genus is known, Platypterygius, and although it had a worldwide distribution, there was little diversity species-wise. This last ichthyosaur genus fell victim to the mid-Cretaceous (Cenomanian-Turonian) extinction event (as did some of the giant pliosaurs), although ironically less hydrodynamically efficient animals like and long-necked flourished. It seems that the ichthyosaurs became the victim of their own overspecialisation and were unable to keep up with the fast swimming and highly evasive new fishes, which were becoming dominant at this time and against which the sit-and-wait ambush strategies of the mosasaurs proved superior (Lingham-Soliar 1999).

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:

  • Phylum:
  • Class: (traditional) / (Benton 2000, 2004)
    • Subclass:
      • Superorder:
        • Order: Ichthyosauria

Author: , 1835

Stratigraphic Range: Early Triassic to Late Cretaceous

Phylogeny: :: + Ichthyosauria. + ( + )

Note: Phylogenetic position still controversial. There has been a good deal of activity recently, both generally at the base of the diapsids and, specifically, in Ichthyosaur phylogeny. In particular the "Ichthyopterygia" name has been dusted off and retooled to cover , , and a few others.

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The following section of this page needs to be . At its simplest, wikifying can just mean adding links to other pages via the edit tab. For help in editing Palaeos org pages, see the page

: Spindle-shaped body, 0.7-15.0 m; lacrimal participates in nares in most forms, but perhaps not primitive for group [C97]; largest orbit of any animal (up to 26 cm); often substantial sclerotic rings [C97]; postorbital does not participate in upper temporal fenestra [C97]; reduced cheek; strong upper temporal fenestra, but lower fenestra lost except in most basal species; supratemporal probably present primitively, but lost in many derived forms [C97a]; quadratojugal present primitively [C97]; teeth in long groove without separate alveoli in later forms; vertebral centra very short & deeply amphicoelous; neural arches separated from centra and without transverse processes(?); caudal fin in J-K species; manus 1 absent, but 2 neomorphic digits in some advanced forms; polydactyly in some, polyphalangy common and very well-developed; length of phalanges reduced; forelimbs used for steering, not propulsion; pelvic girdle not attached to spine (except Shonisaurus of the Middle Triassic); limbs reduced to steering fins; viviparous; diet varies, but squids probably common.

References: Callaway (1997) [C97]; Callaway (1997a) [C97a] . ATW020515

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  • Order ICHTHYOSAURIA
  • Family
  • Suborder
    • (unranked)
      • Family
      • Infraorder ("true ichthyosaurs")
        • Family
        • (Unranked) ("small pelves")
          • Family
          • Infraorder ("tuna lizards")
            • Family
            • Family
            • Family

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Ichthyosauria [Ichthyopterygia]
   |  i. s.: Isfjordosaurus Motani 1999
   |           `--*I. minor (Wiman 1910) [=Pessopteryx minor]
   |         Svalbardosaurus Mazin 1981 (n. d.)
   |           `--S. crassidens
   |         Himalayasaurus Young & Dong 1972 (n. d.)
   |           `--*H. tibetensis Young & Dong 1972 (n. d.)
   |         Tibetosaurus Dong in Young et al. 1982 (n. d.)
   |           `--*T. tingjiensis Dong in Young et al. 1982 (n. d.)
   |         Pessosaurus Wiman 1910 (n. d.)
   |           |--*P. polaris (Hulke 1873) (n. d.)
   |           `--P. suevicus von Huene 1916
   |         Macropterygius von Huene 1922 (n. d.)
   |           `--*M. trigonus (Owen 1840)
   |         Pachygonosaurus von Huene 1916 (n. d.)
   |           `--*P. robustus Maisch & Matzke 1997 (n. d.)
   |--Thaisaurus Mazin, Suteethorn et al. 1991
   |    `--*T. chonglakmanii Mazin, Suteethorn et al. 1991
   `--+--Utatsusaurus Shikama, Kamei & Murata 1978
      |    `--*U. hataii Shikama, Kamei & Murata 1978
      `--+--Grippiidae [Grippidia]
         |    |--Chaohusaurus Young & Dong 1972 (see below for synonymy)
         |    |    `--*C. geishanensis Young & Dong 1972
         |    `--Grippia Wiman 1929
         |         `--*G. longirostris Wiman 1929
         `--+--Parvinatator Nicholls & Brinkman 1995
            |    `--*P. wapitiensis Nicholls & Brickman 1995
            `--+--
               `--Quasianosteosaurus Maisch & Matzke 2003 [Quasianosteosauridae]
                    `--*Q. vikinghoegdai Maisch & Matzke 2003

Chaohusaurus Young & Dong 1972 [incl. Anhuisaurus Chen 1985 (preoc.), Chensaurus Mazin, Suteethorn et al. 1991]

* Type species of generic name indicated


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Callaway, J. M. 1997a. Ichthyosauria: introduction. In Ancient Marine Reptiles (J. M. Callaway & E. L. Nicholls, eds.) pp. 3-16. Academic Press.

