rhamphochromis macrophthalmus is a species of piscivorous cichlid endemic to lake malawi where it prefers open waters at depths of from 30 to 109 metres ( 98 to 358 ft ) .

this species can reach a length of 28.9 centimetres ( 11.4 in ) sl .

it can also be found in the aquarium trade .

rhamphochromis macrophthalmus

there is currently no quality picture available to illustrate rhamphochromis macrophthalmus regan , 1922 , should you have one you would like to contribute , please contact the cichlid room companion editor .
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maréchal , c . , 1991 . rhamphochromis . p . 422 - 424 . in j . daget , j . - p . gosse , g . g . teugels and d . f . e . thys van den audenaerde ( eds . ) check - list of the freshwater fishes of africa ( cloffa ) . isnb , brussels ; mrac , tervuren ; and orstom , paris . vol . 4 . ( ref . 5689 )
this species is most common on shelf zone at 50–100 m , but can also be found between 30–110 m . it can also be found off submerged reefs . juveniles are found in the same habitats as adults . it has a fecundity of 154–202 eggs . the length of maturity is 19 . 9 cm for males and 18 . 6 cm for females . it is commonly caught in bottom and semi pelagic trawl catches in the south as well as by hand line and gillnet catches in the north . sometimes exported by the aquarium trade and is known as
rhamphochromis
. max . size : 29 . 8 cm sl .
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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 .
snoeks , j . ( freshwater fish red list authority ) & darwall , w . ( freshwater biodiversity assessment programme )
justification : endemic to lake malawi where it is widespread with no major widespread threats identified .
a potential threat is over - fishing by the commercial trawlers in the southern part of the lake .
to make use of this information , please check the < terms of use > .
gland , switzerland , 5 july 2018 ( iucn ) – australia’s unique reptiles – including lizards and snakes – face severe threats from invasive species and climate change , with 7 % of th . . .
the value of medicinal and aromatic plant trade has increased three - fold in the past 20 years , but traditional harvesting practices are being replaced by less sustainable alternatives . . . .
a recently released iucn technical brief recommends increasing investments in sustainable land management practices , as well as better cooperation between agriculturalists and conservationists to conse . . .
greek , rhamphos = bill , peak + greek , chromis = a fish , perhaps a perch ( ref . 45335 )
freshwater ; demersal ; ph range : 7 . 5 - 8 . 2 ; depth range 30 - 109 m ( ref . 55949 ) , usually 50 - 100 m ( ref . 55949 ) . tropical ; 20°c - 25°c ( ref . 13614 ) ; 9°s - 15°s
maturity : l m ? , range 19 - ? cm max length : 29 . 8 cm sl male / unsexed ; ( ref . 55949 )
diagnosis : large , big - toothed brownish species ; eyes rather large ; premaxillary pedicel relatively short ( ref . 55949 ) .
common on the shelf zone ; occasionally found off reefs ; but not commonly recorded from the middle of the lake ( ref . 55949 ) . moves in the open water but stays 2 - 5 m above the substrate ; feeds on fish like the small utaka and usipa ( engraulicypris sardella ) ( ref . 5595 ) .
species probably forms breeding leks on or near the bottom ; juveniles are common in the adult habitat ( ref . 55949 ) .
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 5156 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 01000 ( 0 . 00244 - 0 . 04107 ) , b = 3 . 04 ( 2 . 81 - 3 . 27 ) , in cm total length , based on all lwr estimates for this body shape ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 4 . 2 ±0 . 73 se ; based on food items .
resilience ( ref . 69278 ) : medium , minimum population doubling time 1 . 4 - 4 . 4 years ( preliminary k or fecundity . ) .
vulnerability ( ref . 59153 ) : moderate vulnerability ( 36 of 100 ) .
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distinguished by its large eyes ( hence the name ) and its 1 - 2 horizontal line ( s ) . this fish is an important food source for the people living around the lake and is quite abundant in very deep water . their slender bodies are designed for the speedy pursuit of prey .
pronunciation : refer to our pronunciation key for an explanation of the phonetic symbols .
habitat : this is the primary location where the cichlid is found and is a generalization . this does not mean a fish cannot be found in other habitats .
diet : many cichlids specialize in eating one type of food ; notwithstanding , some of these specialized feeders are flexible and can be opportunistic feeders .
temperament : this describes the overall demeanor of a cichlid toward other tankmates that are of a different species . consider that there is variability in temperament due to various factors , including aquarium size , tankmates of similar appearance , stocking levels , and order of introduction . there may even be some variability among individual specimens .
conspecific temperament : this describes the overall demeanor of a cichlid toward other tank - mates of the same species . consider that there is variability in temperament due to such factors as aquarium size , stocking levels and order of introduction . there may even be some variability among individual specimens .
maximum size : this is in regards to total length ( including the tail ) of typical aquarium specimens . wild specimens may not attain this size , or may in fact grow larger than aquarium raised individuals due to various factors . also consider that this is the typical maximum size and there are exceptional individuals that will exceed it .
difficulty : this measure is a relative value , comparing a single species against all other cichlids . this only accounts for maintanence in the aquarium and not breeding considerations . 1 = easy and forgiving , 5 = extremely challenging .
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species profiles presents in depth studies , articles , and information surrounding the numerous species of lake malawi cichlids .
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disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
one of the three syntypes ( bmnh 1921 . 9 . 6 : 217 - 219 ) . drawing from regan ( 1922 : plate vi , fig . 1 ) .
last update : 18 november 1999 web author : m . k . oliver , ph . d . copyright © 1997 - 2018 by m . k . oliver - all rights reserved
pam chin has been replying to cichlid questions for over twenty years . highly respected and experienced aquarist , pam has visited cichlid habitats around the world , and bred in her ' s and her husband gary fish house hundreds of cichlid species . besides her job , she still devotes time to help any person with a cichlid question !
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genner , martin & p . nichols , g . r . carvalho , r . l . robinson , p . w . shaw , a . smith & g . f . turner . 2007 .
evolution of a cichlid fish in a lake malawi satellite lake
. proceedings of the royal society of london b . v . 274 ( n . 1623 ) , pp 2249–2257 ( crc01924 ) ( abstract )
regan , charles tate . 1922 .
the cichlid fishes of lake nyassa
. proceedings of the zoological society of london . 1921 ( pt 4 ) n . 36 ; pp . 675 - 727 ( crc00066 )
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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .


perissodus microlepis is a species of cichlid endemic to lake tanganyika .

this species reaches a length of 11 centimetres ( 4.3 in ) tl .

this species can also be found in the aquarium trade .

it is a scale-eating ' parasite ' on other fish species .

it occurs in two distinct morphological forms .

one morph has mouth parts twisted to the left , enabling it to eat scales off its victim ’s right flank .

in contrast , the other morph , whose mouth is twisted to the right , eats scales off its victim ’s left flank .

the relative abundance of the two morphs in populations is regulated by frequency-dependent selection .

