the tentacled flathead ( papilloculiceps longiceps ) or crocodilefish is a member of the order scorpaeniformes , an order which also includes the scorpionfishes and stonefishes .

it is found in the western indian ocean ; the red sea .

the species now also occurs in the mediterranean , having invaded as a lessepsian migrant through the suez canal .

tentacled flathead

a tentacled flathead ( crocodilefish ) sits on the sand , waiting for prey to swim by .
crocodile fish carpet flathead underwater red sea . amazing relax video about marine tropical animals carpet flathead in world of wildlife .
macro of the eye of a tentacled flathead ( papilloculiceps longiceps ) , . . stock photo , picture and royalty free image . image 8932203 .
macro of the eye of a tentacled flathead ( papilloculiceps longiceps ) , also known as a crocodile fish . taken in the wakatobi , indonesia .
underwater close up flathead scorpion fish . picture of flathead scorpion fish in the tropical reef of the red sea , dahab , egypt .
stock photo - macro of the eye of a tentacled flathead ( papilloculiceps longiceps ) , also known as a crocodile fish . taken in the wakatobi , indonesia .
tentacled flathead ( papilloculiceps longiceps ) , also known as a crocodile fish , camouflaged on a coral reef while looking into the camera . taken in the wakatobi , indonesia .
crocodile fish underwater red sea . amazing relax video about marine tropical animals carpet flathead in world of wildlife .
crocodile fish carpet flathead underwater red sea . amazing relax video about marine tropical animals papilloculiceps longiceps in world of wildlife .
crocodile fish underwater red sea . amazing relax video about marine tropical animals carpet flathead papilloculiceps longiceps in world of wildlife .
crocodile fish carpet flathead papilloculiceps longiceps underwater red sea . amazing relax video about marine tropical animals in world of wildlife .
crocodile fish carpet flathead close - up underwater red sea . amazing relax video about marine tropical animals papilloculiceps longiceps in world of wildlife .
flathead fish in coral underwater red sea . colorful world of wild marine nature on background of beautiful lagoon . awesome video of wildlife in egypt .
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gilberto rodríguez . marine biologist . m . sc . , university of miami , usa . ph . d . , university of wales , uk . emeritus researcher , instituto venezolano de investigaciones científicas ( ivic ) . address : centro de ecología , ivic , apartado 21827 , caracas 1020 - a , venezuela . e - mail : grodrigu @ urltoken
héctor suárez . biologist , universidad de oriente , venezuela . research associate , ivic .
la dispersión antropogénica de crustáceos decápodos es parte de un fenómeno que actualmente adquiere características de un cambio ambiental global . de las 10000 especies conocidas de decápodos se han registrado 58 especies marinas , 8 dulceacuícolas - estuarinas y 21 cangrejos de ríos ( astacura ) que se han desplazado de sus áreas originales de distribución . las migraciones lessepsianas a través de canales , la introducción accidental por transporte marítimo y la importación para acuicultura son los mecanismos responsables de esta dispersión . en venezuela se han introducido 4 especies de camarones peneídeos , el camarón de río
y 3 especies de brachiuros . los crustáceos decápodos están particularmente bien adaptados por su anatomía y fecundidad para colonizar nuevas áreas y establecer poblaciones explosivas .
the anthropogenic dispersal of decapod crustaceans is part of an event that in recent years has acquired characteristics of a global environmental change . from the 10000 species of decapods so far recorded in the world , 58 marine species , 8 freshwater - estuarine species and 21 river crabs ( astacura ) have been recorded outside their original areas of distribution . lessepsian ms through channels , accidental maritime transport and introduction for aquaculture have been the mechanisms responsible for these dispersals . four penaeid species , the freshwater shrimp
, and 3 brachyuran crabs have been introduced in venezuela . due to their characteristic anatomy and high fecundity the decapod crustaceans are particularly apt for the colonization of new areas and the establishment of explosive populations .
a dispersão antropogênica de crustáceos decápodes é parte de um fenômeno que atualmente adquire características de uma mudança ambiental global . das 10000 espécies conhecidas de decápodes foram registradas 58 espécies marinhas , 8 doceacuícolas - estuarinas e 21 caranguejos de rios ( astacura ) que foram deslocados de suas áreas originais de distribuição . as migrações lessepsianas através de canais , a introdução acidental por transporte marítimo e a importação para aqüicultura são os mecanismos responsáveis desta dispersão . na venezuela foram introduzidas quatro espécies de camarões , peneídeos , o camarão de rio
e 3 espécies de brachiuros . os crustáceos decápodes estão particularmente bem adaptados por sua anatomia e fecundidade para colonizar novas áreas e estabelecer populações explosivas .
the suez canal is an unparalleled situation where two biogeographical provinces , previously totally separated , enter into contact and interpenetrate . the pharaonic connection between the red and mediterranean seas from the 13th to the 8th centuries bc , allowed the migration of very few species due to the low salinity there prevailing ( por , 1971 ) . the present canal , projected and built by ferdinand - marie de lesseps ( 1805 - 1894 ) was inaugurated on november 1869 . the channel spans 160 km , from the bay of suez in the red sea to port said in the mediterranean , using the intermediate manzala , timsah and bitter lakes .
there are at present 40 species of decapods in the mediterranean accounted for as lessepsian migrants ( table i ) while , on the contrary , the number of species dispersed in the area through ships ballast water is considered negligible . the occasional presence of the lobster thennus orientalis , first recorded in 1896 in fiume , italy ( elton , 1958 ) , can be explained by transport on ships hulls .
the kiel canal , built between 1887 and 1895 in an extent of 98 km , links the north sea with the baltic sea , from the mouth of the elbe river to the kiel bay , bypassing the detour along the danish peninsula . it is possible that the estuarine mud crab rhithropanopeus harrisii found its way to the baltic through the kiel canal , after its introduction in the netherlands , since its first record in that sea , in 1936 , was from the baltic end of the canal ( wolff , 1954 ) . in other respects , this canal is of little biogeographical relevance .
the panama canal spans 64 km from coast to coast . although its lake - lock design and the presence of a wide freshwater zone supplied by the chagres river precludes any lessepsian migration , at both ends of the present canal several fouling and perforating organisms ( pholadid bivalves , teredos , cirripeds , bryozoans , and other ) usually considered as natives of the opposing ocean , can be observed . these are euryhaline forms that can withstand transport through the freshwater section attached to the hulls of local vessels that have been moored for a long time ( carlton , 1985 ) .
it has been extensively debated whether it is possible that the ballast water discharged at opposite sides of the canal , since its aperture in 1914 , could be an
active
mechanism for the transport . carlton ( 1985 ) recorded 9 invertebrates and 4 fishes for which this mechanism is possible and 5 invertebrates and 2 fishes for which it is probable . among the decapod species , transport in ballast water is considered a possible mechanism for rhithropanopeus harrisii and the freshwater crab neorhynchus alcocki found in pedro miguel lock . eurypanopeus dissimilis , found in the third lock , ranges from florida to brazil and its presence in the panama canal is not unusual .
several species of crabs , such as pachygrapsus transversus , plagusia chabrus , p . depressa , p . tuberculata , planes minutus , carcinides maenas , menippe convexa , etc . , have been observed attached to ships hulls ( wolff , 1954 ) . this was a convenient means of dispersal for crabs when the old wooden hulls were in use , but it became unavailable with the modern metallic hulls , antifouling paints and reduced stowage time . no decapods were detected during a two - year survey of 89 vessels of different types that moored at new zealand ports , although 45 species of sessile invertebrates were found ( skerman , 1960 ) .
ballast water offers the most effective mechanism for the introduction of exotic decapod species , although it has been confirmed only in a few instances ( carlton , 1985 ) . larval stages of decapod crustaceans have been recovered from ballast tanks in viable conditions ( chu et al . , 1997 ) . rees and catley ( 1949 ) proposed this mechanism for the introduction of the larvae of processa equimana , living in the north sea plankton , into the mediterranean and red sea , but carlton ( 1985 ) considered this unlikely . furthermore , the north sea form has been described as a different species , p . modica , with the subspecies p . modica carolii in the mediterranean .
peters and panning ( 1933 ) reviewed the available evidence for the first implant of the chinese crab eriocheir sinensis in the north sea , from 1912 , via ballast water . subsequently large populations of this crab have colonized the european brackish waters and have extended to california and canada , apparently using the same transport mechanism ( cohen and carlton , 1997 ) .
