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 )
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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
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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
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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
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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 ]


lopinga achine , the woodland brown , is a butterfly in the family nymphalidae .

the species is widely distributed , but uncommon and local in continental europe .

most of the suitable habitats for the species have been transformed to gardening .

the woodland brown may be found in warm openings of damp unmanaged mature forests .

lopinga achine

remarks : lopinga achine occurs locally from france across central and eastern europe to northern asia and japan .
woodland brown lopinga achine scopoli 1763 ( nymphalidae : satyrinae ) is one of the threatened butterflies in finland and europe in general .
bergman k . o . ( 2001 ) population dynamics and the importance of habitat management for conservation of the butterfly lopinga achine . journal of applied ecology , 38 , 1303 - 1313
pararge achine var . jezeonsis matsumura , 1919 ; thous . ins . japan . addit . 3 : 727 ; tl : hokkaido
pararge achine ab . vindobonensis kammel , 1913 ; ent . zs . 27 ( 15 ) : 83 ; tl : rohrwald nr wien
polyargia verity , 1957 ; var . géograph . saison . pap . diurn . fr . ( 3 ) : 436 ; ts : papilio achine scopoli
habitat : lopinga achine needs clear , open forests which are rich in undergrowth and grasses ( carex ! ) and show a larger age diversity . for example , it flies in the coppice woodlands or in floodplain forests along rivers . clear and moist spruce forests are also colonized , while dense , low - light spruce forests provide no habitat .
a beautifully marked and rather rare butterfly . i first saw achine at this one location in côte - d ' or , central france , in late june 2007 , a single tired and worn female , probably the last of the colony . i revisited the site again in early june 2008 and was rewarded with several fresh achine flying , not to mention a couple of scarce fritillaries ( euphydryas maturna ) .
we thank d . cizkova , j . dovala , p . dufkova , l . honc , j . patera , j . piszkiewicz , l . spitzer for enjoyable companionship in the field . zuzana veverkova , our fellow zoologist who was born in a village next to the wood , recalled collecting grass in the wood with her grandmother in the late 1970s . lepidopterists l . honc , m . kralicek , l . stiova and j . uricar , and botanists m . chytry , v . grulich and j . rolecek informed us on their observations from historical lopinga achine sites . we acknowledge funding by the grant agency of the czech republic ( 526 / 04 / 0417 ) , czech department of environment ( vav / 620 / 1 / 03 ) and education ( lc06073 ) .
ceu , s . scandinavia , russia , n . asia , amur , ussuri , japan . see [ maps ]
800x494 ( ~ 45kb ) male open pine / birch forest , akademgorodok [ academy town ] , novosibirsk , west siberia , russia . 20th june 1998 , photo © oleg kosterin
682x490 ( ~ 78kb ) upperside male open pine / birch forest , akademgorodok [ academy town ] , novosibirsk , west siberia , russia . 20th june 1998 , photo © oleg kosterin
larva on gramineae , lolium , triticum [ h & r ; ] , agropyron , dactylis , melica , carex spp . [ bru ] , lang , 1884
urals - s . siberia , china , mongolia , korea , japan . see [ maps ]
hipparchia ( pararga ) deidamia eversmann , 1851 ; bull . soc . imp . nat . moscou 24 ( 2 ) : 617 ; tl : irkutsk
pararge deidamia ; [ bow ] : pl . 204 , f . 4 ; [ otakar kudrna ]
800x600 ( ~ 95kb ) underside an open birch / larch forest on a western slope of a mountain massif at the polovinnaya pad ' valley left board few km of the argun ' river left bank 12 km s of the village uryupino , gazimurskozavodskoi district , e chita province , e transbaikalia , siberia , russia . 28th july 1997 , photo © oleg kosterin
1115x856 ( ~ 322kb ) upperside female a road on the akkem river right bank terrace in its lowermost reaches going through a spruce forest , katunskii range northern foot , central [ russian ] altai mts . , ust ' - koksa district , altai republic , west siberia , russia . 7th july 2007 , photo © oleg kosterin
pararge deidamia interrupta fruhstorfer , 1909 ; int . ent . zs . 3 ( 24 ) : 134 ; tl : japan , nikko
pararge deidamia thyria fruhstorfer , 1909 ; int . ent . zs . 3 ( 24 ) : 134 ; tl : c . china ; tsintau
crebeta lehmanni forster , 1980 ; spixiana 3 ( 1 ) : 1 , f . 1 - 2 ; tl : nepal , daulaghiri so - seite , 2700m
[ maps ] warning ! the maps are automatically generated from the textual information , and the process does not always produce acceptable result ; see about maps for more info .