perissodus microlepis

un jeune perissodus microlepis attaque un jeune lepidiolamprologus elongatus , pour lui prélever quelques écailles .
one parent was experimentally removed from brooding pairs of perissodus microlepis in the field . the removal elicited drastic behavioural changes in the remaining parent and young . the remaining parent sometimes showed a sequence of peculiar behaviours which were not observed when guarding the young with its mate . some left the brooding site with young in their mouths and then put the young under the care of another pair of brooding parents .
fifty - four breeding pairs of perissodus microlepis were collected in april 2010 by diving with hand nets at toby veal ' s lodge ( s08°37 . 4′ e031°12′ ) near mpulungu ( zambia ) on the southern tip of lake tanganyika to assess the mating pattern ( kusche , lee , meyer , in revision ) . twenty - one out of these 54 pairs were used for foraging experiments on adult p . microlepis under semi - natural conditions . mouth laterality of each of the pairs was judged by eye in the field by two independent researchers ( h . k . and a . m . ) ( see figure 1 ) .
unknown , but large schools of thousands of juvenile p . microlepis are often seen at sites not far from brooding pairs . this species is very common throughout its habitat , but is never abundant .
thirteen outdoor pools ( 1000 l ) at the shore of lake tanganyika were stocked with one breeding pair of p . microlepis each ( 7 rl , 5 rr and 1 ll pairs ) . also two large community tanks of 4000 l volume ( with 6 l - morphs and 10 r - morphs of p . microlepis , respectively ) were used in these foraging experiments . the cichlid species tropheus moorii ( pair tanks : n = 3–6 ; community tanks : n = 18 and 25 ) was used as prey since it is a preferred natural prey species of p . microlepis [ 22 ] . after 72 hours , all t . moorii were removed from the pools and presence / absence of scars and missing scales and the numbers of bites on both flanks of the prey fish were recorded .
the handedness of the foraging behavior and the associated asymmetry in mouth / head morphology have made the scale - eating cichlid fish , perissodus microlepis , a textbook example [ 16 ] of both , the astonishing degree of ecological specialization and negative frequency - dependent selection [ 8 ] . however , how and when lateralized foraging behavior manifests itself during ontogeny , and whether its association with mouth asymmetry is already apparent in juvenile individual fish had remained untested . here , we report on the strength and individual variation of lateralized foraging behavior as well as its relationship with mouth asymmetry in p . microlepis during its juvenile as well as adult life stages . we find that handed foraging behavior is already prominent at an early age ( e . g . at two - months ) , although the initial morphological asymmetry is less evident , which hints that handed behavior might play a role in bringing about pronounced morphological laterality , considering the potential influences of phenotypic plasticity on mouth asymmetry [ 18 ] .
in this study , we examine the strength and individual variation of lateralized behavior and its interaction with mouth laterality in perissodus microlepis . in semi - natural conditions , we conducted feeding experiments on adult wild - caught scale - eaters with their natural prey to test whether pronounced morphological laterality predicts foraging preferences . we further tested whether laboratory - reared juvenile scale - eaters , which had never encountered prey fish before , displayed lateralized scale - feeding behavior in reference to mouth asymmetry . here we demonstrate relatively strong handedness in foraging behavior in juvenile fish that showed much less mouth asymmetry ( compared to wild - caught adult fish ) and we then discuss the potential role of lateralized foraging behavior in shaping the head asymmetry of this species .
maréchal , c . and m . poll , 1991 . perissodus . p . 367 - 368 . in j . daget , j . - p . gosse , g . g . teugels and d . f . e . thys van den audenaerde ( eds . ) check - list of the freshwater fishes of africa ( cloffa ) . isnb , brussels ; mrac , tervuren ; and orstom , paris . vol . 4 . ( ref . 5678 )
scale - eating cichlid fish , perissodus microlepis , from lake tanganyika display handed ( lateralized ) foraging behavior , where an asymmetric ‘left’ mouth morph preferentially feeds on the scales of the right side of its victim fish and a ‘right’ morph bites the scales of the left side . this species has therefore become a textbook example of the astonishing degree of ecological specialization and negative frequency - dependent selection . we investigated the strength of handedness of foraging behavior as well as its interaction with morphological mouth laterality in p . microlepis . in wild - caught adult fish we found that mouth laterality is , as expected , a strong predictor of their preferred attack orientation . also laboratory - reared juvenile fish exhibited a strong laterality in behavioral preference to feed on scales , even at an early age , although the initial level of mouth asymmetry appeared to be small . this suggests that pronounced mouth asymmetry is not a prerequisite for handed foraging behavior in juvenile scale - eating cichlid fish and might suggest that behavioral preference to attack a particular side of the prey plays a role in facilitating morphological asymmetry of this species .
the frequency of l and r morphs in natural populations of p . microlepis is suggested to be maintained by negative frequency - dependent selection [ 8 ] . over time , the proportion of l and r morphs within populations oscillates around a 50∶50 ratio [ 8 ] , [ 17 ] . the resulting lateralized foraging behavior of p . microlepis is expected to make prey fish more alert to being attacked from the preferred side of the more abundant morph . thus , increased prey vigilance would reduce the predation success of the more abundant morph , and negative frequency - dependent selection would thereby favor the rarer morph in each generation . consequently , the frequency of both morphs is maintained in approximately equal abundances [ 8 ] .
sixty - one more juveniles from three different broods ( 3 young of rl , 6 of ll and 52 of rr parental - pairs ) were obtained by breeding wild - caught fish in the laboratory . lab - reared p . microlepis fish reached sexual maturity at about six to nine months of age . these 61 juveniles , the f 1 fish of wild - caught stock , were used for foraging experiments that examined behavioral handedness and its interaction with mouth asymmetry .
nonetheless , the observed lack of correlation between mouth asymmetry and lateralized foraging behavior in juvenile p . microlepis might also result from the measurement technique used for the quantification of mouth asymmetry ( i . e . mouth bending angle ) not fully capturing the existing true laterality ( i . e . asymmetric skeletal features ) in the mouth / head apparatus of this fish . further tests with cleared and double - stained juvenile fish samples of known behavioral laterality are required to check this possibility .
to statistically analyze if juvenile p . microlepis fish show a bimodal or unimodal distribution in their foraging behavior ( e . g . foraging score , behavioral foraging preference ) , the dip statistic [ 23 ] and a mixture analysis with a parametric bootstrap test ( 1000 iterations ) using the mixtools package [ 24 ] were performed in r [ 25 ] . an anscombe - glynn test [ 26 ] for platykurtosis was further performed for the seven - month old fish only ( see below ) .
mouth asymmetry ( mouth bending angle ) is not significantly correlated with foraging handedness in juvenile p . microlepis . ( a ) two month ( r = 0 . 148 ; p = 0 . 255 ) ; ( b ) three month ( r = −0 . 229 ; p = 0 . 154 ) ; ( c ) seven month old fish ( r = 0 . 069 ; p = 0 . 749 ) . note that 11 fish were tested at two different ontogenetic stages ( i . e . at three and seven months of age ) .
several important questions about this fish , including the bases of its behavioral and morphological laterality remain unanswered [ 20 ] . mouth laterality of p . microlepis has been suggested to be genetically determined by a single mendelian locus with two alleles : the ‘r’ - allele was suggested to be dominant over the ‘l’ - allele and ‘r’ was suggested to be homozygous lethal [ 8 ] , [ 14 ] , [ 21 ] . but a recent review [ 20 ] noted that the data reported so far ( of mouth - morph ratios in the offspring of parents of known laterality ) actually are inconsistent with a single locus mendelian model . this model was further questioned because the distribution of mouth asymmetry was found to be unimodal rather than bimodal [ 18 ] , [ 21 ] ( kusche , lee , meyer , in revision ) . phenotypic plasticity may therefore play a role in shaping mouth asymmetry [ 18 ] and hence , head asymmetry may be governed by both genetic and environmental factors [ 18 ] ( lee et al . , unpublished data ) . the genetic and / or environmental basis of behavioral handedness in p . microlepis , however , remains largely unexplored .
this species belongs to the deep benthic community . it feed upon the scales of a variety of benthic species , chiefly cichlids . they are probably solitary except during periodsof reproduction . young p . microlepis have been only been seen close to the shore . its diet shifts from a variety of micro - organisms and algae when young to an adult diet of mainly fish scales . little is known about its reproduction although it is known to reproduce inshore among rocks and is remarkable for showing features both of mouth - brooding and substrate - guarding , the two major patterns of parental shown in cichlids
