the pencil fishes are a family ( lebiasinidae ) of freshwater fishes found in costa rica , panama , and south america .

they are usually small and are known as ornamental fishes in aquaria , including popular fishes such as the various pencil fish and the splashing tetra .

lebiasinids are small , cylindrical fish , ranging from 2 to 18 cm ( 0.8 to 7.1 in ) in adult length .

they prey on insect larvae , especially those of mosquitos .

the family includes the voladoras ( genera lebiasina and piabucina ) , mostly found in highlands of the north andes , the guiana shield and central america , but the other species are mainly lowland fish inhabiting the orinoco , amazon and paraguay river basins , and rivers of the guianas .

lebiasinidae

evolutionary relationships and cytotaxonomy considerations in the genus pyrrhulina ( characiformes , lebiasinidae ) .
günther ( family lebiasinidae ) . smithson contrib zool . 1975 ; 186 : 1–36 .
new subspecie from peru ( teleostei : lebiasinidae ) . verh ges ichthyol band 2 : 143–154
meinken , 1931 ( teleostei : characiformes : lebiasinidae ) . vertebrate zoology 3 : 283–298 .
evolutionary relationships and cytotaxonomy considerations in the genus pyrrhulina ( characiformes , lebiasinidae ) . - pubmed - ncbi
from the brazilian states of pará and amazonas ( teleostei : lebiasinidae ) . smithsonian contribution zoology 263 : 1–14 .
sp . n . ( pisces : teleostei : lebiasinidae ) . das aquarium . 1997 ; 31 : 12–17 .
sp . n . —ein neuer ziersalmler aus peru ( teleostei : characiformes : lebiasinidae ) . vertebrate zoology 59 : 11–23 .
spec . nov . —ein neuer ziersalmler aus venezuela ( teleostei : characiformes : lebiasinidae ) . vertebrate zoology 2 : 125–137 .
myers , 1956 des nordöstlichen südamerika ( teleostei : characiformes : lebiasinidae ) . vertebr zool . 2011 ; 61 : 13–45 .
( steindachner , 1876 ) einschließlich der beschreibung einer neuen art ( teleostei : characiformes : lebiasinidae ) . zool abh . 2006 ; 56 : 15–46 .
( characiformes , lebiasinidae ) reveals at least two evolutionary units in the rio negro basin , brazil . j fish biol 81 : 110–124 . pmid : 22747807
weitzman sh . osteology and relationships of south american characid fishes of subfamilies lebiasinidae and erythrinidae with special references to subtribe nannostomina . proc u s nat mus . 1964 ; 116 : 127–170 .
citation : marinho mmf , menezes na ( 2017 ) taxonomic review of copella ( characiformes : lebiasinidae ) with an identification key for the species . plos one 12 ( 8 ) : e0183069 . urltoken
netto - ferreira a . l . three new species of lebiasina ( characiformes : lebiasinidae ) from the brazilian shield border at serra do cachimbo , pará , brazil . neotrop ichthyol . 2012 ; 10 : 487–498 .
nannostomus grandis spec . nov . —ein neuer ziersalmler aus brasilien mit bemerkungen zu n . beckfordi günther , 1872 , n . anomalus steindachner , 1876 und n . aripirangensis meinken , 1931 ( teleostei : characiformes : lebiasinidae )
weitzman m , weitzman sh . family lebiasinidae . in : reis re , kullander so , ferraris cj jr . , editors . check list of the freshwater fishes of south and central america . porto alegre : edipucrs ; 2003 . pp . 241–251 .
citation : benzaquem dc , oliveira c , da silva batista j , zuanon j , porto jir ( 2015 ) dna barcoding in pencilfishes ( lebiasinidae : nannostomus ) reveals cryptic diversity across the brazilian amazon . plos one 10 ( 2 ) : e0112217 . urltoken
oyakawa ot , netto - ferreira al . família lebiasinidae . in : buckup pa , menezes na , ghazzi ma , editors . catálogo das espécies de peixes de água doce do brasil . rio de janeiro : museu nacional ; 2007 . pp . 64–65 .
family lebiasinidae ( pencil fishes ) lateral line and adipose fin usually absent . small to moderate - sized predators . south and central america . 7 genera , 61 species . family gasteropelecidae ( hatchetfishes ) deep , strongly compressed body ; pectoral fins with well - developed
sampling sites were chosen at distant points in the amazon basin in order to cover at a minimum , the distribution range of the species [ 33 ] ( fig . 1 ) . two lebiasinids were used as outgroups ( lebiasina colombia and copella nigrofasciata ) because they are members of each of the two subfamilies in lebiasinidae .
