the dugite / ˈdjuːɡaɪt / ( pseudonaja affinis ) is a species of venomous , potentially lethal , snake native to western australia , a member of the family elapidae .

the local nyungar name for the dugite is dobitj .

dugite

dugite - native / package . json at master · desktop / dugite - native · github
perth snake catcher’s joust with fiery 1 . 5m dugite at spearwood factory | perthnow
the dugite is not very aggressive . a member of the brown snake family , the dugite is nowhere near as venomous as its cousin , the eastern brown snake .
dugite snake image by rob ahern - some rights reserved . ( view image details )
workers at the factory spotted a 1 . 5m dugite coiled up outside their kitchen quarters .
snake catcher steve smartt with the fiesty 1 . 5 metre dugite which he later released in bushland .
but the dugite at ms cross ' parents ' home was the first of its kind for the season .
like other brown snakes , dugites are diurnal . the female dugite lays 10 to 20 eggs at a time .
common dugite pseudonaja affinis affinis - notice the broad belly scales on the shed skin . ( photo taken at the west australian reptile park ) an extremely large common dugite pseudonaja affinis affinis - ( photo taken at the west australian reptile park )
the pictures show the dugite wrapping its mouth around the tiger snake ' s body , slowly swallowing the venomous victuals whole .
‘a couple of hours later we saw the culprit by our pond - a 4 ft dugite slithering across the lawn . ’
the fact that the mainland dugite is common in degraded habitats and feeds on house mice suggests that it has a secure future .
kyle releases a rescued brown snake . the dugite is a deadly species of brown snake found in south - western parts of australia .
‘there are several species , the most important medically being the eastern or common brown , the western brown or gwardar and the dugite . ’
dugites are cannibalistic . they eat members of their own species as well as other snakes . an individual dugite can swallow another snake almost as large as itself !
snake handler marcus cosentino , from slithers and sliders who bagged the dugite from mrs cross ' parents ' abode said the area around city beach was notorious for snakes .
well , that question has been settled , sort of , after pictures emerged of a dugite eating a tiger snake near capel in western australia ' s south west .
ms hedzik said she returned to the site later to see if the dugite was still working its way through the tiger snake , but there was no sign of it .
other similar species of brown snake that overlap the dugite’s distribution include the penisular brown snake pseudonaja inframacula , the western brown snake pseudonaja nuchalis and the eastern brown snake pseudonaja textilis .
the 35 - year - old , who is eight months pregnant , said the one metre dugite bolted as soon as it spotted her so she quickly rang a snake handler .
mrs cross , who was aware how deadly a bite from a dugite can be , said at no stage did she fear for the safety of herself or her unborn bub .
they are protected under the australia’s wildlife conservation act 1950 . this means if someone tries to injure or kill a dugite , they can be fined up to aud $ 4000 .
the dugite is a type of brown snake , which is very common around perth and surrounding areas . it is found through the south of western australia through the nullarbor plain and eyre peninsula
in the wild , the dugite eats a variety of vertebrate prey including frogs , lizards , snakes , birds and mammals . however , the two largest groups of prey are lizards and house mice .
the dugite is a highly venomous snake . the adult usually olive green to brown , often with dark spots or flecks . the abdomen may also be dark , unlike that of other brown snakes .
well - known to south - western wa residents , the dugite has made itself at home around urban and semi - rural areas , drawn to the prevalence of its favoured prey – the house mouse .
a heavily pregnant mother - of - one said when she spotted a dugite scurrying across the driveway of her parents ' home , her first thought was to protect her two - year - old daughter .
tiger snake venom activates prothrombin in presence of factor v and calcium , whereas the western brown , brown , dugite , and taipan snakes directly activate prothrombin and may result in very low fibrinogen levels and severe biochemical coagulopathy .
the dugite is a dangerous snake . the venom is neurotoxic although the dugite rarely envenoms when biting people . most people seen in hospital after an attack do not require anti - venom treatment . dugites will avoid people when they can , but will attack if surprised , particularly during mating period . venomous bite symptoms include abdominal pain , breathing and swallowing difficulty , convulsions , hypotension , kidney failure . can be fatal if no anti - venom treatment .
dugites are diurnal , meaning that they aer most active during the day . this is quite common with other brown snakes . in terms of reproduction , a female dugite can lay between 10 to 20 snake eggs at one time .
the dugite is one of nine described species and two subspecies of brownsnake ( genus pseudonaja ) found in australia . the juveniles of all of these are very similar , sharing the dark head - blotch ( or blotches ) . many individuals are adorned with numerous dark crossbands .
the dugite may fall victim to raptors , monitor lizards , feral and domestic cats , and other snakes ( including its own species ) . although vertebrates are probably the main predators of this snake , there is evidence to suggest that spiders occasionally snare and feed on small specimens .
a dugite is an australian species of snake that is highly venomous that can inflict death in simply one bite . it is sometimes refered to as a brown snake and was first described by albert gunther in 1872 . there are 3 sub - species of dugites , and they are :
the dugite was bigger than the tiger - i thought it would have been the other way around , but he must have moved off , even with that big meal . i think anyone who came across that would have done what i ' d done , anyone would be rapt to see that .
as mentioned above , the dugite is an extremely venenous snake . in fact , it is one of the most lethal in the world as it causes coagulopathic and precogulant effects ( meaning that the blood will clot and therefore the victim will die ) . despite this , dugites do avoid biting humans but humans are at risk as the numbers of dugite - human encounters rise when they are more active during the mating season which runs during october and november . in fact , the last death that was attributed to this snake was in 1993 , when an elderly male died in spearwood which is one of perth’s southern suburbs .
because of the snake’s size and highly toxic venom the dugite is considered to be very dangerous to humans . its prevalence in residential areas and nervous disposition have helped make it responsible for approximately 70 % of all snake bites reporting to perth hospitals , but thanks to prompt and effective intervention , there has been only one recorded fatality .
dugites are either brown , green , or grey . however , the colors vary between individual snakes . however , the most distinguishable feature of a dugite is the shape of its head . it is quite small compared to the neck , as it grades gradually into the body . they can grow up to 2m ( 6 . 56 feet ) long .
although naturally shy , the dugite is easily agitated when confronted and will raise its forebody upright in a tight s - shape . it hisses loudly before making a fast snappy strike at the offender , usually aiming high . fortunately the fangs are very small and may not effectively penetrate solid shoes or heavy fabric ; however anyone with a suspected bite should seek immediate medical attention .
description : the rottnest island dugite is genetically different from the mainland population and is generally smaller than the mainland version . it is blackish brown all over . it has a long , slim body with a small head . its blackish brown eyes are large and have a golden orange rim . there is also a defined ridge above the eye . its scales are quite large for its body and give it a semi - glossy look .
pseudonaja species live mainly in australia . about 3000 bites occur per year in australia from all species of snakes , 500 of which require antivenin . the dugite brown snake may be found in the southwestern corner of australia , in western australia and along the south australian border . the speckled brown snake can be found from central queensland to the eastern areas of the northern territory . ingram ' s snake may be found around barkly tableland of the northern territory , and the ringed snake may be found in the arid regions of all the mainland states . the western snake may be found throughout australia and the common brown snake in queensland , new south wales , and from victoria to the southeast of south australia .
the brown snake may be found all over australia . it has extremely potent venom , and although the quantity of venom injected is usually small , this snake causes more snakebite deaths in australia than any other . sudden and relatively early deaths have been recorded . its venom causes severe coagulation disturbances , neurotoxicity , and occasionally nephrotoxicity ( by a direct action of the venom ) , but not rhabdomyolysis . the gwardir is also known as the western brown snake , and the dugite is a spotted brown snake found in western australia . all need brown snake antivenom . see also venom supplies brown snake pages , emedicine brown snake guidelines , urltoken , avru brown snake info and treatment , australian reptile park , wikipedia . some more local pics :
the species occupies a wide variety of habitats including coastal dunes , heathlands , shrublands , woodlands and forests . it seems to cope well with heavily degraded habitats such as golf course , industrial parks and open agricultural country ; in fact , it is one of the few reptile species that has probably increased in numbers with the opening up of natural habitats and the introduction of the house mouse . around the perth area , the dugite is one of the most common reptiles found near buildings . in areas of human habitation the snakes take temporary shelter under refuse such as concrete slabs , fibro sheets , roofing tin and the like , but in more “natural” surroundings they will shelter under rocks and in abandoned termite mounds , abandoned stick ant nests , and rabbit and rodent burrows . dugites also use burrows for overwintering .
the number of species increased from 10 , 711 to 10 , 793 , i . e . an increase of 82 species . 66 new species have been described , 9 species have been revalidated from synonymy and 16 subspecies were elevated to full species . . .
over the past 4 months , the number of species increased from 10 , 639 to 10 , 711 .
the number of species has grown from 10 , 544 in the may release to now 10 , 639 ( + 95 species ) .
overall , 212 new taxa have been added or changed their status or name .
the reptile database is a taxonomic database that provides basic information about all living reptile species , such as turtles , snakes , lizards , and crocodiles , as well as tuataras and amphisbaenians , but does not include dinosaurs .
currently there are more than 10 , 000 species and an additional 2 , 700 subspecies . this is making reptiles the largest vertebrate group after fish ( ~ 25 , 000 species ) and birds ( ~ 10 , 000 species ) , and significantly larger than mammals ( ~ 5 , 000 species ) or amphibians ( ~ 6 , 000 species ) .
the reptile database provides taxonomic information for the catalogue of life and the encyclopedia of life . our taxonomic information has also been used by genbank and many other resources and is the only comprehensive reptile database on the web .
the reptile database can be used to find all species within a certain geographic area ( e . g . all snakes of egypt ) . its collection of more than 2 , 500 images allow users to identify a species or at least get an idea how the species or genus may look like . more than 30 , 000 references provide a guide to further information .
this database is maintained by peter uetz ( database content ) and jakob hallermann , zoological museum hamburg ( new species and updates ) .
the three subspecies are fairly similar in general appearance . the body is long and slender in build . scales are relatively large with a semi - glossy appearance . the head is rather small and indistinct from the neck . there is a strong brow ridge over the large eye , which is blackish brown with a golden orange rim surrounding the round pupil . the mouth lining is pink , while the glottis and trachea are black .
mainland subspecies pseudonaja affinis affinis - the overall colour ranges from brown to olive brown to brownish grey , with irregular black / dark grey spotting ( each spot covering half to a whole scale ) . eastern and southern individuals are more heavily spotted ( sometimes becoming totally dark ) . the ventral surface is pale grey to brown with brownish orange or dark grey blotches .
the two island subspecies pseudonaja affinis exilis and p . a . tanneri are generally smaller and usually a uniform blackish brown above .