the foraging experiments with juvenile fish in the laboratory show that mouth asymmetry does not predict handedness in foraging behavior , possibly due to the small degree of mouth asymmetry . surprisingly , young and still quite small scale - eaters preyed on scales of prey fish and exhibited pronounced handed behavior ( figure 3 ) . even the two - month old fish readily fed on scales of similar - or even slightly larger - sized goldfish . however , the degree of mouth asymmetry in juvenile scale - eaters was rather small [ on average only 2 . 67° ( for two - month : sd = 1 . 92° ) , 2 . 01° ( three - month : sd = 1 . 73° ) and 2 . 12° ( seven - month : sd = 1 . 61° ) ] and the relationship between behavioral bias and morphological asymmetry was always non - significant ( figure 5 ) . note that the degree of mouth asymmetry in those laboratory - reared scale - eaters is indeed substantially lower than in wild - caught adult p . microlepis ( the average = 5 . 07° ; n = 238 ; sd = 3 . 51° ; kusche , lee , meyer , in revision ) . this too supports the hypothesis that handed behavior might play a significant role in shaping the asymmetry of mouths in p . microlepis .
furthermore , whether foraging handedness is expressed earlier during development and induces and thereby facilitates mouth asymmetry via phenotypic plasticity [ 15 ] , [ 18 ] or the reverse – remains unclear . hori [ 8 ] originally suggested that mouth laterality in p . microlepis is a functional ‘prerequisite’ for efficient lepidophagy . he further proposed that mouth asymmetry ( controlled by a single mendelian locus ) precedes and invokes and directs lateralized foraging behavior through natural selection . but , several lepidophagous cichlid species in lake tanganyika lack a pronounced laterality in their heads [ 21 ] and behavioral preferences have not been tested in these species . moreover , handed foraging behavior might actually precede , and even induce mouth asymmetry , given the purported role of phenotypic plasticity in mouth laterality [ 18 ] .
the same clear pattern was found in the 13 pools with one pair of p . microlepis each as predators ( figure 2 c , d ) : seven rl pairs fed from both flanks with similar frequencies ( ratio of attacked left flanks : 40–60 % ; average : 52 % ; median : 50 % ) and produced similar amount of damage onto both flanks ( range of foraging scores on left flanks : 25–67 % ; average : 48 % ; median : 50 % ) . five rr pairs strongly preferred to feed from the left flank ( range : 67–100 % ; average : 88 % ; median : 100 % ) , which caused more bites on that flank ( range : 83–100 % ; average : 94 % ; median : 100 % ) . a single ll pair exclusively fed from the right flanks of their prey fish . differences in foraging patterns such as foraging preference and foraging score among morph pair combinations were both highly statistically significant ( wilcoxon - rank - sum - test with continuity correction : proportion of left flanks affected : w = 35 , p < 0 . 01 ; proportion of foraging scores at the left flank : w = 35 , p < 0 . 01 ) .
juvenile p . microlepis of about two [ n = 61 ; mean standard length ( sl ) = 3 . 2 cm ; sd = 0 . 28 cm ] , three [ n = 47 ; total length ( tl ) = 3–4 cm ] and seven [ n = 24 ; mean total length ( tl ) = 7 . 7 cm ; sd = 0 . 58 cm ] months of age , that had not had an opportunity to eat scales from prey fish before , were tested for lateralized foraging behavior . the older test cohorts ( that were tested at three and seven months ) were caught as 1 - 2 week old fry in the field , whereas the fish tested at two months were bred in the laboratory ( the f 1 fish of the older cohorts ; see above ) . eleven of the three - month old fish were re - tested at seven months , but those individuals could not be traced due to logistical reasons . the scale - eaters were placed individually with a single prey fish ( platy fish , xiphophorus maculatus , for three - month old fish and goldfish , carassius auratus auratus , for two - and seven - month old fish ) in the trial tanks .
neither of the juvenile cohorts showed a significant correlation between mouth asymmetry and lateralized foraging behavior ( two - month old fish : r = 0 . 148 ; p = 0 . 255 ; three - month : r = −0 . 229 ; p = 0 . 154 ; seven - month : r = 0 . 069 ; p = 0 . 749 ; figure 5 ) . unexpectedly , some fish that were morphologically scored as ( slightly ) r - morphs ( with negative values of mouth bending angle of α l − β r ) occasionally even attacked the right side more frequently than the left side , and vice versa ( figure 5 ) . this lack of correspondence suggests that mouth laterality is not a prerequisite for handed foraging behavior for juvenile p . microlepis . also , the level of mouth asymmetry ( i . e . absolute values of mouth bending angles ) of the laboratory - reared juvenile fish did not significantly increase with body size in either two - ( n = 61 , y = 0 . 831 x + 0 . 04 , r 2 = 0 . 015 , p = 0 . 35 ) or seven - ( n = 24 , y = −0 . 496 x + 5 . 394 , r 2 = 0 . 032 , p = 0 . 4 ) month old fish .
through phenotypic effects of “use and disuse” , handed behavior has been shown to drive morphological laterality in different animal groups ( e . g . lobsters [ 37 ] ; snakes [ 4 ] , [ 5 ] ; humans [ 38 ] ) ( reviewed in [ 15 ] ) . lobsters provide a clear example of how claw asymmetry is shaped during development as a function of handed behavior [ 37 ] . laboratory experiments demonstrated that differential use of claws during early juvenile stage induces and facilitates development of a crusher claw [ 37 ] . as such , in p . microlepis lateralized behavior might conceivably lead to an asymmetric remodeling of the structural elements ( e . g . bones ) involved in defining mouth shape [ 39 ] , given that lateralized behavior in fish sometimes has a strong additive genetic component [ 40 ] , [ 41 ] , e . g . , the estimated heritability of laterality of eye preference in the poeciliid fish , girardinus falcatus is 0 . 5 to 0 . 6 ) [ 40 ] . although we are uncertain whether handed scale - eating behavior is genetically programmed ( innate ) , rather than environmentally plastic ( learning ) or both [ 42 ] , the bimodal trait distribution in very young fish ( figure 3 ) speaks for a major genetic locus determining handedness in scale - eating behavior ( [ 18 ] ) .
to more precisely quantify ‘behavioral’ foraging preference in juvenile p . microlepis , a second series of experiments for the seven - and later two - month old scale - eaters was carried out . for each seven - month old individual , its foraging behavior was monitored ( in 3–4 replicates during 1–2 weeks ) by counting the number of attacks to the left and / or right flanks on a single goldfish , until a total of maximally 20 attacks per individual within up to 30 minutes were reached . the scale - eaters showed reported natural foraging behavior , i . e . , they attacked prey from behind [ 8 ] . in only a few cases they attacked from the front , but those attacks were not counted . behavioral foraging preference ( i . e . probability of left attack ) was again found to be consistent among the 3–4 trials ( repeated - measure anova ; f 3 , 45 = 0 . 363 , p = 0 . 78 ) as observed in the three - month old fish . therefore , the handedness scores ( e . g . number of left and right attacks ) were pooled over the trials to calculate behavioral foraging preference for each scale - eater . the total number of attacks observed per fish ranged from 39 to 80 ( mean = 64 ) . for the two - month old scale - eaters , we employed the same procedure as for the seven - month old fish , except that we conducted only one experimental trial per individual . the average number of attacks observed per fish in this test cohort was 19 .
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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 .
snoeks , j . ( freshwater fish red list authority ) & darwall , w . ( freshwater biodiversity assessment programme )
justification : widely distributed throughout lake tanganyika , with no known major widespread threats .
to make use of this information , please check the < terms of use > .
gland , switzerland , 5 july 2018 ( iucn ) – australia’s unique reptiles – including lizards and snakes – face severe threats from invasive species and climate change , with 7 % of th . . .
the value of medicinal and aromatic plant trade has increased three - fold in the past 20 years , but traditional harvesting practices are being replaced by less sustainable alternatives . . . .
a recently released iucn technical brief recommends increasing investments in sustainable land management practices , as well as better cooperation between agriculturalists and conservationists to conse . . .
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 .
citation : lee hj , kusche h , meyer a ( 2012 ) handed foraging behavior in scale - eating cichlid fish : its potential role in shaping morphological asymmetry . plos one 7 ( 9 ) : e44670 . urltoken
copyright : © lee 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 was provided through a travel grant from the international max planck research school ( imprs ) for organismal biology to hk , through the zukunftskolleg postdoctoral fellowship to hl , through grants of the deutsche forschungsgemeinschaft ( dfg ) to hl ( le2848 / 1 - 1 ) and am , the young scholar fund ( ysf ) to hl ( fp 411 / 12 ) and the university of konstanz to am . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
competing interests : axel meyer is a plos one academic editor . this does not alter the authors ' adherence to all the plos one policies on sharing data and materials .
handed behavior has also been frequently reported in fish , e . g . , with respect to eye usage preference ( i . e . visual lateralization ) in a poeciliid fish [ 9 ] , [ 10 ] , swimming - turns in zebra - and goldfish [ 11 ] and foraging in a freshwater goby [ 12 ] and in some african cichlid fishes [ 13 ] , [ 14 ] . lateralized behavior in fish is often correlated with morphological asymmetries . in the herbivorous cichlid telmatochromis temporalis , for example , the right mouth morph uses the right side of the jaw more frequently and the left morph the left side [ 13 ] . a significant correlation between lateralization in swimming and the anatomical bias of the prevalence of different muscle types was found in zebrafish [ 11 ] . however , relatively little effort has been directed towards the exploration of the potential role of handed behavior in facilitating morphological laterality [ 15 ] .