fishes of the neotropical family lebiasinidae occur in central america ( costa rica and panama ) and all south american countries , except chile [ 1 ] . the family includes 75 valid species [ 2 ] . distributed in two subfamilies , lebiasininae and pyrrhulininae , and seven genera : copeina fowler , copella myers , derhamia géry & zarske , lebiasina valenciennes , in cuvier & valenciennes , nannostomus günther , piabucina valenciennes , and pyrrhulina valenciennes [ 1 ] . members of the family can be recognized by having a rather elongate , cylindrical body , large scales , laterosensory canal of head and body reduced , anal fin short - based , no frontal or parietal fontanels [ 1 ] , and the absence of the metapterygoid - quadrate fenestra [ 3 ] .
diagnosis . copella can be easily distinguished from other genera of the lebiasinidae by having the anterior portion of the maxilla triple curved in males ( figs 1a and 2a ) ( vs . approximately straight or convex , fig 2a and 2b ) , and by males being distinctly longer than females ( vs . males and females of about the same size ) . additionally , copella is distinguished from other genera of the family , except nannostomus , by having anterior and posterior nares distant from each other ( fig 2 ) ( vs . juxtaposed , close to each other ) . it is distinguished from nannostomus by having a black spot on the dorsal fin ( vs . dorsal fin hyaline ) , elongate fins , especially on males ( vs . fins not elongate ) , and upturned mouth ( vs . terminal ) .
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urltoken
greek , lebias , - ou = a kind of fish ( ref . 45335 ) .
the list below must not be used as an authority reference synonymy list like those found in scientific published revisions , which must be the source to be used and cited eventually when they exist .
rather , it reflects the current content of fishbase , and the progress with respect to synchronization with the catalog of fishes . however , we think it can be useful for users to assess the quality of information in fishbase , to start new work on the family , or to cross - check with other lists .
but we appreciate to be cited in publications when this list has been of any working value . in particular , for published scientific , we suggest then to cite it in the material and method section as a useful tool to conduct the research , but again , not as a taxonomic or nomenclatural authority reference .
unless it is explicitly precised , the list is not complete , please search all original names published for the family in the catalog of fishes ( genera , species ) , including those with uncertain or unknown status , that are not included in fishbase when they are not attached to a valid species .
this list uses some data from catalog of fishes ( not shown but used to sort names ) .
in the column coff , the digit indicates the status of synchronization with coff : 0 : not checked ; 1 : same status ; 2 : different status ; 3 : other combination ; 4 : synonym in coff ; 5 : species / subspecies issue ; 6 : synonym of another species in coff ; 7 : not in coff ; 8 : should not be in coff . the coff version currently used is the one published on 23 - 07 - 2014 ( ref . 97102 ) .
when subfamilies are recognized , nominotypical subfamily first then other subfamilies by alphabetical order .
type genus of the family first ( or of subfamily when subfamilies are recognized ) then other genera by chronological order of description ( and alphabetical order ) .
type species of the genus first by chronological order ( and alphabetical order ) , with last listed misapplied names in a light gray font .
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 .
laboratório de genética animal , instituto nacional de pesquisas da amazônia , av . andré araújo , 2 . 936 , petrópolis , cep 69067 - 375 , manaus , am , brazil
laboratório de fisiologia comportamental e evolução , laboratório temático de biologia molecular , instituto nacional de pesquisas da amazônia , av . andré araújo , 2 . 936 , petrópolis , cep 69067 - 375 , manaus , am , brazil
laboratório de sistemática e ecologia de peixes , instituto nacional de pesquisas da amazônia , av . andré araújo , 2 . 936 , petrópolis , cep 69067 - 375 , manaus , am , brazil
) reveals cryptic diversity across the brazilian amazon . plos one 10 ( 4 ) : e0123363 .
copyright : © 2015 benzaquem 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 : this study was funded by conselho nacional de desenvolvimento científico e tecnológico ( cnpq ) ( rede brbol # 564953 / 2010 - 5 ) ; fapeam / cnpq / pronex ( call 003 / 2009 ) , cnpq / fapeam ( inct cenbam ) , cnpq / fapeam ( sisbiota rede biopham # 563348 / 2010 - 0 ) . fundação de amparo à pesquisa do estado do amazonas and coordenação de aperfeiçoamento de pessoal de nível superior ( capes ) provided phd fellowship to dcb . conselho nacional de desenvolvimento científico e tecnológico ( cnpq ) provided fellowship to co and jz . dna sequencing core facility was provided by instituto nacional de pesquisas da amazônia ( inpa ) and universidade estadual de são paulo ( unesp ) . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
neotropical ichthyofauna is extremely large and diverse based on the latest survey of the diversity of freshwater fishes of central and south america [ 1 ] , it includes 71 families encompassing 4 , 475 known valid species . moreover , the authors estimate there are 6 , 000 species in the neotropics , close to half the number of freshwater fish worldwide .