midbody scales in 19 rows , ventrals 190 - 230 , anal and subcaudal scales divided .
pseudonaja affinis affinis can reach total lengths of 2m ; however the average is probably around 1 . 5m . a specimen kept for some years in the perth zoo reached a total length of 2130mm . the two island subspecies appear to be smaller than the mainland subspecies , with the maximum recorded snout - vent lengths for pseudonaja affinis exilis and p . a . tanneri being 1005mm and 859mm respectively .
pseudonaja affinis affinis – far southwest corner of western australia , extending eastwards into western coastal south australia .
p . a . tanneri - boxer island and figure of eight island in the recherche archipelago ( wa ) .
the snakes are usually most active in the period between early spring and early autumn ( october through april ) and much less active from mid - autumn through late winter ( may through august ) .
they are known to be cannibalistic in the wild , and individuals can swallow snakes almost as large as themselves .
dugites actively search for prey sheltering in holes , crevices , tussock grasses and under surface debris . the snake employs both venom and constriction to subdue its prey .
the species is diurnal . on hot days , activity occurs mainly in the morning and to a lesser extent in the afternoon .
in captivity , adults can darken over the course of a few months . however , the significance of this colour change is obscure .
in the mainland subspecies , males appear to mature at a snout - vent length of about 581mm and females at about 680mm .
in the wild , male combat and mating occurs in late winter - early spring ( september ) , although male combat has been observed in pseudonaja affinis tanneri in mid - spring on boxer island . in the mainland population male combat involves , at least partly , the combatants loosely entwining their bodies . in captivity , mating can occur year around if the temperature is kept warm enough . after mating , captive females are said to eat very large amounts of food . a captive female weighing 309g when mated weighed 580g 27 days later . this represents an 87 . 7 percent gain and implies a daily rate gain of about 10g per day .
wild caught females from the mainland lay their eggs from late spring to mid - summer ( mid - december to end of january ) . the clutch size ranges from 11 to 35 ( average 21 ) for mainland dugites . in the smaller island pseudonaja affinis tanneri , it ranges from 12 to 15 . eggs from mainland dugites have taken between 53 days ( at 30°c ) and 105 days ( at 23°c ) to hatch .
wind the bandage around the bitten arm or leg , starting from the bite .
seek medical attention as soon as possible . first aid guidelines were correct at time of publication however these guidelines change over time . for up to date first aid information consult medical professionals such as st john ' s ambulance .
mirtshin , p . and davis , r . ( 1991 ) “dangerous snakes of australia” , revised edition , ure smith press
wilson , s . and swan , g . ( 2008 ) “a complete guide to reptiles of australia” , reed new holland
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scarborough mum daile cross was dropping her daughter olivia at her parents ' house in city beach on thursday morning when she noticed something black and skinny on the driveway near a car .
mum still had olivia in her arms so i yelled at my mum to get her out of there ,
she said .
i panicked a little because i have a phobia of snakes – i would rather have a spider fall on my head .
i saw the snake heading towards the garage and i thought , ' you are not going in there ' , so i pressed the button for the garage .
the quick - thinking producer at watoday said she also bellowed at the two painters working at her parent ' s home to keep a look out for the unwelcome reptile .
i ' m a bit of a mumma bear , so i just wanted to protect olivia ,
she said .
dugites are roaming throughout the area , because it is near a dune system and near bold park ,
he said .
there are strips of bushland all around there so you see them all the way down to scarborough .
mr cosentino was blunt when asked what would ' ve happened to mr cross if she was bitten .
if she was bitten she would ' ve obviously been taken to hospital , and i ' m not sure what stress it would ' ve had on the unborn baby ,
he said .
but dugites are dangerous and venomous so essentially a bite could have been fatal .
mr cosentino said with the warmer weather over the last couple of days , he ' s had a number of call outs for tiger snakes .
i usually get about 50 call outs for dugites over the summer , but i usually only grab about 20 , because people forget to keep on an eye on them ,
he said .
last month a 75 - year - old woman died from a suspected snake bite south of fremantle after the warmest august day in 17 years brought the reptiles out of their torpor . follow watoday on twitter
the rare photos were taken by natalie hedzik , who works at iluka resource ' s mining operations near capel , and said it was a chance encounter .
i was driving along a track after taking some water samples at a dam we have and just saw this mass in the middle of the road . i got out and had a look and was just absolutely amazed . i ' m really rapt to have seen something so amazing ,
she said .
adam firstenberg from perth hills reptile removal said ms hedzik ' s photographs have captured an extremely rare event .
i certainly haven ' t ever seen it in the wild before . they [ snakes ] will eat reptiles . . . i ' ve seen a snake eat a bobtail lizard before . but they have rear facing fangs to keep food moving backwards , so it ' s going to take a while for the snake to digest another snake .
australian tiger snakes are widely considered to be among the top 10 most deadly in the world , while dugites are known to use both venom and constriction to subdue their prey .
but mr firstenberg said it ' s unlikely the snakes fought to the death .
snakes are very opportunistic eaters , so if they need a meal and the opportunity is there they will take it ,
he said .
but this is certainly a very rare occurrence . we keep snakes ourselves and we feed them once a week , so this is a pretty meal to take on .
if you need more of a snake fix , here ' s a rare sighting of a snakes ' mating ritual in nsw bushland .
the dinosaurs of zoorassic park are coming back but this time they’re bringing some never before seen friends . australian prehistoric megafauna !
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this dangerous brown snake is slightly less venomous that its eastern counterparts and much less aggressive .
description : dugites are venomous snakes . they vary in colour from grey to olive to brown on the top of their bodies , with an olive or yellowish belly . black scales can be scattered over the body and the head can be paler or darker than the rest of the body . young dugites have black heads .
diet : dugites are carnivores . before human settlement they mostly ate other reptiles , such as lizards and snakes . since then they have adapted well to eating mice and rats .
in the wild : they may be found sheltering beneath logs or rocks or living in abandoned burrows or hollow logs . when disturbed , dugites are very shy and will often slither away , however , they will defend themselves if cornered .
the main natural predators of dugites are birds of prey and monitor lizards . introduced animals such as cats and dogs also attack dugites .
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a highly venomous snake found in south - western australia , similar to the related brown snakes .
‘other examples resembling the general appearance of dugites from the middleback ranges and environs have been found . ’
‘this includes many populous areas , and dugites are a relatively frequent cause of snakebite in this area . ’
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dugites live in abandoned burrows or hollow logs . they can also be found sheltering underneath rocks or logs . in the summer months , they are often seen sunning themselves on roadsides .
dugites are very shy but will attack if they feel threatened . when it attacks , it lifts its head and front part of its body up into an s - shape . it hisses loudly and then makes a sharp strike . this snake has small fangs but highly toxic venom which is dangerous to humans . these snakes should never be approached and should only be watched from a distance . anyone who may have been bitten by a snake should find medical help immediately .
dugites search for prey in holes , cracks , small caves , grasses and under surface debris . it kills its prey by biting it and injecting venom and by wrapping its body around the animal and squeezing . this wrapping and squeezing is called constriction .
not listed under iucn ( 2016 ) . listed as vulnerable under the wildlife conservation act 1950 . all rottnest island’s fauna is protected under the rottnest island authority act 1987 .
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lays 10 - 20 eggs . hatchlings are 15 - 35 cm in length and can be brown , green or yellowish in colour , but always hatch with at least part of the head black .
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- storr et al . ( 2002 ) . snakes of western australia . western australian museum . 309
dangerously venomous . note : even a bite from a ' virtually harmless ' or non - venomous reptile can result in serious complications . play it safe and don ' t get bitten by anything .
note : these are general threats and may not apply to this particular species .
there are many ways you can help our reptiles . here are my top three suggestions :
storr , g . m . ; smith , l . a . ; & johnstone , r . e . ( 2002 ) . snakes of western australia . western australian museum . 309 - search web for this book
note : the links below are automatically generated . some may not take you to useful information .
notes and disclaimer this information may not be complete . while all care is taken to ensure the accuracy of the information in this page , primary sources should always be consulted for definitive information . animals have an endearing habit of disobeying the rules , so the information on this page should be interpreted with a degree of flexibility . the author and site operator accepts no responsibility for any losses or damages incurred through using this web site or the information contained herein . don ' t get bitten by anything ! this page may be cited as : pseudonaja affinis at the australian reptile online database . last updated 2017 - 04 - 22 13 : 20 : 54 . retrieved from urltoken on the 10th of july , 2018 . before citing information contained in arod , please read our citing arod page . copyright notice this page , its content and layout are copyright © 2007 - 2018 stewart macdonald / ug media , unless otherwise stated . all photographs in the australian reptile online database are © the photographer and may not be reproduced in any form without the express written consent of the photographer . no part of the australian reptile online database may be reproduced without written permission from stewart macdonald .
brown snakes are probably the most common cause of significant snakebites in australia . without appropriate antivenom treatment , a significant number of cases may be fatal . main risks are : coagulopathy , acute renal failure , neurotoxic paralysis . target organs : neuromuscular junction , coagulation system , kidneys .
locally : the bite is usually painless , without significant local erythema , bruising or oedema . bite marks may be difficult to see , and vary from a single puncture to multiple punctures or multiple scratches . local secondary infection is unusual . venom may spread to draining lymph nodes with consequent pain and / or tenderness and / or swelling . systemic : headache , nausea , vomiting , abdominal pain , impaired conscious state , occasionally loss of consciousness and convulsions . coagulopathy , rarely with overt bleeding manifestations . progressive neurotoxic paralysis can occur but is unusual . acute renal failure .
monitor coagulation to establish the presence and extent of coagulopathy , and as an index of systemic envenomation . this should be performed at presentation , on development of symptoms or signs of systemic envenomation , regularly thereafter , and 1 - 2 hours after antivenom therapy until sufficient antivenom has been given to reverse coagulopathy . in the absence of a haematology laboratory , whole blood clotting time in a glass test tube is useful . if a laboratory is available , prothrombin ratio , activated partial thromboplastin time , thrombin clotting time , fibrinogen level , and fibrin ( ogen ) breakdown products are most useful . other useful tests are complete blood picture and platelet count , serum electrolytes , creatinine , urea , serum enzymes , especially creatine kinase , urine output and urine myoglobin , and venom detection using csl venom detection kit . best sample is swab from bite site ( sample swab stick in kit ) . if patient has systemic envenomation urine may also be useful sample . blood is not a reliable sample .
venom is produced in paired modified salivary glands , superficially situated beneath the scales , posterior to the eye , and surrounded by muscles , the contraction of which compress the glands , expelling venom anteriorly via venom ducts to the fangs . the paired fangs , situated at the anterior part of the upper jaw , on the maxillary bones . they have an enclosed groove for venom transport , with an exit point near the fang tip . average fang length in adult brown snakes is 2 . 8 mm ( 2 . 0 - 4 . 0 mm ) , and average distance between fangs is 9 mm . average venom yield is 2 - 6 mg , maximum 167 mg ( pseudonaja textilis ) . mean venom injected at first bite ( defensive strike ) is 4 . 5 mg ( 0 . 03 to 9 . 10 mg ) , mean venom left on skin is 0 . 22 mg .
pseudonaja venom is a complex mixture of protein and non - protein components , not all of which have been fully evaluated . ( a ) neurotoxins : both presynaptic ( eg textilotoxin ) and postsynaptic ( pseudonajatoxin a and b ) . ( b ) procoagulants : principally prothrombin converters ( factor xa analogues ) , converting prothrombin to thrombin ( meziothrombin ) . ( c ) nephrotoxins : not conclusively demonstrated experimentally , but strongly suspected on clinical evidence .
