orectoloboides is an extinct genus of wobbegong sharks ( family orectolobidae ) .

it was described by cappetta in 1977 .

a new species , o. angulatus , was described from the cenomanian period of canada by charlie j. underwood and stephen l. cumbaa in 2010 .

orectoloboides

orectoloboides <EOT> copyright © 1999 - 2018 john wiley & sons , inc . all rights reserved

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eugomphodus winkleri , odontaspis aff . winkleri , odontaspis cf . winkleri , odontaspis ( synodontaspis ) aff . winkleri , odontaspis ( synodontaspis ) winkleri , synodontaspis winkleri

gigantichthys pharao , ischyrhiza iwakiensis , onchosaurus cf . pharao , onchosaurus iwakiensis , onchosaurus ( onchosaurus ) pharao , titanichthys pharao

ctenacanthus clarki , ctenacanthus clarkii , ctenacanthus compressus , didymodus compressus , diplodus compressus , dittodus compressus , orthacanthus aff . compressus , orthacanthus cf . compressus , pleuracanthus ( diplodus ) compressus , xenacanthus cf . compressus , xenacanthus compressus

diacranodus texensis , didymodus texensis , orthacanthus ( lebachacanthus ) texensis , xenacanthus aff . texensis , xenacanthus texensis

carcharodon obliquus , lamna obliqua , lamna obliquus , odontaspis taurus obliqua , otodus cf . obliquus , otodus giganteus , otodus plicatilis

carcharocles auriculatus sokolowi , carcharocles cf . sokolowi , carcharocles sokolovi , carcharocles sokolowi , carcharodon angustidens sokolowi , carcharodon sokolovi , carcharodon sokolowi , otodus cf . sokolovi , otodus cf . sokolowi , otodus sokolovi sokolovi , otodus ( carcharocles ) cf . sokolowi , procarcharodon angustidens cf . sokolowi , procarcharodon cf . sokolowi , procarcharodon sokolovi , procarcharodon sokolowi

carcharocles aff . angustidens , carcharocles angustidens , carcharocles angustidens turgidus , carcharocles cf . angustidens , carcharocles megalodon angustidens , carcharocles rectus , carcharocles turgidus , carcharodon aff . angustidens , carcharodon aff . angustidens turgidus , carcharodon aff . turgidus , carcharodon angustidens , carcharodon angustidens angustidens , carcharodon angustidens turgidus , carcharodon arndti , carcharodon cf . angustidens , carcharodon cf . praemegalodon , carcharodon cf . turgidus , carcharodon lanceolatus , carcharodon longidens , carcharodon megalodon robustus , carcharodon praemegalodon , carcharodon rectus , carcharodon robustus , carcharodon simus , carcharodon turgidus , otodus angustidens , otodus angustidens angustidens , otodus angustidens turgidus , otodus arndti , otodus lanceolatus , otodus praemegalodon , otodus rectus , otodus robustus , otodus turgidus , procarcharodon angustidens , procarcharodon angustidens angustidens , procarcharodon cf . angustidens , procarcharodon praemegalodon , procarcharodon turgidus

carcharias giganteus , carcharias grosseserratus , carcharias incidens , carcharias macrodon , carcharias megalodon , carcharias mexicanus , carcharias polygurus , carcharias polygyrus , carcharias productus , carcharias ( prionodon ) incidens , carcharocles aff . megalodon , carcharocles aff . subauriculatus , carcharocles megalodon , carcharocles megalodon megalodon , carcharocles productus , carcharodon arcuatus , carcharodon branneri , carcharodon brevis , carcharodon costae , carcharodon crassidens , carcharodon crassirhadix , carcharodon crassus , carcharodon gibbesi , carcharodon gigas , carcharodon helveticus , carcharodon humilis , carcharodon intermedius , carcharodon latissimus , carcharodon leviathan , carcharodon megalodon , carcharodon megalodon indica , carcharodon megalodon megalodon , carcharodon megalodon polygyra , carcharodon megalodon productus , carcharodon megalodon siculus , carcharodon megalodon yamanarii , carcharodon morricei , carcharodon polygurus , carcharodon polygyrus , carcharodon productus , carcharodon quenstedti , carcharodon rectidens , carcharodon rectideus , carcharodon semiserratus , carcharodon subauriculatus , carcharodon tumidissimus , carcharodon turicensis , megaselachus arcuatus , megaselachus auriculatus falciformis , megaselachus branneri , megaselachus brevis , megaselachus crassidens , megaselachus crassirhadix , megaselachus crassus , megaselachus gigas , megaselachus heterodon , megaselachus humilis , megaselachus incidens , megaselachus leviathan , megaselachus megalodon , megaselachus megalodon indicus , megaselachus polygyrus , megaselachus productus , megaselachus rectidens , megaselachus semiserratus , megaselachus subauriculatus , otodus megalodon , procarcharodon aff . megalodon , procarcharodon megalodon , procarcharodon megalodon megalodon , selache manzonii

host - parasite list / parasite - host list ( version : 01 . 04 . 2015 ) 544 pp , 5 , 37 mb new !

( pdf ) new fossil species of somniosus and rhinoscymnus ( squaliformes : somniosidae ) , deep water sharks from oligocene rocks of western washington state , usa .

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sullivan , r . m . and lucas , s . g . , eds . , 2016 , fossil record 5 . new mexico museum of natural history and science bulletin 74 .

keasey formation crinoid lagerstätte at mist , oregon , and the kirker sandstone at mt . diablo , california . the

sandstone near mt . diablo , california ( welton , 1979 , p . 474 ) were

years ( adnet and cappetta , 2001 ; adnet et al . , 2006 ; t

and adnet et al . ( 2006 ) suggested the chronologic range of the genus

modied from niem et al . ( 1994 ) , and burns et al . ( 2005 ) .

, adnet , et al , 2006 , p . 63 - 64 , designated

, adnet et al . , 2008 , p . 715 - 716 , notes genus invalid ;

large lower teeth ( maximum tooth height 7 . 95 mm ) ; differing from all

et sp . nov . , type and referred specimens were collected . nmmnh and

– biostratigraphic time scales and zonation after berggren et al . ( 1995 ) ,

, oregon . ( a ) , labial view ; ( b ) , lingual view .

upper anteroposterior ( figs . 4 - 5 , 7 ) and lower medial ( fig . 3 ) and anteroposterior teeth ( figs .

6 , 8 - 9 ) , from early oligocene kirker sandstone at mt . diablo , contra costa county , california ( lacm locality 4304 ; figs . 1 - 5 ) , and early

oligocene or latest eocene keasey formation at mist ( nmmnh locality 9105 and lacm locality 4308 ) , columbia county , oregon ( figs . 6 - 9 ) .

) views , selected to represent relative row positions along the dental series . illustrated tooth positions m1 , ap2 ,

right side of the dentition , and have been digitally reversed to mimic teeth from the left side of the jaw . m1 , lacm 1

okes , 1953 ; burns et al . , 2005 ) , earliest oligocene ( hickman ,

al . , 1985 ) , correlative with chron c12r ( 31 . 0 to 33 . 0 ma ; prothero and

stage and the whitneyan north american land - mammal “age” ( fig . 4 ) .

junction , near the town of pittsburg , columbia county oregon ( fig . 1 ) .

referred teeth from the keasey formation ( fig . 7 . 6 - 9 )

oligocene , upper refugian to lower zemorrian ( tipton et al . , 1973 ) ,

holotype and paratype teeth ( fig . 8 ) . the dental series includes medial

distal in the dental series . there is a general decrease in tooth height /

the jaw ( bigelow and schroeder , 1948 ; bass et al . , 1976 ; herman et

8 - ap2 ) . the tooth is 5 . 65 mm high and 3 . 82 mm wide ( rw = tw ) , with

robust , short triangular cusp , inclined distally at an angle of 21 degrees .

and the cusp apex does not extend distally beyond the cusp - heel notch .

trough , and the basal edge is irregular and horizontal . the root is robust ,

the adjacent distal tooth , covers 4 / 5ths of the root height . a

a point , just above a very large upper axial foramen . mesial and distal

by a thin bridge of the lingual root face . mesial labial and distal labial

anteroposterior teeth ( ap1 - ap9 ; figs . 5 . 2 - 5 , 6 . 1 - 5 , 8 ) :

ap ( 1 . 75 - 1 . 36 ) , then remain in the 1 . 3 range

but is estimated to have been wider than high ( 0 . 8 + ) . cusp inclination

is about 1 / 2 the root height . labially , the apron is axially narrow and

developed close to the crown base , or on the distal root face . the mesial

anteroposterior teeth are small ( th 3 . 7 mm and rw = 3 . 0 mm in lacm

ab . 1 ) , approximately 50 to 60 % of the lower anteroposteriors .

in height ( missing mesial and distal root lobes ) . the tooth has a robust

a narrow , short , non - bifurcated apron . enough of the root is preserved

labial groove , and root lobes are unknown . lingually , there is a small ,

anteroposterior tooth ( e . g . , root lobes , lower axial foramen , labial

doubt that these teeth , as suggested by adnet et al . ( 2006 ) , and v

crown gured by vialle et al . ( 2011 , g . 2 - 5 ) compares

the intersection of the distal heel and cusp . in labial view , the smaller

around the apical rim of a large lower axial foramen . there is a very

lee on the solent , hampshire basin , england ( adnet et al . , 2006 , p .