the amazon , home of more than two thousand freshwater fish species , is well known as a diversity hotspot . the complex evolutionary history of amazonian organisms , including fishes , is gradually beginning to be better understood . postulated reasons for the origin and evolution of neotropical fish richness have included tectonic movements and andes orogenesis leading to the rise and fall of biogeographic barriers , rates of speciation , extinction , and taxa dispersal , and the great antiquity of the amazonian ecosystem as a whole that dates to the early cenozoic and cretaceous periods [ 2 – 3 ] .
the identification of amazonian fish species is a challenging using only morphology as a tool , because fish taxonomists face difficulties daily in the process of identifying species . in this context , there are several debates and statements that molecular data should be encouraged as a supplement to species description and diagnosis , but should not replace morphological data [ 4 – 9 ] . indeed , species discriminations or descriptions involving the cytochrome c oxidase subunit i ( coi ) appear to be a trend in fish taxonomy [ 10 – 16 ] .
the barcode of life , referred to as dna barcoding , has recently emerged as a way to identify species by minimal dna sequences . while this approach offers all of the advantages of conventional pcr and dna sequencing , it also connects bioinformatics with the taxonomic identifications in museum and biological repositories , implement a modern and integrated system to ensure that newly reported cryptic species will be described following their discovery [ 17 – 18 ] . this global identification system , based primarily on nucleotide sequences , has been applied to several organisms , and many published works have demonstrated the efficacy of this methodology for species identification [ 19 – 27 ] .
by using dna barcoding , the identification of fish at the species level ranges from 98–100 % [ 18 , 28 – 29 ] and the discriminatory power of identification has been 93 % in freshwater fish and 98 % in marine fish species [ 30 ] .
this worldwide effort often reveals that species richness is still underestimated . as an indicator of cryptic speciation , at least two delimitations have been employed : the “10x rule” and the “cutoff value . ” in the first , barcoded specimens from the same nominal species diverge by 10 times or more from the mean intraspecific variability of the group . the “barcode gap” , that is , the divergence among organisms belonging to the same species being much smaller than the divergence among different organisms , has been clearly demonstrated in different several . in the second , individuals with a genetic distance of 2 % or more are considered much more likely to be congeneric than conspecific [ 30 ] .
despite its economical importance , little is known about biology or history of life of nannostomus species . however , in amazonian streams , they graze on algae on submerged trees and dead trunks , and feed on small invertebrates . it also provides food for animals at higher trophic levels developing an important role in the maintenance of the functioning and health of the whole aquatic system [ 38 ] .
popularly known as the pencilfish , nannostomus has 20 taxonomically valid species [ 31 ] . it is only genus to have undergone two taxonomic revisions , but other poorly defined species complexes probably remain in need of revision , including n . beckfordi günther , n . eques steindachner , n . marginatus eigenmann , and n . trifasciatus steindachner ( a . l . netto - ferreira , unpublished data ) . many of the species have conspicuous color variations depending on their geographic location . indeed , some of these color variants have been described as separate species in the past [ 39 – 42 ] .
recently , molecular analyses were conducted in two nannostomus species ( n . eques and n . unifasciatus ) , both of which are widely distributed throughout the amazon basin [ 43 – 44 ] . strikingly , the divergence time analyses of the lineages of both species allowed the authors to estimate an approximate divergence time in the middle pliocene for lineages in n . eques ( around 2 . 9 mya ) and in the late miocene for n . unifasciatus ( around 8 . 4 mya ) . the presence of distinct phylogroups in both species have been observed in representatives from the negro river basin , showing that due to hidden diversity , those species should not be treated as having a single stock for management purposes . such data may be useful in regulating the exploitation of fish species by the aquarium trade [ 43 – 44 ] .
given the biological and economic importance of these species , and as part of a study of the molecular phylogeny of nannostomus , we report herein an application of dna barcoding to the identification of 12 species of pencilfishes and highlight the deeply divergent clades detected within several nominal species , including the presence of additional cryptic species .
this survey was carried out in strict accordance with the recommendations by the national council for control of animal experimentation and the federal board of veterinary medicine . the protocol was approved by the committee on the ethics of animal use of the instituto nacional de pesquisas da amazônia ( 041 / 2012 ) .