3 . 1 . 2 habitat brown snakes occupy a wide variety of habitats from arid dessert , to scrubland , to moister areas . they are not uncommon in association with man ' s activities , such as in paddocks , around rubbish dumps and farm buildings , and under elevated floors of country homesteads . principally diurnal , though active on warm nights .
3 . 1 . 3 distribution brown snakes are restricted to continental australia and a few adjacent islands but not tasmania ( longmore 1986 ) . p . textilis is also reported from papua new guinea . ( cogger 1975 ) . distribution for each species based on museum records are shown in figures .
venom glands ( paired ) situated superficially in posterior part of head , connected by ducts to forward placed ( paired ) fangs . fangs small , may leave classic single or double puncture in man , or a more complicated array of scratches and other punctures , the latter by non - fang teeth ( white 1987a , c ) . classic bite mark in plaster is shown in figure . actual cases illustrated in figures .
3 . 3 . 1 name : pseudonaja venom ; brown snake venom ; bsv components neurotoxins : textilotoxin ( = textilon ) ( coulter et al 1979 , southcott and coulter 1979 , coulter et al 1983 , su et al 1983 ) tyler et al 1987a ) pseudonajatoxin a ( parnett et al 1980 ) pseudonajatoxin b ( tyler et al 1987b ) coagulants : prothrombin converter ( unnamed ) from p . textilis . ( coulter et al 1983 , masci et al 1987 , white et al 1987 ) .
venom is used both in antivenom production and for laboratory research . the neurotoxins in particular may prove valuable in neuromuscular transmission research , and the procoagulants in further elucidating the mechanism of normal human coagulation .
children : when playing in areas were brown snakes are common , either through accidental encounter ( ie stepping on snake ) or while trying to emulate naturalists ( ie trying to catch snake ) . adults : when living in areas where brown snakes are common , and moving around barefoot and without due care , or while putting hands etc into non - reconnoitred potential snake retreats ( ie hollow logs etc ) . farm workers : when working in areas where brown snakes are common . reptile keepers and snake handlers : if due care is not exercised in catching and handling snakes , including venom milking . recreation seekers : camping or walking or playing sport in areas where brown snakes are common . homes : around homes in brown snake prone areas where water is seasonally scarce and free water is available in the garden or home .
brown snakes are widely distributed in most habitats of mainland australia , but are absent form some southern islands and tasmania . while no high risk geographic regions have been identified , it is evident that brown snakes may readily enter metropolitan fringes , and country towns , and are frequently found around country rubbish dumps and under the floors of elevated country homesteads , and around farm buildings .
no data , but unlikely to be hazardous unless there are open wounds in the gastrointestinal tract .
there is no evidence that venom can be absorbed through intact skin . current first - aid advice is to leave venom on skin for later venom identification .
5 . 5 . 1 bites in human envenomation , venom is always inoculated by the snake biting . owing to the size of the fangs , venom is most likely to be inoculated cutaneously or subcutaneously .
experimentally , venom may be administered to test animals via subcutaneous , intramuscular , intravenous , intraperitoneal , and intraventricular ( cns ) routes .
the rate and amount of absorption will depend on the quantity of venom injected , the depth of injection , site of injection including vascularity , the activity of the victim , and the type , efficiency of application and length of application of first aid . clinical evidence from human cases of envenomation suggests that much initial venom movement is via the lymphatic pathways . this is supported by work in monkeys . ( barnes & trueta , 1941 ; sutherland et al 1975 ; sutherland & coulter , 1977 ; sutherland et al 1981a ) direct intravenous injection , unrecorded in man , obviously allows rapid systemic circulation of venom and may result in different effects from normal routes of inoculation , particularly in regard to coagulation .
it appears that much venom is transported from the bite site via the lymphatic system , then concentrating in draining lymph nodes , before ultimately reaching the systemic circulation . however , experience with numerous human cases of brown snake envenomation shows that symptoms and signs of envenomation may occur within 15 - 30 minutes of the bite , especially in children . such early effects ( eg headache , nausea , abdominal pain , collapse ) may be due to either rapidly systemically circulating venom toxins , or systemically circulating autocoids released at the bite site by the action of venom on local tissue . once in the systemic circulation , venom rapidly reaches high concentrations in the kidneys and excreted in the urine . such venom must also exit the circulation , to enter the extravascular space where it binds within the neuromuscular junction ( presynaptically and / or postsynaptically ) and possibly other nerve junction sites ( eg autonomic system , perhaps causing abdominal pain ) . the kinetics of venom distribution , excretion , and detoxification are incompletely understood . neurotoxic paralysis usually takes at least 2 - 4 hours to become clinically detectable . coagulopathy however may become well established within 30 minues of a bite .
little information is available on the metabolism of venom components in man , but most components are fully active in whole venom , requiring no further modification for activity . venom reaches high concentrations in the kidneys and excreted in urine . ( sutherland & coulter , 1977 a , b ) . the fate of specific venom components , particularly neurotoxins and procoagulants , is unclear . it seems likely that these components are progressively detoxified in situ .
7 . 2 . 1 . 1 adults the human lethal dose for brown snake venom is unknown . without antivenom treatment , a few brown snake bites may be fatal ( fairley 1929a ) , but even in the early part of the 20th century , when neither antivenom nor icu facilities were available , only 8 . 6 % of reported brown snake bites were fatal . despite this , brown snakes are still thought to be the most common cause of fatal snakebite in australia . ( hilton 1989 ) .
7 . 2 . 1 . 2 children no data available , but clearly the smaller body mass of a child compared to available venom ensures that children are more likely to receive a lethal dose than adults .
7 . 2 . 2 animal data ld50 subcutaneous injection of dried venom in mice ( broad et al 1979 . ( mg / kg ) pseudonaja textilis 0 . 053 pseudonaja nuchalis 0 . 473 pseudonaja affinis 0 . 660
8 . 1 . 1 . 1 toxicological analyses for venom detection swab from bite site moistened in sterile saline . if systemic envenomation also collect urine ( 5ml in sterile container ) . for venom analysis ( research only using radioimmunoassay ) : 5ml blood ; 5ml urine , frozen . at autopsy collect vitreous humor , lymph nodes draining bite area , excised bite site . ( for other laboratory tests see 10 . 2 . 1 )
8 . 1 . 1 . 2 biomedical analyses for standard tests ( eg . serum / plasma electrolytes , ck , creatinine , urea ) collect venous blood in a container with appropriate anticoagulant as issued by the laboratory ( usually heparin ) .
8 . 1 . 1 . 3 arterial blood gas analysis collect arterial blood by sterile arterial puncture into a container as issued by the laboratory .
8 . 1 . 1 . 4 haematological analyses for whole blood clotting time as a
bedside
test collect 5 - 10 ml of venous blood without anticoagulant ( either in the collection syringe or from a central line or other venous access line that may have anticoagulant ) and place in a glass test tube . carefully observe the time until a clot appears . for standard tests ( eg . coagulation studies , complete blood picture ) collect venous blood in appropriate containers with anticoagulant as issued by the laboratory ensuring that the right amount of blood is used ( for coagulation studies citrate will usually be the anticoagulant , while edta will be used for complete blood pictures ) .
8 . 1 . 2 . 1 toxicological analyses for samples for standard venom detection : short term ( less than 24 hrs ) ordinary fridge is acceptable ( 4°c ) , in sterile container . long term , store frozen ( - 20°c or lower ) . for samples for venom analysis ( research ) store frozen ( - 200°c or lower ) .
8 . 1 . 2 . 2 biomedical analyses for samples for standard tests refer to laboratory . in general keep at 4°c , particularly for samples for coagulation studies .
8 . 2 . 1 . 2 advanced qualitative confirmation test ( s ) as for 8 . 2 . 1 . 1
8 . 2 . 1 . 3 simple quantitative method ( s ) not applicable .
8 . 2 . 1 . 4 advanced quantitative method ( s ) a radioimmune assay has been developed by staff at the commonwealth serum laboratories , melbourne to detect small quantities of many australian snake venoms . it is primarily a research tool , being too time consuming to be practical in determining emergency treatment of snakebite victims . it has proved useful in demonstrating snake venom either at autopsy or after patient recovery .
8 . 2 . 2 . 1 simple qualitative test ( s ) see 8 . 2 . 1 . 1 .
8 . 2 . 2 . 2 advanced qualitative confirmation test ( s ) see 8 . 2 . 1 . 1 .
8 . 2 . 2 . 3 simple quantitative method ( s ) not applicable .
8 . 2 . 2 . 4 advanced quantitative method ( s ) see 8 . 2 . 1 . 4 .
8 . 2 . 2 . 5 other dedicated method ( s ) no data .
8 . 2 . 3 interpretation of toxicological analyses for venom detection as for 8 . 1 . 1 . 1 subsection ( 12 ) : if the test is positive , it will indicate the presence of snake venom and the species / genus of snake and therefore the appropriate monovalent antivenom to neutralize the effects of that venom . if the test sample was a bite site swab , a positive result does not indicate either the presence of systemic envenomation , or the need to administer antivenom . other clinical criteria are required in this situation ( see sections 9 and 10 ) . if the test sample was urine a positive result indicates present or past systemic envenomation and together with other clinical and laboratory criteria may be used to determine the need for antivenom therapy . for venom analysis refer to the laboratory performing the tests .
8 . 3 . 1 . 1 blood , plasma or serum electrolytes : look for imbalance , particularly evidence of dehydration , hyponatraemia ( inappropriate adh syndrome ? ) , hyperkalaemia ( renal damage , rhabdomyolysis ? ) . urea , creatinine : look for evidence of renal function impairment . ck : if high may indicate rhabdomyolysis , usually greater than 1000 u / l .
