apostichopus japonicus is a species of sea cucumber in the family stichopodidae .

it is found in shallow temperate waters along the coasts of south east asia and is commonly known as the japanese spiky sea cucumber or the japanese sea cucumber .

apostichopus japonicus

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file : japan sea animal , apostichopus japonicus . jpg
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draft genome of the sea cucumber apostichopus japonicus and genetic polymorphism among color variants .
figure 5 : related data from apostichopus japonicus individual levels after each gene silencing .
print version : sea cucumber apostichopus japonicus . ( 9780127999531 / 0127999531 / 900827492 )
figure 4 : related data from apostichopus japonicus primary cultured coelomocytes after each gene silencing .
draft genome of the sea cucumber apostichopus japonicus and genetic polymorphism among color variants . - pubmed - ncbi
expression analysis of immune related genes identified from the coelomocytes of sea cucumber ( apostichopus japonicus ) in response to lps challenge .
expression analysis of immune related genes identified from the coelomocytes of sea cucumber ( apostichopus japonicus ) in response to lps challenge . - pubmed - ncbi
gigadb dataset - doi 10 . 5524 / 100257 - supporting data for
draft genome of the sea cucumber apostichopus japonicus and genetic polymorphis . . .
distribution of the japanese sea cucumber apostichopus japonicus in the intertidal zone of hirao bay , eastern yamaguchi pref . , japan : suitable environmental factors for juvenile habitats
distribution of the japanese sea cucumber apostichopus japonicus in the intertidal zone of hirao bay , eastern yamaguchi pref . , japan : suitable environmental factors for juvenile habitats
distribution of the japanese sea cucumber apostichopus japonicus in the intertidal zone of hirao bay , eastern yamaguchi pref . , japan : suitable environmental factors for juvenile habitats [ 2006 ]
zhang b l , sun d y , wu y q . 1995 . preliminary analysis on the feeding habit of apostichopus japonicus in the rocky coast waters of lingshan island .
bogatyrenko , e . a . , buzoleva , l . s . , and chi , z . , potential probiotics of the far eastern trepang apostichopus japonicus producing digestive enzymes ,
zhang w j , hou h m , zhang g l , li q y , du c m . 2011 . study on diversity of intestine cultivable microorganisms from apostichopus japonicus .
lebedev , a . m . 2000 . fisheries and reserves of the far eastern sea cucumber apostichopus japonicus . russina journal of marine biology 26 ( 4 ) : 296 - 302 .
22 . 6 . problems and prospects in sustainable a . japonicus production in japan
apostichopus japonicus appears to be more tolerant to environmental fluctuations than many other species of sea cucumbers . however , the larval stages remain rather sensitive and less tolerant to variations in their environment .
three color variants of a . japonicus ( green , red , and black ) .
12 . 5 . differences in mitf gene expressions in albino and normal a . japonicus
12 . 6 . differences in astacin gene expressions in albino and normal a . japonicus
gao f , sun h l , xu q , tan j , yan j p , wang q y . 2010 . pcrdgge analysis of bacterial community composition in the gut contents of apostichopus japonicus .
zhang x c , nakahara t , murase s , nakata h , inoue t , kudo t . 2013 . physiological characterization of aerobic culturable bacteria in the intestine of the sea cucumber apostichopus japonicus .
today , in northern china , the sea cucumber apostichopus japonicus is commonly farmed in earth ponds either in an extensive or semi - intensive system . sea rafts are sometimes used , but are not very popular .
selin , n . i . 2001 . vertical distribution of the far east trepang apostichopus japonicus in vostok bay , sea of japan . russian journal of marine biology 27 ( 4 ) : 256 - 258 .
12 . 3 . differences in nutrient and melanin contents between albino and normal a . japonicus
12 . 4 . differences in histological and ultrastructural characteristics between albino and normal a . japonicus
yang z p , sun f x , liu z m , zhang l , cao w , ma y x . 2013 . screening and identification of potential enzyme producing probiotics from gut of sea cucumber apostichopus japonicus .
17 . 3 . bioremediation capability of a . japonicus when combined with bivalve and / or macroalgae
gao f , li f h , tan j , yan j p , sun h l . 2014 . bacterial community composition in the gut content and ambient sediment of sea cucumber apostichopus japonicus revealed by 16s rrna gene pyrosequencing .
li b , rong x j , liao m j , chen g p , zhang z , wang y g , xue t s . 2010 . bacteria community in the intestine and culture environment of apostichopus japonicus in winter .
− 1 d − 1 ) of apostichopus japonicus during the experimental period . means ( n = 4 ) with different letters denoting significant differences ( p < 0 . 05 ) , and bars representing standard deviations of the means .
chen , l . , li , q . and yang , j . 2008 . microsatellite genetic variation in wild and hatchery populations of the sea cucumber ( apostichopus japonicus selenka ) from northern china . aquaculture research 2008 : 1 - 9 .
dubrovskii , s . v . and sergeenko , v . a . 2002 . distribution pattern of far eastern sea cucumber apostichopus japonicus in busse lagoon ( southern sakhalin ) . russian journal of marine biology 28 ( 2 ) : 87 - 93 .
in northern inshore areas of china , many other animals inhabit the eelgrass ecosystems in which a . japonicus is abundant . this study showed that decaying eelgrass debris could act as an important food resource for a . japonicus .
kanno , m . , suyama , y . , li , q . & kijima , a . 2006 . microsatellite analysis of japanese sea cucumber , stichopus ( apostichopus ) japonicus , supports reproductive isolation in color variants . marine biotechnology , 8 : 672 - 685 .
zhao y c , zhang w b , xu w , mai k s , zhang y j , liufu z g . 2012 . effects of potential probiotic bacillus subtilis t13 on growth , immunity and disease resistance against vibrio splendidus infection in juvenile sea cucumber apostichopus japonicus .
citation : du h , bao z , hou r , wang s , su h , yan j , et al . ( 2012 ) transcriptome sequencing and characterization for the sea cucumber apostichopus japonicus ( selenka , 1867 ) . plos one 7 ( 3 ) : e33311 . urltoken
wang , p . , chang , y . , yu , j . , li , c . & xu g . 2007 . acute peristome edema disease in juvenile and adult sea cucumbers apostichopus japonicus ( selenka ) reared in north china . journal of invertebrate pathology 96 : 11 - 17 .
dong , y . , dong , s . , tian , x . , wang , f . & zhang , m . 2006 . effects of diel temperature fluctuations on growth , oxygen consumption and proximate body composition in the sea cucumber apostichopus japonicus selenka . aquaculture , 255 : 514 - 521 .
dong , s . , liang , m . , gao , q . , wang , f . , dong , y . & tian , x . 2010 . intra - specific effects of sea cucumber ( apostichopus japonicus ) with reference to stocking density and body size . aquaculture research , 41 : 1170 - 1178
cheon s ; cho sj ; hong hh ; jin s ; jo j ; kern ema ; lee hg ; lee sg ; oh j ; park c ; park jk ( 2016 ) : supporting data for
draft genome of the sea cucumber apostichopus japonicus and genetic polymorphism among color variants
gigascience database . urltoken
lysenko , v . n . , zharikov , v . v . , and lebedev , a . m . , the abundance and distribution of the japanese sea cucumber , apostichopus japonicus ( selenka , 1867 ) ( echinodermata : stichopodidae ) , in nearshore waters of the southern part of the far eastern state marine reserve ,
citation : liu x , zhou y , yang h , ru s ( 2013 ) eelgrass detritus as a food source for the sea cucumber apostichopus japonicus selenka ( echinidermata : holothuroidea ) in coastal waters of north china : an experimental study in flow - through systems . plos one 8 ( 3 ) : e58293 . urltoken
there are over 1 000 species of sea cucumbers known worldwide . more than 100 species can be found in china , amongst which more than 20 species are considered edible . most of them , e . g . thelenota ananas , stichopus chloronotus , s . variegatus and apostichopus japonicus , are distributed in the southern sea of china .
yuan , x . , yang , h . , wang , l . , zhou , y . , zhang , t . and liu , y . 2007 . effects of aestivation on the energy budget of sea cucumber apostichopus japonicus ( selenka ) ( echinodermata : holothuroidea ) . acta ecologica sinica 27 ( 8 ) : 3155 - 3161 .
zzd is one of the main aquaculture and fishery companies in china , and it is the biggest base for aquaculture and sea ranching of high - valued seafood in north yellow sea . it operates a 2000 km 2 ocean farm , producing about 60 thousand t of scallop patinopecten yessoensis , sea cucumber apostichopus japonicus and abalone haliotis discus hannai annually .
sang , c . 1963 . biology of japanese common sea cucumber , stichopus japonicus selenka . kaibundo , tokyo , japan ( in japanese with english summary )
the yield and growth condition of albino juveniles and thermal resistant juveniles was checked by experts . after the project leader reported the work statement , the experts inquired the key technology and the progress of the project and finally confirmed the achievement on artificial breeding of thermal resistant a . japonicus and albino a . japonicus . according to the results of the project , the thermal resistant strain could terminate the stage of estivation 17 days earlier than normal a . japonicus . furthermore , the thermal resistant strain could endure 1 degree higher temperature than normal a . japonicus . the results of gene expression of heat shock proteins and lethal high temperature test indicated that the thermal resistant strain had better thermostability . at present , the yield of thermal resistant a . japonicus was one million eight hundred thousand , which included eight hundred thousand f1 offspring and one million f2 offspring . in addition , 18 a . japonicus pedigrees were established and each pedigree had 5000 - 30000 individuals . in march 2010 , the artificial breeding of albino a . japonicus was carried out . up to october 2010 , the yield of albino offspring was one million three hundres thousand .