five broods from different parents of determined mouth - laterality ( 3 rl and 2 rr pairs ) were transported to the animal care facility at the university of konstanz . in total , 65 young were raised brood - wise in separate 40 l and later 200 l aquaria with artemia nauplii and flake food . these fish were used for laboratory feeding experiments on juveniles as well as for quantitative measures of mouth asymmetry ( kusche , lee , meyer , in revision ) .
field research was conducted under the study permit ( g . r . no : 2077761 ) granted by the government of the republic of zambia ( immigration department , ministry of home affairs , republic of zambia ) according to their immigration and deportation act . cap123 , section 16 . animal care of the fish and all foraging experiments in the laboratory were approved by the regional board of animal welfare in germany ( regierungspräsidium freiburg , abteilung landwirtschaft , ländlicher raum , veterinär - und lebensmittelwesen ) ( permit number : 35 / 9185 . 81 / g - 10 / 96 ) .
percentages of pooled attacked left ( and right ) flanks of prey fish were calculated for each tank . we considered these estimates as foraging preference for a particular side , given that the scale - eaters had in principle an equal opportunity to attack both flanks . foraging preference was further assessed by taking into account foraging scores reflecting different levels of injury ( i . e . the amount of damage done by the scale - eaters ; 0 bites = 0 scores ; 1–3 bites = 1 score ; 4–6 bites = 2 scores ; > 6 bites = 3 scores ) . the proportion of these scores for the prey fishes ' pooled left ( and right ) flanks was calculated for each tank .
two - tailed fisher ' s exact probability tests were performed to examine whether in both community tanks the left or right side of the prey fish were preferentially attacked and whether the morphs differed in their foraging scores . a mann - whitney - test was performed to test for differences in ratios of affected left flanks of the prey as well as in the amount of foraging scores between rl and rr pairs .
two different methods were used to analyze foraging behavior . for the three - month old fish , one prey fish was added to a 40 l aquarium and after 12 hours , the prey fish was examined for scars and missing scales by two different researchers ( h . l . and h . k . ) . this procedure was replicated ( 2–5 times ) for each individual scale - eater to investigate whether its foraging behavior was consistent across a series of 2–5 experimental trials during a period of 1–2 weeks . because it was impossible to enumerate number of scars and missing scales on the prey fish ( x . maculatus ) , foraging preference was assessed for each scale - eater based on observed presence / absence of scars and missing scales . a foraging score of + 1 was given for fish that attacked only the right side of prey fish in a particular trial , a score of 0 meant that both sides were attacked , and a score of −1 was given for fish that attacked only the left side . the trials where no scars and no missing scales were observed on the prey fish , or where the prey fish died during the experiments were excluded from the analysis . since the estimated foraging score of each individual was found to be constant over the trials ( e . g . 2 trials : wilcoxon - signed - ranks - test , n = 17 , z = −0 . 333 , p = 1 . 0 ; 3 trials : friedman - test , n = 12 , chi - square statistic = 4 . 333 , p = 0 . 189 ) , the mean foraging score was calculated and used in further analyses . note that the foraging score of 34 scale - eaters was calculated from the 2–5 trials , while that of 10 individuals was obtained from a single trial only . only three of the 47 scale - eaters ( 6 % ) tested at three months never fed on scales .
to investigate whether behavioral foraging preference is translated into foraging score ( e . g . number of scales bitten by the scale - eaters ) , surface areas of attacked left and right flanks of prey ( i . e . surface areas of scars and missing scales ) were calculated for a sub - sample ( n = 15 ) of the two - month old fish . because individual fish that exclusively attacked one side of the prey only left scars / missing scales at that flank ( 100 % ) , the 15 fish were selected from individuals that did not forage exclusively from one side ( e . g . 0 . 1 < probability of left attack < 0 . 9 ) . the attacked areas of the prey fish were estimated in imagej 1 . 45r ( urltoken ) from standardized photographs in a lateral view with an implemented scale . a ratio of the attacked areas ( left to right flanks ) on the prey fish was calculated for each scale - eater and linear regression analysis was then conducted using probability of left attack as an independent variable ( predictor ) and the estimated ratio as a dependent ( response ) variable .
the mouth bending angle , ‘α l − β r’ in ° following [ 14 ] was measured to test for a relationship between mouth / head asymmetry and handed foraging behavior in juvenile fish . for this test , each live test fish was photographed from a dorsal view in a standardized upright position using a zeiss axiophot digital microscope ( zeiss , germany ) . the mouth bending angles were then measured in imagej 1 . 45r : on each image , a triangle connecting the most anterior points of the eye sockets and the tip of the snout was drawn to estimate angles ( ° ) , α l ( angle of the vertex by the left eye ) and β r ( angle of the vertex by the right eye ) ( kusche , lee , meyer , in revision ) .
to evaluate the accuracy of the measurements , repeatability of α l − β r was estimated from repeated and blind measurements that were done from two replicate photographs of the same individuals from sub - samples ( n = 20 , 15 and 15 for the two - , three - and seven - month old fish , respectively ) . repeatability , referred to as the proportion of the total variation that is due to variation among individuals , was calculated from one - way anova ( individual = factor ) following [ 27 ] .
correlation analyses were performed between mouth bending angles and foraging score ( for our test cohort of the three - month old fish ) and behavioral foraging preference ( probability of left attack for the two - and seven - month old fish ) to test for the significant relationship between mouth asymmetry and behavioral handedness . linear regression analyses were also carried out to test whether mouth asymmetry amplifies as body size increases in the two - and seven - month old fish . for those analyses , sl ( standard length ) and tl ( total length ) were used as size measures for the two - and seven - month old fish , respectively .
mouth laterality strongly predicted the preferred attack side as well as foraging scores on either side of the prey fish ( figure 2 ; table s1 ) . both l - and r - mouth morphs from the community tanks clearly exhibited opposed foraging preferences ( fisher ' s exact probability test : n = 41 ; p < 0 . 001 ) and yielded more scars / missing scales in foraging from their preferred flanks ( fisher ' s exact probability test : n = 63 ; p < 0 . 0001 ) ( figure 2 a , b ) . r - morphs preferentially attacked left flanks of the prey fish ( 80 % of affected flanks ; 82 % of foraging scores ) . l - morphs preferred to feed from right flanks ( 75 % of affected flanks ; 80 % of foraging scores ) .
mouth asymmetry strongly predicts foraging preferences and foraging scores in community tanks ( a and b ) and pair tanks ( c and d ) of different laterality combinations .
feeding experiments with laboratory - raised juveniles showed that nearly all test fish preyed immediately on scales . scale - eating behavior is already expressed at an early ontogenetic stage ( two - month old : 100 % ; three - month old : 94 % ; seven - month old : 100 % ) . most individuals showed a clear bias to attack only a particular side of their prey and the frequency distribution of the foraging score and behavioral foraging preference clearly exhibited a bimodal distribution ( except in the seven - month old fish ) ( figure 3 ) . foraging behavior of the younger test cohorts ( of two and three months of age ) showed a significant departure from a unimodal distribution ( two - month old : dip statistic = 0 . 114 , p < 0 . 001 ; three - month old : dip statistic = 0 . 136 , p < 0 . 001 ) , whereas the oldest cohort of seven months of age did not ( dip statistic = 0 . 057 , p > 0 . 5 ) . the mixture analyses with the parametric bootstrap tests further showed that two - component normal distributions best fitted foraging behavior data of the two - ( p < 0 . 001 ) and three - month old fish ( p < 0 . 001 ) , while one - component normal distribution statistically best fitted the data of the seven - month old fish with marginal significance ( p = 0 . 057 ) . however , the graphical inspection of the mixture analysis ( figure 3 c ) and a marginal significance of platykurtosis ( p = 0 . 092 ) rather support a weak bimodal distribution [ 28 ] .
frequency distribution of behavioral foraging preference ( for two - and seven - month old fish ) and foraging score ( for three - month old fish ) shows a bimodal distribution . ( a ) two month ; ( b ) three month ; ( c ) seven month old fish . in ( c ) , the graphical inspection of the mixture analysis ( fitting two single - component normal distributions to the data ) is shown , indicating that the distribution better fits to bimodality than to unimodality , despite a marginal statistical significance of one single - component normal distribution ( p = 0 . 057 ) .
there was considerable ‘inter - individual’ variation in the intensity ( strength ) of lateralized foraging behavior ( figure 3 ) . many fish strongly preferred or even exclusively attacked the left or the right sides [ e . g . 7 of 61 ( 12 % ) and 15 of 61 ( 25 % ) of the two - month old fish foraged exclusively from the left and the right sides , respectively ] , while other fish displayed a less pronounced bias in foraging behavior ( figure 3 ) .