clearly a dalatiid shark ( cappetta , 2006 ; adnet et al . , 2006 ) , however

distal labial pit and foramen . the tooth gured by kemp ( 1994 ) differs

notch formed by the intersection of the distal heel and cusp . labially , the

distal , and concave mesial cusp cutting edges , and a more lobate apron .

to establish the identity of this tooth . the very broad - based , erect ,

communication ) . according to heilmann - clausen et al . ( 1985 ) the

point ( molina et al , 2011 ; schnetler and heilmann - clausen , 2011 ) .

ranch no . 1 well , at a depth between 6206 - 6215 feet ( 1892 - 1894 m ) .

and sandstones of the pittsburg bluff formation ( niem et al . , 1994 )

formation ( mcknight et al . , 1992 ; niem et al . , 1994 ) . near the town

mid - shelf depths , and warm - temperate waters ( moore , 1976 ) .

approximately 15 miles ( 25 km ) to the northwest of the type locality ( fig .

okes , 1953 ; burns and mooi , 2003 ; burns et al . , 2005 ) . in addition to

non - calcareous , organic - rich clay ( heilmann - clausen et al . , 1985 ) .

related to periodic low oxygen bottom conditions ( burns et al . , 2005 ;

4e - h ) , from the labguva formation , upper cenomanian , of lithuania .

dental characters : lethaia , v . 34 , p . 234 - 248 .

adnet , s . , cappetta , h . and mertiniene , r . , 2008 , re - evaluation of squaloid

adnet , s . , cappetta , h . and reynders , j . , 2006 , nouveaux genres de squaliformes

, j . d . and kistnasamy , n . , 1976 , sharks of the east coast

paleontologists and mineralogists special publication , v . 54 , p . 129 - 212 .

burns , c . , campbell , k . a . and mooi , r . , 2005 , exceptional crinoid occurrences

burns , c . and mooi , r . , 2003 , an overview of eocene - oligocene echinoderm

cappetta , h . , 2006 , elasmobranchii post - triadici ( index specierumet generum ) .

palaeoecology : bulletin of the geological society of denmark , v . 62 , p .

heilmann - clausen , c . , nielsen , o . b . and gersner , f

denmark : bulletin of the geological society of denmark , v . 33 , p . 287 -

herman , j . , hovestadt - euler , m . and hovestadt , d . c . , 1989 , contributions

stehmann , m . , ed . , part a : selachii . no . 3 : order : squaliformes , families :

sciences naturelles de belgique , v . 59 , p . 101 - 157 .

a . and ivany , l . , eds . , from greenhouse to icehouse : the marine eocene–

johnson , h . d . and baldwin , c . t . , 1986 , shallow siliciclastic seas :

scientic publications , alden press , oxford , england , p . 229 - 282 .

. and moore , g . w . , 1971 , stratigraphic relations of upper

turell , j . , hardenbol , j . , heilmann - clausen , c . , larrasoaña , j . c . ,

luterbacher , h . , monechi , s . , ortiz , s . , orue - etxebarria , x . , payros ,

lutetian stage at the gorrondatxe section , spain : episodes v . 34 , p . 86 - 108 .

oregon [ m . s . thesis ] : corvallis , oregon state university , p . 1 - 370 .

, h . j . and campbell , k . a . , 1994 ,

county , northwest oregon : unpublished m . s . thesis , corvallis , oregon

prothero , d . r . and hankins , k . g . , 2001a , magnetic stratigraphy and tectonic

mineralogists ( society for sedimentary geology ) book 91 , p . 201 - 209 .

prothero , d . r . and hankins , k . g . , 2001b . magnetic stratigraphy and tectonic

journal of geophysical research , v . 105 , p . 16 , 473 - 16 , 480 .

publications in geological sciences , v . 16 , p . 449 - 460 .

schnetler , k . i and heilmann - clausen , c . , 2011 ,

signicance : the paleontological society of japan , v . 81 , p . 24 - 44 .

underwood , c . j . and schlogl , j . , 2013 , deep - water chondrichthyans from the

w . c . and norbisrath , h . , 1946 , stratigraphy of the upper nehalem

united states and british columbia : paleobios , n . 17 , p . 1 - 15 .

from the late eocene of western oregon , u . s . a . , and description of the

welton , b . j . and goedert , j . l . , 2016 , new fossil species of

welton , b . j . , 2016 , a new dalatiid shark ( squaliformes : dalatiidae ) from the early oligocene of oregon and california , usa ; in sullivan , r . m . and lucas , s . g . , eds . , new mexico museum of natural history and science bulletin 74 , p . 289 - 302 .

first report of orthechinorhinus ( squaliformes : etmopteridae ) from the pacific basin ; a new species from early oligocene rocks of oregon , usa ; in sullivan , r . m . and lucas , s . g . , eds . , fossil record 5 : new mexico museum of natural history and science bulletin 74 , p . 303 - 308 .

welton and goedert 2016 new species of somniosus and rhinoscymnus from oligocene of western washingt . . .

first report of orthechinorhinus ( squaliformes : etmopteridae ) from the pacific basin ; a new species . . .

abstract—a decade ago , adnet provisionally placed the genus orthechinorhinus in the echinorhinidae incertae sedis based on its heterodonty and a number of shared general crown and root morphologies , while also noting significant differences in root vascularization . orthechinorhinus possesses a very specialized dentition , unlike that of any known echinorhinid , and is interpreted here to be a . . . [ show full abstract ]

welton , b . j . , 2016 , a new dalatiid shark ( squaliformes : dalatiidae ) from the early oligocene of ore . . .

abstract—isolated teeth of oligodalatias jordani , a new genus and species of extinct dalatiid shark , are described from early oligocene marine deposits of the pittsburg bluff formation , nehalem river basin , northwestern oregon . o . jordani is also known from other eastern north pacific early oligocene marine rocks , including the keasey formation crinoid lagerstätte at mist , oregon , and the . . . [ show full abstract ]

welton , b . j . , 2014 , a new fossil basking shark ( lamniformes : cetorhinidae ) from the middle miocene . . .

abstract . isolated teeth of a middle miocene cetorhinid genus , cetorhinus blainville 1816 , occur abundantly in rocks of the round mountain silt , sharktooth hill bonebed , southeastern san joaquin valley , kern county , california . tooth sets of juvenile and adult dentitions of the sharktooth hill cetorhinus were reconstructed and used as a basis for description of a new species , c . huddlestoni . . . . [ show full abstract ]

welton , b . j . , 2015 , a new species of late early miocene cetorhinus ( lamniformes : cetorhinidae ) from . . .

abstract . microphagous lamniforms of the family cetorhinidae have a significant cenozoic history in the north pacific ocean . the late eocene keasius taylori occurs in the keasey formation of oregon , and k . parvus may occur in the oligocene lincoln creek formation of southwestern washington . the genus cetorhinus has one extant species , c . maximus , and a fossil record , including the middle . . . [ show full abstract ]

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images are presented of fossilised teeth from most of the currently known shark families that can be found in the lower cretaceous , middle and late albian ( gault clay ) of kent in south - east england .

fourteen families and twenty - seven genera are included . apart from the larger sharks , mainly represented by the lamniformes , most teeth are from small species and in the main have been acquired through bulk sampling and sieving of washed clay residues . indeed , most of the gault shark species are represented by tiny teeth , ranging from 1mm up to a few centimetres .

notwithstanding the diversity of the gault shark fauna , fossilised teeth are relatively uncommon and these specimens represent the very best of many years of collecting .

the help and support is gratefully acknowleged of david ward of orpington for making freely available use of his electronic imaging facilities , for help in collecting and processing bulk clay samples and with identification of specimens .


the long-tailed musk shrew ( crocidura dolichura ) is a species of mammal in the family soricidae .

it is found in burundi , cameroon , central african republic , republic of the congo , democratic republic of the congo , gabon , nigeria , and uganda .

long - tailed musk shrew

long - tailed musk shrew <EOT> the asian house shrew ( suncus murinus ) grey musk shrew , asian musk shrew , or money shrew is a widespread , adaptable species of shrew found mainly in south asia but introduced widely throughout asia and eastern africa .

it is a large shrew with a strong musk smell . it is related to the etruscan shrew .

and long - nosed . the teeth are a series of sharp points to poke holes in insect

this species is associated with both lowland and montane tropical moist forest . it is a climbing shrew that probably uses its long tail for balancing ( happold 1987 ) .

advani r , rana bd ( 1981 ) . " food of the house shrew ,

studies on this shrew have suggested its suitability for use in laboratory studies of reproduction and nutrition .

the house shrew has a uniform , short , dense fur of mid - grey to brownish - grey color . the tail is thick at the base and a bit narrower at the tip , and is covered with a few long , bristle - like hairs that are thinly scattered . they have short legs with five clawed toes . they have small external ears and an elongated snout . they also emit a strong odor of musk , derived from musk glands that are sometimes visible on each side of the body . the odor is especially noticeable during the breeding season .

the house shrew has a habit of moving quickly along the edges of the walls when it enters human habitations . as it runs it makes a chattering sound which resembles the sound of jingling money , which has earned them the name ”money shrew” in china . when alarmed , the house shrew makes an ear - piercing , high - pitched shriek , resembling the sound of nails scraping a chalkboard or a metal fork scraping glass , which repels house cats . predators also leave the house shrew alone because of its musky smell and even when they catch one by mistake they will rarely eat it .

this species is locally called chuchunder in india and is mentioned in rudyard kipling ' s jungle book , as a nocturnal inhabitant of houses in india , by the name of chuchundra . however , kipling ' s mistaken use of the name ' musk rat ' has led to confusion with the unrelated north american muskrat ( ondatra zibethicus ) , and the latter species , not found in india , was ( erroneously ) illustrated in the jungle book .

another remarkable habit of this shrew ( shared with the white - toothed shrews of europe ) is that it forms a ”caravan” with its young , that is , the young line up behind the mother and follow it while she walks . the first young will hold on to the mother ' s fur with its teeth , and the subsequent young will do the same with the sibling in front of it .

