it is native to arkansas in the united states , where it is known from only four locations .

it is a federally listed endangered species of the united states .

this crayfish was first described to science as a new species in 1987 .

it is about 48 millimeters ( 1.8 inches ) long .

a troglobite , it lacks pigment , making it white in color , it and has only rudimentary eyes .

it is genetically distinct from other species of cave crayfish .

the crayfish lives at four caves in northern arkansas , three in benton county and one in washington county .

it is not considered an arkansas endemic species because 58 % of one of the benton county cave zones is actually within the neighboring state of missouri .

this crayfish feeds on organic matter washed into the cave from the surface , and on bat guano .

it is sometimes eaten by the banded sculpin ( cottus carolinae ) .

little else is known about its ecology .

threats to this species include direct mortality when they are trampled by cave explorers and trespassers .

gates have been put in place at the caves to protect them but vandalism is still a threat .

pollution of the groundwater in the caves was the main reason the animal was federally listed .

individuals are sometimes washed out of caves during floods , leading to mortality .

cambarus aculabrum

cambarus aculabrum is a rare species of crayfish known by the common name benton cave crayfish . content licensed under creative commons attribution . source : urltoken
cave crayfish ( cambarus aculabrum ) .
beacham ' s guide to the endangered species of north america . . retrieved july 09 , 2018 from urltoken urltoken
cave crayfish ( cambarus aculabrum ) .
beacham ' s guide to the endangered species of north america . . encyclopedia . com . ( july 9 , 2018 ) . urltoken
u . s . fish and wildlife service . 30 october 1996 .
recovery plan for the cave crafish ( cambarus aculabrum ) .
u . s . fish and wildlife service , atlanta , 37pp .
u . s . fish and wildlife service ( usfws ) . 1996 . cave crayfish ( cambarus aculabrum ) recovery plan . u . s . fish and wildlife service , atlanta , georgia . 37 pp .
u . s . fish and wildlife service . 27 april 1993 .
endangered and threatened wildlife and plants ; determination of endangered status determined for the cave crayfish ( cambarus aculabrum ) .
federal register 58 ( 79 ) : 25742 - 25746 .
graening , g . o . , m . e . slay , a . v . brown , and j . b . koppelman . 2006 . status and distribution of the endangered benton cave crayfish , cambarus aculabrum ( decapoda : cambaridae ) . the southwestern naturalist , 51 ( 3 ) : 376 - 439 .
this species is known from two caves in benton county , arkansas ( hobbs , 1989 ; robison and allen , 1995 ; usfws , 1996 ) and just into missouri ( graening et al . , 2006 ) . with all of the effort to look for cave organisms based out of fayetteville , a great deal of search has been focused in the area to find more cambarus aculabrum . it has long seemed confined to the original 2 caves , yet recent discoveries have added 2 additional sites nearby - both based on one or two specimens . missouri has a cave biologist on staff and considerable survey work has been done with no discoveries of the species in that state ( brian wagner , ar game and fish , to cindy osborne , ar heritage , pers . comm . , june 2002 ) .
first form males are distinguished by a fully formed and hardened first pleopod ( reproductive appendages ) . these males can be further distinguished from closely related cambarus setosus and c . tartarus by the absence of a transverse groove separating the proximolateral lobe from the shaft on the first pleopod . it differs from c . zophonastes in that it possesses a longer central projection of the first pleopod that also has a shallow subapical notch .
( < 100 square km ( less than about 40 square miles ) ) this species is known from two caves in benton county , arkansas ( hobbs , 1989 ; robison and allen , 1995 ; usfws , 1996 ) and just into missouri ( graening et al . , 2006 ) . with all of the effort to look for cave organisms based out of fayetteville , a great deal of search has been focused in the area to find more cambarus aculabrum . it has long seemed confined to the original 2 caves , yet recent discoveries have added 2 additional sites nearby - both based on one or two specimens . missouri has a cave biologist on staff and considerable survey work has been done with no discoveries of the species in that state ( brian wagner , ar game and fish , to cindy osborne , ar heritage , pers . comm . , june 2002 ) .
cave crayfish are highly specialized for living in stable cave environments with low light and low temperatures and are unable to cope with changes in their habitats that may be induced by human activities . water quality degradation represents a major threat to c . aculabrum . this species is also vulnerable due to its limited distribution , with only two known populations containing a small number of individuals ; its limited reproductive potential ; and the potential for take by humans .
bear hollow cave contains a small stream approximately 660 ft ( 201 . 2 m ) long and an undescribed pool that are habitat for c . aculabrum . the cave stream flow and depth varies . at times , parts of the stream may dry up leaving tiny pools of water or parts of the stream may completely disappear underground leaving no trace . after some rainfall events the cave may nearly fill up with water , as evidenced by trash found lodged up near the cave ceiling .
the numbers of crayfish observed in logan and bear hollow caves has varied dramatically between cave visits . the greatest number of crayfish observed in a single visit is nine in bear hollow cave and 21 in logan cave . in 14 visits to logan cave , crayfish were observed on only three occasions . in a 1990 survey , three c . aculabrum were observed in logan cave , one of which was dead , and only a single crayfish in bear hollow cave . six crayfish were observed in logan cave during another cave visit in 1995 while four were observed in bear hollow cave . from october 1994 through september 1995 , between seven and 21 cave crayfish were observed in logan cave .
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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 .
livingston , f . , soulsby , a . - m . , batchelor , a . , dyer , e . , whitton , f . , milligan , h . t . , smith , j . , lutz , m . l . , de silva , r . , mcguinness , s . , kasthala , g . , jopling , b . , sullivan , k . & cryer , g .
has been assessed as critically endangered . this species has an area of occupancy of less than 10 km² as each of the four cave pools has a maximum area of 200 m² . the caves are severely fragmented and they are not connected to each other . there is a continuing decline in the extent and quality of habitat due to continued human disturbances and impacts through trampling and groundwater pollution . this species is extremely sensitive to groundwater quality and there is evidence that it is preyed on by banded sculpin (
) . it has a very low population , of only 40 individuals , and longevity of 75 years indicating that there is a late age of reproductive maturity . a recovery plan has been initiated by the usfws in 1996 , which has been implementing measures to protect this species . however , even with this recovery plan in place , the population still remains low and sensitive to disturbance and quality of habitat .
2006 ) . with all of the effort to look for cave organisms based out of fayetteville , a great deal of search has been focused in the area to find further specimens of this species . it has long seemed confined to the original caves , yet recent discoveries have added 2 additional sites nearby - both based on one or two specimens . in missouri , expert survey work has been carried out with no further discoveries of the species in that state ( brian wagner , ar game and fish , to cindy osborne , ar heritage , pers . comm . 2002 ) . the area of occupancy ( aoo ) is estimated to be less than 4 km² .
2006 ) . probably < 200 adults are extant . numbers of crayfish observed vary dramatically between cave visits . the greatest number observed at one time in one cave is nine and in the other cave 21 ( usfws , 1996 ) . the current total observed population size is 40 individuals , based on the latest complete visual census of the second largest occurrence ( n = 6 ) , largest occurrence ( n = 31 ) , small occurrence ( n = 2 ) , and the smallest creek pool discharge occurrence ( n = 1 ) ( graening