as predicted , a highly significant positive correlation was found between behavioral foraging preference and foraging score in the subset ( n = 15 ) of the two - month old juveniles ( y = 1 . 787 x + 0 . 176 , r 2 = 0 . 759 , p < 0 . 001 ; figure 4 ) , suggesting that foraging score is an outcome of behavioral attack preference . this result further indicates that our field data on foraging preference and foraging score of wild - caught adult fish could indeed reflect ‘behavioral’ foraging preference .
lateralized foraging behavior and foraging score ( e . g . number of scales eaten by the scale - eaters ) are highly significantly correlated ( in a sub - sample [ n = 15 ] of the two - month old fish [ y = 1 . 787 x + 0 . 176 , r 2 = 0 . 759 , p < 0 . 001 ] ) .
our measurements of mouth bending angles appeared to be fairly repeatable : estimated repeatability of the mouth bending angles was 0 . 80 , 0 . 77 and 0 . 87 for the two - , three - and seven - month old fish , respectively . those estimates of the repeatability imply that 77 to 87 % of the total observed variation is attributed to underlying ‘true’ variation in the mouth bending angles among individuals and the remaining 13 to 23 % variation is due to measurement error .
the observed strong lateralization in foraging behavior in young scale - eaters ( e . g . bimodal distribution ) that was not accompanied by notable morphological asymmetry ( see also kusche , lee , meyer , in revision ) and the obvious correspondence between mouth orientation and foraging behavior in adult fish might suggest that handed behavior is probably expressed earlier during development . and , it may actually induce and facilitate morphological asymmetry [ 18 ] , if phenotypic plasticity plays a relatively larger role than the genetic determination of this trait ( lee et al . , unpublished data ) . an alternative hypothesis is that both handed behavior and morphological laterality are genetically governed , but expressed at different ontogenetic stages . however , this hypothesis would seem to be rather unlikely given our observation that laboratory - reared fish of now about two - year of age still have a relatively symmetrical mouth ( hl , personal observation ) .
however , we observed a rather more pronounced laterality in foraging behavior among the younger juvenile fish ( e . g . at two and three months ) , compared to the fish at seven months . the observed dwindling laterality in foraging behavior in the older fish might imply that feeding preference is expressed at an early age ( e . g . at two months ) , but the initial level of laterality would diminish over time ( under laboratory conditions ) unless the fish were constantly to feed on scales . yet , whether this trend means the strength of handed behavior truly decreases with age awaits future experiments on “tracked” individuals over a series of ontogenetic stages during their life time .
we here argue that the hypothesis – ‘handed behavior preceding and driving mouth asymmetry’ [ 15 ] , [ 18 ] – seems more strongly supported by evidence than the original hypothesis [ 8 ] that mouth asymmetry precedes and directs lateralized foraging behavior through natural selection acting on a single gene . different lines of evidence support this hypothesis . in a parallel study , we observed a large amount of variation in mouth asymmetry in 238 wild - caught adult specimens and found a continuous and unimodal ( and not bimodal ) trait distribution ( kusche , lee , meyer , in revision ) . if plasticity rather than genetics plays a comparatively larger role ( lee et al . , unpublished data ) , then this unimodal distribution of laterality might simply be the outcome of different levels of lateralization in foraging behavior . this hypothesis is supported by the findings from the foraging experiments of juvenile fish : juvenile fish did not show complete lateralization of foraging preference and even some individuals attacked equally often at both flanks . whether symmetrically attacking fish might have a potential selective advantage over left or right preferentially attacking fish needs to be tested .
a significant role of phenotypic plasticity in the evolutionary origin of novel morphologies has been suggested repeatedly during the last several decades [ 15 ] , [ 29 ] – [ 32 ] . phenotypic plasticity clearly contributes to shaping the morphology of the jaw and the mouth apparatus in teleost fishes , particularly in cichlids [ 33 ] , [ 34 ] . even different food types or diet hardness can induce changes in the external shape of the head during the ontogeny of some cichlids [ 33 ] , [ 35 ] . the teleost skeleton can quickly adapt to changing external factors , so called ‘mechanical adaptation’ , and skeletal phenotypic plasticity in teleosts seems to be rather pronounced and taxonomically widespread [ 36 ] .
we thank christian sturmbauer for support in the field . thanks to frederico henning for statistical advice and julia c . jones , tom j . m . van dooren and richard a . palmer for helpful comments on an earlier version of the manuscript . we thank mr . zyambo , mr . chiti , mr . chansa , mr . musosa and particularly gabriele legant for their superb technical assistance in the field .
conceived and designed the experiments : hl hk am . performed the experiments : hl hk am . analyzed the data : hl hk . contributed reagents / materials / analysis tools : hl hk am . wrote the paper : hl hk am .
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science 09 apr 1993 : vol . 260 , issue 5105 , pp . 216 - 219 doi : 10 . 1126 / science . 260 . 5105 . 216
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greek , perissos = uneven + greek , odous = teeth ( ref . 45335 )
maturity : l m ? range ? - ? cm max length : 11 . 0 cm tl male / unsexed ; ( ref . 5678 )
scale - eating . prefers fish that feed from the algal layer or fish which are inattentive , the usual prey being tropheus and pseudosimochromis ( ref . 7343 ) .
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 7500 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 01514 ( 0 . 00700 - 0 . 03275 ) , b = 2 . 97 ( 2 . 80 - 3 . 14 ) , in cm total length , based on lwr estimates for this ( sub ) family - body shape ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 4 . 2 ±0 . 86 se ; based on food items .
resilience ( ref . 69278 ) : high , minimum population doubling time less than 15 months ( preliminary k or fecundity . ) .
vulnerability ( ref . 59153 ) : low vulnerability ( 13 of 100 ) .
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disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
pronunciation : refer to our pronunciation key for an explanation of the phonetic symbols .
habitat : this is the primary location where the cichlid is found and is a generalization . this does not mean a fish cannot be found in other habitats .
diet : many cichlids specialize in eating one type of food ; notwithstanding , some of these specialized feeders are flexible and can be opportunistic feeders .
temperament : this describes the overall demeanor of a cichlid toward other tankmates that are of a different species . consider that there is variability in temperament due to various factors , including aquarium size , tankmates of similar appearance , stocking levels , and order of introduction . there may even be some variability among individual specimens .
conspecific temperament : this describes the overall demeanor of a cichlid toward other tank - mates of the same species . consider that there is variability in temperament due to such factors as aquarium size , stocking levels and order of introduction . there may even be some variability among individual specimens .
maximum size : this is in regards to total length ( including the tail ) of typical aquarium specimens . wild specimens may not attain this size , or may in fact grow larger than aquarium raised individuals due to various factors . also consider that this is the typical maximum size and there are exceptional individuals that will exceed it .
difficulty : this measure is a relative value , comparing a single species against all other cichlids . this only accounts for maintanence in the aquarium and not breeding considerations . 1 = easy and forgiving , 5 = extremely challenging .
many people know organisms only by the common names , or
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names . unlike scientific names , common names are almost always different for speakers of different languages . they may also vary regionally within a language . this tab shows all the common names provided to eol for this organism from a variety of providers , including eol curators . currently we can only set one preferred common name per language on a given eol page , but all the names should be searchable .
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baerends , g . p . & j . m . baerends - van roon . 1950 . an introduction to the study of the ethology of cichlid fishes . behaviour suppl . 1 : 1–242 .
barlow , g . w . 1974 . contrasts in social behavior between central american cichlid fishes and coral - reef surgeon fishes . amer . zool . 14 : 9–34 .
brichard , p . 1978 . fishes of lake tanganyika . t . f . h . publications , neptune city . 448 pp .
davies , n . b . 1978 . ecological questions about territorial behaviour . pp . 317–350 .
j . r . krebs & n . b . davies ( ed . ) behavioural ecology , and evolutionary approach , blackwell , oxford .
fryer , g . & t . d . iles . 1972 . the cichlid fishes of the great lakes of africa . oliver and boyd , edinburgh . 641 pp .
keenleyside m . h . a . 1978 . parental care behavior in fishes and birds . pp . 3–29 .
e . s . reese & f . j . lighter ( ed . ) contrasts in bahavior , wiley and sons , new york .
keenleyside , m . h . a . 1979 . diversity and adaptation in fish behaviour . springer - verlag , new york . 208 pp .
krebs , j . r . & n . b . davies . 1981 . an introduction to behavioural ecology . blackwell , oxford . 292 pp .
lewis , d . s . c . 1980 . mixed species broods in lake malawi cichlids : an alternative to the cuckoo theory . copeia 1980 : 874–875 .
liem , k . f . & d . j . stewart . 1976 . evolution of the scale - eating cichlid fishes of lake tanganyika : a generic revision with a description of a new species . bull . mus . comp . zool . 147 : 319–350 .


