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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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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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
vernacular
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 .


the mauritius fody ( foudia rubra ) is a rare species of bird in the weaver family .

it is endemic to the island of mauritius .

it is classified by birdlife international as being endangered .

it is also on the united states ' endangered species list with an endangered status .

this bird is 14 centimeters long .

breeding males are olive brown with a red head , breast and rump patch and black lores .

while females , non-breeding males and juveniles are olive brown with white wing bars and a brown bill .

the bird lives in several types of forest , including degraded areas , as well as plantations .

stands of japanese cedar ( cryptomeria japonica ) have replaced native vegetation and now provide protection against predators .

it feeds on insects like grasshoppers , beetle larvae , caterpillars , and also spiders .

it feeds on nectar regularly , using its specialised brush-tipped tongue .

the bird is a weaver , the male and female cooperating to weave each nest , from material like grass , moss and small twigs .

the mauritius fody is threatened by the loss of its habitat and predation from introduced predators .

beginning in the 1970s much of its habitat was lost when the land was cleared for plantations .

by 2001 there were perhaps no more than about 100 breeding pairs .

nests are raided by predators , especially the black rat ( rattus rattus ) and the crab-eating macaque ( macaca fascicularis ) .

this is currently the main cause of the bird 's decline .

some areas of intact habitat have high nest predation , but areas of low nest predation may be poor habitat .

the common myna has also been observed preying on nests .

nest failure may occur when it is infested with tropical nest fly .

the larvae of the fly attack the chicks , latching on and feeding on their blood , causing dehydration and anemia in the chicks .

conservation efforts include the control of rats and macaques .

a captive breeding program carried out by the mauritan wildlife foundation has produced many chicks .

eggs are removed from nests in the wild and hatched in captivity as the wild pairs produce and rear another clutch simultaneously .

nests are treated for tropical nest fly .

supplemental food and water are given .

the population has increased recently due to conservation programs establishing sub-populations on offshore islands .

due to these conservation efforts the species was downlisted from critically endangered to endangered in 2009 .

île aux aigrettes , an islet off the main island of mauritius , is now home to a number of mauritius fodies and other threatened species that have been translocated there .