this species is a cave dwelling species inhabiting subterranean streams and pools ( jacobson 1996 ) . one of the caves from which this species is known is an ozarkian solution channel , within which this species has been observed along the side walls of a pool or at the stream margin ( robison and allen 1995 ) . cave streams in which this species lives are generally less than 50 cm deep .
. ( 2006 ) have assessed continuing threats since the recovery plan . among these , habitat degradation from groundwater pollution is the primary reason for federal listing of the species and remains a serious threat . organic pollutants are present in the groundwater basins of the two initial caves . mean concentrations of nitrate , phosphorous , and faecal bacteria consistently equals or exceeds those of regional surface waters monitored by the national water quality assessment program for the springfield plateau aquifer . over 100 confined animal feeding operations ( poultry and swine ) and cattle ranching operations as well as over 60 residences on septic systems are within the recharge zone of one cave and two confined feeding operations , and at least 200 residences on septic systems are within the discharge zone of the second cave ( graening
. 2006 ) . as for human disturbance , both caves with the largest populations were both formerly popular recreational destinations . vandalism and trespassing continue to be serious management issues at both sites , even after erection of steel channel gates . trampling was formerly considered a threat at all sites but has been alleviated at one cave when a fixed line was bolted to a canyon ledge 3 m above the stream in 2001 ( graening
. 2006 ) . more specifically , hog and poultry operations , fertilization of nearby pasture lands , regional airport expansion , and residential development threaten the water quality of the second cave . residential development is the primary threat to water quality in the largest cave occurrence ( usfws 1996 ) .
compiled by the us fish and wildlife service ( jacobson 1996 ) . this plan has seen the implementation of cave gates on two sites
, and three caves have surrounding land recovered and septic tanks are being upgraded to prevent leakages ( natureserve 2009 ) . one of the localities has been designated as a
. it also qualifies as endangered under the federal environmental species act of 1973 ( jacobson 1996 ) .
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small , white obligate cave - dwelling crayfish with no pigment and reduced eyes .
the cave crayfish is a small , white obligate cave - dwelling crayfish . the body length of this species has been measured to reach up to 1 . 8 in ( 4 . 6 cm ) . this species has no pigment and has reduced eyes . it also has an acute or subacute apex of the antermomedian lobe of the epistome .
the reproductive habits of this species , as well as other sociobiological information , is not known at this time . this species , however , displays similar reproductive characteristics of other decopods . the males probably begin molting into the reproductive state in late summer with copulation occurring in late summer and fall . egg laying likely occurs in late winter and early spring . most males molt back to the nonreproductive form during april .
water quality and clarity of logan cave stream is generally high and many water parameters remain relatively constant for much of the year . most changes in water properties occur when the cave ' s stream flow increases after a storm .
the cave crayfish is most commonly found along the walls of the pool , or along stream edges .
the type locality of this species is logan cave as well as its associated stream and lake . the cave cray - fish is also known from bear hollow cave and its associated stream about 23 mi ( 37 km ) from logan cave . both of these areas are located in benton county , arkansas .
in an observation of other cave crayfish and troglobitic species , small population sizes have been a result of reduced food sources . as an adaptation to this it has also been observed that these species display a lower metabolic rate , increased longevity , delayed maturity and reproduction and decreased fecundity . the otherwise adaptive characteristic could make the cave crayfish highly vulnerable to environmental pollution and limit this species ' ability to recover .
in 1989 the u . s . fish and wildlife service ( fws ) purchased 123 . 9 acres ( 50 . 1 hectares ) at logan cave ( the cave crayfish ' s original locality ) . this attempt will facilitate preservation activities initiated by the fws . the rest of the area , however , is privately owned .
a collecting permit is required for collecting for any species , except for fish bait under state regulations . troglobites are protected from possession and sale by arkansas state law .
potential conservation measures and federal involvement are expected to include : ( 1 ) environmental protection agency : clean water act ' s provisions for pesticide registration and waste management actions . ( 2 ) corps of engineers : inclusion of the cave crayfish in project planning and operation and during permit review . ( 3 ) federal highway administration : consideration of the cave crayfish in bridge and road construction in known areas of occupancy . ( 4 ) farmers home administration : consideration of impacts of the cave crayfish in loan processes . ( 5 ) soil conservation service : inclusion of the cave cray - fish in farmer ' s assistance programs .
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2006 ) . with all of the effort to look for cave organisms based out of fayetteville , a great deal of search has been focused in the area to find further specimens of this species . it has long seemed confined to the original caves , yet recent discoveries have added 2 additional sites nearby - both based on one or two specimens . in missouri , expert survey work has been carried out with no further discoveries of the species in that state ( brian wagner , ar game and fish , to cindy osborne , ar heritage , pers . comm . 2002 ) . the area of occupancy ( aoo ) is estimated to be less than 4 km .
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hobbs , h . h . , jr . and a . v . brown . 1987 . a new troglobitic crayfish from north western arkansas . proceedings of the biological society of washington , 100 ( 4 ) : 1040 - 1048 .
this species is highly imperiled . this species has an extent of occurrence of less than 10 sq . km as each of the four cave pools has a maximum area of 200 m . the caves are severely fragmented and they are not connected to each other . there is a continuing decline in the extent and quality of habitat due to continued human disturbances and impacts through trampling and groundwater pollution . this species is extremely sensitive to groundwater quality and there is evidence that it is predated on by banded sculpin ( cottus carolinae ) . it has a very low population , of only 40 individuals , and a longevity of 75 years indicating that there is a late age of reproductive maturity . a recovery plan has been initiated by the us fisheries and wildlife service in 1996 , which has been putting in place measures to protect this species . however , even with this recovery plan in place , the population still remains low and sensitive to disturbance and quality of habitat .
due to latency between updates made in state , provincial or other natureserve network databases and when they appear on natureserve explorer , for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data . please refer to
this species was previously known only from four caves in / around benton county , arkansas that are 38 km apart and both solution channels in boone formation ( hobbs and brown , 1987 ; usfws , 1996 ) . recent documentation of specimens in one cave recharge zone ( outside the cave itself ) in mcdonald co . , missouri , has increased the current understanding of the range of this species to include this area ( graening et al . , 2006 ) . two caves ( benton co . , arkansas ; and washington co . , arkansas ) are fairly recent range additions with one female sighted in 1999 and one male in 2004 ; plus one specimen expelled during a flood event in 2004 , respectively ( graening et al . , 2006 ) . extensive survey effort nearby revealed no others .