ship cargo also seems to play a role in the transport of organisms . the saber crab platychirograpsus spectabilis was transported in a cargo of cedar logs from tabasco state , mexico , to hillsborough ( st . petersburg ) , florida , where it has become permanently established ( marchand , 1946 ) .
these and similar structures can be slowly towed across the ocean , with the submersed parts unprotected by antifouling paints , and stay anchored for extended periods of time in different parts of the world . in this way was transported the japanese shore crab plagusia dentipes , observed alive in california after a trans - pacific cruise of 61 days on a self - powered drilling platform , which had been working for four years between japan and malaysia ( benech , 1978 ) . similar circumstances are mentioned for the transport of plagusia tuberculata , together with an encrusting community of cirripeds , on an oil platform built in japan and transported to new zealand , after a voyage of 68 days ( foster and williams , 1979 ) .
according to zenkevitch ( 1963 ) two mediterranean shrimps , palaemon adspersus and p . elegans , were accidentally introduced into the aral sea with species of mullets . we have already mentioned the possible introduction of rhithropanopeus harrisii along the atlantic coast of the united states , with oysters , and of carcinus maenas , into san francisco bay , with algae used for packing of living bait ( cohen and carlton , 1997 ) . the dispersal of several species of crayfish throughout the united states , orconectes rusticus for instance ( table iii ) , is attributed to their escape from containers of living bait used by sport fishermen ( taylor and redmer , 1996 ) .
cohen and carlton ( 1997 ) recorded 16 cases of interception of living specimens of eriocheir sinensis at san francisco airport between 1989 and 1995 , with the confiscation of 10 to 50 specimens on each occasion . this species is sold alive in asian food markets in the united states ( lemaitre , 1995 ) .
the trade of organisms for aquaria is another means of dispersal . living specimens of atya scabra and procambarus clarkii can be observed frequently at the pet shops in caracas .
the species of peneaidae most frequently cultured are marsupenaeus japonicus and penaeus monodon , but the possibility exists of an increase in the culture of nine other species , p . semisulcatus , farfantepenaeus aztecus , fenneropenaeus indicus , f . penicillatus , sergestes orientalis , litopenaeus schmitti , l . setiferus , l . stylirostris and l . vannamei ( liao and huang , 1982 ) . farfantepenaeus aztecus is already under experimental culture in aquacop station , in tahiti , since 1975 , and l . stylirostris in corpus christy , texas , and crystal river , florida , from approximately the same date . m . japonicus was the first species cultured in the world after the research by hudinaga in 1934 . by 1942 its full culture had been achieved , but only after world war ii it was established on a commercial basis . in the united states shrimp culture began by the native species l . setiferus and f . aztecus in 1963 , and f . duorarum in 1968 . this same year liao began the culture of p . monodon in taiwan .
although the penaeids have being cultured in shrimp farms for an extended period of time , there is little information in the literature on the escape of these species and its effects on the natural ecosystems . the information available on the dispersal of m . japonicus in the mediterranean sea indicates that , at least under conditions of biological poverty , the species is able to propagate rapidly into the recipient communities .
a vibriosis producing bacterial hemorragic septicemia is frequently found in penaeid cultures in asia and south america . the ethiological agent , vibrio harveyi , has been detected in shrimp farms in venezuela , infesting l . vanamei vanamei and l . stylyrostris , but it is also present in feral populations of l . schmitti and several species of marine and estuarine fishes ( alvarez et al . , 1998 ) .
) . it has been introduced through all the american continent , except in nicaragua , belize , chile , paraguay and bolivia . in the caribbean it is farmed in the dominican republic , puerto rico , dominica , guadalupe , martinica , saint lucia and trinidad - tobago ( fao , 1995 ) .
the crayfishes of the families astacidae and parastacidae are strictly freshwater organisms . according to hobbs et al . ( 1989 ) it would be impossible to mention all the introductions of astacids since in many cases , or in the majority , they have not been recorded ( table iii ) . the largest diversity of species occurs in north america , where hobbs ( 1989 ) reported 271 species and subspecies native to canada , usa and mexico . several species have been transplanted within north america itself . this is the case of orconectes rusticus , which has displaced two other native species of orconectes in illinois ( taylor and redmer , 1996 ) . these displacements can affect the ecosystem through the reduction in the density of macrophytes ( lodge and lorman , 1987 ) and benthonic macroinvertebrates ( houghton et al . , 1998 ) .
another species that has extended its intracontinental distribution is cherax destructor , from australia , which has furthermore reached intercontinentally to washington state . the most notable case of intercontinental expansion is presented by procambarus clarkii , which has spread out to all continents , except australia and antarctica . laurent and forest ( 1979 ) summarized the damages produced by this species as follows :
in japan it was introduced since 1930 ( from 18 specimens ) . a short time afterwards it pullulated , destroying rice fields and ditches , producing populations of 2000 k / hectare and not been used by the people [ as food ] in kenya it produced rapidly enormous populations not exploited by the natives breaking the fishing gears and interfering with the breeding of tilapia
.
authorization for the introduction of marsupenaeus japonicus , p . monodon , litopenaeus stylirostris and l . vannamei was granted by the ministry of agriculture of venezuela on november 1984 . the original areas of distribution of the two former species are in the indian ocean and the western pacific , and of the other two on the pacific coast of america .
this species was introduced in 1980 , into a small shrimp farm in margarita island , by la salle foundation . although its culture in the country has been unsuccessful , pereira et al . ( 1996 ) found a wild population in the orinoco delta in 1996 . they considered that this species passed into the natural environment between 1991 and 1993 .
one specimen of this crayfish ( total length 8 . 2 cm , ivic reference collection ) was captured in a pond of the officers club ( circulo militar ) in caracas . as mentioned before , the species is available at pet shops in the city .
, a species from the indopacific and red sea regions that has migrated to the mediterranean and several localities in the caribbean , has been recorded by hernández and bolaños ( 1995 ) from eastern venezuela . ( 2 )
, recorded by hernández and bolaños ( 1995 ) from eastern venezuela , had not been previously observed outside its original area of distribution , between california and peru , and a misidentification by these authors cannot be ruled out . ( 3 )
was observed by the first time in 1957 in several localities of the tablazo and strait of maracaibo ( rodríguez , 1963 ; rodríguez and morales , 2000 ) . as the species was already very abundant in this estuary in 1957 , it should have been introduced several years before , possibly in the ballast water of oil tankers involved in the intense traffic between the local oil terminals and the eastern coast of the united states .
from approximately 10000 species of decapod crustaceans described in the world , 88 are recorded in the present contribution as migrants - 58 from marine environments , 8 from freshwater - estuarine habitats and 21 are freshwater crayfishes - but still there may be others not yet recorded in the literature . these species have migrated through navigation channels , have been accidentally transported by ships , inadvertently imported with fishery products or introduced for aquaculture . it might be expected that the list would increase in the future .
the effect of invading species on recipient communities could be direct , by displacement of native species or predation on other members of the community . indirectly they could affect the native species through introduction of diseases . up to the present severe damages have been rarely observed , for instance the destruction of rice fields in japan by procambarus clarkii ( laurent and forest , 1979 ) . but as the invasion into coastal zones , rivers and lakes progresses , we can expect the gradual displacement of native forms , with results impossible to foresee at present .
the authors are grateful to jesus eloy conde and jon paul rodríguez for critically reading the manuscript .
2 . abele lg ( 1972 ) introductions of two freshwater decapod crustaceans ( hymenosomatidae and atydae ) into central and north america .
4 . aron wi , smith sh ( 1971 ) ship canals and aquatic ecosystems .
5 . balss h ( 1927 ) bericht über die crustacea decapoda ( natantia und anomura ) . zoological results of the cambridge expedition to the suez canal , 1924 . xiv .
6 . balss h ( 1936 ) decapoda . in : the fishery grounds near alexandria . vii .
8 . ben - tuvia a ( 1966 ) red sea fishes recently found in the mediterranean .
10 . carlton jt ( 1985 ) transoceanic and interoceanic dispersal of coastal marine organisms : the biology of ballast water .
11 . chu kh , tam pf , fung ch , chen qc ( 1997 ) a biological survey of ballast water in container ships entering hong kong .
14 . dall w , hill bj , rothlisberg pc , sharples dj ( 1990 ) the biology of the penaeidae .