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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .

justification : listed as least concern in view of its wide distribution , presumed large population , and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category .

this species is never abundant , but it is widely distributed . the species is infrequently found in surveys .

there is some localised habitat loss over much of this species range . in eastern parts of the species range , people displaced by social instability have established settlements within the forest .

this species has been recorded from several protected areas including the kongana forest in southern central african republic , korup national park in cameroon and kibira national park in burundi .

this errata assessment has been created because the map was accidentally left out of the version published previously .

( errata version published in 2017 ) . the iucn red list of threatened species 2016 : e . t40628a115176367 .

to make use of this information , please check the < terms of use > .

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confused by a class within a class or an order within an order ? please see our brief essay .

to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .

disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .

this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .

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mammal species of the world : a taxonomic and geographic reference , 2nd ed . , 3rd printing

mammal species of the world : a taxonomic and geographic reference , 3rd ed . , vols . 1 & 2

with contributions by bernadette n . graham , adam p . potter , and mariana m . upmeyer

disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .

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has a wide distribution throughout the old world tropics . in most of its range , it was introduced by man . according to burton and burton , it was originally native to the forests of india .

it has been introduced by man to sri lanka , eastern africa , madagascar , islands in the indian ocean ( reunion , comoros ) , pacific ocean ( guam , etc . ) , south - east asia , china , southern japan , malaysia ( peninsular malaysia , sabah , sarawak ) , kalimantan , brunei , indonesia , new guinea , and throughout iran and arabia to egypt and pakistan also .

s . murinus is a commensal species . it is a voracious insectivore with little resistance to starvation . it is active during the night , spending the day in a burrow or hiding place in human habitations . they breed throughout the year , with each female averaging two litters per year . the gestation period is one month . one to eight young are born per litter , usually three young , in a nest made by both of the parents , wherein the young stay until they are nearly adult . it starts breeding when it is around one year old .

it is widespread and found in all habitats , including deserts and human habitations .

the habitat of this species is normally near human settlement , specifically near the house . some also live on the ground in leaf litter and grass . it has been recorded up to 2825 m above sea level near

stuffed specimens , exhibited in the national museum of nature and science , tokyo , japan .

it is often mistaken for a rat or mouse and killed as vermin . in general it is beneficial to humans because its diet consists mostly of harmful insects such as cockroaches , and even house mice . it can therefore be considered as a

despite its use as an insect control , it can be unpopular due to the strong odour of its droppings , which it may deposit in human dwellings behind kitchen cupboards , etc . it can also take to eating human food such as meat in kitchens , or dog or cat food . it is known to occasionally kill young chicks , making it unpopular with farmers , although rats probably kill more chicks , and more quickly . the way it is said to attack chicks , by first biting a tendon , immobilizing it and then killing and eating it ,

could indicate that it has a venomous bite that paralyses , as at least two other shrews species have ( i . e . the

maurice burton , robert burton , international wildlife encyclopedia ” , new york 2002 pp . 1709–1710 . available on google books .

maurice burton , robert burton , international wildlife encyclopedia , new york 2002 pp . 1709–1710 ; see google books .

robert h . schmidt , “shrews” , internet center for wildlife damage management , urltoken . retrieved 12 . 5 . 2013 .

maurice burton , robert burton , international wildlife encyclopedia , new york 2002 pp . 1709–1710 ; available on google books .

rachel sheremeta pepling , “the stunning saliva of shrews , ” on chemical & engineering news website , 2004 , urltoken . dr . werner haberl , “poisonous shrews” urltoken

iucn . ( 1995 ) . eurasian insectivores and tree shrews - status survey and conservation action plan , iucn , gland , switzerland . 108pp

this article is issued from wikipedia - version of the 11 / 6 / 2016 . the text is available under the creative commons attribution / share alike but additional terms may apply for the media files .


the popoche chub ( algansea popoche ) is a species of freshwater fish in the genus algansea .

it is endemic to the lerma river basin and lake cajititlan in mexico .

it is not suitable for human consumption .

popoche chub

popoche chub <EOT> the popoche chub is a species of ray - finned fish in the genus algansea .

have a fact about popoche chub ? write it here to share it with the entire community .

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another local official , erik tapia , suggested that the deaths of the popoche chub fish might be due to overpopulation .

but a local official , ismael del toro , said the deaths of the popoche chub fish happened every year during the rainy season .

local media reported at least 40 tonnes of popoche chub freshwater fish had washed up on the shores of lake cajititlan , in jalisco state .

nearly 50 tonnes of dead popoche chub fish were removed at the weekend from lake cajititlán , a lagoon in the central state of jalisco .

a fisherman collects dead popoche chub fish from lake cajititlán , mexico . nearly 50 tonnes of dead fish were removed . photograph : hector guerrero / afp / getty images

fishermen , firefighters , town hall workers and staff from the state agricultural ministry pulled hundreds of thousands of dead popoche chub fish from the lake and buried them in a pit .

the incident comes after of a series of smaller waves of dead popoche chub in the lake in recent months , including one last week , ensuring that 2014 is already by far the worst year for the species , which has been under attack for the past few years .

froese , rainer and pauly , daniel , eds . ( 2006 ) . " algansea popoche " in fishbase . april 2006 version .

victor hugo ornelas , a reporter from the tlajomulco - based newspaper la verdad , said it also appeared that fertilisers washed into the lake from surrounding cornfields during the rainy season could be a significant factor . the fertilisers appear to fuel the growth of algae near the surface , where the popoche chub swim .

fishermen have been removing thousands of dead fish from a lake near the western mexican city of guadalajara .

one local official described their deaths as a " natural , cyclical phenomenon " that occurred every year .

however , other state and local officials have blamed poor lake management for the deaths of the fish .

the authorities are investigating whether a wastewater treatment plant is to blame for failing to filter out untreated material and thus reducing the amount of oxygen in the water .

last year local fishermen recovered about 200 tonnes of dead fish from lake cajititlan . more than 100 fishermen are involved in this year ' s operation .

" it is a natural , cyclical phenomenon . it happens every year and other species are not affected , " he insisted .

" they do not have any natural predator and they are not suitable for human consumption , " he said .

the foreign secretary quits , telling theresa may " needless self - doubt " is leading to " semi - brexit " .

latin , algensis , - e = somebody that eats algae ; cited by plinius ( ref . 45335 )

lyons , j . , g . gonzález - hernandéz , e . soto - galera and m . guzmán - arroyo , 1998 . decline of freshwater fishes and fisheries in selected drainages of west - central mexico . fisheries 23 ( 4 ) : 10 - 18 . ( ref . 27639 )

phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 5039 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .

bayesian length - weight : a = 0 . 00891 ( 0 . 00387 - 0 . 02054 ) , b = 3 . 06 ( 2 . 85 - 3 . 27 ) , in cm total length , based on lwr estimates for this ( sub ) family - body shape ( ref . 93245 ) .

trophic level ( ref . 69278 ) : 3 . 0 ±0 . 2 se ; based on size and trophs of closest relatives

resilience ( ref . 69278 ) : high , minimum population doubling time less than 15 months ( preliminary k or fecundity . ) .

vulnerability ( ref . 59153 ) : low to moderate vulnerability ( 25 of 100 ) .

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mexicans are baffled at the sudden death of thousands of fish in a lake in the centre of the country , a dramatic intensification of a problem that no one has yet been able to explain .

the authorities in the lakeside town of tlajomulco de zúñiga , about 25 minutes ' drive south of the city of guadalajara , had previously blamed the deaths on " a cyclical phenomenon caused by temperature variations and the reduction of oxygen " .

this weekend , the state ' s environment secretary , magdalena ruiz mejía , ruled out natural causes and blamed " poor management of the body of water " . she pointed to municipal waste water treatment facilities and promised a full investigation .

there have been complaints that a nearby tequila distillery is storing waste in containers that drain into channels that feed into the lake .

