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Mammalia, Cervidae) During the Middle Holocene in the Cave of Bizmoune (Morocco, Essaouira Region
Quaternary International xxx (2015) 1e14 Contents lists available at ScienceDirect Quaternary International journal homepage: www.elsevier.com/locate/quaint The last occurrence of Megaceroides algericus Lyddekker, 1890 (Mammalia, Cervidae) during the middle Holocene in the cave of Bizmoune (Morocco, Essaouira region) * Philippe Fernandez a, , Abdeljalil Bouzouggar b, c, Jacques Collina-Girard a, Mathieu Coulon d a Aix Marseille Universite, CNRS, MCC, LAMPEA UMR 7269, 13094, Aix-en-Provence, France b Institut National des Sciences de l'Archeologie et du Patrimoine, Rabat, Morocco c Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, D-04103 Leipzig, Germany d Aix Marseille Universite, CNRS, LAMES UMR 7305, 13094, Aix-en-Provence, France article info abstract Article history: During the course of archaeological test excavations carried out in 2007 in the cave of Bizmoune Available online xxx (Essaouira region, Morocco), seven archaeological layers yielding Pleistocene and Holocene artefacts and faunal remains were identified. In the layers C4, C3 and C2, respectively from the oldest to the most Keywords: recent, terrestrial Helicidae mollusk shells (Helix aspersa) were dated by 14C. These layers also contained Giant deer many fragments of eggshell, belonging to Struthio cf. camelus, associated with mammal remains such as Extinction Oryctolagus/Lepus, Gazella sp., Sus scrofa, Ammotragus lervia, Alcelaphus buselaphus, Equus sp., Pha- Holocene cochoerus aethiopicus and an undetermined Caprini. Among these remains, an incomplete mandible of North Africa Speciation Megaceroides algericus Lydekker, 1890 with M1 and M2 was found in layer C3. The 6641 to 6009 cal BP Palaeoecology time range attributed to this layer has provided the most recent date known so far for M. -
Fossil Deer Assemblage from Sea Bottom of Bisan
"FOSSIL DEER ASSEMBLAGE FROM SEA BOTTOM OF BISAN-SETO AREA WITH SPECIAL REFERENCE TO THEIR STRATIGRAPHIC POSITIONS (PLEISTOCENE DEER FAUNA IN SETO INLAND SEA, ----PART III)" 著者 "OTSUKA Hiroyuki" journal or 鹿児島大学理学部紀要. 地学・生物学 publication title volume 22 page range 55-87 別言語のタイトル 備讃瀬戸の海底に於ける鹿類化石群集-特にその産 出相に関連して (瀬戸内海の第四期鹿類化石群-そ の3) URL http://hdl.handle.net/10232/5980 FOSSIL DEER ASSEMBLAGE FROM SEA BOTTOM OF BISAN-SETO AREA WITH SPECIAL REFERENCE TO THEIR STRATIGRAPHIC POSITIONS (PLEISTOCENE DEER FAUNA IN SETO INLAND SEA, ----PART III) 著者 OTSUKA Hiroyuki journal or 鹿児島大学理学部紀要. 地学・生物学 publication title volume 22 page range 55-87 別言語のタイトル 備讃瀬戸の海底に於ける鹿類化石群集-特にその産 出相に関連して (瀬戸内海の第四期鹿類化石群-そ の3) URL http://hdl.handle.net/10232/00001718 Rep. Fac. Sci., Kagoshima Univ.,(Earth Sci. & Biol.) No. 22, p.55-87, 1989. FOSSIL DEER ASSEMBLAGE FROM SEA BOTTOM OF BISAN-SETO AREA WITH SPECIAL REFERENCE TO THEIR STRATIGRAPHIC POSITIONS PLEISTOCENE DEER FAUNA IN SETO INLAND SEA, 一一-PART I Hiroyuki Otsuka (Received August 30, 1989) Abstract This paper presents the results of paleontological and geological studies of fossil deer fauna collected from the sea bottom of the Bisan-seto area, in the eastern part of the Seto Inland Sea, West Japan. The fossil deer assemblages in this area are associated with abundant mammals of forest- to grassland habitats such as Naumann's elephant (Palaeoloxodon naumanni) and also with a few Stegodon, wild ox, rhino and water buffalo specimens. As a result of the present study, nine species fossil deer belonging to the family Cervidae were distinguished, among which six species are referred to the genus Cervus and others to the genus Elaphurus and SinamegaFeros. -