15 . enzenross r , enzenross l ( 2000 ) nichmediterrane crustacea - arten in tunesischen gewässern ( decapoda , macrura und brachyura ) .
18 . foster ba , willan c ( 1979 ) foreign barnacles transported to new zealand on an oil platform .
19 . fox hm ( 1926 ) zoological results of the cambridge expedition to the suez canal , 1924 . i . general part .
20 . galil bs ( 1992 ) eritrean decapods in the levant : biogeography in motion .
21 . galil bs ( 1997 ) two lessepsian migrant decapods from the eastern mediterranean .
22 . galil bs , golani d ( 1990 ) two new migrant decapods from the eastern mediterranean .
( fabricius , 1798 ) ( crustacea : decapoda ) a new lessepsian migrant to the eastern mediterranean .
24 . garcía s , le reste l ( 1981 ) life cycles , dynamics , exploitation and management of coastal peneid shrimp stocks .
( gould ) an american crab in the estuary of the mondego river , portugal .
27 . gorgy s ( 1966 ) les pêcheries et le milieu marin dans le secteur méditerraneen de la république arabe unie .
30 . gruvel a ( 1928 ) répartition géographique de quelques crustacés comestibles sur les côtes dégipte et syrie .
32 . hobbs jr hh ( 1989 ) an illustrated checklist of the american crayfishes ( decapoda : astacidae , cambaridae and parastacidae ) .
33 . hobbs iii hh , jas jp , huner jv ( 1989 ) a review of global crayfish introductions with particular emphasis on two north american species ( decapoda , cambaridae ) .
34 . hernández g , bolaños j ( 1995 ) additions to the anomuran and brachyuran fauna of northeastern venezuela . the crustacean society summer meeting , may 25 - 27 , 1995 [ links ] [ abstract ]
35 . holthuis lb ( 1961 ) report on a collection of crustacea decapoda and stomatopoda from turkey and the balkans .
36 . holthuis lb ( 1980 ) fao species catalogue . shrimps and prawns of the world . an annotated catalogue of species of interest to fisheries .
37 . holthuis lb , gottlieb e ( 1958 ) an annotated list of the decapod crustacea of the mediterranean coast of israel with an appendix listing the decapoda of the eastern mediterranean .
( linneaus ) , to british columbia , canada , and washington , usa .
40 . laurent pj , forest j ( 1979 ) donneé sur les écrevisses qon peut rencontrer en france .
( milne edwards , 1867 ) , a nonindigenous portunid crab ( crustacea : decapoda : brachyura ) discovered in the indian river lagoon system of florida .
42 . lewinsohn c , holthuis lb ( 1964 ) new records of decapod crustacea from the mediterranean coast of israel and the eastern mediterranean .
44 . liao c , huang tl ( 1982 ) status and prospect of the culture of two important penaeid prawns in asia .
49 . mccosker je , dawson ce ( 1975 ) biotic passage through the panama canal , with particular reference to fishes .
: grapsidae ) , in its new habitat along the atlantic coast of the united states : geographic distribution and ecology .
51 . monod t ( 1930 ) ueber einige indo - pazifische decapoden der meeresfauna syriens .
53 . naylor e ( 1960 ) a north american xanthoid crab new to britain .
54 . newman wa ( 1963 ) on the introduction of an edible oriental shrimp ( caridea , palaemonidae ) to san francisco bay .
57 . por fd ( 1971 ) one hundred years of suez canal - a century of lessepian m : retrospect and viewpoints .
58 . ramadan se , dowidar nm ( 1972 ) brachyura ( decapoda , crustacea ) from the mediterranean waters of egypt .
60 . rodríguez g ( 1963 ) the intertidal estuarine communities of lake maracaibo , venezuela .
61 . rodríguez g , morales f ( 2000 ) las comunidades bentónicas del sistema de maracaibo . in : rodríguez g ( ed )
62 . skerman tm ( 1960 ) ship - fouling in new zealand waters : a survey of marine fouling organisms from vessels of the coastal and overseas trades .
63 . steinitz h ( 1967 ) a tentative list of immigrants via the suez canal .
67 . wolff t ( 1954 ) occurrence of two east american species of crabs in european waters .
68 . zenkevitch l ( 1963 ) biology of the seas of the u . s . s . r . interscience publishers . new york . 955 pp . [ links ]
calle vera cruz , residencia la hacienda apto . 31 - m las mercedes , caracas 1080 . venezuela telefonos : ( 58 - 0212 ) 992 - 32 - 24 , 991 - 75 - 25 interciencia @ urltoken
latin , papillo , papula = any little like protuberance + greek , kylix , - ikos = cup + latin , ceps = head ( ref . 45335 )
marine ; reef - associated ; non - migratory ; depth range 1 - 15 m ( ref . 9710 ) . tropical
western indian ocean : red sea , including the gulf of aqaba to south africa and madagascar .
maturity : l m ? range ? - ? cm max length : 70 . 0 cm tl male / unsexed ; ( ref . 4315 ) ; common length : 50 . 0 cm tl male / unsexed ; ( ref . 3476 )
dorsal spines ( total ) : 9 ; dorsal soft rays ( total ) : 11 ; anal spines : 0 ; anal soft rays : 11 . body mottled with brownish or greenish above , whitish below ; caudal fin with 3 - 4 dark bands , other fins with large dark blotches ( ref . 4315 ) .
knapp , l . w . , 1984 . platycephalidae . in w . fischer and g . bianchi ( eds . ) fao species identification sheets for fishery purposes . western indian ocean ( fishing area 51 ) . vol . 3 . fao , rome . pag . var . ( ref . 3476 )
) : 24 . 7 - 29 . 2 , mean 28 . 1 ( based on 1815 cells ) .
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 1 . 0000 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 00513 ( 0 . 00222 - 0 . 01187 ) , b = 3 . 04 ( 2 . 83 - 3 . 25 ) , in cm total length , based on lwr estimates for this ( sub ) family - body shape ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 4 . 0 ±0 . 7 se ; based on size and trophs of closest relatives
resilience ( ref . 69278 ) : medium , minimum population doubling time 1 . 4 - 4 . 4 years ( preliminary k or fecundity . ) .
vulnerability ( ref . 59153 ) : moderate to high vulnerability ( 46 of 100 ) .
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english - german online dictionary developed to help you share your knowledge with others . more information ! contains translations by tu chemnitz and mr honey ' s business dictionary ( german - english ) . thanks on that account ! links to this dictionary or to single translations are very welcome ! questions and answers
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stock images of crocodilefish ( papilloculiceps longiceps ) resting on reef . woodhouse reef , sharm el sheikh , south sinai , red sea , egypt . u17515486 - search stock photography , poster photos , pictures , and photo clip art - u17515486 . jpg
stock image of crocodilefish ( papilloculiceps longiceps ) resting on reef . woodhouse reef , sharm el sheikh , south sinai , red sea , egypt .
crocodilefish ( papilloculiceps longiceps ) resting on reef . woodhouse reef , sharm el sheikh , south sinai , red sea , egypt .
crocodile fish ( papilloculiceps longiceps ) . jackson reef , sharm el sheikh , south sinai , red sea , egypt
soft coral ( dendronephthya klunzingeri ) . woodhouse reef , sharm el sheikh , south sinai , red sea , egypt .
risso ' s dolphin ( grampus griseus ) sunbathing on the surface . woodhouse reef , sharm el sheikh , south sinai , red sea , egypt .
bluespotted stingray ( taeniura lymma ) resting in the sand . jackson reef , sharm el sheikh , south sinai , red sea , egypt .
risso ' s dolphins ( grampus griseus ) sunbathing on the surface . woodhouse reef , sharm el sheikh , south sinai , red sea , egypt .
hawksbill turtle ( eretmochelys imbricata ) species critically endangered . woodhouse reef ; sharm el sheikh ; south sinai ; red sea ; egypt
blacktip grouper ( epinephelus fasciatus ) . tower , sharm el sheikh , south sinai , red sea , egypt .
starry puffer ( arothron stellatus ) resting on a mooring drum . na ' ama bay , sharm el sheikh , south sinai , red sea , egypt .
zebra shark ( stegostoma fasciatum ) resting on sandy slope . species near threatened . sha ' ab canoî , ras mohamed national park , sharm el sheikh , south sinai , red sea , egypt .