" it is obvious that there are many sources of pollution around the lake , " ornelas said . " fertiliser runoff in a particularly heavy rainy season could be the straw that is breaking the camel ' s back . "

josé luis castillo , who takes tourists around the lake in his boat , told the newspaper el informador how the fish first attracted attention by swimming even closer to the surface than usual . " after that they died , " he said . " lots and lots have died this year . "

the fish deaths are just the latest in a succession of incidents in which large numbers of creatures have been found dead in particular places , from sea lions in california to seabirds in peru .

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nelson , joseph s . , edwin j . crossman , h . espinosa - pérez , l . t . findley , c . r . gilbert , et al . , eds .

full author list : nelson , joseph s . , edwin j . crossman , héctor espinosa - pérez , lloyd t . findley , carter r . gilbert , robert n . lea , and james d . williams

disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .

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without warning , hundreds of thousands of dead fish have been floating to the surface of lake cajititlan in the mexican state of jalisco over the past week . local authorities claim the deaths are part of a “natural cycle , ” but state authorities disagree , saying that the phenomena is due to “poor management” of the lake . this isn’t the first time this has happened either – 10 tons of fish were found dead in the same lake last october . is this really an annual event , or yet another occurrence caused by the harmful activities of humans ?

state authorities are confident that improper management of sludge produced by local wastewater treatment plants is to blame . the only problem is that they have so far been denied access to the plants , and therefore been unable to gather any concrete evidence .

the argument in favor of natural causes is that between the months of august , september and october , a lack of oxygen in the water , caused by the drag of organic matter during the rainy season causes many of the freshwater fish in the lake to die . while this seems logical , the fact that the number has increased so dramatically since last year ( 10 tons to 50 , and counting ) points to other variables . until there is more evidence , all state authorities have been able to do is issue an environmental alert for the lake .

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brookesia stumpffi , also known as the plated leaf chameleon ( german : stachelchamäleon ) , is a species of chameleon found in some parts of madagascar .

it can be found in nosy bé , north-west madagascar , nosy komba , and nosy sakatia .

brookesia stumpffi

brookesia stumpffi <EOT> brookesia stumpffi boettger 1894 brookesia stumpffi — glaw & vences 1994 : 236 brookesia stumpffi — klaver & böhme 1997 brookesia stumpffi — necas 1999 : 268 brookesia stumpffi— pianka & vitt 2003 : 11 brookesia stumpffi — townsend et al . 2009

brookesia stumpffi has been bred for some years , although not in high amounts .

citation : - brookesia stumpffi . malaysia biodiversity information system ( mybis ) . urltoken downloaded on 10 july 2018 .

schmidt , w . 1986 . brookesia stumpffi boettger , 1894 . sauria 8 ( 2 ) suppl . : 041–042 - get paper here

remark . —this species has been considered before as brookesia sp . “montagne des francais” [ 7 ] and as brookesia sp . nov . [ 37 ] .

schmidt , w . ; henkel , f . w . & böhme , w . 1989 . zur haltung und fortpflanzungsbiologie von brookesia stumpffi boettger 1894 ( sauria : chamaeleonidae ) . salamandra 25 ( 1 ) : 14 - 20 - get paper here

remark . —this species has been considered before as brookesia sp . aff . karchei “ambre” [ 7 ] and as brookesia sp . nov . [ 41 ] .

the aim of my study is to identify morphological and roosting differences in brookesia stumpffi in relation to primary , secondary , and degraded habitats . this can give implications for conservation of chameleons in madagascar and provide more of an idea about dwarf chameleons coping strategies for deforestation . my hypothesis is that brookesia stumpffi have shifted in their morphology due to a change in resources between primary , secondary , and degraded habitats . i am very excited to be conducted this exciting research project over the next six months .

brookesia from madagascar , and the rationale for its listing on cites appendix ii . oryx

the generic name brookesia is in honor of british naturalist joshua brookes . [ 1 ]

( a ) male and ( b ) female of brookesia tristis from montagne des français ; ( c ) male and ( d ) female of brookesia confidens from ankarana ; ( e ) male and ( f ) female of brookesia micra from nosy hara ; ( g ) male and ( h ) female brookesia desperata from forêt d ' ambre .

wgetz ? [ reptilia - species : brookesia * ] [ accessed 10 august 2005 ] .

brookesia are easiest to find at night when they sleep on the leaves of small shrubs .

brygoo , e . r . & c . a . domergue 1971 . notes sur les brookesia ( chaméléonidés ) de madagascar . description d ' une espéce nouvelle , b . antoetrae n . sp . , et des hémipénis de b . stumpffi et b . ebenaui . remarques sur la répartition de b . stumpffi . . bull . mus . nat . hist . nat . paris ( 2 ) 42 ( 5 ) : 830 - 838 . - get paper here

morphometric measurements ( in mm ) of examined type specimens of brookesia tristis and b . confidens .

morphometric measurements ( in mm ) of examined type specimens of brookesia micra and b . desperata .

remark . —this species has been considered before as brookesia sp . “nosy hara” [ 7 ] .

finally got our 2 . 3 group of brookesia so decided to take some video of the unboxing .

all phylogenetic analyses included these outgroup taxa as well as all non - brookesia chamaeleonid taxa from figure 2 .

remark . —this species has been considered before as conspecific with brookesia sp . “montagne des francais” [ 7 ] .

nota bene : a binomial authority in parentheses indicates that the species was originally described in a genus other than brookesia .

boettger , o . 1894 . eine neue brookesia ( chamaeleontidae ) aus nossibé . zool . anz . 17 : 182—185 . - get paper here

brookesia tristis sp . n . zoobank lsid : urn : lsid : zoobank . org : act : 56f36d4d - 1f94 - 49c4 - ac30 - 6e894b6aa998

brookesia confidens sp . n . zoobank lsid : urn : lsid : zoobank . org : act : cb2a9146 - 0161 - 42b7 - a145 - 6ded579f1f21

brookesia micra sp . n . zoobank lsid : urn : lsid : zoobank . org : act : d1a239d6 - 93e8 - 4c34 - a428 - f79a2c8b6405

brygoo , e . r . 1978 . reptiles sauriens chamaeleonidae . genre brookesia et complément pour le genre chamaeleo . faune de madagascar 47 : 1 - 173 .

yeah , it says the stumpffi like 77 to 82 degrees , but cooler and very humid under the leaves , so . . . i don ' t really see how you would keep that high without an incandescent unless you just had a hot room . but if you have a warm room , you don ' t need an incandescent .

it has been noted that most of the 15 brookesia species in northern madagascar show a remarkable degree of microendemism [ 12 ] . the importance of northern madagascar has also been assessed based on an explicit analysis of species richness of brookesia , which was highest in this part of the island [ 14 ] . considering the novel classification and new species of the b . minima group , this trend is now even more obvious , with 21 species of brookesia being confined to northern madagascar [ 12 ] .

brookesia feed on small insects and reply on their cyptic coloration to evade predators . when disturbed , these chameleons will play dead in an effort to resemble a fallen leaf .

. . . and juveniles being found lower than adults ( see razafimahatratra et al . , 2008 ) . however , even species such as brookesia stumpffi , which has relatively high roosts for a leaf chameleon ( mean height , 0 . 43 m ) , still roost 2 m lower than sympatric furcifer angeli ( carpenter and robson , 2005 ) . in two studies of leaf chameleons , no significant differences between sexes were found for roost height ( randrianantoandro et al . , 2007a ; razafimahatratra et al . , 2008 ) . . . .

combined mitochondrial ( nd1 / nd2 / nd4 ) and nuclear ( rag1 / cmos ) data , 12 - partition relaxed - clock bayesian ( rcb ) cladogram with maximum - likelihood ( ml ) phylogram insert . analyses included all ougroup taxa , although for clarity , most nonchameleon taxa are not shown . major clades are color coded , and major clades of “typical” brookesia are numbered ( see text ) . rcb / unrooted bayesian ( ub ) pp > 95 % are denoted by asterisks ( pp between 90 % and 95 % are shown as actual values ) above branches , and ml bootstrap values > 50 % are shown below branches . black diamond indicates alternate attachment point for brookesia stumpffi clade in the ub and ml analyses . dashed terminal branches mark individuals represented by nd4 data only .

in this study , we use brookesia to test the temporal and spatial predictions of 3 species - level diversification hypotheses for madagascar ( mr , wc , and rb ; vences et al . 2009 ) . the first step in this process is to infer a phylogeny of brookesia to identify statistically supported sister - species pairs using dna sequence data from multiple mitochondrial and nuclear protein - coding genes . next , we use divergence dating to statistically test the temporal prediction of both the mr and the wc hypotheses that recent ( pleistocene or possibly pliocene ) climatic cycles are a major force promoting brookesia species diversification .