pyrosequencing was proven to be efficient in rapidly identifying a large set of genes for the sea cucumber a . japonicus . through the large - scale transcriptome sequencing as well as public est data integration , we performed a comprehensive characterization of the a . japonicus transcriptome and identified candidate aestivation - related genes . a large number of potential genetic markers were also identified from the a . japonicus transcriptome . this transcriptome resource would lay an important foundation for future genetic or genomic studies on this species .
some researchers pointed out that the appropriate temperature for the growth of a . japonicus is 5–20°c , while the optimum is 10–16°c [ 40 ] – [ 42 ] . during the experiment , the water temperature ranged from 13 . 5–20 . 8°c ( fig . 2 ) . with an increase in water temperature to 18°c , a . japonicus gradually went into an aestivation state , and the fpr of a . japonicus reduced rapidly ( fig . 2 ) . before entering into aestivation , the fpr of a . japonicus was higher , which were 0 . 33 g·ind . −1 d −1 , 1 . 31 g·ind . −1 d −1 , 1 . 31 g·ind . −1 d −1 , 0 . 63 g·ind . −1 d −1 for treatments es10 , es20 , es40 , and es100 , respectively ( fig . 3 ) . according to the experiment the water temperature had an effect on the sgr and fpr of a . japonicus which fed on the mixed food containing eelgrass detritus and seafloor surface sediment . when the water temperature was between 13 and 17°c , a . japonicus grew faster .
yaqing , c . , changging y . & songxin . 2004 . pond culture of sea cucumbers , apostichopus japonicus , in dalian . in : a lovatelli , c . conand , s . purcell , s . uthicke , j - f . hamel & a . mercier ( eds ) , advances in sea cucumber aquaculture and management , pp . 269 - 272 . fao fisheries technical paper no 463 . fao , rome .
jo , j . , oh , j . , lee , h . - g . , hong , h . - h . , lee , s . - g . , cheon , s . , … park , c . ( 2017 ) . draft genome of the sea cucumber apostichopus japonicus and genetic polymorphism among color variants . gigascience , 6 ( 1 ) , 1–6 . doi : 10 . 1093 / gigascience / giw006
publication date 2015 series developments in aquaculture and fisheries science ; volume 39 note includes index . available in another form print version : sea cucumber apostichopus japonicus : history , biology and aquaculture . london , england : academic press , ©2015 xxiii , 454 pages developments in aquaculture and fisheries science ; volume 39 ( 9780127999531 ) isbn 0127999531 ( trade cloth ) 9780127999531 ( trade cloth ) 9780128004678 ( e - book ) 0128004673 ( e - book ) 9780127999531
in order to verify which factors affect habitat selection for aestivating and in the active adult apostichopus japonicus , animals were tested for their selection of attachment site in an experimental device ( 1 - m pipes ) in which the perceived environmental stimuli ( light intensity , degree of contact with a hard surface , geotaxis ) varied depending on the attachment site . during the aestivating season , the animals showed a strong selection for attachment sites during the daytime and nighttime ; they also showed positive stereotaxis ( thigmotaxis ) , negative phototaxis , and negative geotaxis . the results suggest that ( 1 ) habitats are not suitable for the aestivating adult a . japonicus unless these three environmental requirements are satisfied .
to identify genomic repeat elements in the a . japonicus genome assembly , we ran repeatmasker ( version 4 . 0 . 6 ) [ 19 ] using the repbase transposable element library ( release 20150807 ) [ 20 ] and the de novo repeat library constructed by repeatmodeler ( version 1 . 0 . 8 ) [ 21 ] . approximately 27 . 2 % of the a . japonicus genome was identified as interspersed repeats .
generally , a . japonicus feeds on sediments containing organic matter , which includes microorganisms and the detritus of plants or animals . results in this experiment showed that a . japonicus could use eelgrass detritus as a food resource . according to the present study , a mixture of z . marina debris and seafloor muddy sediments with an organic content of 19 . 6 % ( es40 ) , could lead to a better growth effect .
the collecting of the decaying eelgrass leaves and the invertebrate apostichopus japonicus from swan lake of weihai was permitted by peiliang wang , manager of mashan group co . ltd . no specific permit was required for the collecting of the sediment from jiaozhou bay , qingdao , where is not privately owned or protected . ethical approval was not required for this study because no endangered animals were involved . however , specimen collection and maintenance were performed in strict accordance with the recommendations of animal care quality assurance in china .
recently the institute of oceanology , chinese academy of sciences and shandong oriental ocean sci - tech co . , ltd collaborated on the establishment of sea products breeding and healthy aquaculture lab . these affiliations carried out the research on the breeding of thermal resistant a . japonicus and albino a . japonicus jointly , and finally achieved a series of innovative outcome , which laid the groundwork for popularization and industrialization of thermal resistant strain and albino strain .
according to several sources of information , the output of apostichopus japonicus in dalian was 906 tonnes in 1955 . with the improvement of breeding technology , the quantity of farmed sea cucumbers has increased significantly . in 1999 , the farming area was estimated at around 32 000 hectares and the output at 2 000 tonnes ; the following year , in 2000 , the farming area covered 48 000 hectares , while the output was 3 000 tonnes valued at 200 million yuan . in 2002 , the production reached 8 000 tonnes .
yin - geng , w . , chun - yun , z . , xiao - jun , r . , jie - jun , c . & cheng - yin , s . 2004 . diseases of cultured sea cucumber , apostichopus japonicus , in china in : a lovatelli , c . conand , s . purcell , s . uthicke , j - f . hamel & a . mercier ( eds ) , advances in sea cucumber aquaculture and management , pp . 297 - 310 . fao fisheries technical paper no 463 . fao , rome .
schematic workflow of a . japonicus genome assembly and annotation . the left side represents the genome assembly and the right side represents the transcriptome assembly that was performed in previous publications . to achieve suitable gene prediction , we integrated these two assembly results .
( of stichopus japonicus selenka , 1867 ) selenka , e . ( 1867 ) . beitrage zur anatomie und systematik der holothurien . der philosophischen facultat zu gottingen in december 1866 , als dissertation vorgelegt . : pp . 291 - 374 . [ details ]
the spatial and size - frequency distribution of the japanese sea cucumber , apostichopus japonicus , in the far eastern state marine reserve and unprotected parts of peter the great bay , sea of japan was studied using both scuba diving equipment and a remotely operated underwater sub - fighter 3000 vehicle . the abundance of the japanese sea cucumber and the pattern of its distribution over the southern part of the reserve were determined . it was found that the density of sea cucumber aggregations in the studied unprotected parts of peter the great bay is not lower than that over the major portion of the reserve .
at present , artificial breeding and culture of sea cucumbers is still a work in progress , but the scale of the production is increasing and a number of questions related to the culture techniques are being raised , calling for further studies on the commercial aspects of a . japonicus aquaculture .
figure 1 . the development of a . japonicus . 1 . gastrula ; 2 . early auricularia ; 3 . mid auricularia ; 4 . late auricularia ; 5 . early doliolaria ; 6 . doliolaria ; 7 . early pentactula ; 8 . pentactula ( bar = 100 mm ) .
sgr of a . japonicus in treatment es0 with a negative value was significantly lower than that in other treatments ( all p < 0 . 05 ) ( fig . 1 ) . and sgr of a . japonicus in treatment es40 was significantly higher than those in treatments es10 and es20 ( f = 24 . 08 , df = 7 , p = 0 . 003 ; f = 16 . 57 , df = 7 , p = 0 . 007 , respectively ) . no significant differences were found between the other treatments es10 , es20 and es100 ( all p > 0 . 05 ) .
gut microorganisms play an important role in the digestion of their host animals . the purpose of this research was to isolate and assess the enzyme - producing microbes from the apostichopus japonicus gut . thirty - nine strains that can produce at least one of the three digestive enzymes ( protease , amylase , and cellulase ) were qualitatively screened based on their extracellular enzyme - producing abilities . the enzyme - producing strains clustered into eight groups at the genetic similarity level of 100 % by analyzing the restriction patterns of 16s rdna amplified with mbo i . phylogenetic analysis revealed that 37 strains belonged to the genus bacillus and two were members of the genus virgibacillus . enzyme - producing capability results indicate that the main enzyme - producing microflora in the a . japonicus gut was bacillus , which can produce protease , amylase , and cellulase . virgibacillus , however , can only produce protease . the high enzyme - producing capability of the isolates suggests that the gut microbiota play an important role in the sea cucumber digestive process .