dave , the famous hawksbill turtle ( eretmochelys imbricata ) from sharm el sheikh , rescued after a boat propellor accident , enjoying a meal with his partially fiber cast fixed shell . critically endangered species . shark and yolanda , ras mohamed national par
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video client : patricia martin cabrera review : abdel rahman el gamal ( founder of the video channel ) the video was filmed in damaniyat island ( al batinah region ) , sultanate of oman . the honeycomb moray eel , gymnothorax favagineus shown in the video is a species of marine fish in the family muraenidae and is one of the largest of indo - pacific morays and …
photo credit : sherif sadek ( egypt ) review : abdel rahman el gamal ( founder of the website ) the inserted picture of a brown shrimp specimen was taken in a fish farm in egypt . introduction : brown shrimp ( farfantepenaeus aztecus ) is a species of marine penaeid shrimps which has an important commercial species in the usa . the species has several common …
brown shrimp , brown shrimp and environmental conditions , brown shrimp in the mediterranean sea , description , egypt . , farfantepenaeus aztecus , feeding habits , northern brown shrimp , reproduction habits , usa , utilization
review : abdel rahman el gamal ( founder of the video channel ) this video was filmed in a retail shop located in monterey , california ( usa ) where different fish species and crabs are sold . the dungeness crab is the crab species shown in this video whether sold iced or displayed live . the dungeness crab , metacarcinus magister ( formerly : cancer magister ) , is …
video courtesy : the state archives of florida , usa the title of this video : “florida silent sirens : manatees in peril” and was filmed during 1980s the caption of the video states : “this is an excellent film about the plight of the endangered manatee . it is narrated by leonard nimoy and is full of beautiful underwater …
video credit : patricia martin cabrera ( united arab emirates ) review : abdel rahman el gamal ( founder of the video channel ) introduction : the emperor angelfish ( pomacanthus imperator ) , is a reef - associated marine angelfish and is also called the imperator angelfish or imperial angelfish . the species is one of the most stunning underwater fish . their color and graceful shape make them one …
credit : ajith kumara ( srilanka ) review : abdel rahman el gamal ( founder of the website ) mud crab ( scylla serrata ) which is also called mangrove crab belongs to the family of swimming crabs ( portunidae ) . this is an economically important species of crabs and considered highly esteemed as food whereas the flesh from its claws and walking legs …
video credit : patricia martin cabrera ( united arab emirates ) review : abdel rahman el gamal ( founder of the video channel ) source : urltoken this video was filmed at ras muhammad , sharm el sheikh ( egypt ) . introduction : the video was filmed is the premise of the british navy ship “thistlegorm” which was built in 1940 and got sunk …
photo credit : : patricia martin cabrera ( united arab emirates ) review : abdel rahman el gamal ( founder of the website ) the inserted picture was taken in sharm el sheikh ( egypt ) introduction : blue spotted ribbon tail ray ( taeniura lymma ) is a species of stingray in the family dasyatidae . it is also known as aka blue - spotted fantail rays , blue spotted stingrays , blue spotted rays , and ribbontail …
this video was filmed in the sea world , san diego , usa source : urltoken giant grouper ( epinephelus lanceolatus ) the giant grouper ( epinephelus lanceolatus ) , which belongs to the family serranidae is also known as brindle bass , brown spotted cod , or bumblebee grouper and as the queensland giant grouper in australia . the species is the largest reef - dwelling bony …
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sharm al - sheikh , egypt . 26th october , 2016 . egyptian president abdel fattah al - sisi takes photo with people following the peace marathon in the red sea resort of sharm al - sheikh , in the south sinai governorate , south of cairo , egypt , on oct . 26 , 2016 credit : egyptian president office / apa images / zuma wire / alamy live news
el - sheikh , egypt . 7th nov , 2015 . tourists have fun on yachts along a coast in the resort of sharm al - sheikh , egypt , on nov . 7 , 2015 . russia on friday suspended all flights to egypt as speculation rose over possible terrorist attacks on the russian plane that crashed over sinai peninsula last saturday . credit : xinhua / alamy live news
sharm al - sheikh , egypt . 26th october , 2016 . egyptian president abdel fattah al - sisi rides a bicycle at the street in the red sea resort of sharm al - sheikh , in the south sinai governorate , south of cairo , egypt , on oct . 26 , 2016 credit : egyptian president office / apa images / zuma wire / alamy live news
u . s . secretary of state john kerry sits with egyptian president abdel fattah al - sisi at the congress center in sharm el - sheikh , egypt , on march 13 , 2015 , before a bilateral meeting and their attendance at an egyptian development conference .
sharm al - sheikh , egypt . 26th october , 2016 . egyptian president abdel fattah al - sisi speaks during the peace marathon in the red sea resort of sharm al - sheikh , in the south sinai governorate , south of cairo , egypt , on oct . 26 , 2016 credit : egyptian president office / apa images / zuma wire / alamy live news
egypt , sinai peninsula , gulf of tiran , sharm el - sheikh . snorkeling in the red sea ' s ras mohamed national marine park .
sharm el sheikh , egypt . 8th dec , 2017 . egyptian president abdel - fattah al - sisi ( 2nd r , front ) attends the official opening ceremony of the africa 2017 forum in sharm el sheikh , egypt , on dec . 8 , 2017 . credit : zhao dingzhe / xinhua / alamy live news
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golden - crowned ( tattersall ' s ) sifaka , andranotsimaty , daraina , madagascar . ( propithecus tattersalli )
crested coua ( coua cristata ) , very attractive madagascar birds . ankarafantsika national park , madagascar wildlife and wilderness
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pamizinisaurus is a genus of sphenodontian reptile known from lower cretaceous ( albian ) tlayúa formation of mexico .

a crushed skeleon of a juvenile reptile was found in tlayua quarry , in central mexico .

it was named pamizinsaurus tlayuaensis by reynoso in 1997 , after the name of the quarry of which it was found .

its skull length is 16 millimetres ( 0.63 in ) .

the fossil was covered in small round osteoderms that could have protected it from predators .

pamizinsaurus

pamizinsaurus was a genus of the subfamily sphenodontinae ; grouping it with the modern sphenodon ( better known as the tuatara ) , zapatadon , cynosphenodon , homoeosaurus , sapheosaurus , and ankylosphenodon .
pamizinsaurus was a genus of the subfamily sphenodontinae ; grouping it with the modern sphenodon ( better known as the tuatara ) , zapatadon , cynosphenodon , homoeosaurus , sapheosaurus , and ankylosphenodon .
the taxon pamizinsaurus from the early cretaceous of mexico ( reynoso 1997 ) was identified as a wildcard taxon that strongly influenced the results . this may partially be due to high amounts of missing data ( 64 % ) in this taxon ; however , that several other taxa have even higher amounts of missing data ( eilenodon 68 % ; cynosphenodon : 73 % ; toxolophosaurus : 79 % ) but proved to be less problematic , indicates that the wildcard status of pamizinsaurus might also be due to some character conflict .
reduced strict consensus ( after removing pamizinsaurus ) of sphenodontian interrelationships from 24 most parsimonious trees of 186 steps showing the phylogenetic position of sphenocondor gracilis gen . et sp . nov . bremer support and frequency differences ( gc ) , calculated by symmetric resampling , are indicated above the nodes .
this page is based on the copyrighted wikipedia article pamizinsaurus ; it is used under the creative commons attribution - sharealike 3 . 0 unported license ( cc - by - sa ) . you may redistribute it , verbatim or modified , providing that you comply with the terms of the cc - by - sa
in the strict consensus of the present analysis , the inter - relationships within recovered groupings of derived sphenodontians are poorly resolved , as in other analyses with somewhat similar taxon sampling ( e . g . reynoso , 2000 , 2005 ) . however , this is mainly because of the unstable position of pamizinsaurus , which might be a consequence of the immature condition of the sole known specimen and its peculiar combination of character states ( reynoso , 1997 ) . therefore , pamizinsaurus was excluded from the strict consensus ( but not from the analysis ) , rendering a more resolved topology in the reduced strict consensus ( fig . 5 ) .