mauritius fody

insula franciae = mauritius . the historic distribution of the mauritius fody ranged over much of mauritius , so the type could have been collected anywhere on the island .
the endemic fody bears a close resemblance to the exotic madagascar fody which is very common in mauritius . . the male madagascar fody , however , has completely bright red plumage during breeding season .
the mauritius fody is classified as endangered ( en ) on the iucn red list ( 1 ) .
the population of the mauritius fody suffered a rapid decline in the 1970s due to nest predation and habitat loss .
often mistaken : the madagascar fody ( foudia madagascarensis ) is very common in mauritius . photo © g . gerra & s . sommazzi
the mauritius fody is found only on the island of mauritius . the male mauritius fody boasts a brilliant red head , neck and breast , while females have drabber olive - brown plumage . adults are approximately 14 centimetres long . this solitary songbird prefers native scrub and forest habitat and feeds mainly on insects , supplemented by fruit and nectar .
safford , rj . the annual cycle and breeding behaviour of the mauritius fody , foudia rubra . ostrich , 68 , 58 - 67 .
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clearing mauritius’s forests has been catastrophic to the fody , while predators such as black rats and crab - eating macaques that have been brought to the island have caused almost total breeding failure in most areas . the native fodies may also face competition from the madagascar fody , which has been introduced to mauritius .
the mauritius fody inhabits remaining native forest but also survives in degraded forest that has been invaded by exotics , as well as forests of exotics such as pines .
safford rj . nesting success of the mauritius fody foudia rubra in relation to its use of exotic trees as nest sites . ibis , 139 , 555 - 559 .
safford , r . j . ( 1997 ) the annual cycle and breeding behaviour of the mauritius fody foudia rubra . ostrich , 68 ( 2 ) : 58 - 67 .
the diet of the mauritius fody is comprised primarily of invertebrates , but nectar , fruit , seeds and the eggs of birds and geckos are also consumed ( 5 ) . some females have even been seen feeding on the eggs of other mauritius fodies ( 3 ) .
the mauritius fody has suffered a constant and severe decline in range and numbers since 1975 , and is now found only in southwest mauritius ( 2 ) , and on the small , offshore island of ile aux aigrettes , where captive - bred fodies were released in 2003 ( 3 ) .
peter brown was the first to illustrate , the mauritius fody . he called it the red - headed finch and gave a description of the bird . brown notes that the bird was in the collection of
safford , rj . the destruction of source and sink habitats in the decline of the mauritius fody , foudia rubra , an island - endemic bird . biodiversity and conservation , 6 , 513 - 527 .
the mauritius fody is restricted to south - west mauritius ( see red polygon on map ) where there are 3 small sub - populations . in 2005 , hand reared chicks were released onto the offshore islet ile aux aigrettes where the species apparently occurred historically ( red arrow on map , see also
on january 12 , 1995 , the mauritius fody was designated as endangered in the entire range . within the area covered by this listing , this species is known to occur in : mauritius . the u . s . fish & wildlife service ( null ) is the lead region for this entity .
safford , r . j . ( 1997 ) nesting success of the mauritius fody foudia rubra in relation to its use of exotic trees as nest sites . ibis , 139 ( 3 ) : 555 - 559 .
the mauritius fodies ( foudia rubra ) - also known as mascarene fodies - are only found on the island of mauritius off the southeast coast of africa , where they inhabit forests and plantations .
, an english ornithologist and collector . the fody would have been brought by ship from mauritius to england , where it would have been purchased by tunstall . tunstall kept it in captivity for several years ( fox 1827 ) .
mauritius fodies mostly feed on insects ; but they will also take fruit , nectar , seeds , geckos and - on occasion - the eggs of other birds ( including those of other mauritius fodies ) .
. the illustration in daubenton 1783 is of a male , and the first female to be painted , but buffon does not appear to have written about it . gmelin noted the locality for the mauritius fody as
insula franciae
.
, an english naturalist and illustrator . gmelin did not realise that the same species was involved , due to the slight differences in the artwork . the latin name he provided here was invalid . gmelin noted the locality for this mauritius fody specimen as
insula mauritii
.
gmelin 1789 syst . nat . , 1 ( 2 ) , p . 877 isle de france [ = mauritius ] .
safford , rj . distribution studies on the forest - living native passerines of mauritius . biological conservation , 80 , 189 - 198 .
similar spp . non - breeding male , female and juvenile separated from madagascar red fody f . madagascariensis by darker , less streaked plumage , plumper body and relatively shorter tail .