Dinner in a Dinosaur
Benchmarks DECEMBER 31, 1853: DINNER IN A DINOSAUR he weather in London on Saturday, Dec. 31, 1853, could On New Year’s Eve, 1853, Benjamin Waterhouse Hawkins hosted not have pleased Benjamin Waterhouse Hawkins. a formal dinner in the mold of an Iguanodon. After a relatively warm Friday, the temperature had plummeted, snow had begun to fall, and for the first to celebrate the “triumphs of industry and art,” and hired Ttime in more than a decade, masses of ice floated down the Hawkins to direct the “Fossil Department.” They tasked him Thames River. The snow made the streets so slippery that with populating a vast geologic display with giant monsters injured pedestrians filled the hospitals. of the ancient world, including the first three dinosaurs ever For New Year’s Eve, Hawkins was hosting an elaborate feast described: Megalosaurus, Iguanodon and Hylaeosaurus. at his sculpting studio in Sydenham, 11 kilometers south of Hawkins was uniquely qualified to bring these great ani- London. Would his guests be able to find transportation out to mals to life. He had initially achieved fame for his detailed Sydenham and then across the pastures of muddy swamp that illustrations of animals collected by British explorers, includ- surrounded the wooden building where the dinner would be ing the still relatively obscure naturalist Charles Darwin. held? Hawkins hoped so; he had been planning the meal for Subsequently, Hawkins started to sculpt, and to write and more than a month. It would be the first time that most of his illustrate books on animal anatomy. For his efforts in taking dinner-mates had seen the incredible life-sized dinosaurs that new scientific findings and translating them into words and he was building for the Crystal Palace Exhibition, which Queen images accessible to the general public, Hawkins earned Victoria and Prince Albert would open to the public in June. -
Artiodactyl Brain-Size Evolution
Artiodactyl brain-size evolution A phylogenetic comparative study of brain-size adaptation Bjørn Tore Kopperud Oppgave for graden Master i Biologi (Økologi og evolusjon) 60 studiepoeng Centre for ecological and evolutionary synthesis Institutt for biovitenskap Det matematisk-naturvitenskapelige fakultet UNIVERSITETET I OSLO Høsten 2017 Artiodactyl brain-size evolution A phylogenetic comparative study of brain-size adaptation Bjørn Tore Kopperud © 2017 Bjørn Tore Kopperud Artiodactyl brain-size evolution http://www.duo.uio.no/ Trykk: Reprosentralen Abstract The evolutionary scaling of brain size on body size among species is strikingly constant across vertebrates, suggesting that the brain size is constrained by body size. Body size is a strong predictor of brain size, but the size variation in brain size independent of body size remains to be explained. Several hypotheses have been proposed, including the social-brain hypothesis which states that a large brain is an adaptation to living in a group with numerous and complex social interactions. In this thesis I investigate the allometric scaling of brain size and neocortex size and test several adaptive hypotheses in Artiodactyla (even-toed ungulates), using a phylogenetic comparative method where the trait is modeled as an Ornstein-Uhlenbeck process. I fit models of brain size and neocortex size, absolute and relative, in response to diet, habitat, gregariousness, gestation length, breeding group size, sexual dimorphism, and metabolic rate. Most of the investigated variables have no effect on the relative size of brain and neocortex, but the optimal relative size of the brain and neocortex is 20% and 30% larger, respectively, in gregarious species than in solitary species. -
