<?xml version="1.0" encoding="UTF-8"?><article article-type="normal" xml:lang="en">
   <front>
      <journal-meta>
         <journal-id journal-id-type="publisher-id">PALEVO</journal-id>
         <issn>1631-0683</issn>
         <publisher>
            <publisher-name>Elsevier</publisher-name>
         </publisher>
      </journal-meta>
      <article-meta>
         <article-id pub-id-type="pii">S1631-0683(02)00017-9</article-id>
         <article-id pub-id-type="doi">10.1016/S1631-0683(02)00017-9</article-id>
         <title-group>
            <article-title>The first scorpion fossil from the Cretaceous amber of Myanmar (Burma). New implications for the phylogeny of Buthoidea</article-title>
            <trans-title-group xml:lang="fr">
               <trans-title>Le premier scorpion fossile dans l'ambre Crétacé de Myanmar (Birmanie). Nouvelles implications dans la phylogénie des Buthoidea</trans-title>
            </trans-title-group>
         </title-group>
         <contrib-group content-type="authors">
            <contrib contrib-type="author">
               <name>
                  <surname>Lourenço</surname>
                  <given-names>Wilson R</given-names>
               </name>
               <email>arachne@mnhn.fr</email>
            </contrib>
            <aff-alternatives>
               <aff> Laboratoire de zoologie (Arthropodes), Muséum national d'histoire naturelle, 61, rue Buffon, 75005 Paris, France</aff>
            </aff-alternatives>
         </contrib-group>
         <pub-date-not-available/>
         <volume>1</volume>
         <issue seq="3">2</issue>
         <issue-id pub-id-type="pii">S1631-0683(00)X0002-4</issue-id>
         <fpage seq="0" content-type="normal">97</fpage>
         <lpage content-type="normal">101</lpage>
         <history>
            <date date-type="received" iso-8601-date="2002-01-28"/>
            <date date-type="accepted" iso-8601-date="2002-03-04"/>
         </history>
         <permissions>
            <copyright-statement>© 2002 Académie des sciences/Éditions scientifiques et médicales Elsevier SAS</copyright-statement>
            <copyright-year>2002</copyright-year>
            <copyright-holder>Académie des sciences/Éditions scientifiques et médicales Elsevier SAS</copyright-holder>
         </permissions>
         <self-uri xmlns:xlink="http://www.w3.org/1999/xlink" content-type="application/pdf" xlink:href="main.pdf">
                        Full (PDF)
                    </self-uri>
         <abstract abstract-type="author">
            <p>A specimen belonging to a new genus and species of fossil scorpion, <italic>Palaeoburmesebuthus grimaldii</italic> gen. n., sp. n., is described from the Upper Cretaceous amber of Myanmar (Burma). This is the first scorpion to have been found and described from Burmese amber (± 90 Myr). The new genus and species are unquestionable buthoid elements but they are assigned to an <italic>incertae familiae</italic> until further material may be available for study. </p>
         </abstract>
         <trans-abstract abstract-type="author" xml:lang="fr">
            <p>Un spécimen appartenant à un nouveau genre et une nouvelle espèce de scorpion fossile, <italic>Palaeoburmesebuthus grimaldii</italic> gen. n., sp. n., est décrit dans l'ambre du Crétacé supérieur de Myanmar (Birmanie) (± 90 Ma). Les nouveaux genre et espèce sont sans aucun doute des éléments appartenant aux Buthoidea, mais sont pour l'instant accommodés dans une <italic>incertae familiae</italic>, jusqu'à ce que d'autres spécimens soient disponibles pour étude. </p>
         </trans-abstract>
         <kwd-group>
            <unstructured-kwd-group>scorpion, fossil, Upper Cretaceous, amber, Myanmar (Burma)</unstructured-kwd-group>
         </kwd-group>
         <kwd-group xml:lang="fr">
            <unstructured-kwd-group>scorpion, fossile, Crétacé supérieur, ambre, Myanmar (Birmanie)</unstructured-kwd-group>
         </kwd-group>
         <custom-meta-group>
            <custom-meta>
               <meta-name>miscellaneous</meta-name>
               <meta-value>Communicated by Paul Ozenda</meta-value>
            </custom-meta>
         </custom-meta-group>
      </article-meta>
   </front>
   <body>
      <sec>
         <title>Version abrégée</title>
         <sec>
            <label>1</label>
            <title>Introduction</title>