Callaway, J. M. 1997b. A new look at Mixosaurus. In Ancient Marine Reptiles (J. M. Callaway & E. L. Nicholls, eds.) pp. 45-59. Academic Press.

Ellis, R. 2003. Sea Dragons - Predators of the Prehistoric Oceans. University Press of Kansas

  • Stephen Jay Gould, "Bent out of Shape" in Eight Little Piggies.

Lingham-Soliar, T. 1999. A functional analysis of the skull of Goronyosaurus nigeriensis (Squamata: Mosasauridae) and its bearing on the predatory behavior and evolution of the enigmatic taxon. N. Jb. Geol. Palaeont. Abh. 2134 (3): 355-74

Maisch, M. W., & A. T. Matzke. 2000. The Ichthyosauria. Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) 298: 1-159.

Maisch, M. W., & A. T. Matzke. 2003. Observations on Triassic ichthyosaurs. Part XII. A new Early Triassic ichthyosaur genus from Spitzbergen. Neues Jahrbuch für Geologie und Paläontologie – Abhandlungen 229 (3): 317-338.

McGowan, C. 1992. Dinosaurs, Spitfires and Sea Dragons. Harvard University Press,

McGowan, C., & R. Motani. 2003. Ichthyopterygia. Handbook of Paleoherpetology, Part 8. Verlag Dr. Friedrich Pfeil.

Motani, R. 1997. Temporal and spatial distribution of tooth implantation in ichthyosaurs. In Ancient Marine Reptiles (J. M. Callaway & E. L. Nicholls, eds.) pp. 81-103. Academic Press.

Motani, R. 2000. Rulers of the Jurassic seas. vol.283, no. 6.

Motani, R., N. Minoura & T. Ando. 1998. Ichthyosaurian relationships illuminated by new primitive skeletons from Japan. Nature 393: 255-257.

Motani, R., M. Manabe, & Z.-M. Dong. 1999. The status of Himalayasaurus tibetensis (Ichthyopterygia). Paludicola 2(2):174-181.

Nicholls, E. L. & M. Manabe. 2001. A new genus of ichthyosaur from the Late Triassic Pardonet Formation of British Columbia: bridging the Triassic-Jurassic gap. Canadian Journal of Earth Sciences 38: 983-1002.

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  • - Introduction to the Ichthyosauria
  • (English & Japanese)
  • (after Ryosuke Motani's page, perhaps the most useful site)
  • (German)
  • (Norwegian)
  • (German)
  • (Opthalmosaurus)
  • (Temnodontosaurus)
  • ()