( of stichopus japonicus var . typicus théel , 1886 ) théel , h . ( 1886 ) . report on the holothurioidea dredged by h . m . s . ' challenger ' during the years 1873 - 76 . chall . rep . zool . no . xxxix : 290 pp . [ details ]
at the beginning of the experiment , there were no significant differences in wet body weights of a . japonicus between the five treatments ( f = 1 . 20 , df = 39 , p = 0 . 33 ) ( table 2 ) . by contrast , at the end of the experiment , final wet body weight of a . japonicus in treatment es0 was significantly lower than that in other treatments containing eelgrass debris ( all p < 0 . 05 ) . and final wet body weight in treatment es40 was significantly higher that that in treatments es10 and es20 ( both p < 0 . 05 ) .
the development of a . japonicus includes six major phases : fertilized oocytes , early development ( including cleavage , blastula and gastrula ) , auricularia ( including early auricularia , middle auricularia and late auricularia ) , metamorphosis ( including doliolaria and pentactula ) , juvenile , young sea cucumber and adult ( figure 1 ) .
mixtures of z . marina debris and sediment were used to feed a . japonicus . according to the proportion of z . marina debris , 5 diet treatments in quadruplicate were designed , i . e . , es0 , es10 , es20 , es40 , and es100 , with eelgrass debris accounting for 0 % , 10 % , 20 % , 40 % , and 100 % in dry weight , respectively . the chemical composition of each diet was analyzed in 3 replicates . each diet treatment involved 4 aforementioned pvc boxes , and each box contained 2 individuals of a . japonicus with initial wet body weights of 25 . 12±5 . 79 g·ind . −1 . the experiment was conducted from april 22nd to june 8th , 2009 in the laboratory at the institute of oceanology , chinese academy of sciences , qingdao , p . r . china . during the experiment , specific growth rates and fecal production rates of a . japonicus were measured .
assimilation efficiency ( ae ) of organic matter by a . japonicus had no significant difference in four experimental treatments ( fig . 4 ) , with mean ae being 14 . 2 % , 14 . 3 % , 24 . 2 % , and 21 . 8 % , respectively ( all p > 0 . 05 ) .
although a . japonicus always occurs abundantly in eelgrass - rich meadows in temperate northern coastal areas of china ; yet it has never been reported whether eelgrass detritus can act as a food source for sea cucumbers or not . in this study , decaying eelgrass debris and muddy sediment were mixed as food to feed a . japonicas . the ingestion and growth of the animals were then determined , in order to understand the importance of eelgrass meadows for a . japonicus , and thus clarify the importance of eelgrass - meadow restoration . it is suggested that eelgrass - meadow restoration is imperative and could not only be of huge ecological benefit , but also of potential economic value .
the initial and final wet body weight of a . japonicus was measured . animal feces were collected by siphon every 1–3 days , depending on the amount of produced feces . for estimation of assimilation efficiency ( ae ) of a . japonicus , fresh feces produced within 6 h were collected . fecal samples were dried at 60°c to constant weight and preserved for further analysis . subsamples of the dried food and feces were used to estimate organic material ( om ) by combusting ( 500°c for 3 h ) dried and pre - weighed samples . subsamples were treated with hcl vapor for 6 h to remove carbonates for analyzing organic carbon ( oc ) and nitrogen ( on ) with a perkin elmer model 240c chn analyzer standardized with acetanilide .
in this experiment , the organic contents of food in the five diet treatments were 5 . 6 % , 7 . 6 % , 10 . 9 % , 19 . 6 % , 39 . 9 % , respectively ; and organic matter in the feces of a . japonicus was lower than that in the food ; and organic carbon and nitrogen were the same ( table 1 ) .
in conclusion , a comprehensive collection of ests has been achieved for the sea cucumber a . japonicus using the 454 gs flx platform , permitting gene discovery and characterization across a broad range of functional categories . in addition , our study identified a large number of ssrs and snps , from which genetic markers can be rapidly developed to serve as tools for further genetic or genomic studies on this species .
in comparison with the 454 sequencing data obtained from sun et al . [ 15 ] , 53 % of the hq reads obtained in our study did not find significant matches ( blastn , e < 10 −4 ) with their data . for gene annotation , 94 % of gene annotations obtained in this study were not found in their data . these above results possibly suggest more representative collections of a . japonicus genes in this study .
in this experiment , sgr of a . japonicus in the five treatments were significantly different . sgr was only −0 . 65 % d −1 in the pure sediment diet ( treatment es0 ) , which was significantly lower than that in the mixed diets of z . marina debris and sediment . in contrast , the mean sgr in treatments es10 , es20 , es40 and es100 were 0 . 70 % ·d −1 , 0 . 61 % ·d −1 , 1 . 54 % ·d −1 , and 1 . 20 % ·d −1 , respectively ( fig . 1 ) . the occurrence of negative growth of a . japonicus fed with pure sediment might be due to the lower organic matter content ( 5 . 6 % ) of the diet , which might not satisfy the need for growth . the highest sgr of a . japonicus occurred in treatment es40 with organic matter content being 19 . 6 % and higher than that in es0 , es10 , and es20 ; while in the pure eelgrass - debris diet ( es100 ) with organic matter content as high as 39 . 9 % , the sgr was not significantly higher than that in treatment es40 ( p > 0 . 05 ) . sea cucumbers are deposit feeders , and clay sediment is an important component of ingested material , which is supposed to be helpful for the digestion of food [ 37 ] .
fig . 2 shows variation in fecal production rates ( fpr ; g·ind . −1 d −1 ) of a . japonicus during the experimental period . sea cucumbers in treatment es0 had hardly ingested food during the experiment period , thus only fecal production rates in other treatments were considered comparable with each other . generally , fprs in treatments es20 and es40 were significantly higher than those in treatments es10 and es100 ( all p < 0 . 05 ; fig . 3 ) .
in natural eelgrass ecosystems , deposit feeders feed on organic detritus after eelgrass decomposition , which not only accelerates the cycling of matter but also promotes the health and stability of the ecosystem structure . until now eelgrass meadows in shandong coastal waters with water depth 2–6 m have deteriorated badly , and some eelgrass meadows have even disappeared [ 43 ] . for the restoration of the declining natural resource of a . japonicus , it is suggested that the degraded eelgrass meadows should be restored in the northern inshore areas of china .
with a . japonicus , spawning often occurs between 2100 h and 0200 h . it is important to maintain a quiet environment and keep the tanks in the dark . male sea cucumbers usually are the first to spawn , which in turn induce the females to start releasing the eggs . at this point the number of spawning males should be reduced , only retaining a few strong spawners . following the spawning activity the tanks are drained and all the sea cucumbers removed . the eggs are gently washed and the tanks refilled with clean seawater at a temperature of 22 °c .
to maximize the transcript representation in a broad range of biological processes , eight a . japonicus cdna libraries representing different developmental stages and adult tissues were constructed and used for 454 sequencing ( table 1 ) . after a single - run of 454 sequencing , a total of 1 , 061 , 078 raw reads with an average length of 344 bases were obtained . the size distribution of raw reads is shown in figure 1a . of all these reads , 92 % ( 974 , 004 reads ) passed through our quality filters and represented high - quality ( hq ) reads .
the hq reads combined with the public ests were assembled into 33 , 835 contigs and 199 , 011 singletons . approximately 80 % ( 774 , 993 ) of the hq reads were incorporated into contigs . the size of contigs ranged from 101 to 6 , 323 bp , with an average length of 630 bp . the size distribution of contigs is shown in figure 1b . the sequencing coverage of contigs ranged from 2 to 7 , 429 with a mean of 23 . as expected for a randomly fragmented transcriptome , there was a positive relationship between the length of a given contig and the number of reads assembled into it ( figure 1c ) . contigs were then assembled into 29 , 666 isotigs ranging from 104 to 7 , 697 bp with an average length of 1 , 042 bp ( n50 = 1 , 294 bp ) . the size distribution of isotigs is shown in figure 1d . the average contig coverage for each isotig was 2 . 1 . the isotigs were further grouped into 21 , 071 isogroups , which possibly represents the total number of genes in the a . japonicus transcriptome . the summary statistics for the a . japonicus est assembly are shown in table 2 .
validation of rna - seq results using qrt - pcr . the relative fold changes of 10 genes expressed in a . japonicus coelemocytes at 4 h ( a ) ; 24 h ( b ) and 72 h ( c ) after lps challenge . gene abbreviations are : bf , complement factor b ; c3 - 2 , complement component 3 - 2 ; cat b , cathepsin b ; cd36 , cluster of differentiation 36 ; hsp90 , heat shock protein 90 ; lbp , lipopolysaccharide binding protein ; myd88 , myeloid differentiation primary response gene 88 ; mys , myosin ; rel , nf - κb transcription factor rel ; thy , thymosin β .
eelgrass debris used in this study was from decaying eelgrass leaves . the decaying dark colored eelgrass leaves were collected from swan lake of weihai . muddy sediment was obtained from the non - vegetated seafloor surface in the inshore area of jiaozhou bay , qingdao . both decaying eelgrass leaves and sediment were dried at 65°c , ground and sieved using 0 . 18 mm mesh . the specimens of a . japonicus used in this study were also collected from the cove of swan lake . sea cucumbers were transported to the laboratory where they were acclimatized for a week , after which an experiment was carried out using 20 pvc boxes , with each box measuring 30×40×30 cm in size .
in addition , 52 , 102 high - quality snps and 2 , 083 indels were identified from 10 , 333 contigs ( table 5 ) . the predicted snps included 30 , 366 transitions , 21 , 736 transversions . the overall frequency of all types of snps including indels was one per 393 bp . we randomly chose 32 contigs containing 201 snps to evaluate the preference of snp position in the coding regions . the result showed that 70 % of snps resided in the third codon position , while 18 % and 12 % in the first and second codon position , respectively . experimental validation was also carried out to evaluate the reliability of the predicted snps . fifteen of randomly chosen snps were subject to experimental validation in 48 a . japonicus adults using the high - resolution melting ( hrm ) genotyping method [ 42 ] . it turned out that 80 % of these snps could be validated , suggesting the majority of our predicted snps are likely to be true snps .
the history of sea cucumber fishery dates back for more than 1 000 years . over the last century , and especially the past 20 years , chinese research projects have focused on the breeding , artificial culture and processing of sea cucumber .
after the oocytes have been fertilized , they slowly sink and start their development . when the water temperature ranges from 21 to 24 °c the fertilized eggs require about 45 minutes to begin to divide themselves and finally develop into a blastula ( table 1 ) . the blastula is ciliated and rotates actively . after about 2 to 3 h , the hatched embryo gradually elongates and moves to the surface of the water column . during this ascent the embryo begins to invaginate and gradually develops into gastrula . a fully developed gastrula appears at the surface of the culture tanks 24 to 28 h after fertilization . twelve to twenty hours later the gastrula gradually elongates and begins to fold into the auricularia stage .