            <p>Parmi les arthropodes fossiles trouvés dans l'ambre, les scorpions se situent parmi les groupes les plus rares ; cependant, depuis environ deux décennies, plusieurs spécimens ont été décrits de l'ambre de la République dominicaine, du Mexique et de la Baltique. Les fossiles trouvés dans l'ambre de la République dominicaine et du Mexique sont étroitement associés aux scorpions actuels des régions antillaise et néotropicale. La position phylogénétique des scorpions dans l'ambre de la Baltique est cependant différente. Ces éléments peuvent être classés dans certaines lignées primitives actuelles de micro-buthidés. Cependant, les cinq spécimens décrits depuis 1996 correspondent tous à des genres nouveaux, distincts des genres actuels. Tout récemment, une découverte encore plus étonnante à été celle d'un fossile de l'ambre du Liban, le plus vieux connu (± 125–130 Ma) qui contienne des fragments biologiques. Une nouvelle famille, Archaeobuthidae Lourenço, a été créée pour ce spécimen également rattachée aux Buthoidea.</p>
            <p>Le spécimen étudié à présent est le premier de l'ambre de Birmanie à être décrit ; il correspond au deuxième plus ancien exemplaire de scorpion trouvé dans l'ambre, quel qu'en soit le type (± 90 Myr), étant uniquement supplanté par <italic>Archaeobuthus estephani</italic> Lourenço 2001, décrit de l'ambre du Liban et daté d'environ 125–130 Ma.</p>
            <p>Malgré l'état incomplet du spécimen, une étude détaillée autorise la description d'un nouveau genre et d'une nouvelle espèce. Les caractères diagnostiques définissant le nouveau genre <italic>Palaeoburmesebuthus</italic> sont les suivants : scorpions de petite taille (la taille totale des adultes peut être estimée à environ 10 mm) ; morphologie des parties observées, similaire à celle des Buthoidea ; anneaux du metasoma longs et aplatis, en particulier le cinquième ; carènes simples, mais bien marquées ; arc anal très simple, avec un seul lobe latéral ; telson très allongé, avec la vésicule fortement aplatie, à peine plus haute que le diamètre de la base de l'aiguillon, ce dernier étant pratiquement aussi long que la vésicule, très trapu à la base et avec sa partie distale très aiguë, les caractéristiques du telson étant diagnostiques du nouveau genre (<xref rid="FIG001" ref-type="fig">Figure 1</xref>, <xref rid="FIG002" ref-type="fig">Figure 2</xref>, <xref rid="FIG003" ref-type="fig">Figure 3</xref> and <xref rid="FIG004" ref-type="fig">Figure 4</xref>).</p>
            <p>Étant donné l'état très incomplet du spécimen étudié, il semble difficile d'assigner le nouveau genre à une famille précise au sein des Buthoidea. Pour cette raison, il est placé provisoirement dans une <italic>incertae familiae</italic>. Seule l'étude de nouveaux exemplaires pourra éventuellement justifier la création d'une nouvelle famille pour ce micro Buthoidea du Crétacé birman.</p>
         </sec>
         <sec>
            <label>2</label>
            <title>Palaeogéographie et âge de l'ambre birman</title>
            <sec>
               <p>Selon Zherikhin et Ross <xref rid="BIB007" ref-type="bibr">[7]</xref>, il existe une importante confusion dans la littérature en ce qui concerne l'âge probable de l'ambre birman. De même, une certaine imprécision existe en ce qui concerne les sites en Birmanie, d'où sont originaires les pièces d'ambre. D'après ces auteurs, il existe en Birmanie cinq régions où des résines fossiles ont été trouvées ; cependant, l'ambre birman (ou Burmite) existe exclusivement dans la vallée de Hukawng. Une synthèse des données géologiques sur le bassin de Hukawng peut être trouvée dans <xref rid="BIB007" ref-type="bibr">[7]</xref>.</p>
            </sec>
            <sec>
               <p>Selon <xref rid="BIB007" ref-type="bibr">[7]</xref>, la présence du foraminifère <italic>Nummulites biarritzensis</italic> suggère une datation à l'Éocène moyen des sites contenant de l'ambre. Cependant, les insectes inclus dans l'ambre suggèrent une datation plus ancienne. Cette discordance laisse supposer que l'ambre a été retravaillé et qu'il n'est pas dans son dépôt primitif. Des données de la littérature peuvent appuyer cette hypothèse : ainsi, il semble que deux périodes distinctes de dépôts soient associées aux sites contenant de l'ambre dans la vallée de Hukawng, la plus ancienne datant du Crétacé supérieur (Cenomanian), comme cela peut être attesté par les dépôts du foraminifère <italic>Orbitulina birmanica</italic>. Le lecteur intéressé par la géologie des dépôts de l'ambre birman trouvera de plus amples informations dans la référence <xref rid="BIB007" ref-type="bibr">[7]</xref>.</p>
            </sec>
         </sec>
      </sec>
      <sec>
         <label>1</label>
         <title>Introduction</title>
         <sec>