in a next step , we thus removed pamizinsaurus from the matrix and ran a second analysis with the same settings as above . the analysis resulted in only two most parsimonious trees with a length of 184 steps ( ci 0 . 473 , ri 0 . 654 , rc 0 . 309 ) . the strict consensus tree of these two trees (
vs - m size . see clevosaurs entry for basic skull structure . $ teeth fused to jaw margin ( fully acrodont ) and not replaced ; less than 4 ( 7 ? ) premaxillary teeth ; $ marginal teeth added posteriorly as jaw grows ; dentary fits between maxillary teeth and parallel row of palatal teeth ; meckelian canal runs along midline of jaw & is at least partially open ; broad mandibular symphysis ; prominent coronoid process of dentary , posterior process of dentary ends posterior to coronoid ; with posterior surangular facet ; frontals not fused ? ? ) ; $ lacrimal absent ; large upper temporal fenestra ; complete lower temporal bar ; quadratojugal retained ; rigid quadrate ; well - developed posterior tubercle on posterior margin of ischium ; epiphyses present with determinate growth ; may have osteoderms ( pamizinsaurus ) ; food crushed thoroughly ( insects & birds ) .
in a next step , we thus removed pamizinsaurus from the matrix and ran a second analysis with the same settings as above . the analysis resulted in only two most parsimonious trees with a length of 184 steps ( ci 0 . 473 , ri 0 . 654 , rc 0 . 309 ) . the strict consensus tree of these two trees ( fig . 3 and information s1 ) is considerably better resolved than that resulting from the full analysis , but , importantly , it does not contradict the results of the latter , neither in the strict consensus , nor in any reduced consensus tree . thus , we chose these results to illustrate and further explore the relationships of oenosaurus . bootstrap ( 10000 replicates ) and bremer support analyses were carried out on the pruned matrix , and the following list of synapomorphies is based on the strict consensus tree resulting from the pruned analysis .
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full reference : v . h . reynoso . 1997 . a ' beaded ' sphenodontian ( diapsida : lepidosauria ) from the early cretaceous of central mexico . journal of vertebrate paleontology 17 : 52 - 59
type specimen : instituto de geologãa , universidad nacional autã³noma de mã©xico , a partial skeleton ( severaly crushed skeleton of a juvenile ) . its type locality is tlayua formation , which is in an albian lagoonal / restricted shallow subtidal lime mudstone in the tlayua formation of mexico .
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sphenodontia : diphydontosaurus + ( planocephalosaurus + * : clevosauridae + stem group sphenodontinae ) .
comments : cladistically , this taxon ( equivalent to the standard linnean taxonomic sphenodontidae ) should perhaps be called something like sphenodontoidea , as it includes clevosauridae , pleurosauridae , . crown group sphenodontidae , etc
introduction to the sphenodontidae ; turtle , tuatara , crocodile checklist - - 3 ; jurassic page ' s gallery : the terrestrial wildlife ( sapheosaurus ) ; lebende fossilien , intro , teil 2 ( homoeosaurus ) .
references : evans & sigogneau - russell ( 1997 ) ; reynoso 1996 ) ; reynoso ( 1997 ) ; reynoso & clark ( 1998 ) .
image : kallimodon among a very good collection of fossil photos ) from herbert krause . reproduced by permission .
note : the nomenclature of sphenodont taxa tends to be rather confused . the scheme used here is adopted from the sensible system of mikko haaramo , who credits wu , x - c ( 1994 ) , late triassic - early jurassic sphenodontians from china and the phylogeny of the sphenodontia , in fraser , nc & h - d sues ( eds . ) , in the shadow of the dinosaurs , cambridge univ . press . atw001117 .
1 or more robust
incisor
teeth forming chisel edge ankylosed to premaxilla ; palatine dentition ; marginal dentition acrodont or absent ; shearing ( not crushing or piercing ) bite with simple orthal up - and - down ) jaw motion ; $ long thin posterior process of premaxilla excludes maxilla from margin of nares ; $ lacrimal absent ; may have dentine
lips
; $ antorbital skull very short ( < 25 % of total skull ) ; triradiate postfrontal broadly contacts frontal ; complete lower temporal bar formed by jugal loosely contacting slim quadratojugal ; $ dorsal process of jugal elongate ; $ large ( > 25 % skull length ) lower temporal fenestra ; gap between jugal and quadratojugal small ; proximal caudal vertebrae have large transverse processes ; caudal autotomy septa may be present distally ; humerus strongly bent and expanded at both ends ; femur gently s - shaped .
note : for reasons i have not been able to determine , no one seems to have gotten around to giving this group a latinized family name . atw . update : this has now been rectified . on the basis of new material arantes et al 2009 formalises the family clevosauridae , composed of clevosaurus , brachyrhinodon and polysphenodon . mak101016
links : diapsida ; clevosaurus bairdi ( but server is frequently reduced or absent ! ) ; autapomorphies of diapsid clades ; untitled document ; lecture 12 - early jurassic .
reynoso 1996 ) ; reynoso & clark 1998 ) ; sues et al . ( 1994 ) .
short snout like brachyrhinodon , multiple rows of teeth on the palate . it shows sphenodontian characters such as a groove between the maxilla and palatine for the dentary
comments : known from a partial skull and skeleton . the only known specimen went missing in the 1930s , but a number of casts and plaster moulds are available ( fraser & benton 1989 p . 416 . )
brachyrhinodon taylori , skull in ventral ( a ) , dorsal ( b ) and lateral ( c ) view . from fraser & benton 1989 p . 416 . scale bar 1 cm . note palatal teeth in ( a ) ; teeth on the upper palate ( roof of the mouth ) are a primitive feature found in most basal amniotes
small ( c . 15 cm long ) , short - snouted skull with premaxillary beak . benton 1985
kallimodon pulchellus ( zittel ) , 1887 . late jurassic ; kelheim , bavaria . ventral aspect . from zittel / eastman / woodward 1902 p . 151 . this represents a type similar to or intermediate between homoeosaurus and sapheosaurus
sphenodontoidea : clevosaurinae + * : homoeosaurus + pleurosauridae + ( sapheosaurus + sphenodontinae ) .
vertebrae amphicoelous , sometimes with persistent notochord ; intercentra present in cervical ( neck ) and caudal ( tail ) regions . external nares separated ; interclavicle t - shaped ; dermal scales subrectangular . greatiy enlarged postfrontal bone extending far posteriorly ( to the rear ) on parietal . except for a few teeth on the vomer in juveniles and the palatine row , all other palatal teeth absent ( i . e . teeth on the roof of the mouth , in contrast to primitive amniotes ) . premaxillae each with a small pointed tooth . a single series of enlarged , depressed , triangular , acrodont teeth present on maxillae , mandibles , and outer edge of the palatines ; vomer toothless . pattern of tooth wear facets , tooth ultrastructure , and anteroposterior length of the mandibular articulation , all of which indicate development of the propalinal masticatory movements ( front to rear chewing ) as with the recent sphenodon zittel / eastman / woodward 1902 , gauthier et al 1988 p . 25
kimmeridgian ( lithographic stone ) of bavaria , and cerin , france . kimmeridgian of hanover , and purbeckian ( tithonian ) of england .
from zittel / eastman / woodward 1902 ( copyright expired ) :
attaining a length of between 20 and 40 cm . , and differing from the recent sphenodon in that intercentra are absent between the dorsal vertebrae ribs without uncinate processes , and humerus not pierced by entepicondylar foramen . mandibular rami united at the symphysis by ligaments ; second sacral rib bifid distally .
applies also to sapheosaurus and kallimodon gauthier et al 1988 p . 26 : compared to sphenodontines and sapheosaurs , homoeosaurus is distinguished by smaller size , gracile and elongate limbs , and a broad parietal table ( skull roof above the orbits ( eye holes ) ) , all of which , with the possible exception of the limb proportions , are plesiomorphic ( shared ancestral ) features .
using this material . all material by atw is public domain and may be freely used in any way ( also any material jointly written by atw and mak ) . all material by mak is licensed creative commons attribution license version 3 . 0 , and may be freely used provided acknowedgement is given . all wikipedia material is either gnu open source or creative commons ( see original wikipedia page for details ) . other graphics are copyright their respective owners
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early cretaceous lepidosaurs ( reptilia : d . . . - pdf document ( 15 m )
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today were then held by sphenodontians . there were even several successful groups of aquatic sphenodontians such as
( loss of the tail - tip when threatened ) , and have transverse cloacal slits .
r . l . ditmars , litt . d , says ;
the tuatara resembles in form stout - bodies modern lizards , which we might call iguanas ; this resemblance is further intensified by a row of spines upon the back . it is dark olive , the sides sprinkled with pale dots . the eye has a cat - like pupil . large specimens are two and a half feet long . while superficial resemblance might tend to group this reptile with lizards , its skeleton and anatomy show it to belong to a different part of a technical classification .