[ bru ] ; tuzov , bogdanov , devyatkin , kaabak , korolev , murzin , samodurov , tarasov , 1997 guide to the butterflies of russia and adjacent territories : hesperiidae , papilionidae , pieridae , satyridae butts . russia adj . terr . 1
if you have corrections , comments or information to add into these pages , just send mail to markku savela keep in mind that the taxonomic information is copied from various sources , and may include many inaccuracies . expert help is welcome .
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host plants : the caterpillar feeds on grasses ( cyperaceae and poaceae ) , especially on carex alba , carex brizoides and brachypodium .
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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
van swaay , c . , wynhoff , i . , verovnik , r . , wiemers , m . , lópez munguira , m . , maes , d . , sasic , m . , verstrael , t . , warren , m . & settele , j .
van swaay , c . & cuttelod , a . ( iucn red list unit )
justification : in europe and the eu27 countries , a population decline of more than 30 % falls within the uncertainty limits for this species ' population decline . therefore , this species is considered as vulnerable . it should however be noted that both the distribution and population size of this species in western and central europe have declined severely during the 20th century ( so before the last ten years ) .
this is a central european species , local and very rare . in spain possibly only in biscay , in france except atlantic coast and south , switzerland , north of italy via slovenia to the east , continuous from southeast germany to the south of finland . 200 - 1 , 500 m . it is also found eastwards across northern central asia to korea and japan . the global distribution area of the species is situated both within and outside europe .
changes in woodland or woodland management are the main threats all over the continent . nevertheless agricultural abandonment and land drainage are important threats in some countries , mainly because the habitat was maintained in a successional change by grazing .
the species is listed on the habitats directive annex 4 and bern convention annex 2 . in countries where the species is declining , important habitats should be protected and managed . the effects of conservation actions should be monitored by a butterfly monitoring scheme . it is important to maintain suitable glades by grazing or clearing at regular intervals to prevent overshading of its habitat . to improve overgrown sites small clearings ( 10 - 30 m in diameter ) should be created , wide enough to allow the sun to reach the ground ( bergman pers . comm . ) .
van swaay , c . , wynhoff , i . , verovnik , r . , wiemers , m . , lópez munguira , m . , maes , d . , sasic , m . , verstrael , t . , warren , m . & settele , j . 2010 .
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2017 photographs highlighted in yellow . click on any photograph to go to an enlarged picture , or simply scroll down the page .
it is a very nervous butterfly , remarkably difficult to approach for a photograph , as these distance shots testify ; any closer and it is off very quickly . the underside is magnificent , as can just about be seen from 15945 and 40676 below . in 2009 it was present in good numbers at this same site , but not present in mid - june 2010 , perhaps the flight period being over by then .
were flying but they were so active , even early in the morning , that photography was impossible . however , a trip a few days later to a site in côte - d ' or searching for
still flying there . 37574 very kindly posed for the camera in overcast conditions .
was only just starting to emerge on 28 may . we saw several very fresh but even more nervous males , hence 40676 as a distance shot being the nearest we could get . contrast this with 2017 , when a visit to the same site revealed an
it has a habit , unique among all butterfly species in my experience , of flying into the dense trees and bushes and rarely settling at less than 2 - 3m above the ground . a good photograph of the underside of this fabulous butterfly is still at the top of my wish list for the future .
a female , very much at the end of the flight period , but photographic opportunities are so infrequent with this species , that anything gets included .