inhabits all types of forest , even degraded forest invaded by introduced plant species ( 2 ) . on the mainland , the highest densities of this bird are found in small plantations of exotic trees surrounded by degraded native forest . on ile aux aigrettes , the mauritius fody inhabits lowland ebony forest and coastal scrub ( 3 ) .
endangered . restricted range species : present in mauritius eba . formerly listed as critically endangered , but downgraded to endangered because population , albeit extremely . . .
although threatened by the clearance of upland forest , the mauritius fody has also been suddenly lost from areas of intact habitat ( 2 ) . it is thought that these regions have been affected by particularly high levels of egg predation by both the black rat ( rattus rattus ) and the crab - eating macaque ( macaca fascicularis ) ( 6 ) .
garrett , l . j . h . , jones , c . g . , cristinacce , a . and bell , d . j . ( 2007 ) competition or co - existence of reintroduced , critically endangered mauritius fodies and invasive madagascar fodies in lowland mauritius ? . biological conservation , 140 : 19 - 28 .
safford , rj . a survey of the occurrence of native vegetation remnants on mauritius in 1993 . biological conservation , 80 , 181 - 188 and 84 , 197 .
these brightly colored birds are now only found in southwest mauritius and on the small , offshore island of ile aux aigrettes , where captive - bred birds were released in 2003 .
mauritius fodies generally form monogamous pair bonds ; although males have on occasion been observed with more than one female . pairs maintain a breeding territory that they defend throughout the year .
craig , a . ( 2018 ) . mauritius fody ( foudia rubra ) . 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 mauritius fody is mainly insectivorous , and feeds on grasshoppers , beetle larvae , caterpillars , and also spiders . berries are eaten regularly by some individuals . it feeds on nectar regularly , using its specialised brush - tipped tongue . it forages in dead leaves , and probes bark and epiphytes , and gleans from leaves , for invertebrates . it is usually found singly , in pairs or in family groups .
safford , r . j . and jones , c . g . ( 1998 ) strategies for land - bird conservation on mauritius . conservation biology , 12 : 169 - 176 .
chinese : ? ? ? ? ? ? . . . czech : snovatec mauricijský . . . danish : mauritiusvæver . . . dutch : mauritiuswever . . . finnish : mauritiuksenkutoja . . . french : cardinal de maurice , foudi de l ' ile maurice , foudi de maurice . . . german : mauritius weber , mauritiusweber . . . italian : fody di mauritius , tessitore di mauritius . . . japanese : benihataori , moarishasubeninojiko . . . norwegian : mauritiusvever . . . polish : wiklacz maurytyjski , wik ? acz maurytyjski . . . russian : ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? . . . slovak : fodia maurícijská . . . spanish : fodi de mauricio , fodi de mauritana . . . swedish : mauritiusfody
. however , since 1993 , the population has been stable due to conservation action , the population being about 140 - 170 adults and an estimated 108 pairs in 2012 . it can co - exist with the introduced madagascar red fody .
historically , fodies inhabited the upland areas of southwest mauritius . today , this critically endangered bird is found in just three areas of the island : bassin blanc , macchabee forest road and ile aux aigrettes .
at present , it would be impossible to eliminate all rats and macaques from the whole of mauritius , and therefore conservation efforts have focussed on establishing populations on offshore , predator - free islands ( 3 ) . the release of captive - bred individuals onto the predator–free island of ile aux aigrettes in 2003 has proved successful , with the population , which is provided with supplementary food , increasing to 151 individuals by july 2008 ( 3 ) . other conservation measures in place to help protect the mauritius fody include restoring native forests and establishing plantations of non - invasive , exotic trees such as japanese red cedar ( cryptomeria japonica ) which mammalian predators appear to avoid ( 7 ) .
my thanks go to mr . vikash tatayah of the mauritius wildlife foundation for letting me have access to the conservation area where i was able to take photographs of the endemic birds held at the black river aviaries .
unusually for weaver birds , species of foudia are often monogamous . the mauritius fody generally holds territories of over one hectare exclusively with just one mate ( 4 ) , although some male mauritius fodies have been observed with more than one female ( 3 ) . breeding between late june and early april ( 3 ) ( 4 ) , both sexes help to build the nest , which is then lined by the female only . between two and four eggs are laid and incubated by the female , with the male helping to feed the chicks after hatching for two weeks , before they fledge and leave the nest to disperse . at the end of the breeding season , the adults undergo a complete moult . pairs defend the territory all year , and do not make seasonal migrations ( 4 ) .