Megaloceros Giganteus) from the Pleistocene in Poland
Palaeontologia Electronica palaeo-electronica.org Healed antler fracture from a giant deer (Megaloceros giganteus) from the Pleistocene in Poland Kamilla Pawłowska, Krzysztof Stefaniak, and Dariusz Nowakowski ABSTRACT We evaluated the skull of an ancient giant deer with a deformity of one antler. The skull was found in the 1930s in the San River near Barycz, in southeastern Poland. Its dating (39,800±1000 yr BP) corresponds to MIS-3, when the giant deer was wide- spread in Europe. Our diagnostics for the antler included gross morphology, radiogra- phy, computed tomography, and histopathology. We noted signs of fracture healing of the affected antler, including disordered arrangement of lamellae, absence of osteons, and numerous Volkmann’s canals remaining after blood vessel loss. The antler defor- mity appears to be of traumatic origin, with a healing component. No similar evaluation process has been described previously for this species. Kamilla Pawłowska. corresponding author, Institute of Geology, Adam Mickiewicz University, Maków Polnych 16, Poznań 61-606, Poland, [email protected] Krzysztof Stefaniak. Division of Palaeozoology, Department of Evolutionary Biology and Ecology, Faculty of Biological Sciences, University of Wrocław, Sienkiewicza 21, Wrocław 50-335, Poland, [email protected] Dariusz Nowakowski. Department of Anthropology, Wroclaw University of Environmental and Life Sciences, Kożuchowska 6/7, Wrocław 51-631, Poland, [email protected] Keywords: giant deer; Megaloceros giganteus; paleopathology; Pleistocene; Poland INTRODUCTION -
Историческое Развитие Парнопалых (Artiodactyla) Северной Евразии И Этапы Эв
ÝÂÎËÞÖÈß ÁÈÎÑÔÅÐÛ È ÁÈÎÐÀÇÍÎÎÁÐÀÇÈß 4. ÏÀËÅÎÝÊÎËÎÃÈß È ÝÂÎËÞÖÈß ÑÎÎÁÙÅÑÒ ÓÄÊ 569.73: 415 (551.77) Èñòîðè÷åñêîå ðàçâèòèå ïàðíîïàëûõ (Àrtiodactyla) Ñåâåðíîé Eâðàçèè è ýòàïû ýâîëþöèè èõ ñîîáùåñòâ â êàéíîçîå È.À. Âèñëîáîêîâà Ïàëåîíòîëîãè÷åñêèé èíñòèòóò ÐÀÍ, Ìîñêâà E-mail: [email protected] Ïðîàíàëèçèðîâàíà ýâîëþöèÿ ñîîáùåñòâ ïàðíîïàëûõ â êàéíîçîå è âûäåëåíû îñíîâíûå ýòàïû â èíòåðâàëå îò 56 äî 0,4 ìëí. ëåò íàçàä. Ðîñò ðàçíîîáðàçèÿ ïàðíîïàëûõ òåñíî ñâÿçàí ñ ýâîëþöèåé è äèâåðñèôèêàöèåé ðàñòèòåëüíîãî ïîêðîâà è èçìåíåíèÿìè êëèìàòà è ëàíä- øàôòîâ. Îñíîâíûå ðóáåæè ïðåîáðàçîâàíèÿ ñîîáùåñòâ â ïàëåîãåíå ñèíõðîííû êðóïíåé- øèì áèîñôåðíûì ñîáûòèÿì, à ïëèîöåí-ïëåéñòîöåíîâûå èçìåíåíèÿ îòðàæàþò êðóïíûå êîëåáàíèÿ êëèìàòà â ñåâåðíîì ïîëóøàðèè. Êëþ÷åâûå ñëîâà: ïàðíîïàëûå, ýâîëþöèÿ, îñíîâíûå ýòàïû, áèîðàçíîîáðàçèå, êàéíîçîé, Ñåâåðíàÿ Åâðàçèÿ. Îòðÿä ïàðíîïàëûõ (Artiodactyla) îäíî èç ìàãèñòðàëüíûõ íàïðàâëåíèé ýâîëþöèè ïëàöåíòàðíûõ ìëåêîïèòàþùèõ (Eutheria). Âîçíèêíóâ â ìåçîçîå, îí äîìèíèðóåò ñðåäè êðóï- íûõ ìëåêîïèòàþùèõ ïëàíåòû íàðÿäó ñ ïðèìàòàìè è õèùíûìè. Ýòî äèíàìè÷íî ðàçâèâàâ- øàÿñÿ ãðóïïà ìëåêîïèòàþùèõ õîðîøî ïðåäñòàâëåíà â èñêîïàåìîì ñîñòîÿíèè è óíèêàëüíà äëÿ èçó÷åíèÿ çàêîíîìåðíîñòåé ìàêðîýâîëþöèè è èçìåíåíèÿ áèîðàçíîîáðàçèÿ. Èñòîðèÿ ïàðíîïàëûõ òåñíî ñâÿçàíà ñ ýâîëþöèåé áèîñôåðû è ýâîëþöèåé ðàñòèòåëüíîãî ïîêðîâà Çåìëè è ðàçâîðà÷èâàëàñü íà ôîíå èçìåíåíèé î÷åðòàíèé ìàòåðèêîâ, ñîñòàâà àòìîñôåðû è åå öèðêóëÿöèè, êëèìàòà è ëàíäøàôòîâ. Õîðîøèå èíäèêàòîðû ïîñëåäíèõ, ýòè â îñíîâíîì ðàñòèòåëüíîÿäíûå ìëåêîïèòàþùèå ñ óñïåõîì èñïîëüçóþòñÿ ïðè ðåêîíñòðóêöèè ïðèðîä- íîé ñðåäû ïðîøëûõ ýïîõ è â áèîñòðàòèãðàôèè. Áîëüøîå ðàçíîîáðàçèå è õîðîøàÿ èçó÷åí- íîñòü ïàðíîïàëûõ ïîçâîëÿþò èñïîëüçîâàòü èõ äëÿ ïîçíàíèÿ èñòîðèè íàçåìíûõ áèîöåíî- çîâ êàéíîçîÿ è îñîáåííîñòåé ôîðìèðîâàíèÿ ñîâðåìåííûõ áèîìîâ.  ñòàòüå àíàëèçèðóþòñÿ ñóêöåññèè êîìïëåêñîâ ïàðíîïàëûõ, ñóùåñòâîâàâøèõ â Ñå- âåðíîé Åâðàçèè (íà òåððèòîðèè Ðîññèè, Ìîëäàâèè, Óêðàèíû, Ãðóçèè, Êàçàõñòàíà, Êèð- ãèçèè, Òàäæèêèñòàíà, Ìîíãîëèè è Ñåâåðíîãî Êèòàÿ) îò ýîöåíà äî ñåðåäèíû ñðåäíåãî ïëåéñòîöåíà (560,4 ìëí. -