i had this down as a male , possibly based on behaviour , although the extent of the unh white markings matches the illustration of the female in t & l rather better than the male . the sexes are so similar that maybe this is not a reliable pointer , though .
a male , as mentioned in the narrative , the nearest we could get to this early emerger .
a beautifully marked fresh male , taking salts from the droppings of a wild animal that i am not able to identify . it was a rare opportunity to get a close shot , but even on approaching very gently , it was still enough to spook the subject and , despite considerable patience , it did not return .
a female , photographed at the same site only a week or so later , but already showing some fading , even for a female given that they tend to emerge later than the males .
we thank silja kana , juha pöyry and anu tiitsaar for their expert help in the field , karl - olof bergman for giving directions to field sites , ants kaasik for statistical advice , as well as robert b . davis , toomas esperk , freerk molleman and tiit teder for comments on the manuscript . the study was supported by estonian science foundation grant 9294 , targeted financing project sf0180122s08 and by the european union through the european regional development fund ( center of excellence fibir ) as well as the swedish research council formas and the strategic research programme eko klim at stockholm university .
( nymphalidae : satyrinae ) larvae and ovipositing females : implications for conservation . biol conserv 88 : 69–74
bergman ko ( 2005 ) åtgärdsprogram för bevarande av dårgräsfjäril . rapport nr 5527 , naturvårdsverket
( nymphalidae : satyrinae ) in a fragmented landscape . biol conserv 102 : 183–190
( fabricius , 1787 ) in south - western france ( lepidoptera : satyridae ) . entomol z 116 : 186–188
gotthard k ( 2004 ) growth strategies and optimal body size in temperate pararginii butterflies . integr comp biol 44 : 471–479
gurevitch j , scheiner sm , fox ga ( 2006 ) the ecology of plants . sinauer associates inc , sunderland , usa , p 518
hanski i , singer mc ( 2001 ) extinction - colonization dynamics and host - plant choice in butterfly metapopulations . am nat 158 : 341–353
javoiš j , tammaru t ( 2004 ) reproductive decisions are sensitive to cues of life expectancy : the case of moth . anim behav 68 : 249–255
karlsson b , wiklund c ( 1985 ) egg weight variation in relation to egg mortality and starvation endurance of newly hatched larvae in some satyrid butterflies . ecol entomol 10 : 205–211
( nymphalidae : satyrinae ) : implications for conservation . j insect conserv 16 : 305–313
) in the czech republic : habitat use , demography and site management . j insect conserv 12 : 549–560
kukk t , kull t ( 2005 ) atlas of the estonian flora . maaülikool põllumajandus - ja keskkonnainstituut , tartu
nakamura y ( 2011 ) conservation of butterflies in japan : status , actions and strategy . j insect conserv 15 : 5–22
settele j , feldmann r , reinhardt r ( 1999 ) die tagfalter deutschlands : ein handbuch für freilandökologen . umweltplaner und naturschützer , stuttgart
tammaru t , javoiš j ( 2000 ) responses of ovipositing moths ( lepidoptera : geometridae ) to host plant deprivation : life - history aspects and implications for population dynamics . environ entomol 29 : 1002–1010
van dyck h , van strien aj , maes d , van swaay c ( 2009 ) declines in common , widespread butterflies in a landscape under intense human use . conserv biol 23 : 957–965
van halder i , barbaro l , corcket e , jactel h ( 2008 ) importance of semi - natural habitats for the conservation of butterfly communities in landscapes dominated by pine plantations . biodivers conserv 17 : 1149–1169
van swaay c , warren m , loïs g ( 2005 ) biotope use and trends of european butterflies . j insect conserv 10 : 189–209
zalucki mp , clarke ar , malcolm sb ( 2002 ) ecology and behavior of first instar larval lepidoptera . annu rev entomol 47 : 361–393
zovi d , stastny m , battisti a , larsson s ( 2008 ) ecological costs on local adaptation of an insect herbivore imposed by host plants and enemies . ecology 89 : 1388–1398