1 . water temperature maintained between 21 . 5 and 23 °c . 2 . results obtained in research facilities in yantai and changdao ( china ) and japan .
the auricularia stage lasts from day 7 to day 10 at a water temperature ranging from 18 to 26 °c . early auricularia larvae have a symmetrical bilateral and a length varying from 320 to 600 mm . as the larva grows , the stomach increases in size . when the auricularia reaches 600 to 750 mm , its left cavity , which adjoins the stomach and gullet , begins to grow and gradually takes on a semi circular shape . at this point , the auricularia has reached the middle phase of its development . later , the cilia develop into five pairs of symmetrical circles . at the same time , the primary tentacles begin to develop as the larva reaches the late auricularia stage . typically , the ball shaped body of the larva at this stage is the early sign of the upcoming doliolaria stage .
in the late auricularia phase , the larvae begin to shrink ; the five lipid spheres finally connect while the tentacles enlarge and move to the centre of the body . at this stage the auriculariae are still swimming in the water column ; they move from the surface to the bottom . the settlement plates should be placed in the tanks at this time .
the doliolaria phase usually lasts for 1 or 2 days at the end of which five tentacles reach out of the body . at this moment , the larvae begin to creep along the bottom for an additional 1 or 2 days . the number of cilia on the body surface of the larvae progressively decreases and x - shaped ossicles gradually begin to appear . the body becomes rounder and the first tube - like foot emerges on the left posterior section of the body . the presence of podia is a significant precursor to the juvenile stage .
the primary change in the juvenile stage is the appearance of tube - like feet ( which enables the organism to attach to a substrate ) and the ability to ingest food . the gut , mouth and papillae become clearly visible . in the next couple of months the juvenile sea cucumbers grow to about 1 cm in length assuming the appearance of an adult individual . the body pigmentation changes gradually from a transparent whitish appearance to a red , grey or light green colouration ( figure 2 ) . at this point the individual has reached the young sea cucumber stage .
there is almost no morphological difference between a young and an adult sea cucumber . in a hatchery facility the movement and feeding activity of young sea cucumbers sharply decreases when the seawater temperature exceeds 23 °c . the young specimens move from the surface of the water to the bottom of the tank . for this reason it is important to maintain an optimal temperature level in the culture tanks to ensure adequate growth and reduce the length of this relatively fragile developmental stage .
a large sea cucumber hatchery in northern china typically has the following facilities : a larvae culture volume of 2 600 m 3 divided into tanks of 10 to 20 m 3 with a depth of 1 . 4 m ; a juvenile culture volume of 4 000 m 3 divided into tanks of 30 m 3 with a depth of 1 . 7 m ( figure 3 ) ; a phytoplankton production unit with a total tank volume of 500 m 3 ; 4 filter tanks of 200 m 3 / h filtering capacity ; and 5 overhead troughs of 200 m 3 each . the facility is also fitted with a high quality water supply and drainage system as well as a heater to raise the water temperature when necessary .
the devices used for larval settlement include metal frames each fitted with 5 - 10 polyethylene screens measuring 50 x 50 cm and used as the settlement substrate ( figure 4 ) .
figure 3 . sea cucumber settlement device ( photo : a . lovatelli ) .
figure 4 . sea cucumber hatchery in the vicinity of dalian , liaoning province . juvenile culture tanks ( photo : a . lovatelli ) .
the optimum water temperature in a hatchery is between 23 and 25 °c , even though sea cucumbers can adapt to a range between 10 and 26 °c . the best salinity level is 31 . 6 , but individuals can tolerate salinities from 26 to 33 . the optimal ph is 7 . 8 , but it can range from 7 . 5 to 8 . 2 . the dissolved oxygen should be maintained at > 5 mg / litre .
broodstock harvesting . individuals are generally collected from the 20 th june to the 20 th july in the dalian area . at that time , the seawater temperature varies from 15 to 17 °c . a few individuals are usually sacrificed and dissected in order to adequately determine the maturation stage of the gonads . specimens weighing around 300 g and measuring around 20 cm in length are generally preferred .
considerable care is required during the collection and transportation of the sea cucumber broodstock . it is important to : 1 ) select each specimen individually ; 2 ) keep the selected sea cucumber away from any source of pollution ( e . g . oil spills from the boat engine ) ; 3 ) avoid exposing the sea cucumbers to high temperatures or violent movements during transport ; and 4 ) avoid simultaneous collection of the sea cucumbers and other marine organisms ( e . g . bivalves ) in order to avoid damaging the broodstock .
maintaining broodstock . following collection , the sea cucumbers should be placed in a tank filled with seawater at ambient temperature . the tanks should be clean and the individuals kept at a density of 30 individuals / m 3 for 2 - 3 days . no feed is supplied . at this point the water temperature is gradually raised to 19 °c at a rate of 1 °c per day . after 7 to 10 days the broodstock can be induced to spawn . during this phase it is necessary to maintain the quality of the water in the culture tanks . water exchange should be carried out rapidly , when required .
when the seawater temperature has been raised to 19 °c , the sea cucumber activity should be carefully monitored particularly during the night hours . the bottom of the tanks should be examined for the presence of oocytes as this is an indication that the sea cucumbers are likely to start mass release of gametes . there are two ways to induce spawning : the first method consists of drying the sea cucumber in the shade followed by a jet of violent water ; the second method uses a temperature shock by raising and lowering the water temperature by several degrees .
the clean seawater in the tanks should be stirred gently every 30 minutes ensuring that whirlpools do not form . edta is added at a concentration of 3 - 4 ppm while the larval density is kept at about 5 larvae / ml . the hatching success may exceed 90 % .
selection of the larvae begins before the auricularias tube - like foot is formed , or shortly after . an hour prior to the selection process , stirring is stopped in order to allow the auriculariae to gather at the surface of the tanks . malformed larvae are unable to swim and will sink to the bottom of the tanks . the larvae gathered at the surface are collected from the uppermost 0 . 5 m of the water column with the use of a fine mesh . these are then transferred to a new tank at a density of 300 larvae / litre .
after the appearance of the mouth in the auricularia larvae , feeding should commence immediately . early in the growth of the auricularia , the feeding regime should be in the range of 5 000 to 10 000 microalgae cells / ml . at the intermediate stage , feeding needs to be increased to about 20 000 cells / ml . monitoring the presence of algae in the stomach of the larvae helps the adjustment of the feeding regime . dunaliella is the primary algal species used and is generally mixed with diatoms and chaetoceros sp . the three micro - organisms are added in a proportion of 4 : 1 : 1 . the food mixture should be supplied in small quantities in order to maintain optimal water quality .
rearing during the auricularia stage should be carried out in still water . water should be added to the rearing tank at a rate of 20 cm per day until the tank is filled . this usually takes approximately 3 days . once the tank is filled , one third to a half of the water should be exchanged daily using a siphon . as the larvae grow the water exchange rate should gradually increased . water exchange should be carried out gently in order to avoid injuries or loss of larvae .
the culture of auricularia larvae requires the proper control of all major chemical and physical factors . generally speaking , the ph and salinity values should remain within the range tolerated by the larvae . stirring the water or adding new seawater will control dissolved oxygen levels . it is important to maintain the temperature at around 23 °c . if the larvae are reared in natural seawater , the temperature will rise gradually with the seasonal fluctuations . appropriate measures must be taken to maintain it within the acceptable range .
during the larval culture , the primary harmful organisms are copepods ; however their presence can be contained through a proper water exchange protocol and the use of edta ( 3 to 4 ppm ) , if necessary .
the primary symptoms of disease are a slow development rate of the larvae and the rotten stomach phenomenon observed in late auriculariae . this can be caused by food deficiency or sub - optimal larval densities . the use of quality feeds and maintaining low larvae densities can minimise the outbreak of diseases .
after the larvae have developed for 7 to 10 days , the five lipid spheres on each side of the larvae appear and the hydrocoel begins to develop . the larvae are now transforming into doliolaria . at this stage , the settlement substrates ( see figure 4 ) should be placed in the rearing tanks . the density of the larvae settled on substrate should be maintained between 1 to 2 individuals / cm 2 .
the rearing techniques at this point of the development of the sea cucumber juveniles are very important and should be carefully applied in order to ensure a high rate of survival .
the rearing tanks are fitted with a water flow through system . the presence of food particles and the relatively high water temperature are responsible for the proliferation of undesirable and harmful organisms , such as copepods and bacteria . the water in the tanks should be clean with a daily exchange rate of up to 200 % .
juvenile sea cucumbers grow at different rates and it is therefore important to sort them according to their size . this is done using sieves fitted with different mesh sizes . while sorting , care should be taken not to damage or kill the juvenile sea cucumbers .
raft culture . wooden rafts are usually located in sea areas not exposed to strong winds and tidal action . the sea cucumber cages are usually hung under the rafts or placed directly on the sea floor . during the rearing period , sea cucumbers are fed with sargassum sp . and other macroalgae . this farming method is not very popular due to the high costs involved .