. . . most species of brookesia have very small distribution ranges [ 6 ] , [ 12 ] ; indeed , almost 50 % of the species are known from single localities [ 13 ] . within brookesia , both species diversity and levels of endemism are highest in northern madagascar , and are correlated with elevational and environmental heterogeneity in this area [ 12 ] , [ 14 ] . . . .

when disturbed these chameleons will play dead , resembling a fallen leaf . the brookesia species are often called “leaf chameleons” because their small body and morphology resembling a leaf makes them highly cryptic when on the ground .

necas , p . & schmidt , w . 2004 . mysterious mini - dragons : the stump - tailed chameleons brookesia and rhampholeon . reptilia ( gb ) ( 35 ) : 10 - 21 - get paper here

the following pictures show the habitat of brookesia stumpffi at the end of the rainy season . some are even from finding places of this species . generally , this species has no high demands concerninc specialized habitats , it can cope with various environments . in ankify and ambanja , this species occurs in the thick foliage layer of coffee and cocoa plantations . on nosy be , along the coast of ankify and on nosy komba , they rather inhabit rocky environments with dense undergrowth and mossy soil areas . in contrast the habitat in ankarana is entwined undergrwoth with lots of twigs and thin branches , much foliage and small plants . the ground ist diffused with rocks and stones here , too .

the mrca of the b . nasus – b . lolontany clade and all other brookesia appears early in the history of the group ( although monophyly of neither brookesia nor this subclade is significantly supported ; fig . 2 ) . the remainder of the genus is clearly monophyletic and is divided into 2 well - supported clades . one of these clades corresponds to the extremely miniaturized b . minima group . the other major clade , representing the typical brookesia morphotype , consists of a series of 5 maximally supported subclades ( one of which is a single species ) whose interrelationships are strongly supported by the rcb analysis but less so by the ub and ml analyses ( fig . 2 ) .

necas , p . & schmidt , w . 2004 . geheimnisvolle mini - drachen : die erd - und stummelschwanzchamäleons der gattungen brookesia und rhampholeon . reptilia ( münster ) 9 ( 48 ) : 18 - 27 - get paper here

crottini , angelica ; aurélien miralles , frank glaw , d . james harris , alexandra lima & miguel vences 2012 . description of a new pygmy chameleon ( chamaeleonidae : brookesia ) from central madagascar . zootaxa 3490 : 63–74 - get paper here

one clade of malagasy leaf chameleons , the brookesia minima group , is known to contain species that rank among the smallest amniotes in the world . we report on a previously unrecognized radiation of these miniaturized lizards comprising four new species described herein .

carpenter , a . i . and robson , o . 2005 . a review of the endemic chameleon genus brookesia from madagascar , and the rationale for its listing on cites appendix ii . oryx 39 ( 4 ) : 375 - 380 .

brookesia is a smallest reptiles . members of the genus brookesia are largely brown and most are essentially terrestrial . a significant percentage of the species in the genus were only identified to science within the last three decades , and a number of species that still have not received a scientific name are known to exist . most inhabit very small ranges in areas that are difficult to access , and due to their small size and secretive nature , they have been relatively poorly studied compared to their larger relatives .

. . . these lizards , which appear to form the sister taxon of all other chameleons ( rieppel 1987 ; townsend and larson 2002 ) , constitute one of the largest malagasy reptile groups ( raxworthy and nussbaum 1995 ; glaw and vences 2007 ) . most brookesia species have very small ranges ( raxworthy and nussbaum 1995 ; raselimanana and rakotomamalala 2003 ) , with almost half known essentially from single localities ( carpenter and robson 2005 ) . brookesia can be divided into 3 main groups based on morphology . . . .

beolens b , watkins m , grayson m . 2011 . the eponym dictionary of reptiles . baltimore : johns hopkins university press . xiii + 296 pp . isbn 978 - 1 - 4214 - 0135 - 5 . ( genus brookesia , p . 40 ) .

most species of brookesia have very small distribution ranges [ raselimanana ap , rakotomalala d ( 2003 ) ] , [ raxworthy cj , nussbaum ra ( 1995 ) ] ; indeed , almost 50 % of the species are known from single localities [ carpenter ai , robson o ( 2005 ) ] . within brookesia , both species diversity and levels of endemism are highest in northern madagascar , and are correlated with elevational and environmental heterogeneity in this area [ townsend tm , vieites dr , glaw f , vences m ( 2009 ) ] .

. . . the micro leaf chameleon genus brookesia displays extremes within the realm of microendemism and miniaturization , with 50 % of species in this large genus coming from single type localities and exhibiting patterns of miniaturization in proportion with range ( carpenter and robson 2005 , townsend et al . 2011 ) . as a result , many species of brookesia are listed as threatened or endangered under cites due to small ranges , preference for undisturbed habitat , and extremely specialized niche - filling ( carpenter and robson 2005 , glaw et al . 2012 ) . brookesia miniaturization is a unique trait in madagascar chameleons , mirrored by the dwarf african chameleon genus rhampholeon and rippeleon ( glaw et al . 2012 ) , and most likely developed to fulfill extremely specific niches in small habitat ranges ( gaston 1996 ) . . . .

in my experience ( brevicaudatus ) is better indirect light of other terraium ( present in reptile room ) near pigmy tank . . . the tube on the tank produce noise for the brevicaudatus : it ' s the same for brookesia or they prefer more bright terrarium ?

ted m . townsend , david r . vieites , frank glaw , miguel vences ; testing species - level diversification hypotheses in madagascar : the case of microendemic brookesia leaf chameleons , systematic biology , volume 58 , issue 6 , 1 december 2009 , pages 641–656 , urltoken

b . stumpffi resembles the dead leaves among which they live . the head is flattened and the rear of the casque is adorned with spinose projections . the orbital crests are well developed and their edges are somewhat scalloped . a row of thorn - like projections runs down either side of the dorsum , from a few millimeters behind the casque to the proximal 1 / 4 of the tail . the basic body coloration is typically brown , gray , drab green , or rust but orange specimens are also observed on occasion . indistinct , yellowish spots are also possible . during courtship , males may adopt a mottled , " lichen - like " appearance but otherwise the the color changing ability , as with most members of the brookesia , is not well developed . males may be distinguished by their more slender body shape and the appearance of a hemipenal bulge .

gray je . 1864 . revision of the genera and species of chamæleonidæ , with the description of some new species . proc . zool . soc . london 1864 : 465 - 477 + plates xxxi & xxxii . ( brookesia , new genus , pp . 476 - 477 ) .

townsend , t . m . ; vieites , d . r . ; glaw , f . & vences , m . 2009 . testing species - level diversification hypotheses in madagascar : the case of microendemic brookesia leaf chameleons . systematic biology 58 ( 6 ) : 641–656 - get paper here

brookesia chameleons are some of the world ' s smallest reptiles—one species reaches a maximum length of just over an inch ( 30 mm ) . also known as stumped - tailed or leaf chameleons , these diminutive creatures are found in the leaf litter of rainforests and dry deciduous forests in much of madagascar .

the wc , rb , and mr hypotheses all make general predictions about relative species abundance at different altitudes . following wollenberg et al . ( 2008 ) , we divided madagascar into a coarser grid with cell sizes of 82 × 63 km = 5166 km 2 and used the arcview extension “endemicity tools” ( provided by n . danho ) to calculate brookesia species richness and corrected weighted endemism ( crisp et al . 2001 ) for each cell . we then tested for nonparametric correlation of these values with altitudinal range , defined as maximum–minimum elevations per grid cell ( excluding grids with no occurrence of any brookesia species ) .

males ( left ) and females ( right ) of four brookesia species described in 2012 , all belonging to the b . minima species group : a - b b . tristis , c - d b . confidens , e - f b . micra , g - h b . desperata [ 2 ]

most brookesia are on cites appendix ii , the only exception being b . perarmata on appendix i ( a species also listed as endangered by iucn ) . consequently , a special permit is required to import any of the below species from their native madagascar , and typically no permit is issued for b . perarmata .

glaw , f . ; köhler , j . r . ; townsend , t . m . ; vences , m . ( 2012 ) . salamin , nicolas , ed . " rivaling the world ' s smallest reptiles : discovery of miniaturized and microendemic new species of leaf chameleons ( brookesia ) from northern madagascar " .

glaw f , köhler j , townsend tm , vences m . ( 2012 ) . " rivaling the world ' s smallest reptiles : discovery of miniaturized and microendemic new species of leaf chameleons ( brookesia ) from northern madagascar " . plos one 7 ( 2 ) : e31314 . doi : 10 . 1371 / journal . pone . 0031314 .

madagascar ' s brookesia dwarf chameleons are believed to require relatively intact forest for survival . although they have featured in herpetological surveys , taxonomic reviews and trade assessments , very little is known about their microhabitat requirements or ecology . over a 5 night period in a deciduous forest in western madagascar we recorded the night roosting sites for three sympatric . . . [ show full abstract ]

species richness of brookesia was negatively correlated with the minimum altitude ( p < 0 . 05 ) and positively correlated with the maximum altitude ( p < 0 . 001 ) in each grid cell . species richness and corrected weighted endemism of brookesia furthermore showed a significant correlation with altitudinal range per grid cell ( p < 0 . 001 and p < 0 . 05 , respectively ) . these results are not congruent with the rb and wc hypotheses , both of which predict a higher species richness and endemism at lower elevations where rivers are widest and where river basins forming coes have their headwaters . in contrast , the results are consistent with the mr hypothesis , which predicts more species will be restricted to montane regions where there is a higher probability of isolation during dry periods .