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What about the pill? Although we have shown that PDE inhibitors reduce NOS2 and ARG1, the full mechanisms underlying these effects remain to be elucidated. One putative mechanism involves the impact of these inhibitors on mRNA stability. cGMP destabilizes NOS2 mRNA by reducing the ubiquitous mRNA binding protein, human antigen R (45). As such, destabilization of NOS2 mRNA via PDE5 inhibition would abrogate NO-mediated immunosuppression more effectively than would competitive inhibition of NO itself. However, because ARG1 mRNA does not possess adenylate/ uridylate-rich elements and has not been described to be stabilized by human antigen R, other mechanisms are likely involved in PDE5-mediated down-regulation of ARG1. One possibility is that high levels of cGMP induced by PDE5 blockade reduce the cytosolic Ca2+ concentration (46), leading to a reduction of the calcium-dependent protein kinase C activity (47) that in turn prevents up-regulation of IL-4R (48). The link between IL-4R and ARG1 in MDSCs is supported by recent data (29) demonstrating a direct correlation between ARG1 expression and IL-4R expression. ACT of tumor-primed CD8 T cells completely eradicated a preestablished C26GM tumor in the LysMCreIL-4R–/flox mice in which IL-4R expression has been deleted in neutrophils and macrophages, whereas no effect was seen in the control littermates (29). Our findings support these data by demonstrating that PDE5 blockade down-regulates IL-4R expression on tumor-infiltrating MDSCs (Fig. 6) and synergizes with the adoptive transfer of tumor-primed CD8+ T cells (Fig. 3 and Fig. 4). This effect appears to specifically target MDSCs, because IL-4R expression on isolated CD11b+ cells from tumor-bearing mice is significantly reduced when co-cultured in the presence of sildenafil (Fig. 5). Collectively, these findings underscore the critical role of the IL-4R–ARG1 pathway in MDSCs, as well as the use of PDE5 inhibitors as therapeutically effective drugs to overcome tumor-induced immunosuppression. without viagra prescription By Amanda Gardner Erectile dysfunction (ED), also called impotence, affects most men at some point during their lifetime. However, chronic impotence may indicate a medical condition, such as heart disease, high blood pressure, high cholesterol, kidney disease, or diabetes. While many substances are touted as aphrodisiacs, the modern age of pharmacotherapy began in 1993 when the injection of papaverine, an alpha-receptor blocker that produces vasodilatation, was shown to produce erections when injected directly into the corpora cavernosa. Soon afterwards, other vasodilators, such as PGE1 and phentolamine (Regitine), were demonstrated to be effective either as single agents or in combination. Return to top At least 7 PDE classes are known, many with subtypes identified by structure and function. PDE-5 is cGMP-specific and is a major cGMP-hydrolyzing enzyme in the vascular smooth muscle of the penis. The newer agents, vardenafil and tadalafil, are more specific and potent cGMP inhibitors than sildenafil. Both of the newest agents are PDE-5 inhibitors, which are significantly more selective in their inhibition. For people who are at risk of developing ED due to personal behavior, such as drinking too much alcohol, steps may be taken to prevent its occurrence. However, other causes of ED may not be preventable. Reviewed by Louise Chang, MD To exert a measurable antitumor effect, tumor-specific T cells must be present in sufficient numbers and capable of trafficking to their targets. A direct correlation exists between the number of tumor-infiltrating lymphocytes (TILs) and a favorable clinical outcome, as demonstrated in patients with metastatic ovarian cancer (20). Furthermore, the functional status of TILs has been correlated with a favorable prognosis in various human malignancies (20–24). Because PDE5 inhibition augments antitumor immunity, we asked whether sildenafil treatment altered both the number and activation state of TILs. Histological examination of CT26WT tumors revealed a greater intratumoral cellular infiltrate in the sildenafil-treated mice compared with the untreated controls (Fig. 4 A). To better evaluate these differences, C26GM-bearing mice received either tumor-primed or no T cells followed by sildenafil treatment or no additional therapy. The tumors were excised 9 d later, and single cell suspensions were obtained. The T cell infiltrate was analyzed by flow cytometry for CD4+ and CD8+ T cells. This approach enabled us to accurately examine the entire tumor mass and reliably quantify the infiltrating lymphocytic population. Although no increase in CD4+ T cells was observed with PDE5 inhibition (Fig. 4 B, inset), sildenafil treatment greatly increased CD8+ intratumoral infiltration with up-regulation of the activation markers CD69 and CD25 (Fig. 4 C). There were no differences in activation markers between the sildenafil-treated group and sildenafil + ACT, whereas a significant increase in intratumoral T cells were observed in the sildenafil + ACT–treated group compared with sildenafil alone (Fig. 4 B and Fig. S4, available at http://www.jem.org/cgi/content/full/jem.20061104/DC1). How does erection occur? Penile prosthesis: Patients are instructed in the operation of the prosthesis prior to surgery and again in the postoperative period. The prosthesis is not usually activated until approximately 6 weeks after surgery. This is to allow the edema and pain to subside. The prosthesis is checked in the office before the patient begins to use it. They typically contain stearate, croscarmellose, calcium hydrogen phosphate (anhydrous), and microcrystalline cellulose. After all, this can be not the entire formulation as it is a closely guarded secret of each manufacturer. This work has been supported by grants from the MIUR-PRIN (project no. 2005069853), Cariverona, Bando 2004 "Integrazione tra tecnologia e sviluppo di settore - Bando per progetti di ricerca a indirizzo biomedico," and the Italian Association for Cancer Research. viagra no prescription What is vardenafil (Levitra)? When administered in vitro, PDE5 inhibition induces apoptosis in colon carcinoma (15) and chronic lymphocytic leukemia cells (16). To determine whether similar effects could be observed in vivo, we used various transplantable mouse tumors, including CT26WT (a colon carcinoma; Fig. 1 A), the more aggressive variant C26GM (Fig. 1 B), TS/A (a mammary adenocarcinoma; Fig. 1 C), and the MCA203 fibrosarcoma (Fig. 1 D). PDE5 inhibitors were administered starting on the day of tumor challenge. Sildenafil and tadalafil significantly delayed tumor outgrowth by 50 to 70% in immune-competent mice, although all mice ultimately died (Fig. S1, available at http://www.jem.org/cgi/content/full/jem.20061104/DC1). Similar results were obtained even if sildenafil treatment was started on day 7 after tumor challenge in the CT26WT model (Fig. S2). The fact that no difference in tumor outgrowth was seen between early versus late administration of sildenafil suggests that PDE5 inhibition does not appreciably affect the early phases of tumor uptake but rather influences the later stages of tumor outgrowth. Because the addition of sildenafil to cultured CT26WT cells did not increase their apoptosis or affect their doubling time (unpublished data), we conclude that sildenafil does not have a direct antitumor effect but rather interferes in host–tumor interactions.

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