probably < 200 adults are extant . numbers of crayfish observed vary dramatically between cave visits with the greatest number observed at one time in one cave is nine and in the other cave 21 ( usfws , 1996 ) . the curent total observed population size is 40 individuals , based on the latest complete visual census of the second largest occurrence ( n = 6 ) , largest occurrence ( n = 31 ) , small occurrence ( n = 2 ) , and the smallest creek pool discharge occurrence ( n = 1 ) ( graening et al . , 2006 ) . the maximum historical count ( irrespective of date ) is 56 ( 47 in the largest cave occurrence , 9 in the second largest cave occurrence ) ( graening et al . , 2006 ) .
two of the four sites are considered viable , one with decent viability ( graening et al . , 2006 ) .
a natural mortality factor is expulsion from the subterranean habitat by flooding during storm events . this has been documented on more than one occasion for the two largest occurrences ( into pea ridge hollow , missouri thus expanding range of this species ) ( graening et al . , 2006 ) . natural dispersal of this species is limited by its confinement to a stygobitic habitat in very few localities .
this species cannot survive outside the cave environment ( graening et al . , 2006 ) ; especially since they have been consumed by non - stygobitic predators outside the caves .
albinistic ; eyes without pigment and much reduced ; 2 terminal elements of first pleopod of male bent > 90 degrees , central projection ca . 135 degrees ; body strongly compressed ( hobbs and brown 1987 ) . [ length : to 28 . 5 tcl , to 45 tl ] [ width : to 12 ]
central projection at > 90 degree angle to main axis of pleopod , with notch in apex ; proximolateral groove near base of pleopod ( hobbs and brown 1987 ) .
reproductively active males in dec and jan ; no other data . spawning and actual mating periods unknown .
co - occurs with the rare ozark crayfish , amblyopsis rosae in one cave .
no data ; home range probably not over 50 m ; ability to survive outside cave paractically nil .
one of the caves from which this species is known is an ozarkian solution channel and species has been observed along the side walls of a pool or at the stream margin ( robison and allen , 1995 ) . cave streams in which this species lives are generally less than 50 cm deep .
this species was listed as a u . s . federal endangered species in 1993 and a recovery plan drafted ( usfws , 1996 ) . the primary conservation activity has been land protection . at one cave , usfws purchased 49 . 6 hectares containing the cave entrances and created a national wildlife refuge in 1989 . a cave gate was installed in 1998 by usfws and tulsa regional grotto , national speleological society . the nature conservancy is implementing a voluntary program to upgrate septic systems in the cave recharge zone to reduce nutrient loading of the aquifer . a pilot project at the prperty owner land involved the retrofitting of an old concrete spetic tank system with a recirculating , aerobic digestion trickle filter and fiberglass septic tank which should reduce nutrient and bacteria concentrations in the effluent . another property owner donated 1 . 7 ha of land containing the entrance to a second cave in 1998 to the nature conservancy establishing a natural area and a cooperative clean - up illegal refuse dumps in the discharge zone was performed . a cave gate was installed in the early 1990s . the entrance and surrounding land ( 4 . 1 ha ) of a third cave was donated to the nature conservancy , establishing another nature area and the nature conservancy has begun a reforestation effort of its discharge zone . site conservation and management pland for all 4 caves were developed by the nature conservancy . public outreach projects have not been implemented ( usfws , 1996 ; graening et al . , 2006 ) .
occurrences are based on some evidence of historical or current presence of single or multiple specimens , including live specimens or recently dead shells ( i . e . , soft tissue still attached without signs of external weathering or staining ) , at a given location with potentially recurring existence . evidence is derived from reliable published observation or collection data ; unpublished , though documented ( i . e . government or agency reports , web sites , etc . ) observation or collection data ; or museum specimen information .
separation barriers are based on hydrological discontinuity . additional physical barriers , particularly for secondary and tertiary burrowers , include presence of upland habitat between water connections of a distance greater than 30 m . migration of primary burrowers is generally not hindered by presence of upland habitat unless conditions are very xeric ( dry and desert - like ) ( smith , 2001 ) .
freshwater cave ( troglobitic ) species may occur from near entrances to very deep in cave systems . for cave species , each cave where an observation or collection was recorded ( see minimum eo criteria , above ) constitutes an element occurrence regardless of separation distance unless caves are part of a single hydrological system ( see below ) . occurrences are additionally separated by underground physical barriers to movement . multiple caves within a single hydrological cave system are considered to be a single element occurrence when they are less than one km apart . multiple caves within a single hydrological cave system are considered separate element occurrences when hydrological connections have not been determined or when separated by a distance of at least one km .
use the generic guidelines for the application of occurrence ranks ( 2008 ) . the key for ranking species occurrences using the generic approach provides a step - wise process for implementing this method .
zoological data developed by natureserve and its network of natural heritage programs ( see local programs ) and other contributors and cooperators ( see sources ) .
hobbs , h . h . , jr . 1989 . an illustrated checklist of the american crayfishes ( decapoda : astacidae , cambaridae , and parastacidae ) . smithsonian contributions to zoology 480 : 1 - 236 .
koppelman , j . b . and d . e . figg . 1995 . genetic estimates of variability and relatedness for conservation of an ozark cave crayfish species complex . conservation biology , 9 ( 5 ) : 1288 - 1294 .
mclaughlin , p . a . , d . k . camp , m . v . angel , e . l . bousfield , p . brunel , r . c . brusca , d . cadien , a . c . cohen , k . conlan , l . g . eldredge , d . l . felder , j . w . goy , t . haney , b . hann , r . w . heard , e . a . hendrycks , h . h . hobbs iii , j . r . holsinger , b . kensley , d . r . laubitz , s . e . lecroy , r . lemaitre , r . f . maddocks , j . w . martin , p . mikkelsen , e . nelson , w . a . newman , r . m . overstreet , w . j . poly , w . w . price , j . w . reid , a . robertson , d . c . rogers , a . ross , m . schotte , f . schram , c . shih , l . watling , g . d . f . wilson , and d . d . turgeon . 2005 . common and scientific names of aquatic invertebrates from the united states and canada : crustaceans . american fisheries society special publication 31 : 545 pp .
robison , h . w . and r . t . allen . 1995 . only in arkansas : a study of the endemic plants and animals of the state . university of arkansas press , fayetteville , arkansas .
taylor , c . a . , g . a . schuster , j . e . cooper , r . j . distefano , a . g . eversole , p . hamr , h . h . hobbs iii , h . w . robison , c . e . skelton , and r . f . thoma . 2007 . a reassessment of the conservation status of crayfishes of the united states and canada after 10 + years of increased awareness . fisheries 32 ( 8 ) : 371 - 389 .
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ridgely , r . s . , t . f . allnutt , t . brooks , d . k . mcnicol , d . w . mehlman , b . e . young , and j . r . zook . 2003 . digital distribution maps of the birds of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
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mclaughlin et al . 2005 . common and scientific names of aquatic invertebrates from the united states and canada : crustaceans . american fisheries society special publication 31
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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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 .
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the unicorn crestfish or unicornfish ( eumecichthys fiski ) is a very rare , little-known species of crestfish in the family lophotidae , and the only member of its genus .