divergences amongst species in the b . minima group are known to be uniformly deep ( eocene or early oligocene ) [ 14 ] . although the confidens - tuberculata and tristis - desperata divergences are slightly lower than other sister - taxon divergences within the b . minima group , they are still larger than almost all sister - taxon divergences in other parts of the brookesia tree [ 14 ] .

average uncorrected nd2 sequence divergences amongst all lineages identified by this analysis are quite high . brookesia confidens is 20 % divergent from its sister taxon b . tuberculata , b . tristis is approximately 18 % divergent from b . desperata , and b . micra is about 23 % and 26 % divergent from b . tristis and b . desperata , respectively . the highest uncorrected pairwise divergences reaches 32 . 6 % between b . karchei and b . micra , and the smallest interspecific distance is 18 % between b . desperata and b . tristis . the largest intraspecific distances are within b . exarmata ( 3 % ) , b . tristis ( up to 2 . 6 % ) , and brookesia sp . from betampona ( 2 . 1 % ) ; all other species have maximum divergences among haplotypes ≤1 . 2 % .

maps illustrating ( a ) differential brookesia species diversity ( richness ) across madagascar , quantified as numbers of species present within one - quarter degree square grid cells covering the entire island and ( b ) degree of endemism in different parts of madagascar . endemism is calculated as a measure of range - size rarity , expressed as the percentage aggregated reciprocal range size for all species per one - quarter degree square grid cell .

. . . microendemism on madagascar proper has been theorized to be the result of watershed retreat during the quaternary period ( wilmé et al . 2006 ) , with subsequent effects of low vagility , genetic isolation , low gene flow , and small isolated land area which have been linked with miniaturization ( gaston 1996 ) . the micro leaf chameleon genus brookesia displays extremes within the realm of microendemism and miniaturization , with 50 % of species in this large genus coming from single type localities and exhibiting patterns of miniaturization in proportion with range ( carpenter and robson 2005 , townsend et al . 2011 ) . as a result , many species of brookesia are listed as threatened or endangered under cites due to small ranges , preference for undisturbed habitat , and extremely specialized niche - filling ( carpenter and robson 2005 , glaw et al . 2012 ) . . . .

the reduction of external morphological complexity in these miniaturized representatives of brookesia has led to considerable taxonomic confusion . citing an absence of clear external morphological differentiation , raxworthy and nussbaum [ 12 ] synonymized b . peyrierasi and b . tuberculata with b . minima , and b . ramanantsoai with b . dentata . subsequently , another species of this group was described ( b . exarmata ) [ 20 ] , and it was demonstrated that b . peyrierasi and b . tuberculata are valid species , based on differences in both external and genital morphology [ 5 ] , [ 20 ] . molecular evidence has supported monophyly of the b . minima group , albeit with the inclusion of one slightly larger species , b . karchei [ 14 ] . further , compared to species within other brookesia clades , all described b . minima group species are characterized by unusually high genetic differentiation strongly supporting their specific distinctness .

glaw , f . ; köhler , j . r . ; townsend , t . m . ; vences , m . ( 2012 ) . salamin , nicolas , ed . " rivaling the world ' s smallest reptiles : discovery of miniaturized and microendemic new species of leaf chameleons ( brookesia ) from northern madagascar " . plos one 7 ( 2 ) : e31314 . doi : 10 . 1371 / journal . pone . 0031314 . pmc 3279364 . pmid 22348069 .

within the brookesia minima group , hemipenes have been described for three species ( b . peyrierasi , b . ramanantsoai and b . tuberculata ) [ 5 ] , [ 18 ] , [ 36 ] . we herein provide data on genital morphology for the four newly described species , and for b . minima ( see species accounts below ) . hence , within the group , the genital morphology remains unknown only for b . dentata , b . exarmata and b . karchei .

the newly discovered dwarf chameleon species represent striking cases of miniaturization and microendemism and suggest the possibility of a range size - body size relationship in malagasy reptiles . the newly described brookesia micra reaches a maximum snout - vent length in males of 16 mm , and its total length in both sexes is less than 30 mm , ranking it among the smallest amniote vertebrates in the world . with a distribution limited to a very small islet , this species may represent an extreme case of island dwarfism .

the 26 currently recognized species of brookesia typically dwell and forage on the ground , often within the leaf litter on the floor of rainforest and dry deciduous forest , and climb at night to low perches in the vegetation for sleeping . they are characterized by a typically dull brown or ( rarely ) greenish colour , a short non - prehensile tail that is used as “fifth leg” in walking [ boistel r , herrel a , daghfous g , libourel p - a , boller e , et al . ( 2011 ) ] .

brookesia , calumma , and furcifer constitute the three genera of chameleons occurring on madagascar , an island which harbors almost 80 of the world ' s ca . 185 nominal chameleon species [ 6 ] – [ 8 ] ) . all three genera are endemic to madagascar , with the exception of two furcifer species on the nearby archipelago of the comoros [ 9 ] , [ 10 ] . calumma and furcifer are generally medium - sized to large arboreal chameleons ( ca . 50–295 mm svl and 110–695 mm tl ) , and are often vividly coloured [ 7 ] . on the contrary , the 26 currently recognized species of brookesia typically dwell and forage on the ground , often within the leaf litter on the floor of rainforest and dry deciduous forest , and climb at night to low perches in the vegetation for sleeping . they are characterized by a typically dull brown or ( rarely ) greenish colour , a short non - prehensile tail that is used as “fifth leg” in walking [ 11 ] , and a smaller size of ca . 15–65 mm svl and 25–105 mm tl [ 7 ] .

brookesia distributions span the island , although they are absent from many large areas in the central highlands and the south . three main centers of diversity ( fig . 3a ) are found in the northeast , north , and northwest , respectively . these areas are also coes , with the highest percent values of range - size rarity ( fig . 3b ) . however , there are several areas , mainly in the west and northwest regions of the island , with a high degree of endemism corresponding to several species only known from single locality records .

chronogram from the 12 - partition ( nd1 / nd2 , nd4 , cmos , and rag1 , each partitioned by codon position ) rcb phylogenetic analysis . time units on scale bar in millions of years ago . see supplementary materials for exact divergence time means and 95 % cis for all nodes . maps above chronogram illustrate patterns of species richness ( quantified as numbers of species present within one - quarter degree square grid cells ) for the 3 main clades ( 1 = brookesianasus group , 2 = brookesiaminima group , and 3 = “typical” brookesia group ) .

natural history . — most individuals were found roosting at night on thin branches about 5–20 cm above the leaf litter in deciduous dry forest close to a small forest trail within a small area , surrounded by tsingy outcrops . at this locality , the species was relatively abundant , whereas it was not found at similar localities nearby , suggesting a patchy distribution and a preference for certain microhabitats . this hypothesis is also supported by the fact that earlier herpetological surveys in ankarana [ 38 ] , [ 39 ] , did not record any species of the brookesia minima group .

phylograms ( 50 % majority rule consensus trees ) from bayesian inference analyses of dna sequence alignments of ( a ) the nuclear genes cmos ( 846 bp ) and ( b ) rag1 ( 1522 bp ) . ( c ) phylogram from bi analysis of concatenated dna sequences of the nuclear genes cmos ( 846 bp ) and rag1 ( 1522 bp ) , and the mitochondrial genes nd2 ( 568 bp ) and 16s ( 580 bp ) . ( d ) brookesia portion of the bayesian chronogram derived from the beast analyses of all four concatenated genes ( see supporting information s1 for full chronogram with all outgroups ) . posterior probabilities are indicated above branches , and bars represent 95 % hpds for mean date estimates . units on scale are millions of years . species of the northern clade ( b . tuberculata plus the four new species described herein ) are marked in colour . the trees were rooted with brookesia nasus , and including b . brygooi and b . superciliaris as hierarchical outgroups ( not shown ) . a , b and c refer to major clades as discussed in the text . asterisks denote posterior probabilities of 1 . 0 .

additional analyses using alternative calibration distributions and data sets had little effect on divergence time estimates . analyses performed with normally distributed calibrations ( with standard deviations arbitrarily set at 10 % of the mean ) gave average intrachameleon divergences about 10 % more recent than those using lognormally distributed calibrations . likewise , analyses that excluded mtdna third codon positions and all mtdna data gave respective divergence estimates on average about 15 % and 18 % more recent than the full - data analysis . in all these analyses , brookesia sister - taxon divergences remained solidly miocene in age , and thus , our above conclusions are unaffected .

in their study of biogeographic patterns in the leaf chameleons ( genus brookesia ) of northern madagascar , raxworthy and nussbaum ( 1995 ) recognized 5 rainforest regions delimited largely by altitudinal differences and intervening dry forests and characterized by a high degree of endemism in these lizards . raxworthy and nussbaum ( 1995 ) hypothesized that pleistocene climatic fluctuations had caused fragmentation of the rainforest , resulting in multiple allopatrically distributed sister - taxon pairs ( hereafter referred to as the mountain refuge [ mr ] hypothesis ) . these conclusions were not based on explicit phylogenetic hypotheses ; sister - taxon pairs were assumed based on general morphological similarities .