in 2003 , the mauritius wildlife fund started a program of captive breeding at the black river aviaries . 14 birds were released in the île aux aigrettes nature reserve . but in early 2004 only 9 survived . no trace was found of the other 5 . for the next release , the implant of chips is planned to follow their movements .
the mauritius fody is monogamous and territorial . birds may have long - term pair bonds and remain on their territories year - round . the nest is domed , with a side entrance and often with a porch . nests are built by both sexes using grass with moss , lichens or small twigs . nest material is gathered within 100 m of the nest . the female adds lining , mainly feathers , before egg - laying and up to 8 days into incubation . nests may be strong , able to withstand gale force winds , but others have less secure attachments leading to active nests falling to the ground . nests are built in trees , usually hidden in foliage . many tree species have been recorded as nest sites , but with exotics being used increasingly since the 1970s .
is found only on mauritius island and is listed as endangered since it has an extremely small population . the male in breeding plumage has the head red or orange - red , with a black mask through the eye ; the lower rump and uppertail - coverts are orange - red , and the rest of the body is olive - grey . the female and non - breeding male are dull olive - green , with a yellowish wash below . the juvenile resembles the female , except for a paler brown bill , which may be retained for its first year . the non - breeding male , female and juvenile may be separated from the
in 1993 , roger safford completed an intensive four - year phd study on all of the mauritian passerine species . since 1993 some work has been done on a volunteer basis since we have not yet been able to secure funding to help save our passerines . volunteers and students have been conducting population surveys on all the critically endangered species to gain knowledge of the ecology of each species for future conservation . conservation efforts to help passerines will be more difficult than for the larger , more resilient birds mwf is working with . only one other research station in the world , hawaii , is attempting to restore highly endangered passerine populations through captive breeding . hawaii’s passerine program is still in the preliminary stages and much remains to be learned to develop strategies that will best help conserve these tiny birds . interestingly , the methods used by conservationists in hawaii have been adapted from mwf’s work on pink pigeons . the story will come full circle if we can learn from them on how to care for our passerine species . in 1999 , 21 new territories of olive white - eyes , 8 new pairs of paradise flycatchers and 3 new pairs of mauritius fody were found near newest mwf field station at combo . in addition to many hours of observation resulting in previously unknown nesting behaviour ( male olive white - eyes were seen to share in the care of eggs and nestlings ) , 3 nests of olive white - eyes were found and monitored . previous to this only 2 nests had ever been recorded for this species . unfortunately , all 3 nests were predated by ship
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choose different species from drop - down list and press ' go ' button . see
, a german naturalist . gmelin described many new bird species in a book in 1789 in the style of linnaeus ' publications , giving a brief description in latin and a synonomy , including a reference to the coloured engraving by francois - nicolas martinet in the book edited by
brown ' s painting appeared before that of daubenton , but these paintings were probably based on different specimens , so all that can be said about the collection details is that the type specimen was taken before 1783 by an unknown collector . this is the last weaver described by gmelin .
type specimen no longer exists ; the illustration of daubenton 1783 serves as a type .
the above is based on weaver wednesday 2 , a weekly series about the discovery of each weaver species .
by their darker , less streaked plumage , plumper body , relatively shorter tail , and different calls .
some pairs habitually desert their nests , while others rarely do so . the clutch is 2 - 3 pale blue eggs . incubation is by the female . initially the female feeds the young , and later chicks are fed by both sexes . breeding success is low , with predation being the major cause of nest loss .
the above is based on weaver wednesday , a weekly series about weaver species .
pair bond monogamous breeding season oct - feb nest site placed 2 - 9 m above ground in tree nest building division of work unclear , as both sexes seen to carry material , but in some cases female did most building colony size no information clutch size 2 - 3 eggs egg colour pale blue egg size average size of ten eggs 19 . 2 x 13 . 9 mm incubation incubation apparently by female only , period reported as 14 days chicks and nestling period chicks fed by both sexes , nestling period recorded as 18 days
phown ( photos of weaver nests ) provides valuable info on breeding distribution and colony sizes of weavers . you can contribute by registering and submitting photos at virtual museum webpage .