8 . 3 . 1 . 2 urine output : low output may indicate renal damage or poor fluid input . myoglobin : if present indicates rhabdomyolysis , and may be missed as the red colouration of urine may be mistaken for haematuria ( both may be positive on dip stick testing ) . electrolytes if indicated ( eg . inappropriate adh syndrome )
8 . 3 . 2 arterial blood gas analysis impaired respiratory function is usually secondary to neurotoxic paralysis ; look for evidence of poor oxygenation and its sequelae .
8 . 3 . 5 interpretation of biomedical investigations the interpretation of the above tests should be made in the context of total patient assessment including clinical evidence of pathology such as paralysis , myolysis , coagulopathy and renal damage .
while other investigations are not usually required to make the primary diagnosis of snakebite envenomation , they may be indicated in response to secondary effects of envenomation . if there is either renal failure or severe rhabdomyolysis ( unlikely with brown snake envenoming ) there may be a hyperkalaemia , hence an ecg may be appropriate . if the patient is unconscious , especially in the presence of a severe coagulopathy , then a ct head scan may be appropriate to determine if there is intracranial pathology such as a haemorrhage .
overall interpretation of the results of the above tests will depend on the clinical setting . results should never be interpreted in isolation from an overall clinical assessment . a patient with positive venom detection from either the bite site or urine and a significant coagulopathy clearly is envenomed and will usually require antivenom therapy . a patient with positive venom detection from the bite site only and with no clinical symptoms or signs of envenoming and all other tests negative is not significantly envenomed at that point in time and does not require antivenom therapy . however this situation may change and so careful observation and repeat testing would be indicated . a patient presenting some hours after the bite with positive venom detection from the urine but clinically well and with all other tests either normal or showing a resolved coagulopathy , probably had a minor degree of envenomation , now resolved and will usually not require antivenom therapy . however they should be observed carefully for evidence of relapse .
9 . 1 . 5 parenteral exposure in practical terms , the only likely route of entry , is by s . c . or i . d . injection . early symptoms , usually in the first six hours . local : the wound is often painless , without local erythema , oedema , or ecchymosis ; persistent bleeding from wound , variable ; pain or swelling of draining lymph nodes ( may take 1 - 4 hours to develop ) . systemic : collapse , unconsciousness , convulsions may all occur , especially in children , occasionally as rapidly as 15 minutes after the bite . headache , nausea , vomiting , abdominal pain , and visual disturbance may all occur . early signs of neurotoxic paralysis such as ptosis , diplopia , dysarthria may develop within 1 - 3 hours of the bite , or be delayed for 4 - 12 hours , or may not develop despite other evidence of severe envenomation . coagulopathy may develop within 30 minutes of the bite . delayed symptoms local : local bite site necrosis is not generally seen with brown snake bites but possibly might occur , particularly if first aid left in place more than 4 hours , or if a tourniquet used ( sutherland 1981 , 1983a ; white 1987d ) . systemic : paralysis progressive up to complete paralysis . coagulopathy bleeding from all puncture wounds . renal impairment oliguria or anuria .
9 . 4 . 1 cardiovascular collapse , presumably due to hypotension , is common in the early stages of systemic envenomation , especially in children . the mechanism is uncertain but may be due to release of vasoactive substances by or from the venom . specific cardiac abnormalities due to brown snake envenomation in man are not described , although there has been a suspicion that brown snake venom may cause cardiac anomalies ( sutherland personal communication ) .
9 . 4 . 2 respiratory no primary effects of brown snake venom on the respiratory system in man are reported , with the exception of respiratory muscle paralysis ( see below ) .
9 . 4 . 3 . 1 cns while no direct cns toxins have been reported for brown snake venom , early collapse and convulsions do occur , especially in children . their aetiology remains uncertain .
9 . 4 . 3 . 2 peripheral nervous system see 9 . 4 . 3 . 4
9 . 4 . 3 . 4 skeletal and smooth muscle the effects of brown snake venom at the neuromuscular junction , have been documented experimentally . both presynaptic and postsynaptic neurotoxins present , causing progressive neuromuscular paralysis , up to complete paralysis of all muscles of respiration .


gaussia is a genus of copepods .

it is a " characteristic genus of the mesopelagial " , occurring at depths of 0 – 3,000 metres ( 0 – 9,843 ft ) .

the genus gaussia contains the following species : gaussia asymmetrica t. k. s. björnberg & campaner , 1988 gaussia gadusae sarkar , 2004 gaussia intermedia defaye , 1998 gaussia melanotica wolfenden , 1905 gaussia princeps ( t. scott , 1894 ) ( type species ) gaussia sewelli saraswathy , 1973

gaussia ( copepod )

bioluminescence in the mesopelagic copepod , gaussia princeps ( t . scott ) - sciencedirect
identification of two catalytic domains in a luciferase secreted by the copepod gaussia princeps .
katja schulz selected
gaussia princeps ( copepod )
to show in overview on
gaussia princeps ( t . scott , 1894 )
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as are most other copepod species , the deep - sea copepod is bioluminescent . . . .
identification of two catalytic domains in a luciferase secreted by the copepod gaussia princeps . - pubmed - ncbi
most copepod species are 1 - 3 mm long , but the deep - sea copepod . . .
gaussia luciferase secreted by the copepod gaussia princeps catalyzes the oxidation of coelenterazine to produce blue light . the primary structure of gaussia luciferase deduced from the cdna sequence shows two repeat sequences of 71 amino acid residues , suggesting the luciferase consists of two structural domains . two domains in gaussia luciferase were expressed independently in escherichia coli cells , purified and characterized . we found that both domains have luminescence activity with coelenterazine , and the catalytic properties including luminescence spectrum , optimal ph , substrate specificity and luminescence stimulation by halogen ions ( cl - , br - and i - ) are identical to intact gaussia luciferase . thus , gaussia luciferase has two catalytic domains for the luminescence reaction .
paul merviel added text to
behavior
on
gaussia princeps ( t . scott , 1894 )
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paul merviel added text to
size
on
gaussia princeps ( t . scott , 1894 )
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cloning and expression of cdna for a luciferase from the marine copepod metridia longa . a novel secreted bioluminescent reporter enzyme
one fascinating shrimp - like copepod that god created is the gaussia princeps . gaussia stores its supply of chemicals in glands located on its tail . when a predator approaches this vulnerable little fellow , it shoots out a stream of glowing blue goo . the predator’s attention is immediately drawn away from the copepod to the movement of the blue stream . then suddenly the goo gives off a bright flash of white light . the flash so startles the predator that it is unable to process what is happening . the copepod is able to slip away and avoid becoming a meal .
jennifer hammock set
image of gaussia princeps
as an exemplar on
copepoda milne - edwards , 1840
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in the ocean you may find useful molecules , for instance gaussia luciferase ( gluc ) . this is a pretty novel biomiluminescent reporter encoded by a gene isolated from the marine copepod gaussia princeps . this luciferase does not require atp and catalyzes the oxidation of the substrate coelenterazine in a reaction that emits light ( peak at 470 - 480 nm ) like renilla and
rene noam added the english common name
deep sea coceopod
to
gaussia princeps ( t . scott , 1894 )
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a . barnes & j . f . case ( 1972 ) .
bioluminescence in the mesopelagic copepod , gaussia princeps ( t . scott )
. journal of experimental marine biology and ecology 8 ( 1 ) : 53–71 . doi : 10 . 1016 / 0022 - 0981 ( 72 ) 90056 - 1 .
copepods are the dominant taxa in zooplankton communities of the ocean worldwide . although bioluminescence of certain copepods has been known for more than a 100 years , there is very limited information about the structure and evolutionary history of copepod luciferase genes . here , we report the cdna sequences of 11 copepod luciferases isolated from the superfamily augaptiloidea in the order calanoida . highly conserved amino acid residues in two similar repeat sequences were confirmed by the multiple alignment of all known copepod luciferases . copepod luciferases were classified into two groups of metridinidae and heterorhabdidae / lucicutiidae families based on phylogenetic analyses , with confirmation of the interrelationships within the calanoida using 18s ribosomal dna sequences . the large diversity in the specific activity of planktonic homogenates and copepod luciferases that we were able to express in mammalian cultured cells illustrates the importance of bioluminescence as a protective function against predators . we also discuss the relationship between the evolution of copepod bioluminescence and the aspects of their ecological characteristics , such as swimming activity and vertical habitat .
gaussia’s incredible ability reminds us of world war 2 navy ships that released depth charges that would be dropped into water close to submerged enemy submarines . a time delay allowed the ship to move away while the charges sank down to get close to the submarine before exploding . similarly , gaussia emits its bioluminescent fluid as it swims swiftly in the opposite direction . just as the bluish fluid is about to disappear , a bright flash ensues that “blinds” the predator and distracts him from pursuing the little copepod .
bioluminescent activity of zooplankton homogenates . luminous activity of homogenized copepods was measured by mixing coelenterazine solution and the supernatant of the lysate . total protein concentration was determined to compare specific luciferase activities among copepod species .
like insects on land , crustaceans fill the oceans of the world . they come in many different shapes and sizes . one very common form is the copepod . many copepods are bioluminescent . they typically release their bioluminescent chemicals into the water to produce a glowing cloud of light . this luminous smoke screen serves to distract or blind an attacker . the copepod can then swim quickly away from danger into the darkness .
wurdinger and colleagues reports to be able to assay in just 5 μl of blood ( a very little drop ) the presence of secreted gaussia luciferase , demonstrating that gluc assay is 1000 fold more sensitive than seap assay .
in this study , we isolated and characterized novel luciferase genes from five species of calanoid copepods and conducted the quantitative analyses of copepod bioluminescence using zooplankton homogenates and recombinant luciferases . by quantitative measurements , we could reveal the presence or absence of luminescence of unverified species with confirmation of verified ones . furthermore , we compared the luminous intensity of homogenates among calanoid superfamilies and those of luciferases among genera in augaptiloidea to infer the defensive function and molecular evolution of copepod bioluminescence .
issued from : r . n . wolfenden in die marinen copepoden der deutschen südpolar - expedition 1901 - 1903 , 1911 . [ pl . xxxiii , fig . 4 ] . as gaussia scotti . female : 4 habitus ( dorsal ) .