the wilmé et al . ( 2006 ) wc model predicts that sister species should generally occupy adjacent coes or possibly a coe and an adjacent rdw . to test the fit of our brookesia data to the spatial aspect of this model , we first plotted the distributions of all brookesia sister - species pairs identified in our phylogenetic analyses onto the wilmé et al . ( 2006 ) watershed map ( fig . 1 ) and counted the number of pairs matching the model ' s predictions . next , we randomly assigned each species from all sister - species pairs to one of the watersheds collectively occupied by them and counted the number of pairs fitting the model ' s predictions . we repeated this 10 , 000 times to generate a null distribution of expected number of fits to the model if sister species are actually distributed randomly with respect to relative watershed positions . if less than 5 % of the simulation rounds resulted in a number of matches equal to or greater than the number of matches from the real data , we rejected the null in favor of the wilmé et al . ( 2006 ) wc hypothesis . we repeated this analysis under the interpretation that allopatric sister species occupying the same watershed also fit the model .

locality data for all brookesia species of madagascar were gathered from museum data , our own global positioning system readings , and pertinent literature . single - species maps that almost fully agree with the data used herein are reproduced in glaw and vences ( 2007 ) . small sample sizes of locality records are not appropriate to obtain reliable estimates of potential distribution by modeling . for 9 species , more than 6 ( range 7–39 ) locality records were available . for these species , we defined distributions by potential distribution models ( pruned for overprediction ) , and for the rest of the species , we used point locality data .

all specimens were sequenced for 2 nuclear ( rag1 , ∼1500 bp and cmos , ∼800 bp ) and 3 mitochondrial ( nd1 , ∼ 70 bp ; nd2 , ∼ 1035 bp ; and nd4 , ∼ 700 bp ) protein - coding genes ; we included several nd4 sequences from raxworthy et al . ( 2002 ) that expanded our taxonomic ( 3 species ) or geographic ( 6 species ) coverage . our final brookesia sampling covered approximately 28 species , including all but 2 of the named species . museum / collection and genbank numbers of specimens can be found in the online supplementary material ( available from urltoken ) .

the wealth of new material and the obvious morphological distinctness of some of the new specimens ( especially the nosy hara population ) prompted us to study their genetic differentiation , external morphology and genital morphology in more detail . our integrative data clearly and concordantly indicate that populations d - g listed above represent new , undescribed species , and anticipating their formal description below , we will in the following sections use the new species names erected herein : brookesia tristis ( specimens from montagne des français ) , b . confidens ( ankarana ) , b . micra ( nosy hara ) , and b . desperata ( forêt d ' ambre ) .

this ecomorphological disparity among chameleons is also found in mainland africa , leading to use of the general terms “tree chameleons” and “ground chameleons [ 10 ] . ” african ground chameleons comprise the genera rhampholeon and rieppeleon , and are superficially quite similar to the malagasy brookesia ( which are often further distinguished from other ground chameleons by the name “leaf chameleons” ) . interestingly , neither ground chameleons as a whole , nor the african species , form monophyletic groups . in fact , rieppeleon forms the sister taxon of a tree chameleon genus ( archaius ) endemic to the seychelles islands [ 10 ] , indicating a history of multiple independent evolutionary shifts between arboreal and terrestrial chameleon ecomorphs .

rivers appear to have played a substantial role in the diversification of several lemur groups ( pastorini et al . 2003 ; goodman and ganzhorn 2004 ; louis et al . 2005 ) , some frogs ( vieites et al . 2006 ) , and possibly even some chameleons of the genus furcifer ( raselimanana and rakotomamalala 2003 ) . however , this does not seem to be true for leaf chameleons , at least at the species level and above . it seems unlikely that individual brookesia disperse over large distances , and they certainly cannot swim across rivers . however , given their small size and leaf litter microhabitat , they could quite plausibly float across rivers on debris washed away during heavy rains .

phylogram ( 50 % majority rule consensus tree ) from a bayesian inference analysis of the 568 bp dna sequence alignment of the mitochondrial nd2 gene showing deep divergences among and low differences within species of the b . minima group . species of the northern clade ( b . tuberculata plus the four new species described herein ) are marked in colour . the tree was rooted with brookesia nasus , and including b . brygooi and b . superciliaris as hierarchical outgroups ( not shown ) . a , b and c refer to major clades as discussed in the text . inset maps show known distribution ranges ( typically single localities ) of all species . asterisks denote posterior probabilities of 1 . 0 .

among amniotes , the smallest species are lizards , but they are consistently larger than amphibians , suggesting that different constraints are acting on this group . the maximum recorded svl of sphaerodactylus ariasae ( 18 mm ) is below the maximum value in brookesia micra ( 19 . 9 mm ) . however if using tl as yardstick , the record of the smallest squamate and amniote in fact applies to b . micra with a maximum tl of 29 mm , given that s . ariasae has a tl well above 30 mm [ 1 ] due to its relatively longer tail . the ‘award’ for the lower size record in amniotes , as in amphibians , therefore depends on the type of measurement used for ranking .

. . . results obtained from this project suggest that b . micra has a wider range and habitat tolerance than previously believed , indicating that b . micra could be present elsewhere on the island as well as elsewhere in the archipelago or the mainland . in addition , many behavioral aspects common to thejenkins et al . 2003 ) . carpenter and robson ( 2005 ) believe this to be because of census difficulties and overall low density of the brookesia genus . this study , however , shows that b . micra can be found in incredibly high densities . most likely , this is due to presence of good habitat all along the transect line , particularly in areas of high b . micra abundance . b . micra appears to bene . . .

microendemism is widespread in malagasy animals , but there are few examples in which it is as extreme as in the miniaturized species of the brookesia minima group and in frogs of the genus stumpffia [ 49 ] . this indicates a possible body size effect on the degree of microendemism , which is consistent with the body - range size relationship previously demonstrated in the malagasy mantellid frog radiation [ 58 ] . in general , a correlated decrease in species ' ranges with decreasing body size is a recognized pattern in animals [ 59 ] . a closer analysis of this pattern in madagascar ' s biota may yield new insights and provide at least a partial explanation for the prevalence of microendemic patterns in some groups of animals and the absence of such patterns in others .

on the contrary , in several other species and species complexes of brookesia more convoluted taxonomic situations are found . the widespread eastern b . superciliaris shows strong mitochondrial divergences among populations ; however , this structuring is not mirrored by divergences in one nuclear gene studied nor in morphology , suggesting these populations are conspecific [ 42 ] . the equally widespread b . thieli as well contains several deep genealogical lineages which however do not form a monophyletic group , with the morphologically divergent b . lineata and b . vadoni clustering within the b . thieli clade [ 14 ] . finally , the two morphologically distinct species b . ambreensis and b . antakarana , both microendemic in montagne d ' ambre , do not show any consistent mitochondrial or nuclear dna differentiation [ 14 ] .

we were concerned about the effects that substitutional saturation at deeper levels might have on divergence time estimates , especially with regard to mitochondrial dna ( mtdna ) data ( hugall et al . 2007 ) . however , the mtdna data were needed to confidently resolve more recent branching points ( see below ) , suggesting also that branch lengths within brookesia would be better estimated by including mtdna data . we therefore conducted analyses using 1 ) all data , 2 ) nuclear data plus first and second positions from the mtdna data , and 3 ) nuclear data alone to test the robustness of our divergence time estimates . we also tested our fossil calibrations using the cross - validation method of near et al . ( 2005 ) , which identifies potentially problematic fossils that cause incongruent age estimates of other dated nodes in the tree .

voucher specimens were euthanized using approved methods ( anaesthesia with ketamine , followed by ketamine overdosis ) that do not require approval by an ethics committee . all specimens were fixed in 90 % ethanol . muscle tissue samples for molecular analysis were taken from all specimens and preserved in pure ethanol . definition of measurements and the description scheme of the holotypes follows previous brookesia descriptions [ 12 ] . several additional characters were also scored , and their definition follows a previous morphological revision of the b . minima group [ 5 ] . morphometric measurements were taken by mv with a digital caliper to the nearest 0 . 1 mm , following a previously published measuring scheme [ 5 ] . measurements taken include : tl ; svl ; ( these two were previously defined ) tal , tail length ; hw , maximum head width ; hh , maximum head height ; ed , eye diameter ; forl , forelimb length .

the 3 main clades recovered in our phylogenetic analyses ( figs . 2 and 4 ) show distinct biogeographic patterns . the basally diverging b . nasus clade contains 2 species , b . nasus in the southeast and b . lolontany in the north , suggesting an old north - south connection . the b . minima clade shows the highest degree of endemism ; most of the 11 species are restricted to very small areas in the north and north - central regions . the third and largest clade ( typical brookesia ) spans most of the island , although centers of both diversity and endemism in this clade are in the north . within this clade , b . brygooi , b . bonsi , b . decaryi , and b . perarmata ( all western species ; clade 1 , fig . 2 ) form the sister taxon of the remaining species , none of which are restricted to the western forests .