New Record of Terminal Pleistocene Elk/Wapiti (Cervus Canadensis) from Ohio, USA
2 TERMINAL PLEISTOCENE ELK FROM OHIO VOL. 121(2) New Record of Terminal Pleistocene Elk/Wapiti (Cervus canadensis) from Ohio, USA BRIAN G. REDMOND1, Department of Archaeology, Cleveland Museum of Natural History, Cleveland, OH, USA; DAVID L. DYER, Ohio History Connection, Columbus, OH, USA; and CHARLES STEPHENS, Sugar Creek Chapter, Archaeological Society of Ohio, Massillon, OH, USA. ABSTRACT. The earliest appearance of elk/wapiti (Cervus canadensis) in eastern North America is not thoroughly documented due to the small number of directly dated remains. Until recently, no absolute dates on elk bone older than 10,000 14C yr BP (11,621 to 11,306 calibrated years (cal yr) BP) were known from this region. The partial skeleton of the Tope Elk was discovered in 2017 during commercial excavation of peat deposits from a small bog in southeastern Medina County, Ohio, United States. Subsequent examination of the remains revealed the individual to be a robust male approximately 8.5 years old at death. The large size of this individual is compared with late Holocene specimens and suggests diminution of elk since the late Pleistocene. Two accelerator mass spectrometry (AMS) radiocarbon assays on bone collagen samples taken from the scapula and metacarpal of this individual returned ages of 10,270 ± 30 14C yr BP (Beta-477478) (12,154 to 11,835 cal yr BP) and 10,260 ± 30 14C yr BP (Beta-521748) (12,144 to 11,830 cal yr BP), respectively. These results place Cervus canadensis in the terminal Pleistocene of the eastern woodlands and near the establishment of the mixed deciduous forest biome over much of the region. -
How the 'Seven Deadly Agents Of
Paper presented at the How the ‘Seven Deadly Agents of Destruction’ conference can help preserve the Crystal Palace Dinosaurs Liesa Brierley, Ellinor Michel, Anthony Lewis, Chris Aldhous and Lois Olmstead Introduction Compared to collections kept safe inside museums, outdoor sculpture is affected by additional risk factors. Sculpture in the public realm is exposed to the elements, pollution and direct sunshine. It is more vulnerable to vandalism and the proximity to nature can also take its toll. At the same time, the condition of outdoor sculpture is often not monitored as rigorously as that of museum collections. However, damage doesn’t go unnoticed by the public and, channelled in the right way, the public’s critical eye can be turned to positive pressure, increasing support for conservation work and helping to improve conditions for the displays. To conservation professionals, the Canadian Conservation Institute’s framework of Ten Agents of Deterioration is a familiar and well-established way of describing risk factors to cultural heritage. This paper describes an attempt to translate this framework into a short animated film about risks to outdoor sculpture, using the Crystal Palace Dinosaurs, a renowned Victorian sculpture park in south London, as the stage for the message. The film is aimed at a broad audience of non-professionals of all ages and it is hoped that it will deliver an increased sense of both understanding and ownership of the sculpture park. The Crystal Palace Dinosaurs The Crystal Palace Dinosaurs are the first life-sized sculptural reconstructions of extinct animals, built to engage and enlighten the public on the paradigm- shifting advances in science in the mid-19th century (Owen 1854, McCarthy and Gilbert 1994, Pigott 2004, Bramwell and Peck 2008). -