of the british museum noted features similar to birds , turtles , and crocodiles . he proposed the order rhynchocephalia ( meaning
beak head
) for the tuatara and its fossil relatives .
proposed the sphenodontia to include only tuatara and their closest fossil relatives in 1925 .
ditmars , raymond l . ,
reptiles of the world
the macmillan co . , new york , 1936 , p . xii
v . h . reynoso . 2000 . an unusual aquatic sphenodontian ( reptilia : diapsida ) from the tlayua formation ( albian ) , central mexico . journal of paleontology 74 : 133 - 148
cree , alison . 2002 . tuatara . in : halliday , tim and adler , kraig ( eds . ) , the new encyclopedia of reptiles and amphibians , oxford university press , oxford , pp . 210–211 . isbn 0 - 19 - 852507 - 9
fraser , nicholas ; sues , hans - dieter ; ( eds ) ( 1994 ) .
evans , s . e . , prasad , g . v . r . & manhas , b . k . , 2001 : rhynchocephalians ( diapsida : lepidosauria ) from the jurassic kota formation of india . zoological journal of the linnean society : vol . 133 , # 3 , pp . 309 - 334
rauhut , o . w . m . ; heyng , a . m . ; lópez - arbarello , a . ; hecker , a . ( 2012 ) . farke , andrew a , ed .
a new rhynchocephalian from the late jurassic of germany with a dentition that is unique amongst tetrapods
.
daugherty , ch ; cree , a ; hay , jm ; thompson , mb ( 1990 ) .
neglected taxonomy and continuing extinctions of tuatara (
evans , se ( 2003 ) .
at the feet of the dinosaurs : the early history and radiation of lizards
.
jones meh . 2009 . dentary tooth shape in sphenodon and its fossil relatives ( diapsida : lepidosauria : rhynchocephalia ) . in koppe t , meyer g , alt kw , eds . interdisciplinary dental morphology , frontiers of oral biology ( vol 13 ) . greifswald , germany ; karger . 9–15 .
evans se , jones meh ( 2010 ) the origin , early history and diversification of lepidosauromorph reptiles . in bandyopadhyay s . ( ed . ) , new aspects of mesozoic biodiversity , 27 lecture notes in earth sciences 132 , 27 - 44 .
this article is issued from wikipedia - version of the 11 / 25 / 2016 . the text is available under the creative commons attribution / share alike but additional terms may apply for the media files .
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rhynchocephalians , the sister group of squamates ( lizards and snakes ) , are only represented by the single genus sphenodon today . this taxon is often considered to represent a very conservative lineage . however , rhynchocephalians were common during the late triassic to latest jurassic periods , but rapidly declined afterwards , which is generally attributed to their supposedly adaptive inferiority to squamates and / or mesozoic mammals , which radiated at that time . new finds of mesozoic rhynchocephalians can thus provide important new information on the evolutionary history of the group .
a new fossil relative of sphenodon from the latest jurassic of southern germany , oenosaurus muehlheimensis gen . et sp . nov . , presents a dentition that is unique amongst tetrapods . the dentition of this taxon consists of massive , continuously growing tooth plates , probably indicating a crushing dentition , thus representing a previously unknown trophic adaptation in rhynchocephalians .
the evolution of the extraordinary dentition of oenosaurus from the already highly specialized zahnanlage generally present in derived rhynchocephalians demonstrates an unexpected evolutionary plasticity of these animals . together with other lines of evidence , this seriously casts doubts on the assumption that rhynchocephalians are a conservative and adaptively inferior lineage . furthermore , the new taxon underlines the high morphological and ecological diversity of rhynchocephalians in the latest jurassic of europe , just before the decline of this lineage on this continent . thus , selection pressure by radiating squamates or mesozoic mammals alone might not be sufficient to explain the demise of the clade in the late mesozoic , and climate change in the course of the fragmentation of the supercontinent of pangaea might have played a major role .
citation : rauhut owm , heyng am , lópez - arbarello a , hecker a ( 2012 ) a new rhynchocephalian from the late jurassic of germany with a dentition that is unique amongst tetrapods . plos one 7 ( 10 ) : e46839 . urltoken
editor : andrew a . farke , raymond m . alf museum of paleontology , united states of america
copyright : © 2012 rauhut et al . this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
the recent genus sphenodon is the only living representative of the rhynchocephalia , the sister taxon of the squamata ( lizards and snakes ) within the lepidosauria [ 1 ] . due to the long history of the group , reaching back to at least the middle triassic ( ∼235 million years ) , and the supposedly primitive morphology of the genus , sphenodon is often regarded as a “living fossil” . thus , this taxon is frequently used as a model organism for a basal diapsid in investigations dealing with the evolution of reptiles , even in recent studies ranging from the structure of the pineal organ [ 2 ] to tetrapod locomotion [ 3 ] .
rhynchocephalians were a common component of mesozoic small vertebrate faunas [ 4 ] , [ 5 ] , and research in the past thirty years has shown that they were not only taxonomically and ecologically diverse during this time , but also showed a remarkable morphological variability [ 6 ] . indeed , several of the features that were thought to represent the plesiomorphic condition in sphenodon , such as the closed lower temporal arcade , were recently demonstrated to be apomorphic reversals to a condition resembling the ancestral morphology instead ( e . g . [ 7 ] , [ 8 ] ) . nevertheless , all rhynchocephalians but the most basal forms are characterized by a very specialized dentition pattern , in which the teeth are fully acrodont , no tooth replacement is present in post - hatchling individuals , and additional teeth are added at the posterior end of the tooth row during ontogeny [ 9 ] . although tooth shape itself is quite variable in rhynchocephalians [ 10 ] , this special type of tooth development was thought to limit their adaptive potential [ 11 ] . consequently , the demise of the group in the later mesozoic has been linked to the adaptive radiation of squamates and / or mammals ( see [ 5 ] ) .
here we report on a new taxon of rhynchocephalian from the late jurassic of southern germany that further underlines the morphological and ecological diversity of rhynchocephalians and indicates high adaptive plasticity even in the dentitions of these animals . the specimen was found in the middle parts of the moernsheim formation exposed at mühlheim ( fig . 1 ) , in a section of platy , siliceous limestones overlying lime bank b - h - 5 ( “krebs - bank” ) . the marine tithonian ( upper jurassic ) moernsheim formation covers the famous solnhofen lithographic limestones in the western region of the franconian alb in northern bavaria . in contrast to the underlying solnhofen formation , the sediments of the moernsheim formation are more heterogeneous , consisting of mainly siliceous limestones in alternate bedding with marly and limey sections , and with intercalated banks particularly in the lower parts of the formation .
the data matrix was analysed with paup 4 . 0b10 ( swofford 2003 ) , using a branch and bound search . the initial analyses resulted in 119 trees with a length of 187 steps ( ci 0 . 465 , ri 0 . 649 , rc 0 . 302 ) . the strict consensus tree of these trees showed poor resolution , with oenosaurus representing a member of a polytomy of rhynchocephalians more derived than planocephalosaurus ( information s1 ) ; only pleurosaurs and eilenodontines showed up as monophyletic clades within this polytomy . however , reduced consensus trees of these 119 trees recovered a monophyletic sphenodontinae , including oenosaurus , but without further resolution within this clade ( information s1 ) .
the electronic edition of this article conforms to the requirements of the amended international code of zoological nomenclature , and hence the new names contained herein are available under that code from the electronic edition of this article . this published work and the nomenclatural acts it contains have been registered in zoobank , the online registration system for the iczn . the zoobank lsids ( life science identifiers ) can be resolved and the associated information viewed through any standard web browser by appending the lsid to the prefix “ urltoken ” . the lsid for this publication is : urn : lsid : zoobank . org : pub : 4a00d064 - a136 - 4944 - af8e - 427b941fb38c . the electronic edition of this work was published in a journal with an issn , and has been archived and is available from the following digital repositories : pubmed central , lockss .
urn : lsid : zoobank . org : act : 48c1ba32 - bc6d - 407d - 81e0 - 7ecff22d77a3
urn : lsid : zoobank . org : act : 6648c44f - fdc0 - 48b2 - a6a9 - 399b17e07d38
oenos , [ oinos ] greek , wine , referring to the franconian alb , a famous wine area , where the specimen was found , and saurus , [ sauros ] greek , lizard . species name refers to the village of mühlheim , close to mörnsheim , at the rim of the valley of the altmühl in bavaria .
bayerische staatssammlung für paläontologie und geologie , munich , germany , bspg 2009 i 23 ; partial skull and complete mandibles ( figs . 2 , 3 , 4 ) .