it likely has a worldwide distribution , having been first discovered offshore of kalk bay , south africa , and subsequently reported from the sea of japan , southwest florida , clarion island off mexico , hawaii , and india .

a report from the bering sea may have been in error .

it is found in the bathypelagic zone , at a depth of around 1,000 m ( 3,300 ft ) .

this fish has ribbon-like body measuring up to 150 cm ( 59 in ) in length .

its common name comes from a horn-like supraoccipital process projecting forward of its eyes .

the upper jaw is protrusible , and the jaws contain small conical teeth .

the dorsal fin runs along the entire length of the body and contains 310-392 soft rays ; the first three to five dorsal rays at the tip of the projecting ridge are elongated into a pennant .

the pectoral fins contain 13-15 rays ; the pelvic fins are absent .

the anal fin contains five to 9 rays and in adults is split lengthwise to form two rows of nubbins .

the caudal fin contains 12-13 rays , with the bottommost ray enlarged and bony .

the coloration is silvery with 24-60 dark subvertical bands .

the dorsal and caudal fins are crimson .

eumenichthys is one of three lampriform genera known to possess ink tubes , allowing them to expel a black fluid from their cloacae as a defense against predators .

the ink tube is derived from a primitive gut and runs above and parallel to the intestine .

a known predator of the unicorn crestfish is the longnose lancetfish , ( alepisaurus ferox ) ; a lancetfish 73 cm ( 29 in ) long has been found that had swallowed a unicorn crestfish 55 cm ( 22 in ) long .