intensive farming : intensive sea cucumber aquaculture has only recently developed . large quantities of rocks or other suitable substrates ( e . g . roof tiles ) are placed in the culture ponds to increase the surface area for the juvenile sea cucumbers and to provide shelter from predators . a refrigeration system or underground water supply is used to lower the water temperature in the ponds in the summer months when high temperatures can reduce growth and feeding rates . these pond facilities are expensive and therefore intensive farming is only carried out by large and financially strong companies .
the most suitable sea locations are rocky sites with an abundance of macrophytes and with muddy and sandy bottoms . areas protected from strong winds and tidal actions are usually favoured .
juveniles over 1 cm in length are used when stocking directly in the sea . the larger the initial size of the juveniles , the higher the survival rate . when stocking , the juveniles are placed in bags with a mesh size of 0 . 25 - 0 . 5 cm . a diver then places the bags on the sea bottom in the vicinity of existing rocks or artificially prepared rock piles . subsequently the mesh bags are opened and the juveniles are simply left to crawl away . larger juveniles ( > 5 cm ) can be released into the sea directly .
generally , two years after stocking , the sea cucumbers have reached a marketable size and can be collected . harvesting is often done in the spring ( mid - april to early june ) and autumn ( october to december ) ( figure 5 ) .
figure 5 . sea cucumber harvested off the coast of dalian , china ( photo : a . lovatelli ) .
sea ranching takes full advantage of the natural environment and this method is considered to be ecologically acceptable and therefore worth developing . however , due to a number of seasonal limitations and resource use conflict ( presence of industrial activities ) , this growout method can only be carried out in locations which have the right conditions .
liao , y . l . 1997 . fauna sinica , phylum echinodermata , class holothuroidea . science press , beijing . 334pp .
sui , x . l . & liao , y . l . 1988 . sea cucumber culture and its enhancement . agriculture publishing house , beijing . 288pp .
zhang , f . y . 1958 . the preliminary report of aquaculture and sea ranching of sea cucumber ( apostichopusjaponicus ) . journal of zoology , 2 ( 2 ) : 65 - 73 .
zhang , q . l . & liu , y . h . 1998 . the techniques of sea cucumber culture and its enhancement . qingdao ocean university publishing house , qingdao . 157pp .
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discover a faster , simpler path to publishing in a high - quality journal . plos one promises fair , rigorous peer review , broad scope , and wide readership – a perfect fit for your research every time .
copyright : © 2012 du et al . this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
funding : financial support for this work was provided by the national key technology r & d program of china ( 2011bad13b05 and 2011bad13b06 ) , and the national high technology research and development program of china ( 2012aa10a412 ) . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
( a ) size distribution of raw reads . ( b ) size distribution of contigs . ( c ) log - log plot showing the dependence of contig lengths on the number of reads assembled into each . ( d ) size distribution of isotigs .
regarding gene annotation , all isogroups were searched against the swiss - prot database with an e - value threshold of 1e - 6 . of 21 , 071 isogroups , 8 , 229 ( 39 % ) had at least one hit , which corresponded to 6 , 963 unigene names . the sequences and annotation information of all isogroups are provided in dataset s1 and table s1 . more than half of the isogroups did not match to known genes most likely due to the fact that insufficient sequences are available in public databases from phylogenetically closely related species . the annotation rate in our study is also comparable to those ( 20∼40 % ) reported in the previous de novo transcriptome sequencing studies for non - model organisms [ 26 ] , [ 27 ] , [ 33 ] , [ 34 ] , [ 35 ] .
aestivation is possibly a protection strategy for sea cucumbers to survive from high temperatures [ 10 ] , [ 11 ] . studies on sea cucumber aestivation from different perspectives such as morphology [ 10 ] , [ 11 ] , [ 37 ] , physiology [ 10 ] , [ 11 ] , [ 37 ] , [ 38 ] , and immunology [ 39 ] have been performed . nonetheless , the molecular mechanism of this process is still far from fully understood . identification and characterization of candidate genes involved in aestivation would represent the first step to understand the genetic basis of aestivation in sea cucumbers .
twelve candidate dges were selected for q - pcr validation . q - pcr results verified the differential expression of these genes between the active and aestivating sea cucumbers ( figure 3 ) . however , it should be noted that many genes in the two libraries were lowly expressed and therefore were not included in the comparison . more dges would be determined from these lowly expressed genes by performing deep transcriptome sequencing in future studies .


the black inca ( coeligena prunellei ) is a species of hummingbird found only in colombia .

its natural habitats are subtropical or tropical moist lowland forests , subtropical or tropical moist montane forests and urban areas .

it is threatened by habitat loss .

it was formerly classified as an endangered species by the iucn , but new research has shown it to be not as rare as it was believed .

consequently , it was downlisted to vulnerable in 2008 .

black inca

1st place adult black male huacaya , champion black huacaya male . judge : mrs . liz barlow
british alpaca futurity 2013 1st place intermediate black male huacaya , champion black huacaya male judge : mrs . jill macleod
honiton show 2012 1st place junior black male huacaya reserve champion black male huacaya champion was his sire lillyfiled jack of spades of inca judge : mr . nick harrington - smith
little is known of the life history of this rare species . the black inca is thought to breed between june and october ( 2 ) .
the black inca is endemic to colombia , where it is restricted to the western slopes of the east andes ( 1 ) ( 2 ) .
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please note : inca flagship is now owned by alpakka bromma and alpakka sigdal in norway . please contact them for inca flagship’s availability in norway .
the black inca is classified as vulnerable ( vu ) on the iucn red list ( 1 ) . listed under appendix ii of cites ( 3 ) .
as the shang probably did in china , the black aztec and inca unwisely used their mongol subjects for low - level work and sacrifice . in the inca empire these were the
yanaconas
: when the spanish came , they rebelled and joined forces with the spanish to bring down the inca empire .
inhabits humid montane forests , particularly where oaks are dominant ( 2 ) . black inca individuals have also been recorded in parkland and riverine forests ( 2 ) .
bas national show 2014 1st place adult black male huacaya , champion black huacaya fleece . judges : mrs . jenny jackson , mr . nick harrington - smith and mr . rob bettinson
c . 13·5 cm ; 6·4–7 g . male has long , straight , black bill , legs pale flesh - coloured ; almost entire plumage purplish - black ; throat patch small , greenish . . .
bas national fleece show 2013 1st place juintermediate black huacaya fleece judge : mr . nick harrington - smith
inca flagship is the culmination of 5 generations of careful selective breeding with the goal of achieving an outstanding black male . we here at inca alpaca are proud of our achievement with this male as he is the best jack of spades son that we have ever produced . inca flagship has a pedigree full of the most famous genetic lines found in australia and europe . his sire is our champion male lillyfield jack of spades of inca who is not only a champion himself but has and continues to sire the black champions of the uk show circuit . the dam of inca flagship is our own female inca anais who is not only a multi black show champion herself but she is from our strongest family line – jannarie rhanee . conformationally inca flagship shows strength , presence and perfect alignment of his limbs and jaw . he is heavy chested and exudes great capacity through his frame . the fleece on this male is where his value really becomes apparent . he is super fine , soft , bright and there is not one white fibre to be seen . each fibre is well crimped and forms a staple that is thin and perfectly aligned . the very high density and staple length this male exhibits resulted in the maximum cutting weight of all our yearling males . inca flagship is the ultimate in black males and seeing is believing . with an amazing pedigree which is full of champions and an outstanding phenotype inca flagship is the diamond standard for any discerning black alpaca breeder .
2 ) the holy roman emperor and spanish ruler , charles v , ( carlos quinto ) was a black man .
11 cm . dark hummingbird with long , needle - like bill . mainly black with conspicuous white patch on each side of chest and postocular spot . glittering blue shoulders . small greenish - blue throat patch . white - edged undertail - coverts . black and forked tail . long , slender , straight black bill . rosy - red legs . female slightly duller overall .
black inca is endemic to colombia , where it is distributed in the eastern cordillera in the departments of boyacá , santander and cundinamarca , between 1200 and 2800 m . this species inhabits humid montane forests , primarily forests that are dominated by oak . it is a medium - sized hummingbird with a unique color pattern : purple black with a white patch on each side of the breast , shoulders contrasting with dark blue metallic . black inca currently is listed as endangered ( en ) in colombia and globally its conservation status is rated as vulnerable ( vu ) , due to drastic declines of habitat in its narrow geographic range .
here at inca alpaca we are committed to the long term success of the british alpaca industry . we have been breeding black and grey alpacas for over twenty years and in this time have helped many new owners establish and develop their own enterprise .
we will use these following two papers to try and glean an understanding of life in the inca homeland after the europeans came .
ayllu - a clan / network of families , that constituted the basic socioeconomic unit , and local government , of inca society .
the main threats affecting the black inca include habitat loss and degradation , largely as a result of human settlement and the clearance of the forest for wood and for agricultural land , including coffee and sugarcane plantations ( 2 ) . much of the remaining habitat is greatly fragmented and isolated ( 2 ) .
yanaconas - in the inca empire yanacona was the name of the servants to the inca elites . it is important to note that they were not forced to work as slaves . some were born into the category of yanacona ( like many other professions , it was a hereditary one ) . they were to care for the herds of the nobles , do fishing , and were dedicated to other work , like the making of pottery , construction , and domestic service . yanaconas were sometimes given high positions in the inca government .
the black inca is listed under appendix ii of the convention on international trade in endangered species ( cites ) . it occurs within a nature sanctuary in one part of its range , and so receives a level of protection in this area . there is currently a need to carry out surveys in some parts of the range and to study the life - history and breeding behaviour of the species ( 2 ) .