            <p>Scorpions are rare among the Arthropods fossilised in amber. Nevertheless, several specimens have recently been described from Dominican, Mexican and Baltic amber <xref rid="BIB003" ref-type="bibr">[3]</xref>, <xref rid="BIB004" ref-type="bibr">[4]</xref> and <xref rid="BIB005" ref-type="bibr">[5]</xref>. Even though some of these descriptions may require confirmation, there is little doubt that the amber fossils found in Hispaniola and Mexico are closely related to the extant Buthidae of the Caribbean and Neotropical regions. The position of the Baltic amber scorpions is different, because the known specimens, which also belong to the family Buthidae, can be classified among some extant but primitive lineages of micro-buthids. Five of these specimens have been described in recent years and represent distinct new genera that can be associated only approximately with extant groups. Two other specimens, which were described during the last century, retain an enigmatic status. They can, however, almost certainly be regarded as belonging to the category of primitive micro-buthids <xref rid="BIB003" ref-type="bibr">[3]</xref>, <xref rid="BIB004" ref-type="bibr">[4]</xref> and <xref rid="BIB005" ref-type="bibr">[5]</xref>. The history of amber scorpions, particularly those from the Baltic region, is complex. It has been described in detail by Lourenço and Weitschat <xref rid="BIB003" ref-type="bibr">[3]</xref>, <xref rid="BIB004" ref-type="bibr">[4]</xref> and <xref rid="BIB005" ref-type="bibr">[5]</xref>.</p>
         </sec>
         <sec>
            <p>More recently, a remarkable scorpion fossil was described from Lebanese amber <xref rid="BIB002" ref-type="bibr">[2]</xref>. Lebanese amber is the oldest one known to contain biological fragments. A new family, Archaeobuthidae Lourenço, was created for this specimen, which can be classified among the Buthoidea.</p>
         </sec>
         <sec>
            <p>Lebanese amber scorpions are much less common than those found in Baltic, Mexican or Caribbean amber. The type specimen of <italic>Archaeobuthus estephani</italic> Lourenço 2001 (family Archaeobuthidae) is not merely the first Lebanese amber scorpion to have been discovered, but is undoubtedly the oldest scorpion ever found in amber of any kind (± 125–130 Myr). The specimen described below is the first ever to have been found in Burmese amber, and is the second oldest amber fossil so far discovered (± 90 Myr). It is supplanted in age only by <italic>Archaeobuthus estephani</italic> Lourenço 2001.</p>
         </sec>
      </sec>
      <sec>
         <label>2</label>
         <title>Material and method</title>
         <sec>
            <p>The specimen investigated is trapped in a very clear block of yellowish amber that measures 7×6 mm and has been embedded in epoxy under vacuum <xref rid="BIB006" ref-type="bibr">[6]</xref>. The scorpion is rather incomplete, since the piece of amber had been broken. Only the metasoma with segments III, IV, V and telson are present. Both sides of the metasoma are clearly visible and allow detailed investigation.</p>
         </sec>
      </sec>
      <sec>
         <label>3</label>
         <title>Palaeogeography and the age of Burmite</title>
         <sec>
            <p>According to Zherikhin and Ross <xref rid="BIB007" ref-type="bibr">[7]</xref>, there is considerable confusion in the literature as to the probable age of Burmese amber. Also some confusion may exist regarding the precise sites in Myanmar from where the amber pieces originated. According to these authors there are five regions in Myanmar where fossil resins have been found, however Burmite only occurs in the Hukawng Valley. A synthesis of the geological data of the Hukawng Basin can be found in Zherikhin and Ross <xref rid="BIB007" ref-type="bibr">[7]</xref>.</p>
         </sec>
         <sec>
            <p>According to these authors, the presence of the foraminiferan <italic>Nummulites biarritzensis</italic> suggests a Middle Eocene (Upper Lutetian–Bartonian) age for the amber-bearing deposits, but the insects included in the Burmite suggest an older age for the amber. This could imply that the amber has been reworked, and some evidence can be found in the literature to support this. Therefore, from the evidence currently available, there appears to be two periods of deposition related to the amber bearing deposits in the Hukawng Valley. The first one took place during the Upper Cretaceous (Cenomanian), which is attested by the deposition of the Foraminiferan <italic>Orbitulina birmanica</italic>. For precise information on the geology of Burmese amber deposits, refer to Zherikhin and Ross <xref rid="BIB007" ref-type="bibr">[7]</xref>.</p>
         </sec>