the above is based on weaver wednesday 3 , a weekly series about range changes in south african weaver species .
durrell wildlife conservation trust is a member of the association of jersey charities , membership number 69 . patron : hrh the princess royal . founder : gerald durrell , obe , lhd . durrell wildlife conservation trust - uk is registered in england and wales . a charitable company limited by guarantee . registered charity number : 1121989 . registered company number : 6448493 . registered office : c / o elian corporate services ( uk ) limited , 35 great st . helen ' s , london ec3a 6ap
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del hoyo , j . , collar , n . j . , christie , d . a . , elliott , a . , fishpool , l . d . c . , boesman , p . and kirwan , g . m . 2016 . hbw and birdlife international illustrated checklist of the birds of the world . volume 2 : passerines . lynx edicions and birdlife international , barcelona , spain and cambridge , uk .
14 cm . medium - sized , brown forest weaver . vermilion - red head , neck and breast with black loral area . dark brown back , wings and tail streaked with buff . reddish rump and uppertail - coverts .
this species is classified as endangered because it has an extremely small population . the population has been stable since the early 1990s but then increased following an island translocation , and if it remains stable or increasing with over 250 mature individuals it may warrant downlisting to vulnerable in the future .
, suffered rapid population declines since 1975 , descending from 247 - 260 pairs to c . 108 - 122 pairs in late 2001 ( nichols 2002 )
; no doubt following long - term historic population declines owing to heavy predation by invasive mammals . between 1975 and 1993 , a 55 % decline in both population size and area of occupancy occurred ( safford 1997c )
, and the number of locations fell from six in 1975 to three ( r . nichols
, and an increase in range has been recorded in the main breeding subpopulation ( c . jones
. the long term viability of these small subpopulations is in doubt ( r . nichols
in 2005 , 45 hand reared chicks were released onto ile aux aigrettes ( where the species apparently occurred historically , v . tatayah
2010 ) , where two pairs from previous releases have raised chicks ( anon . 2005 )
. following that success , in 2006 , c . 40 young fledged on ile aux aigrettes and the total population on the island stood at 140 individuals in 2008 ( l . garrett
2012 ) . the most recent population estimate is 180 - 200 individuals in 2011 - 2014 ( v . tatayah , c . jones and n . zuel
. 2015 ) . it is hoped that the population will increase further in the coming years as it expands on ile aux aigrettes . a second translocation to round island began in 2010 ( v . tatayah
however of 32 birds released here between october 2010 and january 2011 all had died by early 2012 , most or all having been predated by round island boas ( v . tatayah
the mainland population was estimated at 216 - 244 individuals in 2002 , with a minimum of 108 breeding pairs , and the population was estimated to have remained at 108 pairs in 2012 ( v . tatayah
2012 ) . this is roughly equivalent to 140 - 170 mature individuals . the latest population estimate for the mainland population in 2011 - 2014 is 240 - 330 individuals ( v . tatayah , c . jones and n . zuel
. 2015 ) , equivalent to 160 - 220 mature individuals . it has since increased following the release of birds on ile aux aigrettes in 2006 with a total of 110 adults there in 2012 and an estimated 180 - 200 individuals in 2011 - 2014 ( v . tatayah , c . jones and n . zuel
. 2015 ) . the pre - release population estimate will be retained until the population is confirmed to have sustained itself above 250 mature individuals for five years .
the population declined rapidly until the early 1990s since which time it has been stable or possibly increasing on the mainland ( cristinacce 2007 , v . tatayah
2011 ) . following translocations to ile aux aigrettes it increased but has since dropped and then stabilised ; translocations to round island have so far been unsuccessful .
it holds territories in all types of native forest , including degraded forest , and it shows an increasing reliance on non - native plantations that afford some protection from nest predators ( c . jones
2010 ) , c . 2 months earlier than the mainland population ( safford , 2013 ) . breeding may be terminated if there is a prolonged period of rain ( > 1 month ) , but dry conditions will not cause an interruption ( safford , 2013 )
historically , clearance of upland forest , particularly for plantations in the 1970s , catastrophically affected this species . introduced predators , notably black rat
and nest predation is regarded as the major cause of present - day decline in this species ( r . switzer
. released birds on round island in 2010 suffered predation by round island boas , with none surviving by early 2012 ( v . tatayah
rats and macaques were controlled as part of a programme to rehabilitate plots of native vegetation ( safford and jones 1998 ) , however macaque control has since dropped to nearly nil ( v . tatayah