unicorn crestfish

the unicorn crestfish ( eumecichthys fiski ) is the only species in the . . .
a fossil unicorn crestfish ( teleostei , lampridiformes , lophotidae ) from the eocene of iran .
# fishaday †babelichthys olneyi unicorn crestfish from iran , named for the babelfish . ht @ gombessagirl urltoken
a fossil unicorn crestfish ( teleostei , lampridiformes , lophotidae ) from the eocene of iran . - pubmed - ncbi
rt @ hannahoish : # fishaday †babelichthys olneyi unicorn crestfish from iran , named for the babelfish . ht @ gombessagirl urltoken
head of a unicorn crestfish , eumecichthys fiski . source : eric woroch , us nmfs - piro observer program / wikimedia commons . license : public domain
a rare deepsea oarfish relative with an extremely long silvery ribbon - like body , a horn - like process extending froward on the head , bright red fins , and 24 - 60 darker vertical bands along the sides . as a defence against predators , the unicorn crestfish can expel black fluid from around the anus .
the unicorn crestfish ( eumecichthys fiski ) is the only species in the genus eumecichthys , which is one of just two genera in the oceanic family lophotidae ( the crestfishes ) . this species is rare and known only from the deep sea ( possibly around 1000 m ) , but is apparently widely distributed from south africa to india , japan , hawaii , and mexico . the head and body are silvery with 24 to 60 dark vertical bands and the fins are red . the highly elongated body may reach 130 cm or more ( body depth is around 1 / 25th of length ) . the first three or four dorsal rays are produced into a long , narrow pennant that extends far forward of the mouth . the dorsal fin includes 310 to 393 rays , the anal fin 5 to 9 rays , the pectoral fin 13 to 15 rays , and the caudal fin ( tail ) 12 to 13 rays . ( robins and ray 1986 ; heemstra 2003 ; froese and pauly 2011 ) the common name , unicorn crestfish , comes from the distinctive projecting supraoccipital , a bone on the dorsal ( upper ) side of the skull . a unique feature shared by members of the lophotidae ( lophotes and eumecichthys ) is the presence of an ink tube ( or ink sac ) . the ink tube allows its bearer to expel black fluid from the cloaca as a defense against predators ( robins and ray 1986 ; honma , ushiki , and takeda , 1998 ) .
lophotidae , or crestfishes , is a family of rare deep - sea teleosts characterised by an enlarged horn - like crest on the forehead . they are poorly represented in the fossil record , by only three described taxa . one specimen attributed to lophotidae has been described from the pelagic fauna of the middle - late eocene zagros basin , iran . originally considered as a specimen of the fossil lophotid † protolophotus , it is proposed hereby as a new genus and species † babelichthys olneyi , gen . et sp . nov . , differs from the other fossil lophotids by its relatively long and strongly projecting crest , suggesting a close relationship with the modern unicorn crestfish , eumecichthys . this new taxon increases the diversity of the deep - sea teleost fauna to which it belongs , improving our understanding of the taxonomic composition of the early cenozoic mesopelagic ecosystems .
in the present paper , † babelichthys olneyi , a new genus and species of lophotidae from the eocene of iran is described . few fossil representatives of taeniosomi , an elusive group of deep - sea teleosts , are known and only one of them has been previously described in detail ( bannikov , 1999 ) . † babelichthys is potentially the only currently known fossil close relative of the unicorn crestfish eumecichthys . this discovery is also significant because it expands the diversity of the middle - late eocene ilam fauna . modern lophotids are found in mesopelagic environments ( olney , 2002 ) , so the presence of at least two representatives of the family in the fauna that is mostly composed by relatives of modern deep - sea teleosts ( arambourg , 1967 ; afsari et al . , 2014 ; přikryl , brzobohatý & gregorová , 2016 ) reinforces its potential as a valuable glimpse of the otherwise poorly known early cenozoic deep - water ecosystems .
the distinction between an almost horizontal “crest” projecting anteriorly and a more vertical and relatively shorter “crest” distinguishes † babelichthys from † protolophotus ( see above , taxonomic justification ) , but also from the extant lophotus and the other known lophotid fossil taxa ( table 1 ) . conversely , in the eumecichthys specimen that is examined , the crest is strongly projected anteriorly ( angle of 72 . 4° ) and relatively very long ( table 1 ) . another element is the apparent absence of vomerine fang - like teeth in † babelichthys ( it is however possible that they were present , but not preserved in the fossil ) , like in eumecichthys , while they are present in lophotus ( olney , johnson & baldwin , 1993 ) . since only one specimen is available , it is impossible to perform a thorough comparison of head morphologies at various growth stages and between individuals . nevertheless , it seems on the basis of preserved elements that head morphology in † babelichthys is closer to the one observed in eumecichthys than in lophotus , corroborating the proposition of oelschläger ( 1979 ) that it represents a potential fossil sister group to eumecichthys . it would then be the first known fossil unicorn crestfish . nevertheless , † babelichthys also differs from eumecichthys : its crest is less strongly projecting and relatively shorter ( table 1 ) . moreover , no other lophotid , fossil or extant , has such an extreme enlargement and expansion of the dorsal - fin pterygiophores , the second one in particular .
lampridiforms are strange spiny - rayed teleosts , found in mesopelagic environments in every ocean of the world ( olney , johnson & baldwin , 1993 ; olney , 2002 ) . their most famous representatives are the endothermic opah ( lampris guttatus ) and the gigantic , serpentine oarfish ( regalecus glesne ) , the longest known teleost . along with these iconic taxa , lampridiforms include equally weird ribbon - like and elongate animals , characterized by a silver - coloured skin and long , bright red fins : the taeniosomes . the 15–18 extant species of the clade taeniosomi include oarfishes ( regalecidae ) , ribbonfishes ( trachipteridae ) , the tapertail ( radiicephalidae ) and lophotidae , the crestfishes ( regan , 1907 ; walters & fitch , 1960 ; olney , 1984 ; roberts , 2012 ) . lophotids are characterized by unique anatomical structures , such as an ink gland ( walters & fitch , 1960 ; honma , ushiki & takeda , 1999 ) not found anywhere else in teleosts ( except in the closely related radiicephalids ; harrisson & palmer , 1968 ) . the most conspicuous osteological feature of lophotids is a well - developed horn - like crest , formed by an anteriorly projecting expansion of the frontal and supraoccipital bones of the cranium ( oelschläger , 1979 ; oelschläger , 1983 ; olney , johnson & baldwin , 1993 ) . this crest is closely associated with the anterior pterygiophores supporting the dorsal fin , and as a result , the dorsal fin expands over , and sometimes anterior to the cranium . lophotids are represented in modern fauna by one to three lophotus species and by the unicorn crestfish , eumecichthys fiski ( walters & fitch , 1960 ; craig , hastings & pondella , 2004 ) . their fossil record consists in at least three monotypic genera ( bannikov , 1999 ; carnevale , 2004 ) . the present article is a revision of an anatomically distinctive fossil specimen attributed to lophotidae . arambourg ( 1943 ) and arambourg ( 1967 ) first described the specimen from a rich late eocene fauna located near ilam , zagros basin , iran . the ilam fauna comprises numerous representatives of teleost taxa such as beryciformes , gadiformes , ophidiiformes and stomiiformes , typical of the modern deep - sea pelagic environments ( arambourg , 1967 ; afsari et al . , 2014 ; přikryl , brzobohatý & gregorová , 2016 ) .
probably worldwide . western atlantic : southeastern florida in usa . southeast atlantic : false bay , south africa ( ref . 4165 ) . northwest pacific : japan ( ref . 559 ) . eastern central pacific : hawaii and mexico ( ref . 4165 ) . indian ocean : india ( ref . 4165 ) .
maturity : l m ? range ? - ? cm max length : 150 cm tl male / unsexed ; ( ref . 7251 )
dorsal spines ( total ) : 0 ; anal spines : 0 ; anal soft rays : 5 - 9 . head and body silvery in color with 24 - 60 dark sub vertical bands ; dorsal and caudal fins crimson in color ( ref . 4165 ) . dorsal fin with 310 - 392 soft rays .
a rare ( ref . 4165 ) , mesopelagic species found at 1 , 000 m depth ( ref . 5213 ) .
heemstra , p . c . , 1986 . lophotidae . p . 402 - 403 . in m . m . smith and p . c . heemstra ( eds . ) smiths ' sea fishes . springer - verlag , berlin . ( ref . 4165 )
) : 7 . 7 - 15 . 1 , mean 10 . 6 ( based on 338 cells ) .
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 1 . 0625 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 00115 ( 0 . 00043 - 0 . 00306 ) , b = 3 . 07 ( 2 . 84 - 3 . 30 ) , in cm total length , based on lwr estimates for this ( sub ) family - body shape ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 4 . 1 ±0 . 6 se ; based on size and trophs of closest relatives