issued from : r . n . wolfenden in die marinen copepoden der deutschen südpolar - expedition 1901 - 1903 , 1911 . [ pl . xxxiii , fig . 3 ] . as gaussia scotti . male : 3 , habitus ( dorsal ) . :
we have isolated nine novel copepod luciferases using the homology - based pcr cloning strategy . a blast search using the amino acid sequences of copepod luciferases as the query revealed no highly similar proteins in the database , except for other calanoid luciferases ( gluc , mluc , and mpluc ) . this may be due to the limited number of deposited sequences of copepod proteins . so far , all of copepod luciferase genes , including gluc and mluc reported elsewhere , have been isolated from species in the augaptiloidea superfamily in calanoida . we have tried to isolate luciferase or luciferase - like genes from other superfamilies of calanoida , such as pseudocyclopoidea or centropagoidea using degenerate pcr primers . the result was no amplification or no luciferase - like sequences in the amplified dna product ( data not shown ) . however , we assume that luciferases or luciferase - like genes may exist , not only in augaptiloidea but also in other superfamilies , because the luminous assay of zooplankton lysate from these superfamilies all had a measurable , though very weak , level of activity compared with that of negative control samples ( fig . 2 ) . further screening of the calanoid luciferase or luciferase - like genes by different cloning strategies may reveal the structure and evolution of a direct ancestral copepod luciferase gene .
issued from : r . n . wolfenden in die marinen copepoden der deutschen südpolar - expedition 1901 - 1903 , 1911 . [ pl . xxxiii , figs . 6 - 7 ] . as gaussia scotti . male : 6 , p5 ; 7 , right p5 .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 182 , fig . 7 ] . zoogeographical distribution of gaussia princeps , g . asymmetrica and g . sewelli .
consensus amino acid residues and domain structure were revealed by an alignment of 13 copepod luciferases obtained in this study ( fig . 4a ) . highly conserved amino acid residues , c - x ( 3 ) - c - l - x ( 2 ) - l - x ( 4 ) - c - x ( 8 ) - p - x - r - c , which are present in both domains , would be one of the criteria for the copepod luciferase , although the reason for this cysteine - rich area in the conserved sequence remains obscure . because the similarity in the structure of the two domains was found in all of the copepod luciferases isolated in this study ( fig . 4b ) , we assume this similarity in the structure of the two domains is very characteristic of the luciferases isolated from augaptiloidea species .
luminous activity of zooplankton homogenates varies greatly among the copepods ( fig . 2 ) . for instance , luminous activity of an m . pacifica homogenate was ∼500 - fold higher than that of a h . tanneri homogenate , even though both species belong to the same superfamily . one possible explanation is very different levels of luciferase expression among bioluminescent copepod species . the other is a substantial divergence of specific activity of copepod luciferases . to compare the specific activity of copepod luciferases isolated in this study , 11 luciferases were expressed in hek293 cells as recombinant proteins tagged with hexahistidine at the c - terminus , concentrated from culture medium by a nickel - chelating column , and then tested for their protein expression and activity . specific activity was estimated by dividing the luminous activity of the culture media by signal intensity of polyhistidine - tagged copepod luciferases detected by an anti - polyhistidine antibody ( fig . 6a ) . results showed that there is a large difference in specific activity among copepod luciferases ( fig . 6b ) but only a slight difference in their expression levels ( fig . 6a ) . in several experiments , we consistently observed that mpluc1 showed the highest level of specific activity among tested luciferases , which was approximately 10 6 - fold higher than that of htluc2 . the specific activity of luciferases isolated from copepods in the family metridinidae ( mpluc1 , mpluc2 , moluc1 , moluc2 , paluc1 , and paluc2 ) was notably higher than that of luciferases from the heterorhabdidae family ( htluc1 , htluc2 , hmluc1 , and hmluc2 ) . loluc , isolated from l . ovaliformis , which belongs to the lucicutiidae family , showed intermediate levels of activity . these results are consistent with the luminous activity obtained by homogenate assays of calanoid copepods ( fig . 2 ) . thus , species differences in copepod bioluminescence was reflected in the substantial divergence of specific activity of the expressed luciferases .
bioluminescence in all experiments , except for analyses of the spectra and kinetics , was measured using a minilumat lb 9506 luminometer ( berthold , japan ) . the basic procedure for the luciferase assay was as follows : 5 μl of protein solution containing each of the luciferases was transferred into the luminometer tube and then placed in the luminometer . a 10 - s measurement was initiated immediately after injecting 10 μl of 1 ng / μl coelenterazine in 20 mm tris–hcl ( ph 8 . 0 ) with 50 mm mgcl 2 into the protein solution . luminescent spectra of recombinant copepod luciferases were measured using an ab - 1850c spectrofluorometer ( atto , japan ) ( slit width , 1 mm ; spectral resolution , 0 . 5 nm ) for 1 min after mixing 20 μl of copepod luciferases expressed by cultured cells and 200 ng of coelenterazine in a 0 . 2 - ml pcr tube . to determine thermostability of copepod luciferases , 20 μl of copepod luciferases expressed by cultured cells was incubated in a block heater at 60 or 80 °c for 30 min and then cooled on ice for 5 min . bioluminescence was measured by injecting 10 μl of coelenterazine solution ( 1 ng / μl ) into 5 μl of heat - treated luciferases at room temperature using a minilumat lb 9506 luminometer .
issued from : r . n . wolfenden in die marinen copepoden der deutschen südpolar - expedition 1901 - 1903 , 1911 . [ pl . xxxiii , fig . 5 , 8 ] ] . as gaussia scotti . female : 5 , last thoracic segment and urosome ( lateral , left side ) ; 8 , forehead ( lateral ) .
issued from : r . stephen , 2007 [ data sheets of nio , kochi , india ( on line ) ] . after m . saraswathy , 1973 . as gaussia scotti . female ( from w indian ) : a , last thoracic segment and urosome ( dorsal ) . male : last thoracic segment and urosome ( lateral ) .
( a ) immunoblot of polyhistidine - tagged copepod luciferases expressed by hek293 cells . proteins were concentrated from culture media on a ni - chelating column . the same volume of eluents was loaded on sodium dodecyl sulphate–polyacrylamide gel electrophoresis , blotted , and detected by anti - polyhistidine tag antibody . ( b ) specific activity of secreted copepod luciferases expressed by hek293 cells . the initial light intensity was counted for 10 s and then normalized to the protein expression level obtained by immunoblotting shown in figure 5a . luciferase activities are expressed as percentages of normalized luminescence activity ( light intensity / protein expression level ) , using the value of mpluc1 defined as 100 % . the data represent the mean for triplicate measurements .
in phylogenetic analyses of aligned amino acid sequences of copepod luciferases , 48 substitution models for proteins used in ml estimates were tested by mega5 to find the most appropriate model . the best model showing the lowest bayesian information criterion score was whelan and goldman plus gamma ( gamma distribution ) , therefore , an ml bootstrap consensus tree was generated with the model (
issued from : r . n . wolfenden in die marinen copepoden der deutschen südpolar - expedition 1901 - 1903 , 1911 . [ pl . xxxiii , figs . 9 - 12 ] . as gaussia scotti . female : 9 , p2 ; 10 , p3 ( distal segment of exopod ) ; 11 , p4 ( distal segment of exopod ) ; 12 , p5 .
one strong piece of evidence for the presence of bioluminescence in a particular organism is the molecular identification and functional analysis of luciferase , which is the enzyme directly involved in producing luminescence in an organism . in 2002 , the luciferase gene was cloned from calanoid gaussia princeps ( family metridinidae ) , which is a large ( ∼10 mm ) gray - black–pigmented copepod usually found in the mesopelagic zone ( at the depths from 200 to 1 , 000 m ) ( verhaegen and christopoulos 2002 ) . a novel feature of gaussia luciferase ( gluc ) is its relatively small size ( ∼20 kda ) and secreted luminescence , not only in planktonic expression but also when expressed in mammalian and insect cells , with robust stability and great luminescent activity . gluc catalyzes the oxidation of a luciferin called coelenterazine to produce the light . luciferase genes have also been found in metridia longa ( mluc ) ( markova et al . 2004 ) and metridia pacifica ( mpluc1 and mpluc2 ) ( takenaka et al . 2008 ) , both of which belong to the metridinidae family just as g . princeps does . these gluc - homologues also react with the substrate coelenterazine to produce light . the bioluminescence in these species was confirmed by identification and functional analysis of their luciferases .
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 41 , table 2 ] . male : comparison of pore number of right a1 of gaussia princeps from the caribbean ( gpc ) , the indian ocean ( gpl ) from soh & al . , 1998 , and g . sewelli ( gsi ) from saraswathy & bradford , 1980 from the indian ocean . a1 segments grouped in 3 sectors : 1st segment , segments 2 - 5 , and 6 - 12 . nota : the number and general distribution of the pores ( gland openings ) have been found to express important differences between congeners in gaussia . the caribbean specimens showed a pore pattern that differs in number with that described from specimens from the indian ocean and japan ( soh & al . , , 1998 ) on different appendages
santos , e . , yeh , r . , lee , j . , nikhamin , y . , punzalan , b . , punzalan , b . , perle , k . , larson , s . , sadelain , m . , and brentjens , r . ( 2009 ) . sensitive in vivo imaging of t cells using a membrane - bound gaussia princeps luciferase nature medicine , 15 ( 3 ) , 338 - 344 doi : 10 . 1038 / nm . 1930 .
wurdinger , t . , badr , c . and tannous , b . ( 2008 ) . gaussia luciferase blood level as an index of cell growth and proliferation . nature methods , wurdinger t , badr c , pike l , badr c , de klein r , weissleder r , breakefield xo , tannous ba . ( 2008 ) a secreted luciferase for ex vivo monitoring of in vivo processes . nature methods 5 : 171 - 173 doi : 10 . 1038 / nmeth . 1177
( a ) multiple amino acid sequence alignment of 13 copepod luciferases . positions of two putatively similar domains are underlined . outlined letters represent at least 90 % identical ( dark shade ) or chemically similar ( light shade ) amino acid residues . white letters in the black boxes are highly conserved residues in both domains . arrowhead indicates proposed cleavage sites of n - terminal secretion signals of copepod luciferases . ( b ) multiple amino acid sequence alignment of tandem repeat domains of luciferases . residues are darkly shaded and outlined if they are conserved at a level of more than 90 % among all aligned sequences . similar residues are lightly shaded and outlined . the distance between the two domains in each species is indicated as number of amino acids in parentheses at the end of the sequence . those amino acids that are conserved in both domains ( consensus ) , domain 1 only ( consensus _ domain1 ) or domain 2 only ( consensus _ domain2 ) , are indicated in the last three rows .
adult male and female gaussia princeps ( t . scott ) produce bright , extracellular luminescent displays usually coupled with the initiation of rapid swimming . glands of the head , mandibular palps , urosome , furca , and underlying at least 14 body pores contribute to the luminous response which , in free swimming copepods , is characterized by a sudden discharge to the exterior of luminous blue material that remains at peak intensity for 1–3 sec before undergoing an 80 sec decay as 8 or less discrete , low intensity points of light .