( a ) –skull in dorsal view . ( b ) –skull in ventral view . ( c–d ) –right mandible in lateral ( c ) and medial ( d ) views . ( e ) –left mandible in lateral view . ( f ) –stereophotographs of the skull in ventral view . abbreviations : a , angular ; ar , articular ; bsp , basisphenoid ; bt , basal tubera ; co , coronoid ; cos , coronoid shelf ; d , dentary ; ect , ectopterygoid ; eo , exoccipital / opisthotic ; f , frontal ; fo , foramen ; j , jugal ; m , maxilla ; mf , mandibular foramen ; mg , meckelian groove ; oc , occipital condyle ; pa , parietal ; pal , palatine ; po , postorbital ; pof , postfrontal ; prf , prefrontal ; pt , pterygoid ; sa , surangular ; sy , symphysis ; tp , tooth plate . scale bars are 10 mm .
( a ) –left maxillary tooth plate in ventral view ( anterior is to the left ) . ( b ) –enlargement of posterior part of left maxillary tooth plate , showing compound structure of the plate . ( c ) –further enlargement of left maxillary tooth plate , showing closely packed , worn teeth with central pulp cavities . scale bars are 1 mm ( a , b ) and 0 . 5 mm ( c ) .
posterior is to the right . abbreviations : bc , basal cavity ; f , foramen . small arrows point to bifurcating dentine channels . scale bar is 5 mm .
mörnsheim formation ( lower tithonian ) [ 16 ] , “krautworst naturstein” quarry in mühlheim near mörnsheim , central bavaria , germany ( fig . 1 ) .
small rhynchocephalian with the following autapomorphic characters : maxilla with a medial process at the posterior end ; ectopterygoid with a secondary lateral process that contacts the medial side of the maxilla ; palatines with a midline contact in ventral view ; strongly pronounced lateral longitudinal groove on the dentaries , housing several large foramina ; very high coronoid process , the anterior border of which forms an angle of approximately 90° with the tooth row ; coronoid with a pronounced shoulder medially ; dentition composed of extensive tooth plates formed by a multitude of fused , small , pencil - like teeth .
the skull of oenosaurus was preserved in ventral view , with the mandibles slightly displaced from the upper jaws . the skull is diagenetically dorsoventrally compressed and much of the dorsal skull roof is missing , as is the roof of the braincase . however , enough is preserved to record the general shape of the skull and the position and size of the orbitae ( fig . 2a ) .
the ectopterygoid attaches anterolaterally to the pterygoid wing . it is unusual in that it has two lateral processes , one that contacts the posterior end of the maxilla , as in other rhynchocephalians , and a more anterior one that extends anterolaterally lateral to the palatine and contacts the medial side of the maxilla ( fig . 2b , f ) . this process forms the posterior border of the suborbital fenestra , which is surrounded by the palatine , maxilla , and ectopterygoid . thus , the pterygoid is excluded from the rim of this fenestra , as in many other sphenodontids , but not more basal rhynchocephalians . in contrast to clevosaurs [ 21 ] , [ 22 ] , [ 23 ] there is no contact between the palatine and the ectopterygoid lateral to the fenestra . together with the medial process of the maxilla , the secondary anterolateral process frames a large foramen lateral to the palatine ridge .
only the floor of the braincase and the ventral parts of the occiput are preserved . the basisphenoid is narrow at the basicranial articulation , but rapidly widens posteriorly . the basipterygoid processes seem to be stout and short , but are mostly hidden by the pterygoids . the basisphenoid has a broad , shallow depression on the ventral side between the basipterygoid processes and the low , widely separated basal tubera . the occipital condyle is broad and not separated from the basioccipital body by a constricted neck , similar to the situation in the modern sphenodon .
the lower jaw is 33 mm long and is especially notable for its high coronoid process , which exceeds the tooth row by approximately 8 . 7 mm in the right mandible and is thus almost 1 . 5 times higher than the body of the dentary ( fig . 2c , d ) . the dentary accounts for most of the length of the mandible , and , as in all rhynchocephalians , a posteroventral process of the dentary reaches posteriorly beyond the coronoid process to the level of the anterior margin of the mandibular articulation . a large incision in the posterodorsal margin of the dentary marks the enlarged mandibular foramen ( fig . 2c , e ) . a pronounced groove with several large foramina is present on the lateral side of the dentary , and its dorsal margin seems to be formed by a secondary bone skirt [ 5 ] , [ 23 ] , which probably accounts at least partially for the strong transverse thickening of the tooth - bearing part of the dentary . anteriorly , the ventral margin of the bone extends ventrally to form a small , triangular “chin” , as it is present in many rhynchocephalians ( e . g . [ 20 ] , [ 24 ] – [ 26 ] ) . there is no splenial .
the postdentary bones are entirely restricted to the posterior half of the mandible . the surangular makes up the posterior margin of the ventral part of the coronoid process and continues posteriorly to the end of the mandible . the mandibular articulation is developed as a longitudinal dorsal ridge on the articular . the ridge is sharp - edged dorsally and slightly displaced to the lateral side . medially , it is flanked by a slightly anteroposteriorly concave medial bulge that becomes more pronounced posteriorly . this morphology is similar to the condition in derived rhynchocephalians , possibly indicating a propalinal movement of the lower jaw . the retroarticular process is short and stout . the medial side of the coronoid is thickened and forms a pronounced shelf below the tip of the coronoid process .
the most unusual character of the new taxon is the dentition . unlike the situation in any other rhynchocephalian , broad tooth plates are present in both the maxillaries and dentaries . the maxillary tooth plates ( fig . 2b , f , 3a ) are broad posteriorly and narrow in their anterior half . they are 13 mm long and 4 mm wide at their widest part . the lateral edge of the tooth plates is slightly raised and shows a few small bumps . the dentary tooth plates are elongate and oval to subrectangular in outline , with a length of 13 mm and a maximum width of 3 . 5 mm . ct data shows that the central part of the tooth plate is placed in a broad depression on the dorsal surface of the mandible , though the tooth implantation can still be regarded as acrodont rather than protothecodont , as it is the case in some basal rhynchocephalians [ 27 ] .
under closer inspection , the surfaces of the tooth plates show small , but clearly defined , round , oval , or angular subunits , made up of concentric dentine layers with a small central cavity ( fig . 3 ) . fine striations radiate outwards from the central cavity . micro ct - images of the dentary tooth plate show that these dentine tubes extend over the entire height of the plate ( fig . 4 ) . in the basal part , the central cavities become larger and form large , elongate cavities at the base of the tooth plate . in some instances , the central cavities of the dentine tubes bifurcate in their course from the base to the occlusal surface . tooth enamel seems to be absent , but the dentine tubes are tightly cemented together towards the occlusal surface . at the occlusal surface , the central cavities of the tubes also seem to be filled with cement . the dentine tubes are generally somewhat larger in the central parts of the tooth plate , but become smaller towards the margins ( fig . 3a ) . the dentine tubes reach further ventrally in the anterior part of the dentary than in the posterior part , which might indicate a later addition of posterior dentine tubes during ontogeny , as it is the case with individual teeth in other rhynchocephalians [ 9 ] .
despite its apomorphic cranial morphology and highly unusual dentition , cladistic analysis ( see information s1 ) places oenosaurus well within the rhynchocephalia ( fig . 5 ) , because the taxon shows numerous synapomorphies of this clade and more exclusive ingroups , such s sphenodontia and sphenodontidae . these characters include a pronounced ridge or tooth row laterally on the palatine , a high coronoid process in the mandible , a posterior process of the dentary that extends posterior to the coronoid process , an enlarged mandibular foramen , the lack of a splenial , and an acrodont dentition [ 18 ] , [ 22 ] , [ 26 ] , [ 28 ] – [ 30 ] . within rhynchocephalians , oenosaurus even falls within the clade that includes the recent form , the sphenodontinae [ 5 ] , [ 24 ] , though support for this placement is weak ( see information s1 ) , and further material of this taxon ( and others , which , with the exception of sphenodon , are generally poorly known [ 5 ] ) might lead to changes in this part of the tree . nevertheless , the phylogenetic position of the new taxon within sphenodontidae ( sensu [ 24 ] ) is well supported .