caerois chorinaeus is a butterfly of the nymphalidae family .

it was described by johan christian fabricius in 1775 .

it is found in suriname , the guianas and peru .

caerois chorinaeus

george arents collection , the new york public library . caerois chorinaeus . retrieved from urltoken
george arents collection , the new york public library .
caerois chorinaeus .
the new york public library digital collections . urltoken
acrylic diptych painting on canvas of the gold and brown caerois chorinaeus butterfly . the diptych is meant to be framed separately and hung 2 inches apart .
george arents collection , the new york public library .
caerois chorinaeus .
new york public library digital collections . accessed july 9 , 2018 . urltoken
< ref name = nypl > { { cite web | urltoken | title = ( still image ) caerois chorinaeus . } } | author = digital collections , the new york public library | accessdate = july 9 , 2018 | publisher = the new york public library , astor , lennox , and tilden foundation } } < / ref >
devries pj , kitching ij , and vane - wright ri . 1985 . the systematic position of antirrhea and caerois , with comments on the classification of the nymphalidae ( lepidoptera ) . syst . ent . 10 : 11 - 32 .
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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
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a neotropical genus of two species dwelling in the forest understorey . the undersides of the wings are remarkably leaflike .
correspondence regarding this page should be directed to andrew v . z . brower at
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each tol branch page provides a synopsis of the characteristics of a group of organisms representing a branch of the tree of life . the major distinction between a branch and a leaf of the tree of life is that each branch can be further subdivided into descendent branches , that is , subgroups representing distinct genetic lineages .
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curators : konstantin efetov , vasiliy feoktistov , svyatoslav knyazev , evgeny komarov , stan korb , alexander zhakov .
species catalog enables to sort by characteristics such as expansion , flight time , etc . .
wahlberg , n . ; leneveu , j . ; kodandaramaiah , u . ; peña , c . ; nylin , s . ; freitas , a . v . l . ; brower , a . v . z . ( 2009 ) nymphalid butterflies diversify following near demise at the cretaceous / tertiary boundary . proceedings of the royal society b : biological sciences 276 ( 1677 ) : 4295–4302 . doi : 10 . 1098 / rspb . 2009 . 1303
all nymphalidae have only 4 fully - functional legs , while pieridae have 6 . beware of pierids with broken legs , or not using all for standing on ( eg leucidia , sometimes ) .
sex unknown gamboa - pipeline road area , colon , panama 75 m . 2018 - may04 david geale
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the haploid chromosome numbers for the south american representatives of the nymphalid subfamilies charaxinae , morphinae ( including brassolini ) and satyrinae . question marks indicate uncertain data
subfamily satyrinae ( nomenclature according to peña et al . , 2006 ; fig . 1 )
the nomenclature follows the list of lamas et al . ( 2004 ) , except for the tribal and subtribal division of the satyrinae , where we follow peña et al . ( 2006 ) ; note that the species names used in some original publications may differ from the names used here . a comma between chromosome numbers shows that the numbers come from different individuals and a dash indicates variation within individuals . in a few cases , a single individual had different chromosome numbers in different cells ; in these cases , the chromosome numbers have been separated with a semicolon . additional data : voucher codes , the name of the specimen in the original reference and an exact reference to the locality are given in urltoken .
localities are grouped by region ; a number at the end of locality codes indicates the number of populations sampled within a region . a letter in parentheses indicates previous work ( a , de lesse , 1967a ; b , de lesse , 1967d ; c , de lesse , 1970a ; d , de lesse , 1970b ; e , de lesse & brown , 1971 ; f , wesley & emmel , 1975 ; g , maeki & remington , 1960a ; h , maeki & remington , 1960b ; i , t . c . emmel , pers . comm . ) . numbers with an asterisk without locality and number of individuals are derived from the unpublished notes left by the late dr h . de lesse .
locality codes : ac , acre ( extreme western brazil ) ; am , amazonas ( north - western brazil ) ; an , andes of north - central colombia ; ba , bahia ( eastern brazil ) ; cc , chocó ( western colombia ) ; cm , chanchamayo ( central peru ) ; ct , catatumbo ( north - western venezuela ) ; cz , canal zone ( central panamá ) ; df , brasília ( central brazil ) ; dr , dominican republic ; eb , eastern bolivia ; ee , eastern ecuador ; es , espírito santo ( eastern brazil ) ; go , goiás ( central brazil ) ; mg , minas gerais ( central brazil ) ; mt , mato grosso ( central brazil ) ; ox , oaxaca ( southern mexico ) ; pa , pará ( northern brazil ) ; pe , pernambuco ( extreme eastern brazil ) ; pn , paraná ( southern brazil ) ; pr , puerto rico ; pt , putumayo ( southern colombia ) ; rg , aragua , northern venezuela ; rj , rio de janeiro ( south - eastern brazil ) ; ro , rondônia ( western brazil ) ; sc , santa catarina ( southern brazil ) ; sp , são paulo ( south - eastern brazil ) ; tv , táchira ( south - western venezuela ) ; vc , valle de cauca ( western colombia ) ; vv , villavicencio , meta ( eastern colombia ) ; we , western ecuador .
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peppers pride ( foaled march 24 , 2003 , in new mexico ) is a retired , undefeated , multiple stakes winning american thoroughbred race horse .