( a ) ml phylogeny of copepod luciferases based on an amino acid sequence alignment of domains 1 and 2 with a linker sequence . the tree is midpoint rooted , and numbers on the nodes indicate % bootstrap values from 1 , 000 replicates . luciferases isolated and analyzed in this study are shown in shaded boxes . scale bar indicates evolutionary distance and is in the units of the number of amino acid substitutions per site . ( b ) gene structures of moluc1 and moluc2 . introns and exons are shown as open boxes and lines , respectively .
, a calanoid copepod , measuring about 6 mm from head to telson , making it a giant among hawaiian copepods . copepods are the most abundant and diverse group in the planktonic community . they are crustaceans , related to crabs and lobsters . they are members of the phylum arthropoda , as are other crustaceans , insects , and spiders . there are planktonic , benthic , and parasitic copepods , appearing in freshwater and salt water . their distribution around the world and success in so many niches earns them the title of
the insects of the sea
.
pleuromma princeps t . scott , 1894 b ( p . 42 , descr . m , figs . m ) ; metridia scotti giesbrecht & schmeil , 1898 ( p . 107 ) ; no sewell , 1913 ( p . 354 ) ; gaussia melanotica wolfenden , 1905 a ( pl . 2 , figs . 1 - 5 ) ; gaussia scotti wolfenden , 1905 a ( p . 5 ) ; 1911 ( p . 290 , figs . f , m ) ; lysholm & al . , 1945 ( p . 32 ) ; saraswathy , 1973 a ( p . 191 , figs . f , m ) ; stephen , 2007 [ data sheets of nio , kochi , india ( on line ) ] . female metridia atra esterly , 1906 a ( p . 70 , juv . 5 : f ) ; c . b . wilson , 1950 ( p . 263 , rem . ) ; guérédrat , 1969 a ( p . 362 ) ; no g . princeps : sewell , 1932 ( p . 270 , figs . f , m ) ; 1947 ( p . 173 ) ; ? tanaka & omori , 1967 ( p . 251 )
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 42 , table 3 ] . male : comparison of pore number of p1 - p4 of gaussia princeps from the caribbean ( gpc ) , the indian ocean ( gpl ) from soh & al . , 1998 , and g . assymmetrica ( gasa ) from the south - western atlantic ocean ( björnberg & campaner , 1990 ) , and g . intermedia ( ginp ) from the north pacific ( defaye , 1998 ) cx = coxa ; bp = basipodite ; enp = endopodite ; exp = exopodite .
the reactivity of homogenates with coelenterazine indicates the expression of luciferase in these luminous species tested ( fig . 2 ) , as previously reported molecular identification of luciferase genes from single gaussia and two metridia species has indicated ( verhaegen and christopoulos 2002 ; markova et al . 2004 ; takenaka et al . 2008 ) . we amplified full - length luciferase genes from five copepod cdnas with degenerate primers designed based on the nucleotide sequences of the g . princeps , m . longa , and m . pacifica luciferases . at least two forms of luciferase genes , as identified in m . pacifica ( mpluc1 and mpluc2 ) ( takenaka et al . 2008 ) , were also identified from m . okhotensis , p . abdominalis , h . tanneri , and h . major cdnas . the gene names and biochemical features of these proteins are summarized in table 2 . they are relatively small proteins , ranging from 19 . 0 to 23 . 5 kda , and they are efficiently secreted into the culture medium when they are expressed by cultured mammalian cells . the n - terminal 17–22 amino acid sequences of all novel luciferases seem to have the features of a consensus sequence that signals secretion based on the prediction by psort ii ( urltoken ) . moluc1 , moluc2 , paluc1 , and paluc2 retained more than half of their initial activity even after 30 - min incubation at 60 °c but were largely inactivated at 80 °c . thermostability of htluc1 , htluc2 , hmluc1 , and hmluc2 could not be determined correctly because of their low initial activity . the identity of the amino acid sequence of these luciferases was notably high at the internal and c - terminal regions rather than n - terminals ( fig . 4a ) . inouye and sahara ( 2008 ) reported the identification of two catalytic domains in gaussia luciferase , and we confirmed the presence of two short repeat sequences in the primary structures of these novel copepod luciferases . alignment of two repeat sequences consisting of 62–64 amino acid residues ( referred as domains 1 and 2 , respectively ) from 12 luciferases revealed consensus sequence of c - x ( 3 ) - c - l - x ( 2 ) - l - x ( 4 ) - c - x ( 8 ) - p - x - r - c ( x , amino acid residue ) in both domains ( fig . 4b ) . substitution of all cysteine residues with alanine in domains 1 and 2 of mpluc1 resulted in complete loss of activity when it was expressed in escherichia coli ( data not shown ) , suggesting an essential role of cysteine residues in luciferase activity .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 178 , fig . 5 ] . female ( from suruga bay ) : a , p2 ( anterior ) ; b , p3 ( anterior ) ; c , p4 ( anterior ) ; d , p5 ( posterior ) . female ( from the indian ocean ) : e , distal seta on 3rd exopodal segment of right p5 . gaussia sewelli saraswathy , 1973 , female ( from the indian ocean ) : g , distal seta on 3rd exopodal segment of right p5 . arrowheads indicate gland openings .
dvm is a copepod behavior to stay in darker deeper layers during daytime and migrate upward to food - rich surface layers at night for feeding . a most important function of copepod dvm is considered to be avoidance from predation by visual - feeding fishes ( gliwicz 1986 ; neill 1992 ) . species characterized with strong bioluminescence ( metridinidae ) are all medium - sized suspension feeders known to perform strong nocturnal ascent dvm ( haury 1988 ; hays 1995 ) . the dominance of metridinidae in oceans world - wide can be explained by the fast swimming speed and strong dvm intensity ( mauchline 1998 ) . because of its numerical dominance , metridinidae is selectively captured by various mesopelagic lantern fishes ( merrett and roe 1974 ; hopkins and sutton 1998 ) . the bioluminescent behavior of copepods is considered to have a function of avoidance from vision - dependent predators ( herring 1988 ; widder 1992 ) . to distract or blind a predator of mesopelagic lantern fish predator , metridinidae with reinforced luciferases would have a selective advantage for their stronger bioluminescent system ( mpluc , moluc , and paluc ) . we suggest that mpluc from m . pacifica ( fig . 1 ) shows the highest bioluminescence ( fig . 6b ) because the habitat depth of m . pacifica is the epipelagic layer ( 0–200 m ) ( table 1 ) where a large number of predators exist .
issued from : m . saraswathy in handbook to the intern . zoopl . coll . , cochin - 18 , kerala state , india , 1973 . [ p . 191 , fig . 1a , 1b , 2 , 3 ] . as gaussia scotti . female ( from world - wide ) : 2 , p5 . male : 1a , right a1 ( segments 10 to 16 ) ; 1b , glandular structure on segment 12 ) ; 3 , p5 . nota : proximally directed spine on 3rd segment of p5 blunt , short and stumpy . terminal segment of right leg with prominent undulating inner border , and 4 setae ; distance between 1st and 2nd and 2nd and 3rd setae almost equal .
this study was supported by an advanced industrial science and technology research grant ( y . t . , y . s . , n . t . , and m . t . ) , japan foundation for applied enzymology ( y . s . ) , and the nig cooperative research program ( t . g . , 2010 - a66 ) . the authors are greatly indebted to the following individuals : prof . susumu ohtsuka , hiroshima university , for critical discussion for the copepod evolution ; dr steven haddock and ms lynne christianson , monterey bay aquarium research institute , for kindly providing genomic dna of g . princeps ; dr tadashi imanishi and dr akiko ogura noda for introducing mega5 software to calculate ml phylogeny ; and prof . leslie sargent jones , appalachian state university , for valuable discussions .
the origin of the bioluminescence of calanoid copepods must be carefully verified with an assessment of bioluminescence , including nonluminescent primitive species , such as the pseudocyclopoidea , and luminescent oncaea species . nonetheless , we postulate that bioluminescence in calanoid copepods could have originated from an ancient species between the families pseudocyclopoidea and augaptiloidea ( fig . 7 ) . the ancient augaptiloidea may have ventured into the surface water , which was probably caused by unexpected food or oxygen demands ( bradford - grieve 2002 , 2004 ) , and then encountered a number of difficulties for adapting to the pelagic habitat . until myelinated nerve axons evolved , the earlier evolution of bioluminescence might have been a highly efficient strategy for them to avoid surrounding predators . further analysis of copepod bioluminescence in the world ' s oceans could be a clue for understanding the ecology and evolution of marine organisms .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 180 , fig . 6 ] . male : ( from the indian ocean ) : a , a1 ( segments i - viii ) ; b , idem ( segments ix - xix ) ; c , idem ( segments xx - xxviii ) ; d , e ( holotype ) , anterior part of a1 segment xiv ; f , p5 ( posterior ) ; g , medial process on 2nd exopodal segment of left p5 ( anterior ) . gaussia sewelli saraswathy , 1973 ( from the indian ocean ) male : h , p5 ( posterior ; distal two exopodal segments of right p5 torted ) . arrowheads indicate inner process proximally directed on 2nd exopodal segment . p : process . s : spinule .
the phylogeny of 13 calanoid luciferases suggests a different evolution of the metridinidae and heterorhabdidae families ' luciferases ( fig . 5a ) . in the clade containing metridinidae luciferases ( clade 1 ) , two proteins in a single species are likely to have evolved independently . the similarity of primary structures among mpluc1 , moluc1 , and paluc1 ( type - i ) luciferases were greater than those of their counterparts ( mpluc2 , moluc2 , and paluc2 ) ( type - ii ) luciferases ( fig . 4a ) . the mrna coding for moluc1 or moluc2 would be transcribed from two different loci or alleles since studies of genomic sequences revealed the presence of different lengths and positions of the introns in moluc genes ( fig . 5b ) , as seen in mpluc genes ( takenaka et al . 2008 ) . gene duplication events of luciferase probably happened in the ancestral species of the metridia and pleuromamma families , thereafter , paralogous luciferases might have diverged independently . conversely , it would be expected that specialization of heterorhabdidae species preceded the duplication and diversification of luciferase genes since pairs of luciferases ( htluc1 and htluc2 ; hmluc1 and hmluc2 ) are monophyletic ( clades 6 and 7 ) within species . nucleotide identity between coding sequences of hmluc1 and hmluc2 is 91 . 8 % , whereas it is 62 . 6 % between hmluc1 and htluc1 , which were isolated from species of the same heterorhabdidae family . extensive genome survey of two lineages of metridinidae and heterorhabdidae families may shed light on the multigene structure of luciferase genes and how copepod luciferases have evolved . from the present study , we could not interpret any evolutionary novelty derived from gene duplication of luciferases in calanoid copepods . because phenotypic observation on live copepod bioluminescence is difficult without special video equipment and relevant technical skills , we are trying to evaluate any possible evolutionary novelty of the gene duplication by revealing different biochemical characteristics between type - i and type - ii luciferases identified in this study .