resilience ( ref . 69278 ) : very low , minimum population doubling time more than 14 years ( ) .
vulnerability ( ref . 59153 ) : very high vulnerability ( 80 of 100 ) .
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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 : this species is likely distributed worldwide to depths of 1 , 000 m . there are no known species - specific threats and there are no known species - specific conservation measures . this species is assessed as least concern .
this species is likely distributed worldwide . in the western atlantic it is known from southeastern florida in usa . it has also been described from southern and central brazil . in the southeastern atlantic this species is known from false bay , south africa ( heemstra 1986 ) and from the gulf of guinea . in the northwest pacific this species is known from japan ( masuda et al . 1984 ) . in the eastern central pacific , this species is known from hawaii and mexico ( heemstra 1986 ) . in the indian ocean , it is known from india ( heemstra 1986 ) . it is found to depths of 1 , 000 m .
angola ; anguilla ; antigua and barbuda ; bahamas ; barbados ; bermuda ; brazil ; british indian ocean territory ; christmas island ; cocos ( keeling ) islands ; comoros ; congo , the democratic republic of the ; dominica ; dominican republic ; equatorial guinea ; france ( clipperton i . ) ; french guiana ; french southern territories ; gabon ; guadeloupe ; guatemala ; guyana ; india ( andaman is . , nicobar is . ) ; indonesia ; iran , islamic republic of ; japan ; kenya ; kiribati ( kiribati line is . , phoenix is . ) ; madagascar ; maldives ; martinique ; mauritius ; mayotte ; mexico ; mozambique ; myanmar ; namibia ; nigeria ; oman ; pakistan ; philippines ; russian federation ( central asian russia , kuril is . ) ; saint kitts and nevis ; saint martin ( french part ) ; sao tomé and principe ; seychelles ; sint maarten ( dutch part ) ; somalia ; south africa ; sri lanka ; suriname ; taiwan , province of china ; tanzania , united republic of ; trinidad and tobago ; turks and caicos islands ; united states ( hawaiian is . ) ; united states minor outlying islands ( howland - baker is . , johnston i . ) ; virgin islands , british ; virgin islands , u . s . ; yemen
there is little species - specific population information available for e . fiski . it is represented by six lots in museum collections ( fishnet2 database searched june 2014 ) .
this is a rare mesopelagic species which is found to depths of 1 , 000 m ( fischer et al . 1990 ) .
there is no species - specific use and trade information available for this species .
there are no known species - specific conservation measures in place for e . fiski . it is likely found in marine protected areas throughout its range .
fischer , w . , sousa , i . , silva , c . , de freitas , a . , poutiers , j . m . , schneider , w . , borges , t . c . , feral , j . p . and massinga , a . 1990 . fichas fao de identificaçao de espécies para actividades de pesca . guia de campo das espécies comerciais marinhas e de águas salobras de moçambique . publicaçao preparada em collaboraçao com o instituto de investigaçao pesquiera de moçambique , com financiamento do projecto pnud / fao moz / 86 / 030 e de norad . food and agricultural organization of the united nations ( fao ) , rome , italy .
heemstra , p . c . 1986 . trachichthyidae . in : m . m . smith and p . c . heemstra ( eds ) , smith’s sea fishes , pp . 410 - 413 . springer - verlag , berlin , germany .
iucn . 2015 . the iucn red list of threatened species . version 2015 - 4 . available at : urltoken . ( accessed : 19 november 2015 ) .
masuda , h . , amaoka , k . , araga , c . , uyeno , t . and yoshino , t . 1984 . the fishes of the japanese archipelago . tokai university press , tokyo , japan .
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in australia , the only known specimen was washed ashore at cheynes beach , albany , in 1989 . elsewhere the species is very rare , but thought to occur worldwide in depths of around 1000 m .
lophotes fiski günther 1890 , proc . zool . soc . london 1890 ( 2 ) : 244 , pls 19 - 20 . type locality : cape of good hope , kalk bay , south africa .
bray , d . j . 2008 . family lophotidae . pp . 298 in gomon , m . f . , bray , d . j . & kuiter , r . h . ( eds )
. the iucn red list of threatened species 2015 : e . t190107a60791470 . urltoken downloaded on 06 may 2017 .
olney , j . e . 1999 . families veliferidae , lamprididae , stylephoridae , lophotidae , radiicephalidae , trachipteridae , regalecidae . pp . 1966 - 1975 in carpenter , k . e . & niem , v . h . ( eds )
the living marine resources of the western central pacific . fao species identification guide for fisheries purposes
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this is an open access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , reproduction and adaptation in any medium and for any purpose provided that it is properly attributed . for attribution , the original author ( s ) , title , publication source ( peerj ) and either doi or url of the article must be cited .
the taxonomic status of the lophotid specimen studied here is currently unclear ( walters , 1957 ; oelschläger , 1979 ; bannikov , 1999 ) , and it lacks a proper anatomical description . given the rarity of fossil material attributed to taeniosome lampridiforms , a detailed description and revised taxonomy of this material is needed in order to improve our understanding of the morphological evolution and fossil record of this peculiar group .
the material described herein , mnhn . f . eip11 ( figs . 1 and 2 ) , was discovered during excavations near ilam ( zagros basin , western iran ) by camille arambourg in 1938–1939 . the specimen was chosen to be the paratype of † lophotes elami ( arambourg , 1943 ) , along with the holotype mnhn . f . eip10 ( fig . 3 ) . on the basis of osteological differences from extant lophotids , such as the well - ossified pelvic girdle in a ventral position observed in the holotype of † lophotes elami , walters ( 1957 ) assigned it to a distinct new genus † protolophotus ( fig . 3 ) . oelschläger ( 1979 ) later proposed that mnhn . f . eip11 differs sufficiently from mnhn . f . eip10 to be classified in a different genus . he related the specimen to the extant eumecichthys and gave it the name †‘ protomecichthys ’ . however , the genus †‘ protomecichthys ’ lacks both a designated type species and a formal description . thus , it fails to meet the requirements of article 13 . 3 of the international code of zoological nomenclature ( international commission on zoological nomenclature , 1999 ) and should be considered a nomen nudum ( bannikov , 1999 ) .
( a ) mnhn . f . eip11d . ( b ) counterpart mnhn . f . eip11g . scale bars = 20 mm .
photograph ( detail of the head ) and interpretative drawing . legend : achy , anterior ceratohyal ; bra , branchiostegal ; bsp , basisphenoid ; den , dentary ; dfr , dorsal - fin ray ; dhhy , dorsal hypohyal ; dpt , dorsal - fin pterygiophore ; enpt , endopterygoid ; fr , frontal ; hyo , hyomandibula ; iop , interopercle ; lac , lachrymal ; let , lateral ethmoid ; mpt , metapterygoid ; mx , maxilla ; osp , orbitosphenoid ; pal , palatine ; pchy , posterior ceratohyal ; pmx , premaxilla ; pop , preopercle ; psp , parasphenoid ; qu , quadrate ; soc , supraoccipital ; soc - sp , spine of the supraoccipital ; spl , splint of the first dorsal - fin ray ; vhhy , ventral hypohyal ; vo , vomer . scale bar = 10 mm .
† eolophotes lenis , pin 1413 / 86 ; eumecichthys fiski , usnm 164170 ( radiographs ) ; lophotus lacepede , nhmuk 1863 . 8 . 27 . 1 ( radiographs ) ; † oligolophotes fragosus , pin 3363 / 121 ; † protolophotus elami , mnhn . f . eip10 .
the electronic version of this article in portable document format ( pdf ) will represent a published work according to the international commission on zoological nomenclature ( iczn ) , and hence the new names contained in the electronic version are effectively published under that code from the electronic edition alone . 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 : b677ba4f - ccf4 - 4678 - a8a8 - 502f059704d2 . the online version of this work is archived and available from the following digital repositories : peerj , pubmed central and clockss .
the specimen was examined with a stereomicroscope equipped with a camera lucida drawing arm . the interpretative drawing was produced with adobe illustrator cs6 from the camera lucida drawings and from photographs . measurements were taken with a compass or with the software imagej 1 . 5 from radiographs ; angles were also measured with imagej . the method for estimating the degree of projection of the crest is modified from craig , hastings & pondella ( 2004 ) : it is based on the angle between the straight line from the tip of the crest to the proximal end of its anterior margin ( instead of the tip of the upper jaw , due to varying jaw positions in fossils ) and the vertical line drawn perpendicular to the main axis of the parasphenoid ( instead of the vertebral column , not preserved in mnhn . f . eip11 ) . the relative length of the crest is the ratio between the crest length ( distance between the tip of the crest and the proximal end of its anterior margin ) and the head length without the crest ( from the anterior margin of the ethmoid region to the posterior margin of the neurocranium ) . all extinct taxa are indicated with a dagger ( † ) .