lindman , l . , johansson , b . , gotthard , k . et al . j insect conserv ( 2013 ) 17 : 375 . urltoken
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benes j , konvicka m ( 2006 ) denní motýli v národních maloploškách : první poznatky z celostátní inventarizace . ochrana prirody 61 : 145–150
benes j , kuras t ( 1997 ) dlouhodobé změny diverzity heliofilních motýlů ( lepidoptera ) opavské pahorkatiny a nízkého jeseníku ( česká republika ) . i cas slez muz opava ( a ) 46 : 135–158
benes j , cizek o , dovala j , konvicka m ( 2006 ) intensive game keeping , coppicing and butterflies : the story of milovicky wood , czech republic . forest ecol manag 237 : 353–365
benes j , konvicka m , dvorak j , fric z , havelda z , pavlicko a , vrabec v , weidenhoffer z ( 2002 ) butterflies of the czech republic : distribution and conservation i . som , praha
borhidi a ( 1995 ) social behaviour types , the naturalness and relative ecological indicator values of the higher plants in the hungarian flora . acta bot hung 39 : 97–181
braun - blanquet j ( 1964 ) pflanzensoziologie , grundzüge der vegetationskunde , 3rd edn . springer , wien
chytry m . , kucera t , koci n ( eds ) ( 2001 ) katalog biotopů české republiky . aopk , prague
ellenberg h , weber he , düll r , wirth v , werner w , paulissen d ( 1992 ) zeigerwerte von pflanzen in mitteleuropa , 2nd edn . scr geobotanica 18 : 1–258
fric z , konvicka m ( 2007 ) dispersal kernels of butterflies : power - law functions are invariant to marking frequency . basic appl ecol ( in press )
greatorex - davies jn , sparks th , hall ml , marrs rh ( 1993 ) the influence of shade on butterflies in rides of coniferized lowland woods in southern england and implications for conservation management . biol conserv 63 : 31–41
hofmeister j , mihaljevic m , hosek j , sadlo j ( 2002 ) eutrophication of deciduous forests in the bohemian karst ( czech republic ) : the role of nitrogen and phosphorus . forest ecol manag 169 : 213–230
horváth f , dobolyi z , morschhauser t , lokos l , karas l , szerdahelyi t ( 1995 ) flora adatbázis 1 . 2 . —taxonlista es attributum allomany . hungarian academy of sciences , vacrátót
höttinger h , pennerstorfer j ( 1999 ) rote listen ausgewählter tiergruppen niederösterreichs—tagfalter ( lepidoptera : rhopalocera & hesperiidae ) , 1 fassung 1999 . nö landesregierung abt naturshcutz , st pölten
ide j , kondoh m ( 2000 ) male - female evolutionary game on mate - locating behaviour and evolution of mating systems in insects . ecol lett 3 : 433–440
konvicka m ( 1999 ) macrolepidoptera of the litovelské pomoraví protected landscape area—i . cas slez muz opava ( a ) 48 : 41–64
( lepidoptera : papilionidae ) in the litovelske pomoravi , czech republic . j insect conserv 3 : 211–223
kralicek m , gottwald a ( 1984 ) motyli jihovychodni moravy i [ butterflies of southeast moravia i ] . okresni museum & ov csop , uhersky brod & uherske hradiste
kralicek m , gottwald a ( 1987 ) motyli jihovychodni moravy iiii [ butterflies of southeast moravia i ] . okresni museum & ov csop , uhersky brod & uherske hradiste
kudrna o ( 2002 ) the distribution atlas of european butterflies . oedippus 20 : 1–342
lastuvka z ( 1994 ) motýli rozšířeného území chko pálava . agronomická fakulta vsz , brno
lastuvka z , marek j ( 2002 ) lepidoptera of the moravian karst—diversity , communities and protection . korax , blansko
lebreton jd , burnham kp , clobert j , anderson dr ( 1992 ) modeling survival and testing biological hypotheses using marked animals—a unified approach with case studies . ecol monogr 62 : 67–118
lekes v ( 2000 ) motýli ( lepidoptera ) rožd’álovické tabule . práce muzea kolíně , řada přírodovědná 4 : 45–148
leps j , smilauer p ( 2003 ) multivariate analysis of ecological data using canoco . cambridge university press , cambridge uk
pollard e , woiwod ip , greatorex - davies jn , yates tj , welch rc ( 1998 ) the spread of coarse grasses and changes in numbers of lepidoptera in a woodland nature reserve . biol conserv 84 : 17–24
rolecek j ( 2005 ) vegetation types of dry - mesic oak forests in slovakia . preslia 77 : 241–261
schneider c ( 2003 ) the influence of spatial scale on quantifying insect dispersal : an analysis of butterfly data . ecol entomol 28 : 252–256