issued from : j . j . childress in mar . biol . , 1977 , 39 . [ p . 23 , fig . 3 ] . gaussia princeps adult males ( in experimental proceedings ) . interpolation of data to predict oxygen - consumption in the field . solid line shows oxygen - consumption rate based on temperature and pressure considerations only ; dotted line takes into account limitations of oxygen uptake by low oxygen and indicates maximum oxygen - consumption possible in the region of this limitation ; hatched area represents that part of the metabolic needs which must be accounted for by anaerobic metabolism . the two scales on the right show the temperature and oxygen characteristics of a typical area ( san clemente basin , southern california ) where the animals are found . crosses indicate primary , non - interpolated data , while dots are interpolated points . nota : gaussia princeps is found only below 400 m in the daytime and is present down to the greatest depth sampled ( 900 m ) . at night an appreciate fraction of the population migrates to depths shallower than 400 m . even at night , however , many remain below 400 m . the individuals clearly do not need to migrate above this depth on a daily basis . this species ' consumption was measured at 3 . 5 , 7 and 10°c and 1 , 14 , 28 , 61 , 121 and 181 atm of hydrostatic pressure ( 1 atm corresponds about 10 m of depth ) . hydrostatic pressure is shown to have significant effects on the oxygen - consumption rate at pressures as low as 28 atm . at all temperature and pressure combinations , the species displays a very low metabolic rate compared to shallow - living copepods . the critical oxygen partial pressure for this species is shown to be about 10 to 1 , mm hg o2 at 10 , 7 and 5 . 5 °c . this shows a higher oxygen - consumption rate at the nighttime depths and a much lower , partially anaerobic metabolism at the daytime depths .
the full coding sequences of the copepod luciferases were amplified by pyrobest dna polymerase ( takara bio ) from the template plasmids that harbor the full - length cdnas and subcloned into pcdna3 . 1 / v5 - his - topo ( invitrogen ) to express luciferase proteins tagged with 6× histidines at their c - terminus in mammalian cultured cells . all expression vectors were purified using an endofree plasmid maxi kit ( qiagen , japan ) . hek293 cells ( health science research resources bank , japan ) were maintained in megacell mem ( sigma - aldrich , japan ) supplemented with 5 % fetal bovine serum at 37 °c in a humidified atmosphere of 5 % co 2 . for transfections , 5 × 10 5 cells per well were seeded in six - well culture plates , and fugene hd ( roche applied science ) was used according to the manufacturer ' s instructions . the culture medium was incubated for 72 h after transfection and stored at −30 °c until immunoblotting and luciferase assay .
concerning the bioluminescent copepods other than calanoida , oncaea belonging to poecilostomatoida is known to have bioluminescence ( herring et al . 1993 ) . since many of the bioluminescent copepods are predatory and live moderately deep , it is unique that the tiny poecilostomatoid copepod oncaea is one example that is relatively shallow water habitat and has pseudoplanktonic behavior ( böttger - schnack and schnack 2005 ) . the most notable feature of oncaea bioluminescence is their nonsecretory and repetitive fast flashes . the reason for this different characteristic of oncaea bioluminescence compared with those of other bioluminescent copepods might lie in their smaller size and reduced motility . throwing off bioluminescence as a decoy against potential predators is not an effective protection for oncaea , which cannot escape rapidly from surrounding luminescent materials ejected from themselves , unlike calanoid copepods ( herring et al . 1993 ) . furthermore , the bioluminescence emission spectra of oncaea ( ∼470 nm ) are slightly different from those of other calanoid copepods ( 480–500 nm ) , possibly due to the structural difference of their luciferase and / or luciferin .
copepods are the most numerous taxa of zooplankton fauna in the ocean worldwide . their bioluminescent abilities and characteristics have been investigated for over a 100 years ( giesbrecht 1895 ; clarke et al . 1962 ; haddock et al . 2010 ) . luminescence of calanoid copepods is produced by glandular cells in the luminous glands , which are largely located on the caudal rami and legs ( fig . 1 ) , although their number and location vary considerably among families . most of the luminous species reported to date belong to the superfamily augaptiloidea , which contains the families arietellidae , augaptilidae , heterorhabdidae , lucicutiidae , metridinidae , and nullosetigeridae ( herring 1988 ) . within these families , the luminescent behavior of metridinidae and most heterorhabdidae , lucicutiidae , and augaptilidae are well known ( herring 1988 ) . based on a previous cladistic analysis of morphological features , the augaptiloidea is believed to have diverged very early in the evolution of calanoid copepods ( bradford - grieve et al . 2010 ) . except for the augaptiloidea superfamily , a single species in megacalanoidea ( megacalanus princeps ) was reported as luminous ( herring 1988 ) with very limited evidence , and it remains unverified . it is also unclear whether species in other superfamilies , such as centropagoidea , clausocalanoidea , and eucalanoidea , are luminous . this indicates that among the ten calanoid superfamilies , bioluminescent copepods are found almost exclusively in the superfamily , augaptiloidea . concerning bioluminescence behavior other than calanoid copepods , that of tiny pseudoplanktonic poecilostomatoid oncaea spp . is only notable ( herring et al . 1993 ) . one reason for the limited information on copepod bioluminescence is that correct copepod identification is very difficult . in addition to the species identification challenges , zooplankton sampling , especially in deep water , requires special equipment , including large - sized plankton nets , to collect sufficient numbers of specimens . furthermore , macro - and microscopic detection of weak and flash - type bioluminescence from tiny copepods is also difficult , especially if the specimen was damaged during the collection or manipulation by an investigator . therefore , a device , such as a luminometer , to detect weak bioluminescence of zooplankton is needed .
issued from : a . t . barnes & j . f . case in j . exp . mar . biol . ecol . , 1972 , 8 . [ p . 59 , fig . 3 ] . distribution of fluorescent glands , luminescent glands , and luminous pores of adult male and female gaussia princeps . a : female ; b : male ; c : lateral view of 5th thoracic segment and abdomen of female ; d : lateral xiew of female . f : fluorecent gland ; l . g . : luminescent gland ; l . p . : luminous pore . ( modified after wolfenden , 1911 , and sewell , 1932 ) . nota : adult male and female produce bright , extracellular luminescent displays usually coupled with the initiation of rapid swimming . glands of the head , mandibular palps , urosome , caudal rami , and underlying at least 14 body pores contribute to the luminous response which , in free swimming copepods , is characterized by a sudden discharge to the exterior of luminous blue material that remains at peak intensity for 1 - 3 sec before undergoing and 80 sec decay as 8 or less discrete , low intensity points of light .
copepod luciferases , isolated in this study and previous reports , are all derived from the augaptiloidea superfamily in calanoida . there are few to no reports of bioluminescence in other calanoid superfamilies . the reason for the difference in the luminous ability between the augaptiloidea and other superfamilies could be explained by their ecology ( e . g . , motility , habitat , and body size ) . although precise evolutionary positions of the pseudocyclopoidea and epacteriscoidea are not determined yet , these superfamilies are believed to be the most primitive calanoida ( fig . 7 ) ( bradford - grieve et al . 2010 ) . comparing with other eight superfamilies illustrated in figure 7 , they are highly specialized because of their adaptation to the epibenthic habitats . they are found only at shallow to shelf depth ( ohtsuka et al . 1999 ) or in submarine caves ( ohtsuka et al . 2002 ) , in contrast to the augaptiloidea , which has a greater diversity of distribution within the pelagic zones ( yamaguchi et al . 2004 ) . these observations imply little change , if any , of habitat in the evolutionary history of the pseudocyclopoidea and epacteriscoidea ( bradford - grieve 2004 ) . living on - bottom– or in - bottom–dwelling habitat would be a very effective means to find nutrients and hide from predators . there is no report , so far , describing the bioluminescence of these primitive copepods , probably because of the very limited research on these minor species in calanoida so far . however , considering their stable benthic habitat and very small body size ( below 1 mm ) ( ohtsuka et al . 1999 ) , evolving bioluminescence as a defense against the predators seems to be unnecessary for them .
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davis , 1949 ( p . 50 , figs . f ) ; c . b . wilson , 1950 ( p . 235 , figs . f , m ) ; brodsky , 1950 ( 1967 ) ( part . , p . 312 , figs . np : f , m , non figs . f , m : ind . oc , rem . ) ; vervoort , 1965 ( p . 103 , rem . ) ; owre & foyo , 1967 ( p . 74 , figs . f , m ) ; barnes & case , 1972 ( p . 53 , bioluminescence ) ; gardner & szabo , 1982 ( p . 348 , figs . f , m ) ; björnberg & campaner , 1988 ( p . 351 , 354 , fig . f , rem . ) ; hulsemann , 1988 ( p . 188 , rem . ) ; baessa - de - aguiar , 1989 ( 1992 ) ( p . 117 , figs . f , m ) ; bowlby & case , 1991 ( p . 440 , bioluminescence ) ; soh & al . , 1998 ( p . 171 , redescr . f , m , figs . f , m , juv . , fig . 7 : carte , rem . ) ; bradford - grieve & al . , 1999 ( p . 884 , 948 , figs . f , m ) ; bradford - grieve , 1999 b ( p . 111 , figs . f , m , rem . , figs . 177 , 192 ) ; vives & shmeleva , 2007 ( p . 3 , figs . f , m , rem . ) ; vives & shmeleva , 2007 ( p . 357 , figs . f , m , rem . ) ; suarez - morales , 2007 ( p . 34 , figs . m , tabl . 2 , 3 , 4 , rem . ) ; mercier & al . , 2013 ( p . 760 , neural ultrastructure )
issued from : j . m . bradford - grieve in the marine fauna of new zealand : pelagic calanoid copepoda . national institute of water and atmospheric research ( niwa ) . niwa biodiversity memoir , 111 , 1999 . [ p . 111 , fig . 74 ] . female ( from 39°49 . 9 ' s , 178°26 ' e ) : a , habitus ( dorsal ) ; b , p5 . male : c , habitus ( right lateral side ) ; d , p5 ( l = left leg ; r = right leg ) . female characteristics : - spinous prolongations of posterolateral corners of last thoracic segment divergent . - proximal part of genital segment asymmetrically inflated , with a prominent curved process usually on right side . - blunt processes on caudal rami are prominent . - penultimate segment of p5 as broad as long . male characteristics : - processes on caudal rami as in female ; - proximally directed spine on segment 3 of left p5 blunt , short and stumpy ; terminal segment of right leg with a prominent undulating border , and 4 setae ; the distance between setae 1 and 2 and 2 and 3 almost equal . remarks :
in the southwest pacific males right p5 terminal segment with the distance between spines 1 and 2 ( numbering from proximal part of segment ) is almost twice that of the distance between spines 2 and 3 ; spines 3 and 4 much shorter than spines 1 and 2 .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 173 , fig . 1 ] . female ( from suruga bay ) : a , habitus ( dorsal ) ; b , idem ( lateral left side ) ; c , forehead ( lateral ) ; d , genital compound somite ( dorsal ) ; e , idem ( lateral ) ; f , idem ( ventral . c : copulatory pore ; cd : copulatory duct ; g , gonopore .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 143 , fig . 2 ] . female : scanning electon microscope micrographs of genital compound somite ; aventral ( copulatiory pores damaged ) ; b , right gonopore ; c , left gonopore . scale bars = 0 . 3 mm ( a ) ; 0 . 05 mm ( b , c ) .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 175 , fig . 3 ] . female : a , left a1 ( segments i - xiv ) ; b , idem ( segments xv - xx ) ; c , idem ( segments xxi - xxviii ) ; d , idem ( segments xiv - xv ) ; e , idem ( segments xxvii - xxviii ) ; f , a2 ; g , 2nd endopodal segment of a2 ; h , labrum and paragnaths ; i , md . arrowheads indicate gland openings .