urn : lsid : zoobank . org : act : 86986e5e - 5fff - 465d - a987 - e475fbf02966
etymology . hellenization of the name of the “babel fish” , the teleost - like , ear - dwelling , polyglot extra - terrestrial species from douglas adams’ 1979 book the hitchhiker’s guide to the galaxy , in reference to the very peculiar , almost alien - like , appearance of the genus .
diagnosis . a lophotid differing from † eolophotes , lophotus , † oligolophotes and † protolophotus by the relatively longer , strongly projecting crest ; and from eumecichthys by the relatively shorter , deeper and less strongly projecting crest , with strongly expanded anterior dorsal - fin pterygiophores .
urn : lsid : zoobank . org : act : d2540d1f - f169 - 40de - b910 - 7302810615e7
1943 † lophotes elami arambourg , p . 287 , pl . x , fig . 1
holotype . mnhn . f . eip11d / g , almost complete articulated cranium and anterior portion of the dorsal fin , in part and counterpart ( figs . 1 and 2 ) . this is the only specimen known for the genus and species .
etymology . species named in honour of the late john e . olney , in recognition of his work on the anatomy and ontogeny of lampridiformes .
type locality and horizon . near ilam , zagros basin , western iran . this teleost fauna , part of the pabdeh formation , was erroneously aged cretaceous by priem ( 1908 ) , and rupelian ( oligocene ) by arambourg ( 1943 ) and arambourg ( 1967 ) . it is more accurately middle to late eocene in age ( afsari et al . , 2014 ; and references therein ) .
mnhn . f . eip11 consists only of the head of the animal , along with the associated anterior portion of the dorsal fin . the specimen is mostly articulated , except for the left ventral portion of the hyoid arch that is upturned and preserved ventral to the rest of the cranium . the limits of most bones are poorly preserved , probably due to their low degree of mineralization in life as is the case in modern taeniosome lampridiforms .
total head length : 104 mm ; head length ( without the crest ) : 44 mm ; crest length ( anterior margin ) : 51 . 5 mm ; head depth : 25 . 5 mm ; orbit diameter : 23 mm .
the neurocranium of mnhn . f . eip11 is highly modified . the frontal develops a dorsal lamina that projects anterior to the jaws . throughout approximately its anterior half , it is in contact with an enlarged laminar process of the supraoccipital , delimited dorsally by a strong supraoccipital spine . together , they form a conspicuous “crest” , long and strongly projecting anteriorly ( at an angle of 64 . 5° ) . alone , the crest contributes to 58 % of total head length .
the frontal makes up approximately 60 % of the anterior margin of the crest . both the frontal and the supraoccipital show radial ornamentation on the crest ; it radiates from the posterior end of the frontal and the distal tip of the supraoccipital . the supraoccipital spine borders the dorsal margin of the bone , and narrows towards the tip .
the ethmoid region is poorly preserved , with an probable enlarged lateral ethmoid that hides the mesethmoid . an enlarged lachrymal is nested in the antero - ventral corner of the orbit ; it is parallel to the parasphenoid ventrally , and curves dorsally along the posterior edge of the lateral ethmoid . the orbitosphenoid runs along the dorsal margin of the orbit and has a conspicuous process pointing ventrally . the posterior wall of the orbit is delimited ventrally by a robust and straight basisphenoid . otherwise , the sphenoid , otic and occipital regions are too poorly preserved to distinguish the individual bones . the parasphenoid is robust and slightly curves dorsally at its anterior end . the junction between the parasphenoid and the vomer is not discernable . there is no evidence of vomerine teeth .
the premaxilla is relatively small , with a well - developed ascending process , and a barely visible alveolar process . the maxilla bears a conspicuous and pointed process at its antero - dorsal end . the posterior end is expanded dorsoventrally , forming a rounded lamina . neither the premaxilla nor the maxilla bear visible teeth . there is no evidence of a supramaxilla . the anterior margin of the dentary , slightly concave and bearing no visible teeth , forms a strong angle with the ventral margin . the posterior margin of the dentary forms an interosseous space with the anguloarticular , which is mostly hidden by overlaying bones .
only the proximal , single - headed articulation of the hyomandibula is clearly visible ; the distal end of the bone seems to be preserved in close association with the metapterygoid . the latter is roughly triangular and is one of the best preserved bones of the suspensorium . the symplectic is rod - like , narrows slightly anteriorly and inserts in a notch on the postero - ventral margin of the quadrate . the triangular quadrate bears an antero - ventral condyle that articulates with the anguloarticular . the anterior portion of the suspensorium is poorly preserved , and it is difficult to outline the limits of the endopterygoid , ectopterygoid and palatine bones . the dorsal and posterior portions of the endopterygoid are preserved , suggesting that the bone forms two laminae , the dorsal one along the parasphenoid , and the ventral one contacting both the quadrate and the metapterygoid .
both the left and right ventral hyoid arches are visible . one is preserved in life position : its posterior end overlaps the operculum , but its dorsal margin is hidden by the lower jaw , suggesting it corresponds to the right ventral hyoid arch . the left one is displaced and upturned , and lies ventral to its counterpart . the posterior ceratohyal is triangular and articulates with the anterior ceratohyal with an interdigitated suture . the anterior ceratohyal shows a strong ventral concavity at midlength ; its dorsal margin is much less concave . the anterior end of the anterior ceratohyal forms a rounded condyle , over which the curved ventral hypohyal articulates . the dorsal hypohyal lies dorsally over the anterior ceratohyal . there are six branchiostegals : the anterior two are shorter and articulate with the anterior ceratohyal at the level of its ventral concavity ; the four others articulate more posteriorly ( due to the faint distinction between both ceratohyals , it is difficult to determine on which one they articulate ) ; they are very long ( the posteriormost being the longest ) and curved posteriorly over the ventral margin of the interopercle . the branchiostegals of the left hyoid arch are disarticulated .
the preopercle is wide and angled at mid - length . the interopercle is an elongate bone rounded at its extremity that forms the ventral margin of the opercular series . it has a smooth ventral margin , closely associated with the posterior branchiostegals . the potential presence of parts of the opercle , in contact with the preopercle , is unclear .
the dorsal fin is only partially preserved , with only the most anterior pterygiophores and dorsal - fin rays visible . its most striking feature is the extremely elongated and enlarged first dorsal - fin ray , which is 10 times as wide as the more posterior fin rays ( at their base and greatest width ) . it does not bifurcate distally , lacks any visible segmentation , and a groove runs throughout its length . a rounded splint protrudes at its anterior base ; it is unclear whether it constitutes a separate dorsal - fin element or not . fifteen other dorsal - fin rays are preserved posteriorly . their distal end is missing in most cases , but they all seem to be of a similar length , except for the second and third dorsal - fin rays that are noticeably longer . they do not bifurcate distally , and no segmentation is clearly visible .
ten dorsal - fin pterygiophores are unambiguously preserved . they are strongly inclined anteriorly , which causes the dorsal fin to originate at the tip of the crest , and to run along the entire head of the animal . the first two dorsal - fin pterygiophores are greatly enlarged and in close contact with the crest . both also show a conspicuous flange at their posterior margin . the first pterygiophore is narrow posteriorly , where it does not contact the supraoccipital , and widens in its distal end . the second one is much wider and slightly narrows at its distal extremity . it is in close contact with the first pterygiophore throughout its entire length . the third and fourth preserved pterygiophores are in close contact with the second one throughout almost all of their lengths . the more posterior pterygiophores have a mostly straight shaft that curves slightly at its distal extremity . the most posterior ones are less inclined than the anterior ones . the proximal ends of all preserved pterygiophores converge at the same point : the base of the crest—thus they insert anterior to the ( not preserved ) first neural spine . the elongated and enlarged first dorsal - fin ray inserts on the first pterygiophore . it is unclear if the rays two to eight insert on pterygiophores that are mostly hidden or not preserved , or in supernumerary association with the enlarged second pterygiophore . the rays 9–16 each insert serially on a corresponding pterygiophore .