schtickzelle n , baguette m , le boulenge e ( 2003 ) modelling insect demography from capture - recapture data : comparison between the constrained linear models and the jolly–seber analytical method . can entomol 135 : 313–323
schtickzelle n , le boulenge e , baguette m ( 2002 ) metapopulation dynamics of the bog fritillary butterfly : demographic processes in a patchy population . oikos 97 : 349–360
stiova l ( 1973 ) vyskyt dennich motylu v oblasti oderskych vrchu , jeseniku a hlucinske pahorkatiny . entomol zpravodaj ( ostrava ) 3 ( 2 , 3 ) : 1–20 , 1–14
tybirk k , strandberg b ( 1999 ) oak forest development as a result of historical land - use patterns and present nitrogen deposition . forest ecol manag 114 : 97–106
van swaay cam , warren ms ( 1999 ) red data book of european butterflies ( rhopalocera ) . nature and environment 99 strasbourg , council of europe publishing
. iii . population dynamics and the effect of habitat management . j appl ecol 24 : 499–513
warren ms , key rs ( 1991 ) woodlands : past , present and potential for insects . in : collins nm , thomas ja ( eds ) the conservation of insects and their habitats . academic press , london , pp 155–212
whigham df ( 2004 ) ecology of woodland herbs in temperate deciduous forests . ann rev ecol evol syst 35 : 583–621
konvicka , m . , novak , j . , benes , j . et al . j insect conserv ( 2008 ) 12 : 549 . urltoken
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a review of genetic algorithms based methods relevant to the estimation of the distribution of the woodland brown and similar organisms is given together with an outline of possible applications with this particular finnish example species .
it is included in the list of endangered flora and fauna compiled by the bern convention ( council of europe , 1993 ) and in the habitats directive ( annex iv ; van helsdingen et al , 1996 ) . there are only a few occurrences known in finland ( marttila 1990 ) .
e . g . in sweden there is only one mainland occurrence left ( k - o bergman 2001 ) .
some of these reasons constrain the possible habitat , which seem to have certain microclimate and vegetation requirements .
it seems that the optimal environment of woodland brown resembles more a diverse pastoral idyll of the past slash and burn cultivation ( kaskiviljely ) combined with graze of the resulting wasteland and natural forest fires than the current highly efficient
timberfields
of quite uniform vegetation .
as a result the woodland brown inhabits only a few small patches more or less well connected to a metapopulation ( hanski 1991a ) .
, which is considerably larger area than the previously known largest occurrence in finland in hattula having area of less than 20 . 000m
in all there are about 50 suitable sites of which most were inhabited by woodland brown ( fig . 3 ) .
the first inventory is based on a very short visit ( 5 to 15min ) to most suitable sites by the author during two visits in july 2008 .
currently it seems that there are two close by clusters of habitats ( fig . 3 ) , but that might be just due to lack of observations .
the number of suitable sites has certainly been decreasing due to very active digging of brooks ( ojitus ) to dry the once abundant wetlands .
only the smallest , uneconomical , and most difficult to dry marshes were left surrounded by dense forests .
luckily the found occurrence has a variable topography offering suitable bowls for small marshy glades between low rocky hills ( fig . 2 ) .
there might be also some other geological factors , like nutrients and ph , that has made this area one of the last resorts for the woodland brown in finland .
the small marshy glades are visible in ordinary ( civil ) satellite and aerial images , which gives one way to search for more suitable sites .
it is interesting to notice that in sweden ( and elsewhere ) the mainland occurrence is not related to wetland ( k - o bergman 1999 , 2001 ) .
therefore the marsh itself is not a key factor of a suitable habitat . also the plants that the caterpillar eats are quite common . the adult butterfly does not live long and does not seem to have any special nutritional requirements .