Patterns of Late Quaternary Megafaunal Extinctions in Europe and Northern Asia
Cour. Forsch.-Inst. Senckenberg | 259 | 287 – 297 | 2 Figs | Frankfurt a. M., 13. 12. 2007 Patterns of Late Quaternary megafaunal extinctions in Europe and northern Asia With 2 figs Anthony J. STUART & Adrian M. LISTER A b s t r a c t This paper summarizes the results so far of our ‘Late Quaternary Megafaunal Extinctions’ project, focussing on an assessment of latest available dates for selected target species from Europe and northern Asia. Our approach is to directly radiocarbon-date material of extinct megafauna to construct their spatio-temporal histories, and to seek correlations with the environmental and archaeological records with the aim of estab- lishing the cause or causes of extinction. So far we have focussed on Mammuthus primigenius, Coelodonta antiquitatis, and Megaloceros giganteus, and are accumulating data on Panthera leo/spelaea, Crocuta crocuta and Ursus spelaeus. Attempts to date Palaeoloxodon antiquus and Stephanorhinus hemitoechus (from southern Europe) were largely unsuccessful. The pattern of inferred terminal dates is staggered, with extinctions occurring over ca. 30 millennia, with some species previously thought extinct in the Late Pleistocene – M. primigenius and M. giganteus – surviving well into the Holocene. All species show dramatic range shifts in response to climatic/vegeta- tional changes, especially the beginning of the Last Glacial Maximum, Late Glacial Interstadial, Allerød, Younger Dryas and Holocene, and there was a general trend of progressive range reduction and fragmenta- tion prior to final extinction. With the possible exceptions of P. antiquus, S. hemitoechus and Homo nean- derthalensis, extinctions do not correlate with the appearance of modern humans. However, although most of the observed patterns can be attributed to environmental changes, some features – especially failures to recolonize – suggest human involvement. -
Evolution of Mammals Classifying Mammals
Outline 20: Evolution of Mammals Classifying Mammals • Paleontologists recognize at least 5 major groups of mammals. Only 3 are still living: –Monotremes: lay eggs –Marsupials: poorly developed at birth –Eutherians or Placentals: well developed at birth 5 Major Groups: 3 Living Defining Mammals • Warm blooded • Fur • Milk glands • Can lay eggs or have some form of live birth. Recognizing Fossil Mammals • Our definition of mammals doesn’t work with fossil bones. • How do we recognize the first mammals? –Reptiles have 3 bones in lower jaw. –Mammals have 1 bone in lower jaw –Mammal teeth are specialized. Dinosaurs have 3 bones in lower jaw 2 3 1 Mammals have 1 bone in lower jaw Hadrocodium, a lower Jurassic mammal with a “large” brain (6 mm brain case in an 8 mm skull) Eomaia, oldest placental mammal, 125 my old, Lower Cretaceous, China Eomaia, oldest placental mammal, 125 my old, Lower Cretaceous, China Eomaia Mammal fossil from the Cretaceous of Mongolia Jaw bones • Reptiles have 3 bones in their jaw: dentary, articular, and quadrate. • Articular and quadrate bones of reptile jaw became the hammer and anvil bones of the mammalian inner ear. • Marsupials are born with a reptilian jaw, which quickly changes before they eat solid food. = articular of = quadrate of Human Ear Bones, or lower reptile upper reptile Auditory Ossicles jaw jaw Cochlea Mammal Teeth • Teeth make excellent fossils. • Reptile ancestors had simple, cone- shaped teeth they regularly replaced. • Mammal teeth are specialized into incisors, canines, pre-molars and molars. • Mammals have only two sets of teeth during their lifetime. A Nile crocodile. -