. a captive - rearing programme implemented by the mauritian wildlife foundation , the gerald durrell endemic wildlife santuary and the national parks and conservation service is proving highly effective and in 2005 produced 47 individuals from captive parents ( 25 ) and hand reared wild born chicks ( 22 ) ( anon . 2005 )
. protocols for captive husbandry and artificial propagation were developed to facilitate the translocation objectives ( r . switzer
2010 ) . the black river national park partly covers its range , and the habitat around bassin blanc , not originally included within the boundary , is a priority for compulsory government purchase in the future ( jones and hartley 1995 , r . safford
, but this had not happened by 2012 . research into the species ' s ecology is ongoing , and prospective surveys to assess the suitability of round island for translocation were conducted ; initial releases on the island began in 2010 ( l . garrett
and native forest . develop a conservation management area at combo , stocked with favoured nectar - producing plants and with predator controls . increase breeding productivity by supplemental feeding , double clutching and captive - rearing of harvested wild clutches ( c . jones
. identify possible native habitat refuges on both the mainland and offshore islands ( safford 2013 ) . continue releases on offshore islands , including flat island when management allows , and monitor the population on ile aux aigrettes . advocate for the compulsory purchase of land separating bassin blanc from the black river national park and ensure that the national park boundary is extended and appropriate management activities implemented .
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a 1999–2001 survey revealed that the number of fodies in the wild had shrunk substantially since 1975 , from 247–260 pairs to just over 100 . thanks to intensive management , including hand - rearing and translocations to offshore islands , the population has stabilized and is increasing . according to the most recent population estimates , there are 160–220 individuals on the mainland , with an additional 180–200 individuals on offshore island ile aux aigrettes . in 2009 , the species was downlisted from critically endangered to endangered .
wildlife preservation canada saves animals on the brink of extinction . since 1985 , we’ve been saving critically endangered species – species whose numbers in the wild are so low that a great deal more than habitat protection is required to recover them .
has been thought to include † f . delloni as a race . monotypic .
14 cm ; 16–20 g . male breeding has forehead , crown , nape , chin , throat , cheek , ear - coverts and breast red or orange - red , black mask through eye ; mantle and back greyish - . . .
song of male an extended series of disyllabic notes , generally terminating in buzzy sound ; female . . .
remaining native forest ; also degraded forest invaded by exotics , as well as forests of exotic . . .
primarily insectivorous ; seen to take grasshoppers ( orthoptera ) , beetle larvae ( coleoptera ) , caterpillars ( lepidoptera ) , also spiders ( . . .
season oct–feb . monogamous . occupies permanent territory 4–5 ha in size in sparsely populated areas , or down to as small as 0 . . .
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this map displays aggregated data from ibc and my birding ( anonymously ) ; markers do not indicate precise localities .
breeding males of this medium - sized forest weaver bird are highly distinctive , with a bright red head , neck and breast , and dark olive - brown back , wings and tail streaked with pale brown ( 2 ) ( 3 ) . the rump is also reddish ( 2 ) , and two white bars pattern the wings ( 3 ) . in the non - breeding season , males lose their vivid red colouration and resemble females . the slightly hooked beak of males is black , while that of females and juveniles is horn - coloured ( 3 ) .
invertebrates animals with no backbone . monogamous having only one mate during a breeding season , or throughout the breeding life of a pair . territory area occupied and defended by an animal , a pair of animals or a colony .
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recommended citation birdlife international ( 2018 ) species factsheet : foudia rubra . 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 .
foudia rubra male at the black river aviaries of the mauritian wildlife foundation . photos © peter m . c . werner 2 - 2004
foudia rubra female . photo © peter m . c . werner 2 - 2004
the sexes are different , the female being rust - brown , the male basically grey .
the preferred habitat includes native scrub vegetation with a few scattered taller trees , but the species also inhabits low native scrub and native forest .
its diet is comprised primarily of insects , but also fruit and nectar . occasionally robs nests of other birds , such as the pink pigeon , sucking the eggs
clearance of upland forest has catastrophically affected this species . introduced predators , notably black rat rattus rattus and crab - eating macaque macaca fascicularis and mongoose , have caused almost total breeding failure in most areas . introduced f . madagascariensis may compete and restrict its range .