2 . conservation : how to prevent this occurrence from extinction . the occurrence is already surrounded and split by many different main infrastructures .
3 . theory ( biology ) why some species are very abundant while others are very rare , if known at all .
hot topic : a . m . kuris et al , nature 454 , 515 - 518 24 . july 2008 .
one possible factor causing the rarity and patchy occurrences of wood land brown may be some host - parasitoid interactions .
it seems as if woodland brown likes places where there is not many other butterfly species , which might share some parasitoid species with it .
a parasitoid , wasp or fly usually , always kills its host , which seems to lead to highly unstable host - parasitoid population dynamics .
however , there seems to be factors like patchy occurrence that may stabilise this dynamics .
there are certainly differences in the ability of moving from one glade into another between the host and its parasitoids .
there is also an obvious asymmetry of host and parasitoid : while host benefits from finding an unoccupied site the parasitoid has to find a site also already occupied by the host .
host - parasitoid dynamics can be simulated and under some assumptions also be mathematically analysed .
host - parasite model has been used also in ga based optimisation ( hillis 1992 , olssen 1996 ) .
it was therefore natural to consider gas as one tool set to analyse ecology , biodiversity , and distribution related problems .
david stockwell ' s and ian noble ' s garp ( genetic algorithm for rule - set prediction ) has been used in tens of biodiversity and distribution estimation projects gaecolbib .
ecological ga applications have been done also in finland , actually finland is at the very top of most active countries in applying ga based methods in environmental and ecological problems ( table below ) .
to find that there was already at least one study applying gas to the prediction of woodland brown ' s distribution ( h . romo 2006 ) was certainly at least as surprising for the author that it was at first hand for him to find the new occurrence .
i . hanski has developed metapopulation model concept for species having several loosely connected habitats ( hanski 1991 ) .
dramatic change in patch occupancy probability resembling phase transition happens by reduced number of habitats or increased distance between them ( hanski 1991 ) .
( genetic algorithm for rule - set production ) ( stockwell 1992 , 1999 , 2006 ) .
recently new versions of garp have been used in quite many biodiversity and distribution prediction project . we will briefly review some of them below .
desktopgarp is a software package for biodiversity and ecologic research that allows the user to predict and analyze wild species distributions .
- - -
a . t . peterson and his research group ( e . martinez - meyer , m . ortega - huerta , r . anderson ) , have helped to define software requirements , establish short and long - term goals , and test the software .
a set of geographic layers representing the environmental parameters that might limit the species ' survival ( climate , geology ) .
normalized difference vegetation index ( ndvi ) is a much used measure of multi - spectral satellite images .
it has been used with garp for neotropical species of genus coccocypselum distribution prediction ( amaral 2007 ) .
j . bond et al and a . stockman et al have studied the extinction of populations of endemic californian trapdoor spiders apomastus ( tarantula ) ( bond 2006 , stockman 2006 ) .
ecology is closely related to economy . garp has used with estimating conservation economy ( fuller 2007 ) .
estimation of biodiversity in europe has been done using garp ( thuiller 2003 ) .
while many species has difficulties in surviving others are busy invading to new areas potentially causing problems . . .
invading species distribution prediction has also been done with garp ( sanchez - cordero 2000 , christenhusz 2008 ) .
image : centers for disease control and prevention , 1600 clifton rd , atlanta , ga 30333 , u . s . a
phillips et al have compared their maxent entropy model and garp ( phillips 2004 ) .
y . wang et al have also compared garp and the entropy model maxent ( wang 2007 ) .
there seems to be a lively debate about the merits of garp vs . maxent : ( peterson 2007 , phillips 2008 etc )
wildlife planning comparison with eight heuristics including ga have been done by p . bettinger
finally h . romo et al have compared desktop garp and domain by estimating the distribution of thirteen threatened or rare butterflies , including woodland brown in ibero - balearic area .