züchner , t . & boesman , p . ( 2018 ) . black inca ( coeligena prunellei ) . in : del hoyo , j . , elliott , a . , sargatal , j . , christie , d . a . & de juana , e . ( eds . ) . handbook of the birds of the world alive . lynx edicions , barcelona . ( retrieved from urltoken on 9 july 2018 ) .
the black inca ( coeligena prunellei ) is a dark - coloured hummingbird , which has a long straight bill ( 2 ) . the plumage is generally black , with a greenish - blue throat patch ; on each side of the chest there is a white patch , and the shoulders are iridescent blue ( 2 ) . as with most hummingbirds , the female is somewhat drab in colour compared to the male ( 4 ) . the narrow wings are adapted for hovering and the legs and feet are small and weak , a feature hinted at by the name of the order to which hummingbirds and swifts belong , ‘apodiformes’ , a term that means footless ( 4 ) .
from the text above , it is clear that the inca could not possibly have been a mongol type people : simply by the mere fact that a
full - blooded , unmixed inca
could
pass
for a spaniard , simply by learning the language and dressing and acting the part . that simply would not be possible for a person of mongol phenotype . the corollary is that blacks were still prominent in spain at that time .
abstract we present two nesting records ( habitat , nest , eggs and chicks ) of black inca ( coeligena prunellei ) , a threatened and ende - mic bird species , from two localities in the eastern andes of colombia . the nests were cup - shaped , constructed with scales from tree - fern fronds , fibers , moss and spiderweb . both nests were found in the understory of oak ( quercus humboldtii ) forest , indicating the possible importance of this habitat for the nesting and conservation of this species .
mita - mita was mandatory public service in the society of the inca empire . mita was effectively a form of tribute to the inca government in the form of labor , public service was required in community - driven projects such as the building of their extensive road network and military service . all citizens who could perform labor were required to do so for a set number of days out of a year . overseers were responsible to make sure that a person after fulfilling his duty in the mita still had enough time to care for his own land and family .
macana - garcía , d . c . , j . e . zuluaga - bonilla , a . sua - becerra & s . chaparro - herrera . 2012 . primeros registros de anidación del inca negro ( trochilidae , coeligena prunellei ) . ornitología colombiana 12 : 61 - 64 .
mitimaes - i s a term commonly associated with yanaconas , but its meaning is different , as the mitimaes were used as labor for large projects . yanaconas were specifically not a part of an ayllu and were relocated individually instead of in large labor groups . an example of the differences of the classes is that mitimaes were labor that built machu piccu , but yanaconas lived and served the inca there .
historians jeffrey cole and enrique tandeter testify to corrupt local officials . cole says ,
many of the indians who came to potosi were yanaconas - artisans , former inca retainers , and others who were not affiliated with an ayllu - men who had been displaced by the conquest . tandeter adds ,
even more surprising is the fact that more than half of the forasteros of oruro were not exempt from the mita either . perhaps surprising , nonetheless it was common practice , especially if the spaniards wished to maintain and increase mining productivity .
tandeter states ,
in the area subject to the mita , the census showed a pronounced population decline of 45 percent since the toledo inspection . on the following graph cook demonstrates the total number of tributaries and mitayos in several regions encircled around the mitas .
many of the indians who came to potosi were yanaconas - artisans , former inca retainers , and others who were not affiliated with an ayllu - men who had been displaced by the conquest . tandeter adds ,
even more surprising is the fact that more than half of the forasteros of oruro were not exempt from the mita either .
once the indios had been relocated from their native hilltop dwellings to valley reservations they were put to work and taxed . the type of work the indios were forced to do varied to some extent , but if it was not working in the mines , it was agricultural or domestic work . the produce of the work would in turn be taken by spanish officials as payment for the tribute . often after paying the tribute indios were left with very little and in some cases they could not work enough to pay the tribute . establishing the tribute and the mita was a devious plan by the spaniards , who tried to understand inca history and use it for their advantage .
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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 .
sacc . 2005 and updates . a classification of the bird species of south america . available at : urltoken .
vulnerable b1ab ( i , ii , iii , v ) ; c2a ( i ) ver 3 . 1
this species has has a larger range and population than previously thought . nonetheless , its range is still highly fragmented and habitat patches are decreasing in size and quality through ongoing degradation and clearance for agriculture . it is therefore considered vulnerable .
velasquez - tibata et al . ( 2005 ) used a habitat model to estimate the population at 4 , 070 - 8 , 720 individuals , and so it is placed in the band 2 , 500 - 9 , 999 mature individuals . this equates to 3 , 750 - 14 , 999 individuals , rounded here to 3 , 500 - 15 , 000 individuals . trend justification : the degradation of remaining habitat patches within the species ' s range continues , hence its population is suspected to be declining at a slow to moderate rate .
2007 ) . most observations have been at 1 , 675 - 2 , 500 m , but it is known between 1 , 000 and 2 , 800 m ( schuchmann 1999 , t . züchner
. 1999 ) . breeding is thought to take place between june and october .
. 1999 ) , with the significant exception of guanentá - alto río fonce ( wege and long 1995 ) . however , there are still extensive forests that are poorly known to ornithology in the serranía de las quinchas , west boyacá ( stiles
. 1999 ) . prepare a management plan for the species ( t . züchner
. 1999 ) . augment conservation activities in guanentá - alto río fonce fauna and flora sanctuary ( p . g . w . salaman
. 1999 ) . protect areas of the favoured habitat holding significant populations ( p . g . w . salaman
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spoon - billed sandpiper eurynorhynchus pygmeus was uplisted to critically endangered in birdlife ' s 2008 update to the iucn red list . photo : choi soon kyoo
one in eight of the world’s bird species is globally threatened , and the fortunes of some 200 critically endangered species are now so perilous that they are at risk of imminent extinction . birdlife aims to improve the conservation status of all the world ' s birds , and hence species are often the starting point for our scientific research .
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since the seventies , millions of north american birds have disappeared and a third of species are now of conservation concern , a new report reveals .
scientists and ngos are calling for a ban on veterinary diclofenac after finding it could kill as many as 6 , 000 vultures per year in spain , home to 95 % of the griffon vulture population .
deforestation since the turn of the century has driven at least 500 species of mammals , birds and amphibians closer to extinction , according to a new scientific study published in conservation biology journal .
) , version 1 . 0 . in neotropical birds online ( t . s . schulenberg , editor ) . cornell lab of ornithology , ithaca , ny , usa .
birdlife international 2003 birdlife’s online world bird database : the site for bird conservation version 2 . 0 . cambridge , uk : birdlife international : urltoken
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endemic a species or taxonomic group that is only found in one particular country or geographic area . montane forests forest occurring in the montane zone , a zone of cool upland slopes below the tree line dominated by large evergreen trees .
birdlife international 2003 birdlife’s online world bird database : the site for bird conservation version 2 . 0 . cambridge , uk : birdlife international . ( march , 2004 ) urltoken
erritzoe , j . ( 1993 ) the birds of cites and how to identify them . the lutterworth press , cambridge .
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recommended citation birdlife international ( 2018 ) species factsheet : coeligena prunellei . downloaded from urltoken on 09 / 07 / 2018 . recommended citation for factsheets for more than one species : birdlife international ( 2018 ) iucn red list for birds . downloaded from urltoken on 09 / 07 / 2018 .
hbw alive contains information on descriptive notes , voice , habitat , food and feeding , breeding , movements , status and conservation plus a list of bibliographical references for this species account .
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luis eduardo urueña , mauricio rueda , phillip edwards , dusan m . brinkhuizen , juanjaimemg , joe tobias , oswaldocortes , greg griffith , lars petersson , manakin nature tours , megan , dubi shapiro , diego calderon - colombia birding .
genetic data indicate that present species and c . wilsoni are sisters , together forming a sister - group to c . coeligena # r . taxonomic status of proposed form c . assimilis uncertain : could represent a valid race of present species or a variant ; further study required . described form c . purpurea ( popayán , in sc colombia ) may be hybrid of present species and c . coeligena . monotypic .
nc colombia on w slope of e andes ( se santander , w boyacá , w cundinamarca ) and on both slopes of serranía de los yariguíes . specimen supposedly from c andes of quindío was incorrectly labelled # r .
rather silent . calls include a strident single “tsit” or doubled “tsi - tsit” , often in longer series . . .
probably may–oct , based on gonadal condition . no further data available , nest undescribed .
vulnerable . cites ii . restricted - range species : present in colombian east andes eba . rare to locally fairly common . recent records near virolín indicate that it is . . .
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recent molecular study found that most of the genera in the following sequence , from haplophaedia to heliodoxa , formed a monophyletic group # r . present family - group name selected over docimastini by first revisers # r .
recent molecular studies found this genus to represent a monophyletic group , with heliodoxa as sister # r .
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the only time i actually heard the species , during foraging . species common here .