she is by desert god , by fappiano .

desert god 's dam , blush with pride , was also the dam of better than honour , a broodmare that sold for $ 14 million at fasig-tipton in early november 2008 .

peppers pride 's dam is the chili pepper pie mare lady pepper that placed in the 1990 bluebonnet stakes .

peppers pride was her last foal .

joe allen purchased desert god from the university of arizona .

peppers pride

peppers pride at richland hills farm near midway , ky . anne m . eberhardt
peppers pride wins new mexico state racing commission handicap . eugene o ' neill / coady photo
peppers pride cruises to win number 16 in the russell & helen foutz distaff handicap . coady photography
not much has changed about peppers pride from her racing days to her current career as a broodmare .
peppers pride won her 15th consecutive race march 23 , capturing the $ 100 , 000 sydney valentini handicap at sunland park . coady photography
indeed , the memories of peppers pride’s remarkable streak have not faded , for her fans and for those most closely associated with the mare .
peppers pride , on the verge of tying the record of 16 consecutive victories , is set to make her next start april 26 at sunray .
carlos madeira rode peppers pride in each of her 19 victories , which included wins in the new mexico classic cup juvenile fillies . peppers pride proved time and again that she was capable of meeting and defeating the best fillies and mares in new mexico , but no one will ever know how she would have done against those on other circuits . pepper’s pride was retired after her 19
joe allen’s peppers pride has entered a six - furlong allowance race at zia park on saturday afternoon in her attempt to set the record for consecutive wins .
attempting to explain the mare’s success , marr also noted that consistency in the barn’s routine and attention to detail may have helped peppers pride reach her full potential .
spectacular bid , secretariat , nijinsky ii , pepper ' s pride - oct . 4th
here is record breaking filly peppers pride at her home in kentucky , working as a brood mare after her amazing racetrack career of 19 wins from 19 starts . this is from may 2009 .
taylor made farm announced friday afternoon that the undefeated new mexico - bred mare peppers pride has been confirmed in foal to triple crown champion american pharoah . the announcement was made via the farm ' s
peppers pride , who won all 19 of her starts and earned $ 1 . 07 million , has produced three foals , including two by distorted humor , and is in foal to hard spun .
“she has a few of the mental characteristics that [ peppers pride ] had , ” marr said of funny pepper . “she’s not stubborn , but she’s very determined , i guess i would say . ”
although these days she lives a quiet life outside of the public eye – and her win streak was never afforded mass media attention outside of the racing industry – fans have not forgotten about peppers pride .
in each of the past two years , marr used an allowance race as a prep race for peppers pride before the new mexico cup program . that is the same strategy he is using this year .
peppers pride’s owner , joe allen , took the criticism in stride . he stated that he was well aware that peppers pride was not a horse that could be compared to cigar or citation , and said that he was simply glad to have her . there is a refreshing aspect to this that some horseracing fans miss . allen and trainer joel marr did an excellent job of managing the mare’s career , placing her in races where she had the best chance to win .
just as peppers pride’s mental attributes and personality made her an ideal racehorse , that disposition has endeared the mare to her caretakers at taylor made farm , where she is boarded for allen and michael stinson , who now co - owns her .
peppers pride was foaled in new mexico on march 24 , 2003 . her sire was desert god , a horse of moderate success on the racetrack . her dam , lady pepper , placed in the 1990 bluebonnet stakes , but her career was otherwise unspectacular . peppers pride is an example of how breeding does not always serve to precisely indicate how well a horse will perform on the racetrack . while the mare certainly had successful horses in her lineage , there was little on paper that would have inspired bloodstock agents to take note .
marr said peppers pride is back in foal to hard spun . as on the racetrack , the mare has settled comfortably into a routine at taylor made . during foaling season , expecting mares spend the night in the barn and are turned out during the day . once a foal is at a mare’s side , however , the routine becomes the opposite . now , peppers pride and her 4 - month - old colt come to the barn in the morning to be looked over and attended to before being turned out in a group for the rest of the afternoon and night .
all of peppers pride’s races came in her native new mexico , where she dominated her division for four seasons . she captured the new mexico cup juvenile fillies stakes as a 2 - year - old , the new mexico cup championship fillies stakes during her sophomore season , and won back - to - back editions of the event’s fillies and mares stakes in 2007 and 2008 . her list of stakes triumphs included two editions each of the new mexico state racing commission handicap and the sydney valentini handicap . peppers pride carried weights as high as 127 pounds , an impost she won under three times .
peppers pride is an example of brilliant management by the team of owner and trainer . they only raced in events they thought they could win , and as a result the mare broke the record for consecutive wins in north america . camarero , a horse that raced in puerto rico , won an amazing 56 in a row !
peppers pride ( usa ) dkb / br . m , 2003 { 2 - h } dp = 7 - 7 - 8 - 0 - 0 ( 22 ) di = 4 . 50 cd = 0 . 95 - 19 starts , 19 wins , 0 places , 0 shows career earnings : $ 1 , 066 , 085
the joel marr - trained peppers pride has won each of her 16 career wins and shares the record for consecutive wins with triple crown winner citation , two - time horse of the year cigar , santa anita derby winner mister frisky and the louisiana - bred sprinter hallowed dreams . “she’s doing well , ” said marr . “she hasn’t raced since april , but has had two works . ” the daughter of desert god won the foutz distaff handicap at sunray park on april 26 . she has remained in training since that win in marr’s stable . peppers pride was scheduled to make her record attempt on two occasions this summer at ruidoso downs . she was slated to run in the lincoln handicap on july 27 but that racing card was cancelled after the remnants of hurricane dolly damaged ruidoso downs . the lincoln handicap was rescheduled for august 31 , peppers pride was entered and then marr elected to scratch her due to an off track . she has only raced on fast racing surfaces . this means that marr has twice prepared to race and then the 5 - year - old mare has not competed , not the optimal training regimen for a racehorse . “it probably has had an effect and we’ll find out saturday , ” marr said . “but , she has been training well . ” peppers pride has raced at zia park each of the last three years and is five - for - five over the hobbs oval . as a 2 - year - old in 2005 , peppers pride impressed when she made a four - wide rally on the turn to win the new mexico classic cup juvenile fillies , the premier race for two - year - old females during new mexico cup day .
began their careers training quarterhorses in the state . the thoroughbred competition in new mexico , however , is typically considered to be among the weakest in the united states . this would become a source of controversy after peppers pride set the record for consecutive wins . many pointed out that cigar and others with long win streaks had accomplished their wins against much tougher competition .
peppers pride did everything right during her racing career , too . from july 16 , 2005 , to dec . 14 , 2008 , the mare answered the call to the post 19 times and emerged victorious on each occasion , the longest undefeated career by a modern north american thoroughbred . the streak included 14 stakes wins , and she earned $ 1 , 066 , 085 .
peppers pride produced her third foal , a colt by malibu moon , on feb . 22 . her first two foals are by distorted humor – a filly foaled in 2011 and a yearling colt . the filly , named funny pepper , is at a training center in new mexico , preparing to join marr’s string to see if she can follow in her mother’s large hoofprints .
now boarded at kentucky ' s taylor made farm , peppers pride has settled into her life as a broodmare . “she ' s doing great ; she ' s having nice foals , ” taylor made vice president frank taylor told the daily racing form in 2013 . “she ' s just really easy to get along with and never causes any problems . she fits right in . ”
lesson that should be learned from the career of peppers pride is the factor owners and trainers play in a horse’s career . many times when a horse performs poorly it is because they have been entered in races that are too difficult . horse betting requires a handicapper to look at a horse’s past performances to determine whether or not they belong in the race they are in today .