according to the result got they recommend domain even if the results got were widely coincident ( romo 2006 ) .
there are also other ga applications in addition to the above and quite many with garp implementations .
a relative old report for environment australia by s . ferrier and g . watson evaluates the effectiveness of several modelling techniques , including ga based rule generation system in predicting the distribution of biological diversity for forested north east new south wales ( ferrier 1997 ) .
their ga was from d . peters ' and r . thackway ' s cortex system ( peters 1998 ) .
d . hughell and j . roise have done simulation studies with ga for management of timber and wildlife ( 1997 ) .
the precision of forest classification has been studies by m . katila using also ga ( 2006 ) .
finnish forest experts have analysed tropical rain forest and their biodiversity using ga and remote sensing ( rajaniemi 2005 ) .
finns have also used ga to optimize remote sensing classification ( holmström 2003 , tomppo 2004 ) .
segmentation of aerial and satellite remote sensing images is a popular application area of gas . landcover classification has been done in ( palaniappan 2000 ) .
h . fang et al have used ga to retrieve leaf area index from satellite images ( fang 2003 ) .
neural networks and fuzzy logic are also popular soft computing methods used with ga in environmental monitoring ( schleiter 2001 ) .
tutorial of machine learning methods for ecologists is given in ( olden 2008 ) .
bayesian classification and ga in plant species distribution modeling for uk has been done by m . termansen et al ( 2006 ) .
gas to optimise land usage for species having conflicting habitat requirements can be found in ( holzkamper 2006 ) .
when building new infrastructure , wildlife concerns can be modelled by optimisation methods including wildlife hazard minimization , which is a new engineering management point of view ( kalafallah 2006 ) .
a review of species distribution forecasting machine learning methods has been done by m . b . araujo and m . new ( 2006 ) .
a review of gas in ecology is given by d . morrall ( 2006 ) . in ( recknagel 2006 ) .
for a bibliography of garp and other biodiversity and ecology related contributions see bibliography gaecolbib .
chemometry and remote sensing image processing have tools that seem to be suitable for species distribution estimation .
we have used ga to both chemometrical spectral analysis wave length selection and medical image segmentation ( välisuo & alander ; 2008 ) .
there has been surprisingly many studies related to application of genetic algorithm based methods in wildlife conservation studies .
in figure 4 you can see the number of papers using gas in ecology related topics compared to the number of all ga papers .
based on the literature review given in this paper the author plans to analyse the site more carefully with ga based methods .
it would also be interesting to compare the site of the occurrence to those few existing in finland and elsewhere in europe .
plan for creating new suitable sites , including sites in heavily processed areas ( parks , gaspipeline ) . in this particular case the occurrence already overlaps a jogging path network .
consideration of other , easier to monitor species for metapopulation studies : solitary wasp and bees needing special nesting environment and some of which are also threatened by lack of suitable biotopes .
a gaspipe will divide the occurrence quite precisely at the middle and it was actually the construction work that triggered the subsequent set of actions that finally lead to considering woodland brown , gas , and remote sensing .
uppvärmning , försurning , övergödning och syrefria bottnar är några av de förändringar som pågår i världens oceaner . nu lanserar ett internationellt forskarteam en modell för att förutse och få bukt med kommande förändringar i världshavens kustområden .
många djur har en iögonenfallande färgteckning som signalerar till t . ex . artfränder och potentiella fiender . frågan om hur det gått till när de fått sina signaler har studerats i över 150 år . nu har forskare vid zoologiska institutionen med hjälp av höns visat att en psykologisk mekanism vid inlärning hos signalmottagaren förklarar evolutionen av dessa färger , och kanske också andra signalsystem i naturen .
rike stelkens har utsetts till wallenberg academy fellow 2017 av knut och alice wallenbergs stiftelse . hon kommer att att undersöka om den genetiska mångfalden som härrör från hybridisering hjälper jäst att överleva i nya miljöer .
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