issued from : h . y . soh , s . ohtsuka , h . imabayashi & s . sawamoto in species diversity , 1998 , 3 . [ p . 177 , fig . 4 ] . female : a , mx1 ; b , mx2 ; c , mxp ; d , p1 ( anterior ) . arrowheads indicate gland openings .
issued from : t . k . s . björnberg & a . f . campaner in hydrobiomogia , 1988 , 167 / 168 . [ p . 352 , figs . d , e , f ] . female ( from chile ) : d , forehead ( dorsal ) ; e , idem ( lateral ) ; f , metasomal corners and genital segment ( dorsal ) .
issued from : t . k . s . björnberg & a . f . campaner in hydrobiomogia , 1988 , 167 / 168 . [ p . 353 , figs . b - f ] . female : b , genital segment with attached couplers ( lateral right side ) ; d , idem ( ventral ; e , right coupler with spermatophore after removal of filling material ) ; c , genital segment ( lateral right side ; another specimen ) .
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 35 , fig . 1 ] . male ( from w caribbean ) : a - b , habitus ( lateral and dorsal , respectively ) ; c , forehead lateral ; ( showing frontal process and depression ) ; d , 5th pedigerous and 1st urosomites ( dorsal ) ; e , anal somite and caudal rami ( dorsal ) ; f , same ( lateral ) . scale bars : 3 mm ( a , b ) ; 0 . 7 mm ( c - f )
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 36 , fig . 2 ] . male : a , left a1 ( segments 1 - 10 ) ; b , same ( segments 11 - 16 ) ; c , same ( segments 17 - 23 ) ; d , right a1 ( segments 1 - 10 ) ; e , same ( segments 11 - 20 ) ; f , same ( detail of modified seta and aesthetascs ) ; g , detail of segment 17 and distal acute process ( position of integumental pores indicated by solid circles . scale bars : 0 . 7 mm ( a - e ) ; 0 . 05 mm ( f ) ; 0 . 35 mm ( g ) .
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 37 , fig . 3 ] . male : a , a2 ; b , md ( mandibular palp ) ; c , md ( mandibular blade showing detail of teeth and blade ornamentation ) ; d , mx2 ; e , mxp ; e , p1 ; g , rostal area ( ventral ) . position of integumental pores indicated by solid circles . scale bars : 0 . 7 mm ( a , b - g ) ; 0 . 2 mm ( c ) .
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 38 , fig . 4 ] . male : a , p2 ; b , detail of hook - like and spiniform processes on 1st endopodal segment of p2 ; c , p3 ; d , p4 ; e , p5 ( anterior ) ; f , basipod and exopod of left p5 ( lateral view ) ; g , detail of processes of 2nd exopodal segment of left p5 ; h , detail of distal exopodal segment of right p5 ( showing ornamentation ) . position of integumental pores indicated by solid sircles . scale bars : 0 . 7 mm ( a , c - e , h ) ; 0 . 3 mm ( b , f , g ) .
issued from : e . suarez - morales in zootaxa , 2007 , 1621 . [ p . 42 , table 3 ] . male : comparison of pore number of p5 of from the caribbean ( gpc ) , the indian ocean ( gpl ) from soh & al . , 1998 , and g . sewelli ( gsi ) from saraswathy & bradford , 1980 from the indian ocean . cx = coxa ; bp = basipodite ; enp = endopodite ; exp = exopodite ; r = right ; l = left .
issued from : c . b . wilson in bull . u . s . natn mus . , 100 , 14 ( 4 ) . [ pl . 25 , figs . 377 - 378 ] . as metridia atra . female ( from 13°37 ' 40 ' ' n , 120°39 ' e ) : 377 , endopod of p2 ; 378 , p5 .
issued from : c . b . wilson in bull . u . s . natn mus . , 100 , 14 ( 4 ) . [ pl . 11 , figs . 117 - 119 ] . female ( from pacific ) : 117 - 118 , habitus ( dorsal and lateral ) . male : 119 , habitus ( dorsal ) . nota : the color is very dark , almost black , with the ventral surface of the genital segment a reddish brown .
issued from : m . saraswathy in handbook to the intern . zoopl . coll . , cochin - 18 , kerala state , india , 1973 . [ p . 194 ] . comparison between g . scotti ( = g . princeps ) and g . sewelli .
issued from : h . b . owre & m . foyo in fauna caribaea , 1 , crustacea , 1 : copepoda . copepods of the florida current . 1967 . [ p . 73 , fig . 479 ] . female : 479 , p2 .
issued from : h . b . owre & m . foyo in fauna caribaea , 1 , crustacea , 1 : copepoda . copepods of the florida current . 1967 . [ p . 17 , fig . 47 ] . female : 47 , p5 .
issued from : h . b . owre & m . foyo in fauna caribaea , 1 , crustacea , 1 : copepoda . copepods of the florida current . 1967 . [ p . 20 , fig . 74 ] . male : 74 , p5 .
sewell , 1948 ( part . , p . 330 , 503 , 516 , 521 , 530 , 548 , 558 , 561 ) ; grice , 1963 a ( p . 496 ) ; fleminger , 1967 a ( tabl . 1 ) ; grice & hulsemann , 1967 ( p . 17 ) ; 1968 ( tab . 2 ) ; morris , 1970 ( p . 2301 ) ; lee & al . , 1971 ( p . 1151 , 1152 ) ; gueredrat & friess , 1971 ( p . 187 , fig . 1 ) ; roe , 1972 ( p . 277 , tabl . 1 , tabl . 2 ) ; björnberg , 1973 ( p . 338 , 387 ) ; lee & barnes , 1975 ( p . 265 , lipids ) ; childress , 1975 ( p . 787 , respiratory rate ) ; 1977 ( p . 19 , pressure , temperature v . s . oxygen consumption ) ; vives , 1982 ( p . 293 ) ; petipa & borichenko , 1985 ( tab . 2 ) ; madhupratap & haridas , 1986 ( p . 105 , tab . 1 ) ; lozano soldevilla & al . , 1988 ( p . 59 ) ; heinrich , 1990 ( p . 18 ) ; bowlby & case , 1991 ( p . 329 ) ; 1991 a ( p . 440 ) ; suarez - morales & gasca , 1998 a ( p . 110 ) ; razouls & al . , 2000 ( p . 343 , appendix ) ; haury & al . , 2000 ( p . 69 , table 1 ) ; lenz & al . , 2000 ( p . 338 ) ; fields & al . , 2002 ( p . 173 ) ; fernandes , 2008 ( p . 465 , tabl . 2 ) ; galbraith , 2009 ( pers . comm . ) ; inouye & sahara , 2008 ( p . 96 , luciferase ) ; takenaka & al . , 2012 ( p . 1669 , fig . 3 , table 1 , bioluminescence ) ; teuber & al . , 2013 ( p . 1 , table 2 , fig . 5 , respiration rates ) ; in calcofi regional list ( mdo , nov . 2013 ; m . ohman , comm . pers . )
issued from : m . saraswathy in handbook to the intern . zoopl . coll . , cochin - 18 , kerala state , india , 1973 . [ p . 193 ] . localities from g . scotti ( = g . princeps ) ( black circle ) and g . sewelli ( white circle ) obtained .
sub - antarct . ( se pacif . ) ( in björnberg & campaner , 1988 ) , de trindade is . , off st . paul is . , off gabon , g . of guinea , cape verde is . , off morocco - mauritania , canary is . , caribbean sea , belize , florida , w sargasso sea , s . cape hatteras , indian , bay of bengal , philippines , japan , s . hokkaido , off n new caledonia , new zealand ( e north island ) , pacif . ( equatorial ) , ne pacif . ( queen charlotte is . ) , hawaii , off n hawaii , california ( off monterey bay , san pedro , santa catalina , san clemente , velero and tanner basins ) , ne easter is . , sw galapagos , pacif . ( se tropical ) , off juan fernandez is . , peru , chile ( n & s , sub - antarct . )
( 10 ) f : 10 , 5 - 10 ; ( 16 ) f : 10 , 75 - 10 , 5 ; m : 10 , 4 - 9 , 4 ; ( 59 ) f : 12 - 9 ; m : 12 - 9 ; ( 94 ) f : 10 . 5 - 11 . 4 [ indian ] ; 10 . 0 - 10 . 7 [ atlant . ] ; 10 . 0 - 11 . 6 [ pacif . ] ; m : 9 . 6 - 10 . 6 [ indian ] ; 9 . 1 - 10 . 4 [ atlant . ] ; 10 . 1 - 11 . 1 [ pacif . ] ; ( 199 ) m : 10 , 08 - 9 , 92 ; ( 770 ) f : 10 , 47 [ indien ] ; 10 , 9 [ japon : suruga bay ] ; m : 9 , 54 [ indien ] ; ( 909 ) f : 10 , 45 - 11 , 06 ; m : 9 , 39 - 9 , 85 [ n - z ] ; ( 986 ) m : 10 , 1 ; { f : 9 , 00 - 12 , 00 ; m : 9 , 00 - 12 , 00 } the mean female size is 10 . 589 mm ( n = 19 ; sd = 0 . 6712 ) , and the mean male size is 10 . 058 mm ( n = 22 ; sd = 0 . 7233 ) . the size ratio ( male : female ) is about 0 . 94 ( n = 9 ; sd = 0 . 0347 ) .
razouls c . , de bovée f . , kouwenberg j . et desreumaux n . , 2005 - 2018 . - diversity and geographic distribution of marine planktonic copepods . sorbonne université , cnrs . available at urltoken [ accessed july 09 , 2018 ]