cladistic analysis of 70 osteological characters in 19 rhynchocephalian and two outgroup taxa resulted in the recovery of 2 trees with 184 steps ( see information s1 ) . a strict consensus tree shows oenosaurus well nested within sphenodontine rhynchocephalians , as a close relative to the recent sphenodon . 1 , rhynchocephalia ; 2 , sphenodontinae .
an interesting problem is the evolution of these tooth plates . the phylogenetic position of oenosaurus shows that this taxon is well nested within taxa that have the specialized acrodont dentition typical for sphenodontids . the very peculiar plates are therefore derived from such a dentition and demonstrate a surprising evolutionary plasticity of such a seemingly highly specialized tooth anlage [ 9 ] . a possible , but currently untestable , hypothesis of the evolution of these tooth plates might be that they formed by fusion and modification of the hatchling dentition , which consists of small , peg - like teeth with active tooth replacement in several rhynchocephalians [ 41 ] , including the modern sphenodon [ 42 ] , [ 43 ] . however , further studies and , possibly , further discoveries of this or other taxa with similar dentitions are necessary to test this hypothesis .
despite their specialized dentition , rhynchocephalians are quite variable in their tooth morphology . rhynchocephalian teeth can be referred to three basic functional types [ 10 ] : basal forms usually have small , conical teeth that are mainly used for piercing , most sphenodontids have anteroposteriorly elongate teeth that are used for cutting and slicing , and opisthodontians [ 18 ] have transversely expanded teeth capable of grinding and shredding . these different tooth types can roughly be equated with insectivory , generalized carnivory ( or piscivory , in the marine pleurosaurs ) , and herbivory , respectively [ 10 ] . the tooth plates of oenosaurus now demonstrate a further type of dentition in rhynchocephalians , a crushing dentition , indicating a durophagous diet . therefore , the new taxon adds a further trophic adaptation and a previously unknown ecotype to the already recognized ecological diversity of mesozoic rhynchocephalians [ 10 ] .
character list , data matrix , analytical procedures , consensus trees , and list of synapomorphies at internal nodes for the cladistic analysis mentioned in the text .
we thank roland pöschl , who found the fossil , and the owners and operators of the krautworst naturstein quarry , ulrich leonhardt , roland pöschl and uwe krautworst , for donating the specimen to the bspg . ulrich leonhardt , stefan sónyi , and renate liebreich are thanked for preparation of the fossil . the paper benefited from discussions with marc jones and alexander nützel . furthermore , critical comments by marc jones and nick fraser considerably helped to improve the paper . ct scans were carried out at the steinmann institute of the university of bonn with the help of irina ruf , which is greatly appreciated .
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oliver w . m . rauhut , 1 , 2 , * alexander m . heyng , 2 adriana lópez - arbarello , 1 and andreas hecker 3
* e - mail : ed . nehcneum - inu . zrl @ tuhuar . o
this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are properly credited .
here we report on a new taxon of rhynchocephalian from the late jurassic of southern germany that further underlines the morphological and ecological diversity of rhynchocephalians and indicates high adaptive plasticity even in the dentitions of these animals . the specimen was found in the middle parts of the moernsheim formation exposed at mühlheim (
) , in a section of platy , siliceous limestones overlying lime bank b - h - 5 ( “krebs - bank” ) . the marine tithonian ( upper jurassic ) moernsheim formation covers the famous solnhofen lithographic limestones in the western region of the franconian alb in northern bavaria . in contrast to the underlying solnhofen formation , the sediments of the moernsheim formation are more heterogeneous , consisting of mainly siliceous limestones in alternate bedding with marly and limey sections , and with intercalated banks particularly in the lower parts of the formation .
) is considerably better resolved than that resulting from the full analysis , but , importantly , it does not contradict the results of the latter , neither in the strict consensus , nor in any reduced consensus tree . thus , we chose these results to illustrate and further explore the relationships of oenosaurus . bootstrap ( 10000 replicates ) and bremer support analyses were carried out on the pruned matrix , and the following list of synapomorphies is based on the strict consensus tree resulting from the pruned analysis .
was preserved in ventral view , with the mandibles slightly displaced from the upper jaws . the skull is diagenetically dorsoventrally compressed and much of the dorsal skull roof is missing , as is the roof of the braincase . however , enough is preserved to record the general shape of the skull and the position and size of the orbitae (
the skull seems to have been rather robust . as preserved , it is 24 . 3 mm long . unfortunately , the exact skull length cannot be determined , since the premaxillae are missing , but these bones probably did not account for more than an additional 2–3 mm . thus , the skull was slightly broader ( 28 mm ) than long , as in many rhynchocephalians , but unlike the slender , elongate skull of the marine pleurosaurs
. premaxillae and nasals are missing . the frontals are elongate and narrow , and flanked by the prefrontals anterolaterally (
) . there is no trace of a lacrimal , though it cannot be ruled out completely that this might be due to preservation . the posterior part of the skull roof is poorly preserved and nothing can be said about the shape and size of the temporal fenestrae . a small fragment of the parietals shows that these bones were fused and formed a narrow sagittal crest medially . the postfrontals and postorbitals formed a broad shelf anterior to the dorsal temporal fenestra . the jugal was broad and reached anteriorly to almost half - length of the orbit . the maxillae are massive and broad , but rapidly narrow anteriorly . posteriorly , the maxilla has a stout lateral process for the contact with the jugal and a smaller medial process that slots into the forked lateral process of the ectopterygoid .
) . the vomers were small . their posterior ends separate the anterior ends of the palatines , but do not reach the pterygoids posteriorly , so that the palatines meet at the midline . such a midline contact of the palatines was illustrated for the marine pleurosaurs by carroll and wild
, but not described in the text , and the line of contact is dotted in their reconstruction , so that some uncertainty of the condition in these animals remains . the palatines contact the maxillae laterally in a broad suture at about one third of the length of the latter bone , and form the posterior border of the internal choanae . the bones flank the pterygoids laterally over most of the length of the anterior process of the latter bone , taper posteriorly , and end a short way posterior to the posterior end of the maxillae . as in all sphenodontids , there is a raised ridge along the lateral edge of the posterior half of the palatines , but , due to preservation , it is uncertain if this ridge bore teeth , as it is the case in most taxa
, the ridge diverges from the maxillary posteriorly and is more or less parallel to the midline of the skull . the pterygoids taper anteriorly and broaden abruptly posteriorly towards the ventrolaterally extending pterygoid wings . there is no interpterygoid vacuity , but the pterygoids form a midline suture up to the basicranial articulation . posteriorly , the slender , but well - developed , quadrate wings of the pterygoids diverge away from the ventral side of the braincase . a small fragment of bone lateral to the remnants of the left quadrate wing of the pterygoid might represent a remain of the pterygoid wing of the quadrate . unlike basal rhynchocephalians , there are no teeth on the pterygoid . no suture between the quadrate wing of the pterygoid and the pterygoid wing of the quadrate is visible in the preserved portions , so the latter bone , which is not preserved , might have had a rather short pterygoid wing .
the ectopterygoid attaches anterolaterally to the pterygoid wing . it is unusual in that it has two lateral processes , one that contacts the posterior end of the maxilla , as in other rhynchocephalians , and a more anterior one that extends anterolaterally lateral to the palatine and contacts the medial side of the maxilla (
) . this process forms the posterior border of the suborbital fenestra , which is surrounded by the palatine , maxilla , and ectopterygoid . thus , the pterygoid is excluded from the rim of this fenestra , as in many other sphenodontids , but not more basal rhynchocephalians . in contrast to clevosaurs
there is no contact between the palatine and the ectopterygoid lateral to the fenestra . together with the medial process of the maxilla , the secondary anterolateral process frames a large foramen lateral to the palatine ridge .
the lower jaw is 33 mm long and is especially notable for its high coronoid process , which exceeds the tooth row by approximately 8 . 7 mm in the right mandible and is thus almost 1 . 5 times higher than the body of the dentary (
) . the dentary accounts for most of the length of the mandible , and , as in all rhynchocephalians , a posteroventral process of the dentary reaches posteriorly beyond the coronoid process to the level of the anterior margin of the mandibular articulation . a large incision in the posterodorsal margin of the dentary marks the enlarged mandibular foramen (