Deer from Late Miocene to Pleistocene of Western Palearctic: Matching Fossil Record and Molecular Phylogeny Data
Zitteliana B 32 (2014) 115 Deer from Late Miocene to Pleistocene of Western Palearctic: matching fossil record and molecular phylogeny data Roman Croitor Zitteliana B 32, 115 – 153 München, 31.12.2014 Institute of Cultural Heritage, Academy of Sciences of Moldova, Bd. Stefan Cel Mare 1, Md-2028, Chisinau, Moldova; Manuscript received 02.06.2014; revision E-mail: [email protected] accepted 11.11.2014 ISSN 1612 - 4138 Abstract This article proposes a brief overview of opinions on cervid systematics and phylogeny, as well as some unresolved taxonomical issues, morphology and systematics of the most important or little known mainland cervid genera and species from Late Miocene and Plio-Pleistocene of Western Eurasia and from Late Pleistocene and Holocene of North Africa. The Late Miocene genera Cervavitus and Pliocervus from Western Eurasia are included in the subfamily Capreolinae. A cervid close to Cervavitus could be a direct forerunner of the modern genus Alces. The matching of results of molecular phylogeny and data from cervid paleontological record revealed the paleozoogeographical context of origin of modern cervid subfamilies. Subfamilies Capreolinae and Cervinae are regarded as two Late Miocene adaptive radiations within the Palearctic zoogeographic province and Eastern part of Oriental province respectively. The modern clade of Eurasian Capreolinae is significantly depleted due to climate shifts that repeatedly changed climate-geographic conditions of Northern Eurasia. The clade of Cervinae that evolved in stable subtropical conditions gave several later radiations (including the latest one with Cervus, Rusa, Panolia, and Hyelaphus) and remains generally intact until present days. During Plio-Pleistocene, cervines repeatedly dispersed in Palearctic part of Eurasia, however many of those lineages have become extinct. -
Variation and Mechanisms of Life History Evolution in Insular Dwarfism As Revealed
bioRxiv preprint doi: https://doi.org/10.1101/2020.12.23.424186; this version posted December 24, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY 4.0 International license. 1 Title: 2 Variation and mechanisms of life history evolution in insular dwarfism as revealed 3 by a natural experiment 4 5 Author affiliation: 6 Shoji Hayashi1,2,*,**, Mugino O. Kubo3,*,**, Marcelo R. Sánchez-Villagra4, Hiroyuki 7 Taruno5, Masako Izawa6, Tsunehiro Shiroma6, Takayoshi Nakano2, Masaki Fujita7 8 9 1 Faculty of Biosphere-Geosphere Science, Okayama University of Science, Kita-ku 10 Ridaicho 1-1, Okayama, 700-0005, Japan; 2 Division of Materials and Manufacturing 11 Science, Graduate School of Engineering, Osaka University, Yamada-Oka, Suita, Osaka, 12 565-0871, Japan; 3 Department of Natural Environmental Studies, Graduate School of 13 Frontier Sciences, The University of Tokyo, 5-1-5 Kashiwanoha, Kashiwa, Chiba 277- 14 8563, Japan; 4 Paläontologisches Institut und Museum der Universität Zürich, Karl 15 Schmid-Strasse 4, CH-8006 Zürich, Switzerland; 5 Osaka Museum of Natural History, 16 Nagai Park 1-23, Higashi-Sumiyoshi-ku, Osaka, 546-0034, Japan; 6 Faculty of Science, 17 University of the Ryukyus, Okinawa 903-0213, Japan; 7 Department of Anthropology, 18 National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba-shi, Ibaraki 305- 19 0005, Japan 20 * Both authors contributed equally to this work. 21 ** Authors for correspondence: Shoji Hayashi e-mail: [email protected], Mugino 22 O.