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this is a specific subject page , dealing exclusively with , or primarily with , the subject in the title . because of need , there are many such pages at rhww : usually , but not always , linked to primary pages . for those in a hurry , they enable a quick summary of many important subjects . the menu for these pages is here :
1 ) the incas and the peruvian rulers before them , were not mongol type people .
the moche ruled in peru from about 250 b . c . to 750 a . d .
the amerindians were later
double - crossed
by the spanish . they then tried to sue the spanish in spanish courts for redress . see :
litigation over the rights of “natural lords” in early colonial courts in the andes
below .
corruption , the reforms of francisco de toledo and the backlash of indio social changes in sixteenth & seventeenth spanish peru . by jeffrey benson , western oregon university ,
in 1569 francisco de toledo was dispatched from spain by king phillip ii to assume the position of viceroy of spanish peru . toledo was expected to install political reforms that would further subordinate the natives especially in the andes , provide adequate workers for the mines , and increase the overall revenue for the royal treasury . although toledo legislated several reforms his three most influential were :
3 ) establishing a regimen of forced labor to support the silver mines of peru and alto peru { bolivia } . the reforms were readily accepted by the local officials , but not because they wished to improve the status of the spanish crown ; rather it gave them a chance to further corrupt and reap the benefits . prior to the reforms of francisco de toledo , political and economic corruption was already in motion . after toledo ' s reforms were instituted , royal officials , clergy , entrepreneurs and even kurakas ( local native chieftains ) did what they could to take advantage of the system and profit . toledo ' s reforms permitted a larger share of spaniards to exploit the indios ( indigenous peoples ) productivity , increase their revenues and appease the crown . to some degree , both indirectly and directly , the reforms encouraged the indios to assimilate , adapt , change or hide in order to escape the obligations of the indio caste system .
when the spanish conquistadors arrived in modern - day peru , the yanaconas declared themselves “friends of the spaniards” , as most peasant societies are very sensitive to changes in power balances . they then assisted the spaniards to take control of the empire . after conquest , the yanacona population exploded with people leaving ayllus in correspondence with mining . spaniards favored the individual yanaconas ( as they were an alternative labor force ) instead of the ayllu - based encomienda system , so the population continued to increase .
potosi - is a city in bolivia ; it is one of the highest cities in the world by elevation at a nominal 13 , 420 ft . and it was the location of the spanish colonial mint . potosí lies at the foot of the cerro de potosí , sometimes referred to as the cerro rico (
rich mountain
) . a mountain popularly conceived of as being
made of
silver ore , which has always dominated the city . the cerro rico is the reason for potosí ' s historical importance , since it was the major supply of silver for spain .
two things to consider about the local officials were their capability to perform and their integrity . a . m . fuentes writes about the inefficiency of tribute collection of the yanaconas by the corregidores ( spanish mayors ) at the potosi .
he writes ,
in the secular government i referred to your majesty about the sustenance and origin of these people , the very moderate tax they pay in some regions , how the corregidores and royal officals collect it , the value of yanacona tribute at potosi , and the silver which is consigned to the foot soldiers of government . the rest is of the royal treasury but of little importance owing to the poor collection , which you should rectify .
toledo ' s reforms for collecting tribute clarified that tributarios ( indios who held and cultivated native property / land ) from the ages of 18 to 50 were to be subjected to the tribute system .
in rural society , toledo ' s reorganization created imposing networks of authority , formal and informal , in which the corregidores stood at the center , armed with the police powers of the colonial state with that authority and protection corregidores began to take advantage of the system . the police powers were real enough , for corregidores and other officials jailed and whipped people , and impounded their belonging , under the guise of enforcing laws and punishing criminals . the corregidores also withheld tribute monies belonging to the crown in order to finance their own local business ventures .
in addition to intimidating royal opposition and embezzling tribute , the corregidores also took advantage of the indios ' service . spanish officials also required andeans to serve as mitayos ( conscripted laborers ) in textile mills , on coca farms , and on public works projects , despite imperial and local laws forbidding the practices . when kurakas attempted to protest or take the local magistrates to court , the corregidores and their allies among the parish priest usually conspired to intimidate , abuse , jail , or even replace the ethnic leader with a more pliable candidate .
the lure of riches , though , did not solely attract spaniards , but also kurakas . there is not sufficient evidence that all kurakas took advantage of the toledan system , however ; andrien offers one specific example that permits historians to identify the vast amount of wealth that a kuraka could acquire . he writes ,
the fabulously wealthy and powerful diego caqui , kuraka of tacna . . . when diego caqui died in 1588 , his will specified that the kuraka owned an estate worth 260 , 000 pesos , including a coastal vineyard with forty thousand plants and three ships engaged in coastal trading . the example offered by andrien demonstrates the potential for kurakas to take advantage of toledo ' s reforms . andrien does go to clarify that diego caqui used his wealth to promote festivals and distribute gifts among the indios , however ; that cannot be said of all the kurakas that took advantage of toledo ' s reforms and of their kinsmen .
john v . murra notes a 1559 meeting , in which settlers were asked to discover the history of the indios . murra records the instructions of a royal official to several members of his staff ,
you will inquire if in olden times there were corporal services and in what form so that if these had prevailed , one would understand in all fairness what they could and should pay .
in theory , mitayos were to work for one year then be paid and return home not having to serve again for about another seven years . zulawski notes ,
each worker was to remain for a year in potosi and be paid for his labor . after his turn in the villa imperial , he could return to his village and theoretically was not to serve again for about seven years . unfortunately , the decline of originarios and mass migration away from the mita regions caused the seven years away from the mita to become three years . tandeter comments ,
viceroy toledo had determined that each village would send to potosi each year a fixed number of indian men between the ages of eighteen and fifty , selected from a list of villages located in sixteen provinces centered in the altiplano ( also known as andean plateau ) , but reaching to the east and north , to the dividing line between collao and cusco .
however , the bulk of the pressure fell upon the kurakas and the poor mitayos . the kurakas were in charge of supplying the local officials with a yearly quota of mitayos and if they were unable to accomplish this task then they and their family would lose their social status , be imprisoned and in some cases sent fo the mines . wightman further captures the intense amount of pressure bestowed upon a kuraka ,
as a parish priest explained in 1689 , ' there is no indian who wants to be kuraka , because of the problems to be faced in the fulfillment of the different obligations ; and the corregidores and their assistants force the richest indian to take this office , to serve as kuraka of these ayllus , even though he may not be an originario of the town .
wightman states ,
the toledo reforms , however , were particularly vulnerable to manipulation because the per - capita basis for tribute and mita assessments led indian leaders to underreport their base population . however falsifying census records was extremely risky for kurakas because if caught they ' d be demoted , forced to pay a tribute and handed over to public works . in addition , for indios that were not properly recorded in the census , their houses would be demolished , they would be taken out by force , fined and handed over to public works .
for the few indios that did survive their turn at the mita , when they return to their homeland , often they found that their land had been occupied by another party . wightman explains ,
indians who did return to their home communities often found that their lands had been seized by spaniards , taken by neighbors , or occupied by migrants from other communities . this unfortunate turn of events is not surprising in that the percentage of survivors was fewer than 15 % . it is likely that the spaniards or neighboring indios did not expect . the return of the majority mitayos .
toledo ' s reforms clarified who was exempt and who wasn ' t , but he was unable to supervise local officials , so he was helpless to fully supervise his reforms . most indians didn ' t want to participate in the mita . so great was their distaste for the mita that they desired by whatever means to escape the mita service . cole comments ,
rather , the indians responded to the worsening situation in the mines by using every available means to evade the mita . however , were a huge mass of the work force to change their civil status , then the local officials would suffer . to avoid such cases the local officials simply ignored toledo ' s specifications .
in addition to the mita , indios were required to pay a tribute to the royal crown and local officials . each civil status group within the indios caste system had to pay tribute with the exception of the kurakas . andrien states ,
kurakas were exempt , but members of the community clan structure ( tributarios ) paid the largest sums . those outside the ayllu or kin structure ( yanaconas ) and recent migrants ( forasteros ) paid lesser amounts . after the collection of the tributes the kurakas would transfer the goods over to the corregidores who would then deposit the sum to various offices .
toledo ' s statement clearly shows that one of the spanish intentions for replacing the indios in reducciones was to christianize them and teach them of the catholic faith . whether their intentions were genuine or not is debatable , but what is noticeable is the official approval to christianize the indios and change their religious upbringings . one method of assimilating the indios into the catholic faith was rewarding them with exemption from the mita .
the priest could bestow exemptions from the mita upon his favored lay assistants , and heap abuses upon the troublesome by accusing them of idolatry . thus , the more assimilated an indio became , culturally , the less he / she was punished , taxed , or forced to work , technically .
most of the cultural changes , though , geographical , economic , horticultural , and religious were examples of forced change . indios who changed culturally were not guarantee better treatment , tax relief or mita exemption . as demonstrated earlier many forasteros and yanaconas were still being subjected to mita service even though they had changed their social economic status . that is why , as some historians have argued , the indios further changed socially in order to ascend beyond the indio caste system by establishing themselves as mestizo status .
in order to do this the indios aimed to change every social aspect from indio to spanish . jackson argues ,
individuals consciously changed their behavior to be able to move to another and usually higher racial status within the caste system . indios , for example , could escape tribute obligations and service in labor drafts such as the andean mita by passing as mestizos . methods of change included clothes , language , surname , occupation , activities , architecture , religion , baptism , catholic marriage , location of home and accumulation of wealth . jackson continues ,
indios could change their mode of dress , learn to speak spanish , move to a city or away from their place of birth , take up a profession generally not associated with the indigenous population , and be reclassified as mestizos exempt from the unique legal obligations of the indigenous population .
however , once the outward appearance has changed in an attempt to climb the social ladder , there still remained the legal documentation of identification .