it has unexpectedly disappeared from areas of apparently intact habitat , possibly regions of severe nest - predation , previously sustained by relatively predator - free areas which have now been degraded and can no longer supply new recruits .
it was already becoming fairly rare at the turn of the twentieth century , although it was still not considered uncommon as late as the 1950s .
between 1971 to 1974 , the clearing of an upland forest at les mares on plaine champagne had a devastating effect on the species , decreasing its population by more than 50 % . since then , its habitat has been better protected , as remaining native forest habitat received almost complete protection with the creation of the macabe / bel ombre nature reserve in 1974 .
foudia rubra has suffered rapid population declines since 1975 , descending from 247 - 260 pairs to c . 105 - 125 pairs in late 1999 . from 1975 to 1993 , a 55 % decline in both population and area of occupancy occurred . however , since 1993 , the decline rate has slowed , and an increase in range has been recorded in the main breeding subpopulation .
the tiny mare aux vacoas subpopulation has remained stable ( four pairs ) , but numbers and range have continued to decline in the bel ombre subpopulation ( seven pairs ) . research indicates that there is no population fragmentation during winter .
two or three pale - green spotted eggs laid in a rough , cup - shaped nest of thin plant parts bound by cobwebs and placed high in trees . both parents sit on the eggs . nest predated by rats .
an attempt was made to introduce the species to the island of reunion , but it was unsuccessful it is uncertain whether further attempts will be made . the introduction and establishment of two species of nectar - producing shrubs on the high plateau has been suggested .
the main goal should be to eliminate the introduced rats and mice that plague the species .
illustrated on a 10 cent stamp and a calendar published by the state commercial bank .
safford rj and jones cg . did organochlorine pesticide use cause declines in mauritian forest birds ? biodiversity and conservation , 6 , 1445 - 1451 .
i thank particularly mrs . frédérique koenig , aviararies manager , who was a helpful , friendly and competent guide .
this small island is a great place to see this rare species , introduced here where there are no land predators , though i did regrettably see a house crow fly over .
i don ' t know which species makes the high pitched call ( for exemple : 00 : 18 ) . there was a female - like bird next to the male foudia rubra , so maybe a ( female ? ) call of that species as well .
i don ' t know which species makes the high pitched call ( for exemple : 00 : 10 ) . there was a female - like bird next to the male foudia rubra , so maybe a ( female ? ) call of that species as well .
sonogram images © xeno - canto foundation . sonogram images share the same license terms as the recording they depict .
overview , distribution , range and status . . . alternate ( global ) names
these rare weavers are in danger of becoming extinct . in 2011 , less than 170 individuals were believed to still have survived in the wild .
as part of conservation efforts , the mauritian wildlife foundation removed eggs from active nests prompting the parents to get started on a new clutch . those eggs that were taken were artificially incubated and the resulting chicks were handraised ; hence doubling the number of offspring per pair . thirty - two of these hand - raised birds were then released back into the wild between october 2010 and january 2011 . unfortunately , most or even all of them are believed to have perished .
their sharp declines are attributed to habitat loss and predation by introduced mammals ( black rats and macaque monkeys ) and the native round island boa ( snake ) .
breeding males : plumage olive - brown , except black lores ( area between the beak and the eyes ) ; red head , chest and rump patch ; two white wingbars ; and pale - brown streaked wings and tail . after the breeding season , the male molts into the plainer
winter plumage
( described below ) .
most breeding occurs from late june to early april . both parents weave the nest ; but only the female lines it . the average clutch consists of 2 - 4 eggs which are incubated by the female for about two weeks to hatching . the male feeds the female and the young .
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throughout history , crows , ravens and other black birds were feared as symbols of evil or death . …
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conservationists believe that between 1975 and 1993 the foudia ruba suffered a loss of 55 percent both in terms of population size and area of occupancy .
in 1994 its status on the iucn red list was categorised as critically endangered .
now due to massive conservation efforts its population is stable and its status downlisted to endangered .
a captive - rearing programme is implemented by the mauritian wildlife foundation , the gerald durrell endemic wildlife sanctuary and the national parks and conservation service .