      </sec>
      <sec>
         <label>4</label>
         <title>Systematic description</title>
         <sec>
            <p>
               <bold>Superfamily.</bold> Buthoidea C.L. Koch, 1837.</p>
         </sec>
         <sec>
            <p>
               <bold>Family.</bold>
               <italic>Incertae familiae</italic>.</p>
         </sec>
         <sec>
            <label>4.1</label>
            <title>Genus <italic>Palaeoburmesebuthus</italic> gen. n.</title>
            <sec>
               <p>
                  <bold>Diagnosis.</bold> Total length about 9 to 10 mm, based on the length of the metasomal segments III to V, which is about 3.9 mm long. General morphology of the metasoma somewhat similar to that of a few genera of micro-buthoids. The new genus can be distinguished from other known genera placed in the Buthoidea by the unusual shape of its telson, which is extremely long and flattened. The vesicle is long and flattened, while the aculeus is almost as long as the vesicle and the diameter of its base is almost as great as the vesicle depth; the distal half of the aculeus becomes extremely thin and has the shape of a nail; two glandular conduits can be observed in the proximal half of the aculeus (<xref rid="FIG001" ref-type="fig">Figure 1</xref>, <xref rid="FIG002" ref-type="fig">Figure 2</xref>, <xref rid="FIG003" ref-type="fig">Figure 3</xref> and <xref rid="FIG004" ref-type="fig">Figure 4</xref>).</p>
            </sec>
            <sec>
               <p>
                  <bold>Type species.</bold>
                  <italic>Palaeoburmesebuthus grimaldii</italic> gen. n. and sp. n.</p>
            </sec>
            <sec>
               <p>
                  <bold>Holotype.</bold> A juvenile (sex unknown). It can be included among the microbuthoid scorpions.</p>
            </sec>
            <sec>
               <p>
                  <bold>Type locality and horizon.</bold> Upper Cretaceous, Myanmar (Burma), Kachin: Tanai Village (on Ledo Rd. 105 km northwest of Myitkyna), coll. Leeward Capitol Corp., 2000. Collection AMNH Bu-710.</p>
            </sec>
         </sec>
         <sec>
            <label>4.2</label>
            <title>
               <italic>Palaeoburmesebuthus grimaldii</italic> gen. n. and sp. n.</title>
            <sec>
               <p>
                  <bold>Diagnosis.</bold> As for new genus.</p>
            </sec>
            <sec>
               <p>
                  <bold>Derivatio nominis.</bold> Patronym in honor of Dr David Grimaldi (AMNH), who authorised the study of the type specimen.</p>
            </sec>
            <sec>
               <p>
                  <bold>Coloration.</bold> The general colour is reddish yellow, with some portions of the vesicle and metasomal segments dark reddish. The amber is very clear and no parts of the specimen are covered with the usual milky white substance, which often prevents more precise observation of coloration.</p>
            </sec>
            <sec>
               <p>
                  <bold>Morphology.</bold> Carapace, keels and furrows unknown. Median ocular tubercle and lateral eyes unknown. Sternum unknown. Mesosoma: tergites I to VII unknown. Venter: genital operculum, pectines and sternum unknown. Sternites and spiracles unknown. Metasoma: segments I unknown. Segment II unknown. Segments III and IV with eight keels. Segment V very flattened with five keels. All keels crenulate. Lateral inframedian keels on segment III vestigial, represented by 2–3 granules; absent from segment IV. Ventrolateral and submedian keels crenulate on segments III–IV; dorsal keels with one distal slightly spinoid granule. Segment V with dorsolateral keels crenulated; ventrolateral keels with small spinoid granules, which become stronger near to the anal arch. Intercarinal spaces weakly granular, shagreened. Telson weakly granular laterally and dorsally, with some strong granules ventrally, and with three well developed ventral keels. Aculeus strongly curved and exceptionally long in comparison with the length of the vesicle, and very wide at its base; vesicle with several very long setae. Subaculear tooth absent. Cheliceral dentition unknown. Morphology of pedipalps unknown. Trichobotriotaxy unknown. Morphology of legs unknown.</p>
            </sec>
            <sec>
               <p>
                  <bold>Measurements (in mm) of the holotype of <italic>Palaeoburmesebuthus grimaldii</italic> gen. n. and sp. n.</bold> Total estimated length: 9.000/10.000. Metasoma. Segment III: length 1.000, width 0.452, depth 0.435; segment IV: length 1.161, width 0.420, depth 0.420; segment V: length 1.774, width 0.387, depth 0.322. Telson length 1.450; vesicle: length 0.740, width 0.290, depth 0.258; aculeus: length 0.710, diameter at the base 0.162.</p>