within each of the ml , ub , and rcb analysis sets , phylogenetic results were broadly congruent across the different genomes and genes , with topological conflicts involving only poorly supported nodes ( i . e . , ml bootstraps < 70 % , bayesian posterior probabilities [ pps ] < 95 % ) . one exception to this pattern involves strongly conflicting nuclear and mitochondrial results for the placement of 1 specimen of brookesia brygooi . this discrepancy is probably due to ancient mtdna introgression , and the nuclear topology almost certainly reflects the true organismal history ( see online supplementary material ) . mitochondrial data for this specimen were therefore excluded in all further phylogenetic analyses . in general , the mtdna data alone gave strong support for relationships within more terminal clades , but poor support for basal nodes , and the nuclear data showed the opposite pattern . all results presented here are from analyses of the combined data set ( fig . 2 ) . this data set and trees from this study can be found at urltoken under accession number sn4455 .

the newly discovered species appear to be restricted to single , mostly karstic , localities in extreme northern madagascar : brookesia confidens sp . n . from ankarana , b . desperata sp . n . from forêt d ' ambre , b . micra sp . n . from the islet nosy hara , and b . tristis sp . n . from montagne des français . molecular phylogenetic analyses based on one mitochondrial and two nuclear genes of all nominal species in the b . minima group congruently support that the four new species , together with b . tuberculata from montagne d ' ambre in northern madagascar , form a strongly supported clade . this suggests that these species have diversified in geographical proximity in this small area . all species of the b . minima group , including the four newly described ones , are characterized by very deep genetic divergences of 18–32 % in the nd2 gene and > 6 % in the 16s rrna gene . despite superficial similarities among all species of this group , their status as separate evolutionary lineages is also supported by moderate to strong differences in external morphology , and by clear differences in hemipenis structure .

nossi be = nosy bé , nw madagascar , nosy komba , nosy sakatia .

type locality : restricted to insel nossibé ( = nosy bé ) , off n . w madagascar

note : tdwg regions are generated automatically from the text in the distribution field and not in every cases it works well . we are working on it .

andreone f . , f . glaw , r . a . nussbaum , c . j . raxworthy , m . vences , and < br / > j . e . randrianirina 2003 . the amphibians and reptiles of nosy be ( nw madagascar ) and nearby islands : a case study of diversity and conservation of an insular fauna . journal of natural history 37 ( 17 ) : 2119–2149

beolens , bo ; michael watkins , and michael grayson 2011 . the eponym dictionary of reptiles . johns hopkins university press , baltimore , usa - get paper here

d ' cruze , neil ; jeremy sabel , katie green , jeffrey dawson , carlie gardner , janine robinson , georgina starkie , miguel vences , and frank glaw . 2007 . the first comprehensive survey of amphibians and reptiles at montagne des francais , madagascar . herp . cons . biol . 2 ( 2 ) : 87 - 99 - get paper here

durkin , louise ; mark d . steer , and elise m . s . belle 2011 . herpetological surveys of forest fragments between montagne d ' ambre national park and ankarana special reserve , northern madagascar . herp . cons . biol . 6 ( 1 ) : 114 - 126 - get paper here

d’cruze , n . ; köhler , j . ; franzen , m & glaw , f . 2008 . a conservation assessment of the amphibians and reptiles of the forêt d’ambre special reserve , north madagascar . madagascar conservation & development 3 ( 1 ) : 44 - 54

glaw , f . 2015 . taxonomic checklist of chameleons ( squamata : chamaeleonidae ) . vertebrate zoology 65 ( 2 ) : 167–246 - get paper here

hyde roberts , s . & c . daly 2014 . a rapid herpetofaunal assessment of nosy komba island , northwestern madagascar , with new locality records for seventeen species . salamandra 50 ( 1 ) : 18 - 26 - get paper here

müller , r . & hildenhagen , t . 2009 . untersuchungen zu subdigital - und subcaudalstrukturen bei chamäleons ( sauria : chamaeleonidae ) . sauria 31 ( 3 ) : 41 - 54 - get paper here

müller , rolf & thomas hildenhagen 2009 . zeigt her eure füßchen - bemerkungen zu sohlen - und schwanzstrukturen bei chamäleons . chamaeleo 19 ( 1 ) : 13 - 24 - get paper here

necas , p . 1995 . chamäleons - bunte juwelen der natur . edition chimaira bei bücher - kreth ( frankfurt / m . , germany )

necas , p . & schmidt , w . 2004 . stump - tailed chameleons . miniature dragons of the rainforest . edition chimaira , frankfurt , 256 pp . [ review in elaphe 14 ( 1 ) : 24 ]

necas , petr 1999 . chameleons - nature ' s hidden jewels . edition chimaira , frankfurt ; 348 pp . ; isbn 3 - 930612 - 04 - 6 ( europe ) < br / > isbn 1 - 57524 - 137 - 4 ( usa , canada )

rakotoarison , andolalao ; jesse erens , fanomezana m . ratsoavina , miguel vences 2015 . amphibian and reptile records from around the betsiboka delta area in north - western madagascar herpetology notes 8 : 535 - 543 - get paper here

raxworthy , c . j . ; enrique martinez - meyer ; ned horning ; ronald a . nussbaum ; gregory e . schneider ; miguel a . ortega - huerta & a . townsend peterson 2003 . predicting distributions of known and unknown reptile species in madagascar . nature 426 : 837 - 841

rovatsos , michail ; marie altmanová , martina johnson pokorná , petr velenský , antonio sánchez baca orcid and lukáš kratochvíl 2017 . evolution of karyotypes in chameleons genes 8 ( 12 ) : 382 ; doi : 10 . 3390 / genes8120382 - get paper here

schmidt , w . ; tamm , k . & wallikewitz , e . 2010 . chamäleons - drachen unserer zeit . natur und tier verlag , 328 pp . [ review in reptilia 101 : 64 , 2013 ] - get paper here

this database is maintained by peter uetz ( database content ) and jakob hallermann , zoological museum hamburg ( new species and updates ) .

this species occurs in northwestern madagascar and offshore islands including nosy be , nosy komba and nosy sakatia ( andreone et al . 2003 , glaw and vences 2007 ) . this species has an elevation limit of 150 m above sea level . the lizard ' s extent of occurrence is estimated to be 61 , 884 km .

continent : indian - ocean distribution : nossi be = nosy bé , nw madagascar , nosy komba , nosy sakatia . type locality : restricted to insel nossibé ( = nosy bé ) , off n . w madagascar

this chameleon has been recorded from primary rainforest habitat , dry forests and disturbed forest areas ( glaw and vences 2007 , carpenter and robson 2005 ; durkin et al . 2011 ) . this species is terrestrial during the day , where it inhabits leaf litter , and at night it is found perched on branches approximately 30 - 80 cm above the ground ( glaw and vences 2007 ) .

the following is a representative barcode sequence , the centroid of all available sequences for this species . there is 1 barcode sequence available from bold and genbank .

below is the sequence of the barcode region cytochrome oxidase subunit 1 ( coi or cox1 ) from a member of the species .

listed as least concern as it has an extent of occurrence of 61 , 884 km , appears somewhat adaptable to disturbed habitats , and there are no major threats resulting in a significant decline at present . however , as this species is used in the pet trade , and habitat loss and degradation is occurring , monitoring of harvest levels and population trends is needed .

this species is reported to be common in primary rainforest , dry forests and secondary vegetation ( glaw and vences 2007 ) .

forests in this region are being lost as a result of land clearance and the effects of widespread slash - and - burn farming . however , due to this species ' relatively large distribution and broad range of known habitat types , these should not be considered major threats at this time . small numbers of this species are exported for the pet trade , though this is unlikely to be a major threat .

there are several protected areas within this lizard ' s range . international trade of thespecies in this genus is managed under cites appendix ii . similarly , under ec regulation no . 709 / 2010 , an amendment to ec regulation 338 / 37 ( european union wildlife trade regulation ) , species of the genus are listed in annex b . the harvest levels and population trends of this species need to be monitored .

six times , initially by boettger in 1894 . it was later described by glaw and vences in 1994 : 236 , klaver and böhme in 1997 , necas in 1999 : 268 , pianka and vit in 2003 : 11 , and most recently by townsend

glaw and veneces found the species on small islands of madagascar away from the main land mass of the country in 2007 .

can be found up to a height of 150 metres ( 490 feet ) above sea level , and can be found over an area of 61 , 884 kilometres ( 38 , 453 miles ) .

can grow up to 9 cm ( 3 . 5 in ) , and has a life expectancy of at least three years . during reproduction , this species of

lays between three and five eggs , which hatch between 60 and 70 days later , provided they are at a temperature of 23 °c ( 73 °f ) .

( waxworms ) , and grasshoppers . during the day , the body temperature of

is between 22 and 25 °c ( 72 and 77 °f ) , and is 20 °c ( 68 °f ) during the night .

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