oelschläger ( 1979 ) proposed that mnhn . f . eip11 is different enough anatomically from the other lophotids , fossil and extant , to justify its attribution to a new genus . indeed , it differs from the holotype of † protolophotus , found in the same geological levels , by the relative development of the crest . in mnhn . f . eip11 , the crest is projecting anteriorly with an angle of 64 . 5° , and the ratio between the lengths of the crest’s anterior margin and of the head without the crest is of 1 . 17 to 1 . in the holotype of † protolophotus , mnhn . f . eip10 ( fig . 3 ) , the anterior margin of the crest is almost vertical ( degree of projection : 20° ) , and it is relatively shorter ( margin of the crest / head length without the crest : 0 . 67 / 1 ) . mnhn . f . eip11 also shows a much stronger first dorsal - fin ray , and its two anterior dorsal - fin pterygiophores are much more developed . body size is known to affect relative crest size and degree of projection in extant lophotus ( craig , hastings & pondella , 2004 ) , which could be misleading when trying to differentiate taxa based on crest morphology . however , this bias can probably be ruled out in the case of mnhn . f . eip11 and mnhn . f . eip10 : both individuals have similar head lengths without the crest ( 42 and 44 mm , respectively ) , suggesting that they are at a similar growth stage . it then seems that classifying mnhn . f . eip11in a different genus and species , † babelichthys olneyi , is justified from a morphological point of view .
the monophyly of lampridiformes ( excluding stylephorus , sensu nelson , grande & wilson , 2016 ) is well - supported by molecular phylogenetic analyses ( wiley , johnson & dimmick , 1998 ; miya et al . , 2007 ; betancur et al . , 2013 ; near et al . , 2013 ) and by numerous morphological features ( olney , johnson & baldwin , 1993 ; davesne et al . , 2014 ; davesne et al . , 2016 ; delbarre , davesne & friedman , 2016 ) . several of these character states are unambiguously found in † babelichthys : the premaxilla and dentary are toothless , the frontal and the supraoccipital are both involved in a sagittal crest , the anterior ceratohyal forms a condyle that articulates with the ventral hypohyal , and the first dorsal - fin pterygiophore inserts anterior to the neural spine of the first abdominal vertebra .
the phylogenetic studies that explore lampridiform intrarelationships with a sufficient sampling all recover a monophyletic taeniosomi ( wiley , johnson & dimmick , 1998 ; grande , borden & smith , 2013 ; martin , 2015 ) . the taeniosome character states found in † babelichthys include the absence of supraneurals , and anterior dorsal - fin pterygiophores that are enlarged and inclined over the neurocranium ( olney , johnson & baldwin , 1993 ) . † babelichthys then clearly shows a character state combination that confirms its identification as a taeniosome lampridiform .
olney , johnson & baldwin ( 1993 ) proposed that the enlarged supraoccipital process , projecting anteriorly over the frontals ( forming the “crest” as described herein ) and supporting the first dorsal - fin pterygiophore , is a synapomorphy of lophotidae . since it is not found elsewhere in lampridiforms , this character confirms the attribution of † babelichthys to lophotidae . it has to be noted that in the yet unpublished phylogenetic analysis of martin ( 2015 ) , the monophyly of lophotidae is ambiguous , with one parsimonious tree finding lophotus more closely related to the other taeniosomes than to eumecichthys , while in the other both genera are sister groups . given this ambiguity , lophotidae is considered monophyletic in this discussion .
angle ( ° ) between the straight line from the tip of the crest to the proximal end of its anterior margin and the line drawn perpendicular to the main axis of the parasphenoid .
distance ( mm ) between the tip of the crest to the proximal end of its anterior margin .
distance ( mm ) between the anterior margin of the ethmoid and the posterior margin of the neurocranium .
taeniosome lampridiforms are known by several fossil representatives . the oldest unquestionable occurrences are all attributed to lophotidae : the diminutive † eolophotes lenis ( fig . 4a ) , from the lutetian ( eocene ) of georgia ( daniltshenko , 1962 ; daniltshenko , 1980 ) and † protolophotus elami ( fig . 3 ) , found in the same middle - late eocene formation as † babelichthys ( see above ) . an additional , younger fossil lophotid is † oligolophotes fragosus ( fig . 4b ) from the early oligocene pshekha formation of adygea , northern caucasus , russia ( bannikov , 1999 ) . the taeniosome fossil record also includes the trachipterid † trachipterus mauritanicus from the messinian ( late miocene ) of algeria ( carnevale , 2004 ) , and a fragmentary possible oarfish ( regalecus ) from the pliocene of italy ( bronzi , 2001 ; roberts , 2012 ) . there is no known fossil radiicephalid . finally , the small and distinctive † bajaichthys elegans , from the ypresian ( early eocene ) of bolca , italy , has been classified as a taeniosome or close relative due to its mobile jaws , elongate body and reduced caudal fin ( sorbini & bottura , 1988 ; bannikov , 2014 ) . however , it can be confidently classified in zeiformes , a separate teleost clade ( davesne , carnevale & friedman , 2017 ) . in total , five entirely fossil taeniosome species are currently known ( four lophotidae , one trachipteridae ) , a diversity expanded by the present description of † babelichthys .
( a ) † eolophotes lenis , holotype pin 1413 / 86 ; scale bar = 5 mm . ( b ) † oligolophotes fragosus , holotype pin 3363 / 121 ; scale bars = 10 mm .
national museum of natural history , smithsonian institution , washington d . c . , united states
the author thanks gaël clément ( mnhn ) for allowing access to the specimen of study , alexandre bannikov ( pin ) for his useful information on the lampridiform fossil record and for suggesting the redescription of this specimen , as well as giorgio carnevale ( università degli studi di torino ) for insightful anatomical discussion . sandra raredon ( usnm ) kindly sent radiographs of extant lophotids for comparison , while lilian cazes , philippe loubry ( mnhn ) and av mazin ( pin ) provided high - quality photographs of the fossil specimens . finally , the editor jérémy anquetin and referees alexandre bannikov and dave johnson are thanked for their invaluable reviews and comments .
donald davesne conceived and designed the experiments , performed the experiments , analyzed the data , wrote the paper , prepared figures and / or tables , reviewed drafts of the paper .
the measurements in table 1 are the raw data . the material described herein , mnhn . f . eip11 is stored at mnhn , muséum national d’histoire naturelle , paris , france .
publication lsid : urn : lsid : zoobank . org : pub : b677ba4f - ccf4 - 4678 - a8a8 - 502f059704d2 .
babelichthys gen . nov . : urn : lsid : zoobank . org : act : 86986e5e - 5fff - 465d - a987 - e475fbf02966 ,
babelichthys olneyi sp . nov . : urn : lsid : zoobank . org : act : d2540d1f - f169 - 40de - b910 - 7302810615e7 .
the author was supported by the natural environment research council ( grant ne / j022632 / 1 ) and by the leverhulme trust ( grant rpg - 2016 - 168 ) . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
résultats scientifiques de la mission c . arambourg en syrie et en iran ( 1938–1939 ) . ii . les poissons oligocènes de l’iran
ontogeny and systematics of fishes , special publication number 1 , american society of ichthyologists and herpetologists .
the living marine resources of the western central atlantic . volume 2 : bony fishes part 1 ( acipenseridae to grammatidae ) .
systematics , biology and distribution of the species of the oceanic oarfish genus regalecus ( teleostei , lampridiformes , regalecidae ) .
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yearsley , g . k . , last , p . r . & morris , g . b . 1997 ,
codes for australian aquatic biota ( caab ) : an upgraded and expanded species coding system for australian fisheries databases
, pp . 15 pp . + appendices
günther , a . 1890 ,
description of a new species of deep - sea fish from the cape ( lophotes fiski )
, proceedings of the zoological society of london , vol . 1890 , no . 2 , pp . 244 - 247 pls 19 - 20
urn : lsid : biodiversity . org . au : afd . taxon : 300d6bfc - 8d7c - 4324 - be6d - 169fccc93ccc
urn : lsid : biodiversity . org . au : afd . taxon : 7f8740be - b135 - 4e67 - 853b - 775fec3e7b63
urn : lsid : biodiversity . org . au : afd . name : 342582
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pmid : 28674642 pmcid : pmc5493034 doi : 10 . 7717 / peerj . 3381
† babelichthys olneyi , gen . et sp . nov . holotype mnhn . f . eip11d .
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