            </sec>
         </sec>
      </sec>
      <sec>
         <label>5</label>
         <title>Discussion</title>
         <sec>
            <p>According to some of the characters proposed in the description – shape of the metasomal segments and telson, structure of carinae, and especially the presence of a simple anal arch with one lateral lobe – the specimen is unquestionably a member of the Buthoidea. The assignment to one of the three families presently accepted within the Buthoidea: Archaeobuthidae Microcharmidae and Buthidae is, however, complicated on account of the incompleteness of the specimen. Therefore, the new genus is assigned to an <italic>incertae familiae</italic> until further material becomes available.</p>
         </sec>
         <sec>
            <p>It is important to notice that no extant buthoid genera have a telson morphology like that of the single <italic>Palaeoburmesebuthus</italic>. The reduced size of the vesicle in contrast with the very strong aculeus would suggest some kind of mechanical use in the technique of predation. This suggestion has already been discussed in the case of <italic>Archaeobuthus</italic> from Lebanese amber <xref rid="BIB002" ref-type="bibr">[2]</xref>.</p>
         </sec>
         <sec>
            <p>The new genus shares, with the Microcharmidae and some primitive buthids, a low evolutionary level. This places it in the first two gradients of the four/five that were defined by Lourenço <xref rid="BIB001" ref-type="bibr">[1]</xref> as being characteristic of genera of the Buthoidea. This situation is similar to that observed in Baltic amber scorpions <xref rid="BIB003" ref-type="bibr">[3]</xref>, <xref rid="BIB004" ref-type="bibr">[4]</xref> and <xref rid="BIB005" ref-type="bibr">[5]</xref> and, in particular, to that observed for <italic>Archaeobuthus</italic>
               <xref rid="BIB002" ref-type="bibr">[2]</xref>, suggesting that several more recent groups may have evolved since the Late Mesozoic and Early Cenozoic.</p>
         </sec>
      </sec>
   </body>
   <back>
      <ack>
         <title>Acknowledgements</title>
         <p>Thanks are due to D. Grimaldi (AMNH), for facilitating the study of the specimen, to D. Geffard for technical assistance, and to John L. Cloudsley-Thompson (London) for reviewing the manuscript.</p>
      </ack>
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   </back>
   <floats-group>
      <fig id="FIG001">
         <label>Figure 1</label>
         <caption>
            <p>
               <italic>Palaeoburmesebuthus grimaldii</italic> gen. n., sp. n. Metasomal segments IV–V and telson, lateral aspect.</p>
            <p>
               <italic>Palaeoburmesebuthus grimaldii</italic> gen. n., sp. n. Anneaux du metasoma IV–V et telson, vue latérale.</p>
         </caption>
         <graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="main.assets/gr001.jpg"/>
      </fig>
      <fig id="FIG002">
         <label>Figure 2</label>
         <caption>
            <p>
               <italic>Palaeoburmesebuthus grimaldii</italic> gen. n., sp. n. Telson, lateral aspect.</p>
            <p>
               <italic>Palaeoburmesebuthus grimaldii</italic> gen. n., sp. n. Telson, vue latérale.</p>
         </caption>
         <graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="main.assets/gr002.jpg"/>
      </fig>
      <fig id="FIG003">
         <label>Figure 3</label>
         <caption>
            <p>
               <italic>Palaeoburmesebuthus grimaldii</italic> gen. n., sp. n. Aculeus, lateral aspect, showing glandular conduits.</p>
            <p>
               <italic>Palaeoburmesebuthus grimaldii</italic> gen. n., sp. n. Aiguillon, vue latérale, avec, en détail, les conduits glandulaires.</p>
         </caption>
         <graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="main.assets/gr003.jpg"/>
      </fig>
      <fig id="FIG004">
         <label>Figure 4</label>
         <caption>
            <p>
               <italic>Palaeoburmesebuthus grimaldii</italic> gen. n., sp. n. Schematic drawing of metasomal segments III–V and telson, lateral aspect.</p>
            <p>
               <italic>Palaeoburmesebuthus grimaldii</italic> gen. n., sp. n. Dessin schématique des anneaux III–V du metasoma et telson, vue latérale.</p>
         </caption>
         <graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="main.assets/gr004.tif"/>
      </fig>
   </floats-group>
</article>