“i think peppers pride is putting plenty of bone on [ her foals ] , and they’re athletic - looking , ” taylor said . “i really love the distorted humor yearling she’s got this year . he’s just a really big , strong , good - looking horse . he’s got a lot of class to him , i think . i think people will really like him a lot . ”
this entry was posted in bloodstock and tagged american pharoah , frank taylor , joel marr , new mexico , pepper ' s pride , taylor made farm by paulick report staff . bookmark the permalink .
in 2006 , peppers pride was nearly defeated when she won her prep allowance race by a nose with a late run . she then won the new mexico cup championship for fillies by a length as the 11 - 10 favorite . last year , when the winning streak was gaining national attention , she won her 12th straight race with a seven - length allowance win at six furlongs and then captured the new mexico cup championship for fillies and mares with a late rally to win by one - and - a - half lengths . peppers pride will point to the new mexico cup championship for fillies and mares after saturday’s allowance race , providing she comes out of the allowance race in good order . new mexico cup day is november 9 and the $ 2 million in purses makes that program the richest day for any state - bred racing program .
any thoroughbred trainer will tell you that winning just one race is hard . there are so many things that can go wrong in the running of a race . to win , a horse must be talented but also possess racing luck . peppers pride , a filly that raced solely on the new mexico racing circuit , had both , and she broke the legendary cigar’s record for consecutive wins when she captured her 19 th straight victory on december 14 , 2008 .
“she was a tremendous athlete , not only physically , but mentally , ” said joel marr , who trained peppers pride for owner and breeder joe allen . “you don’t do that by just getting lucky 19 times in a row , obviously , or it would be done all the time . she had that sense that most horses probably don’t have – she knew how to win . she had that ability and that sense and that determination . she knew exactly what she was doing .
owned by michael stinson and her breeder joe allen , the 13 - year old mare holds a record of 19 wins from 19 starts , including 14 stakes wins . the daughter of desert god earned over $ 1 million in her three - year career for trainer joel marr , and retried with her perfect record intact at the end of 2008 . peppers pride held the north american record for the longest undefeated streak in history , alongside champion zenyatta , until rapid redux took over that title in 2012 with 22 wins .
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2 - year - old gelding broke his maiden july 8 at gulfstream park .
blame filly has now won a pair of graded stakes for new trainer bob baffert .
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as a sports fan , i love winning streaks , and i es . . .
as the end of the 2011 year draws near i decided t . . .
winx ' s staying power as one of the world ' s top rac . . .
jason servis is on the best run of his career , wi . . .
t . j . comerford , assistant trainer for aidan o ' brie . . .
“she’s really a classy mare to be around , ” said frank taylor , vice president of boarding operations at his family’s taylor made farm , the nicholasville , ky . , facility where the 10 - year - old daughter of desert god resides . “she just does everything right . ”
“we trained her to be fit and to do the best job she could , but she ultimately won the races , ” marr added . “it was just something that she had . i can’t explain it . just the ability she had within her and . . . it was just something that she possessed way before we touched her . ”
“she was just one in a million , ” marr said . “and a lot of it was mental . obviously , she was physically a superb athlete and had tremendous abilities . and people will say , ‘well , you ran her in new mexico . ’ well , we did , and [ a new mexico - bred is ] what she was . but it really doesn’t matter . ”
“all the people around her – i was her only trainer , ” marr said . “ [ carlos madeira ] was her only rider . she had the same farrier , the same vet , the same groom . she stayed in the same barn at every racetrack , the same stall . she went to post with the same pony every time . just a lot of things like that . horses are creatures of habit . they need to be fed at the same time ; they need to be exercised at the same time . they need a routine , and she had that for the four , almost five years that we had her . she went to the track at the same time every morning , with the same person on her back .
“joe allen was the breeder and the owner and the partial stallion owner . it was really special . it wasn’t something we went out and bought . she was raised there . it made it a lot more special just to be around her . ”
“she’s doing great ; she’s having nice foals , ” taylor said . “she’s just really easy to get along with and never causes any problems . she fits right in . ”
the yearling colt is expected to be offered at this year’s keeneland september sale , where he could be among the featured horses in the catalog .
“we get some regular calls from people wanting to come visit and come see her , ” taylor said . “we probably get [ a few ] a month , fans wanting to know how she’s doing or wanting to come see her . ”
“i wouldn’t trade it for anything , ” marr said . “it was tremendous to be a part of it . and you look back and you can say that you were part of something like her [ winning streak ] . not very many people get to say that . you knew there was something special about her , just the way she held herself , the way she looked , the look in her eye . i was just lucky to be a part of it . ”
won new mexico cup juv . fillies s . ( zia , 6 . 5f ) , rio grande senorita futurity ( rui , 6 . 5f )
won new mexico cup championship fillies s . ( zia , 6 . 5f ) , la senora h . ( sun , 6f ) , la coneja h . ( sun , 5 . 5f )
at 4 : won sydney valentini h . ( sun , 8f ) , new mexico cup f & m ; championship , new mexico racing commission h . ( zia , 6f )
won sydney valentini h . ( sun , 8f ) , russell & hellen foutz distaff h . ( srp , 6 . 5f ) , new mexico cup fillies & mares s . ( zia , 8f ) , new mexico racing commission h . ( zia , 6f )
april 26 , 2008 : equaled north american record for 16 consecutive wins , a record shared by cigar , citation , hallowed dreams , and mister frisky .
broke the consecutive win record on october 4 , 2008 , in a six furlong allowance race at zia park .
joel marr was retained by the mare’s owners to train her for racing in her home state of new mexico . with several notable tracks , new mexico is perhaps more known for quarterhorse racing . many famous trainers such as
why would a trainer enter a horse in a race they cannot win ? there are several reasons . first , the trainer may be pressured by the owner to enter a specific race . this does not happen as much at the highest levels with trainers like steve asmussen or todd pletcher . those trainers are likely to refuse an owner’s request . at the smaller tracks , however , this happens all the time . the trainer must often give in to the owner’s request or lose the horse to another trainer that will .
sometimes , a horse is entered in a race to prepare or “prep” for another , more important race . in this case , winning may not be the goal . the trainer may simply want the horse to have a good showing and increase its fitness level . finally , a horse may simply have no other place to compete and need to race .
are used to the activity of a race , and many of them need regular racing in order to keep sharp . unfortunately , especially on the smaller circuits , there may not always be a suitable race for a horse’s class level . in that case , the horse is often entered in a race that is beyond their skill level just to give them an opportunity to run .
? you can take advantage of great bonuses and get access to a full menu of betting options at the most popular tracks .
bloodstock agent tom mcgreevy , who likes to work exclusively for one client , has signed on to buy yearlings and 2 - year - old at sales for texas owner mike stinson .
the auction offers horseshoes worn by afleet alex and first samurai , a vip three chimneys tour , and autographed race photos . all proceeds help with the mission of racehorse retirement and rescue .
as california chrome prepares for a shot at redemption in this year ' s dubai world cup ( uae - i ) , interest in the horse as a stallion prospect continues to grow , much to the satisfaction of taylor made farm .
new mexico - based trainer joel marr tallied his 1 , 000th career thoroughbred victory by winning the 11th race nov . 11 at zia park in hobbs , n . m . , with sweet diamond .
the keeneland september sale ' s second book 1 session sept . 10 proved as bullish as the previous day ' s opener as far as $ 1 million horses were concerned , with four seven - figure horses exiting the ring again .
undefeated mare to stay in new mexico for mark - tying 16th straight victory attempt .
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