one method of escaping the indio caste by legal documentation was to simple state that you had some spanish background and were , thus , mestizos . this was such the case for antonio and agustin carrillo who won exemption from the mita based on their claim to spanish heritage in 1603 . many local officials advised against the judicial conclusion fearing an onslaught of social claims . stern records ,
the audiencia of la plata ruled in favor of the carrillos despite the objections of its fiscal , who counseled the tribunal that a ruling in favor of the brothers would open a pandora ' s box of problems for the mita , for the judicial system would soon be clogged with petitions from indians claiming some degree of spanish ancestry . jackson concurs adding this excerpt ,
the indians change their name , and declare themselves mestizos and yanaconas , they dress in the spanish way and work as artisans or in the convents with the intention of not complying with their obligations .
claims of spanish ancestry were usually accompanied with wills , marriage records , baptismal records or testimony from a known spaniard . jackson discusses baptisms by stating ,
this was particularly the case with parish priests who recorded the racial status of newborn children . if the parents of a newborn child were members of the catholic church they could try to claim spanish ancestry at the baptism or provide a generous bribe to the local priest so that their son or daughter could be classified as a mestizo . in some cases the priest would even baptize infant boys as girls so that they could escape the mita service . another method that didn ' t involve using the church was the testimony of a spaniard , usually a hacienda owner . a hacienda owner who employed indios had to do one of two things when it came time to pay tribute ; 1 ) pay the tribute for the indios or 2 ) permit the indios to leave for a certain duration of time so that they could earn enough money to pay the tribute . if , however , a indio suddenly changed identity to a mestizo then the hacienda owner would no longer have to burden himself with his / her tribute situation . jackson notes ,
hacienda owners , for example , conspired to have their workers removed from the tribute rolls . in this way , the hacendados would not have to contend with paying their workers ' tribute , or allow the workers time off to work elsewhere to earn money to cover the tribute payments .
in a 1582 manuscript from la biblioteca nacional de lima titled
limpieza de sangre ,
contains the process by which a witness might have had to justify the social identity of another . in this particular manuscript dona juana fernandez de ugarte is the person of interest , who is trying to solidify her social class standing and the witness is martin hurtado de aviento .
there are several questions the court asks martin and in the manuscript he provides adequate answers to assure the social identity of dona juana . provided are several of those questions along with martin ' s replies :
question 1 : first to be asked if they know the parents and the francisco de yrarrazabal , so the father and mother and grandfather and parents of dona lorenza de zarate said his wife , so the father and mother contained in this memorial and if they know who is the legitimate child of such parents .
response : the first question he said , who knows that don francisco de yrarrazabal , twenty - five years now , what was known in this city , and has news of dona lorenza de zarate said his wife , because although he has not seen public knows or noticeable thing is his wife , and there were , this witness had known and understood , as a country and try and recruit very familiar with many relatives of the don francisco .
question 5 : and who knows if the said dona lorenza zarate woman said don francisco de yrarrazabal is legitimate daughter of these parents and that she and they and their grandparents and those from father and mother every of them have been and are christians and clean blood and clean without spot or race or indian descent , moors and converts or another sect of newly converted and that these have been incurred and taken and if it would otherwise have been known rumor or what they know or have heard about .
response : to the fifth question he said , who knows what is contained in this question because it has been treated and chat for a striking thing to be clean people , old christians , gentlemen . sons , content on this question , heard or understood without knowing anything to the contrary and also knows that to be the legitimate daughter of dona lorenza said parents , who saw her as such in seville in his mother ' s house .
response : in the sixth question he said , that never understood or heard anything contained in this question , if any , this witness know what you think and could not be less . the play out of the interrogation creiltes an environment of uncertainty . the questions tend to represent a common procedure , however ; the responses are not very clear and direct , but are somewhat witty and taunting in discourse . if the court is to make a decision against dona juana they must explain their reasoning with evidence , otherwise , the social status of many could be at jeopardy . to approve of the social status of dona juana , the court has to clarify that the evidence provided is not falsified and the honest testimony of the witness was absolute . the atmosphere of uncertainty was not to be impeded with one trial . if indios had more success than failure at changing their identity through the spanish courts then there was no reason to not attempt social change and escape the spanish colonial corruption .
toledo ' s reforms permitted a larger share of spaniards to exploit the indios productivity , increase their revenues , appease the crown to some degree and indirectly encourage the indios to assimilate , adapt , change and hide in order to escape the obligations of the indio caste system . the indios were exhausted physically and economically and they used all sorts of methods to change socially ; language , clothes , lifestyle , occupations , food , migration of forastero status , etc . they took great risk to challenge the spanish court system and justify their spanish ancestry using wills , genealogy records , baptismal records , marriage records , and testimony from known spaniards . they risked much in order to liberate themselves from the corrupted reforms of toledo , the hardships of the indio caste system and from the abusive relationships with the local and royal officials . it was these atrocities that grew with fervor with the toledo reforms that caused the indios to seek ways of socially changing their identity and become mestizos .
in the earliest days of the european invasion , when inka resistance , potentially so threatening , turned out to be virtually absent ( lockhart 1972 ) , the pizarros acquired a steadfast ally , the wanka / cañaris lords . it was in their territory , xauxa , that the europeans established their first capital . along with thousands of soldiers and bearers , the canaris provided the newcomers with strategic information , plus the food and weapons stored in hundreds of warehouses built by the inka and filled locally ( polo de ondegardo 1940 [ 1561 ] ) . in one region where the inka had managed to cobble together some resistance , as at huánuco , the europeans had to call on canaris troops to help them put down “the rebellion . ” all this assistance provided the europeans was recorded with care on a khipu kept by the canaris lords . this record was first described by cieza de león , some fifteen years after the invasion . such bookkeeping later became the subject of litigation initiated at the viceregal court , at lima , by one of the lords who in 1532 had opened the country to the troops of charles v ( murra 1975 ) . this man , don francisco cusichac , felt betrayed by the ill treatment of his people and the neglect of his own privileges .
the only other group to be treated so harshly by toledo were the descendants of another wing of the andean elite who also sided from the earliest days with the invaders . these were the “sons” and heirs of pawllu thupa , the one inka “prince” to make peace , early and openly , with the europeans . pawllu had helped them through extreme difficulties , particularly almagro’s invasion of chile . the efficiency of that thrust south was attributed by many to pawllu thupa’s ability to mobilize the lords of charcas , the region known today as bolivia . for his services , pawllu had been allowed to keep “his indians , ” coca - leaf terraces , food - producing fields , and much other inka wealth . a test came in 1550 at pawllu’s death : various europeans attempted to deprive the “indian’s” heirs of these lands and people , but the emperor’s representative , bishop lagasca , resisted such claims . for the next two decades , pawllu’s many sons were a distinguished and rich lineage in cuzco . they spoke spanish , invested in the long - distance coca - leaf trade to the mines at potosí , and employed europeans in their various enterprises . the main heir , don carlos , was married to a european woman . thirty - five years after the invasion , pawllu thupa’s heirs were the one group of inkas at cuzco who had managed to hang on to both status and wealth ( glave 1991 ) .
when toledo reached cuzco on his way to the mines at potosí , he selected pawllu’s lineage for special attention . as at xauxa , the lords were ordered to these grants are transcribed from the originals in the archivo general de indias , seville : section lima , legajo 567 , lib . 8 , fols . 107v–108r ; see also other grants cited by espinoza soriano ( 1972 ) . letters from francisco de toledo to philip ii , found in the biblioteca nacional , madrid . see pawllu thupa’s testament published in revista del archivo histórico del cuzco ( 1950 : 275 , 286 ) .
litigation over the rights of “natural lords” display the credentials testifying to their services to the spanish crown . the papers were publicly burned . don carlos and his kin were accused of maintaining illicit contacts with those inka who had taken refuge at vilcabamba , in the eastern lowlands ( kubler 1946 ) . some twenty of pawllu thupa’s heirs were put on trial for subversion ; during the proceedings , which lasted many months , the princes were kept in animal corrals , exposed to the elements . the testimony was conducted in quechua even though many of the accused spoke spanish ; a mestizo , one gonzalo gómez ximénez , “interpreted” for the only record kept of the proceedings , despite continuous protests by the accused . ximénez’s version of what they had “confessed” became the official transcript . the “natural lords” were sentenced by gabriel de loarte to the loss of “their” indians and of their coca - leaf fields , which were granted by toledo to loarte . some twenty inka , including aged princes , don carlos , and several children , were deported on foot to lima . from there they were supposed to be shipped into exile to mexico .
of the twenty , seven survived . they were able to rally support from some of the judges at the audiencia who were hostile to the viceroy . toledo remained in the highlands for almost another decade , the only viceroy to devote such personal attention to the andean population . he sponsored many institutional innovations ; some of them were consistent with ideas to end the las casas “benevolent” approach to indian affairs , which he brought with him from court . he tried to put an end to the influence of bishops gerónimo de loaysa of lima and domingo de santo tomás in charcas , men from another era , who spoke quechua and had earlier corresponded with las casas ( las casas 1892 ) .
of the people toledo consulted , the best informed were two salamanca trained lawyers—juan de matienzo and juan polo de ondegardo—who gave him diametrically opposed advice . matienzo , a crown justice at the audiencia of charcas , was frequently active away from his court . even before toledo’s arrival in 1569 , matienzo had argued for the “extirpation” of the inka lineage that had taken refuge in the forest at vilcabamba . the high court in lima was betting on a reduction policy , resulting in the conversion of the refugee princes and their resettlement at cuzco . matienzo thought such a policy was dangerous . resettlement expanded the number of “natural lords”at cuzco—a loss of revenues for the spanish crown and the threat implicit in an additional focus of traditional loyalty ( matienzo 1967 ) . after toledo’s arrival , he and matienzo formed an intimate alliance broken only by the judge’s death in 1579 .