all specimens for this study were collected in accordance with brazilian laws under a permanent scientific collection license issued in the name of dr . jorge i . r . porto and approved by the brazilian institute of environment and renewable natural resources ( ibama ) through the system authorization and information on biodiversity ( sisbio # 11489 - 1 ) .
we sampled 14 species of nannostomus ( n . beckfordi , n . britiskii , n . digrammus , n . eques , n . espei , n . harrisoni , n . limatus , n . margnatus , n . marylinae , n . morthenhaleri , n . nitidus , n . rubrocaudatus , n . trifasciatus and n . unifasciatus ) . after , each specimen was identified , it is anesthetized by immersion in eugenol in water and preserved in 96 % ethanol for the molecular studies . upon identification , morphological vouchers were deposited in the zoological collection at the national institute for amazonian research , inpa , manaus , brazil .
map of the study area showing the sampling sites the pencilfishes collected in this study .
all specimens were identified with the help of taxonomists and identification keys [ 34 , 37 ] end all procedures complied with the recommendations of local ethics committees . voucher specimens were deposited in the collection of the instituto nacional de pesquisas da amazônia , manaus , am , brazil .
genomic dna was isolated from muscle tissue based on standard techniques and protocols [ 45 ] . the partial mitochondrial coi gene ( 648pb ) was amplified by pcr using a combination of primers fishf1 , fishr1 , fishf2 fishr2 ) [ 20 ] or cock - tail c _ vf1lf _ t1 - c _ vr1lr _ t1 under conditions previously described [ 46 ] .
polymerase chain reactions were performed in a total volume of 15μl ( ∼10–50 ng dna template , 1x buffer ( 750 mm tris - hcl , ph 8 . 8 , 200 mm ( nh 4 ) 2 so 4 ) , 1u taq polymerase ( thermo scientific , waltham , usa ) , 0 . 2 mm dntps , 0 . 2 μm of each primer , 2 mm mgcl 2 , and ultrapure water . pcr cycling was performed with the initial denaturation for 2 min at 95°c followed by 35 cycles of 30s at 95ºc , 30s at 52ºc - 54ºc , 1 min at 72ºc and with a final extension for 10 min at 72ºc . pcr products were resolved on 1 % agarose gels and purified using polyethylene glycol 8000 ( usb , cleveland , usa ) . the bi - directional sequencing was performed utilizing an abi bigdye tm terminator v . 3 . 1 cycle sequencing ready reaction kit and an abi 3130xl dna analyzer ( applied biosystems , foster city , usa ) .
data sequences , collection sites , primers details and trace files were submitted to the barcode of life database ( bold ; http / / www . boldsystems . org ) in under project “barcoding of lebiasinids . ”
consensus sequences for the coi gene were generated using the bioedit program [ 47 ] , and after editing the sequences , the final matrix was 574bp .
all sequences were analyzed using mega 5 to check the occurrence of deletions , insertions , and stop codons . search tools with local alignment were used to identify the sequence in genbank and the bold . sequences were aligned using clustal w [ 48 ] , and the program dnasp version 4 . 0 [ 49 ] was used to determine the nucleotide composition , number of polymorphic sites , and haplotypes diversity .
the genetic distance among and within observed clusters was calculated using the kimura - 2 - parameter ( k2p ) model . a bayesian phylogenetic analysis was conducted using mrbayes 3 . 2 [ 50 ] . for this analysis , markov chain monte - carlo sampling was conducted every 120 , 000 generations until the standard deviation of split frequencies was < 0 . 01 . a burn - in period equivalent to 25 % of the total generations was necessary to recapitulate the parameter values and trees . the parameter values were evaluated based on 95 % credibility levels to ensure a sufficient number of generations had been run for the analysis . a neighbor - joining ( nj ) tree of k2p distances was created to provide a graphic representation of the relationships among specimens and clusters with mega 6 . 0 [ 51 ] . bootstrap resampling [ 52 ] was applied to assess the support for individual nodes using 1000 pseudo - replicates . the program haploviewer [ 53 ] was used to construct a tree - based haplotype network . independent networks were regarded as unconfirmed candidate species .
coi sequence data were obtained for 110 specimens of nannostomus representing 14 species - level taxa . a mean of 8 individuals ( range 1–17 ) represented each species , with only n . harrissoni represented by a single specimen .
the amplified product was approximately 650 bp but only 573 nucleotides were considered for this analysis , of which 234 were variable and parsimony informative . these variations defined 68 haplotypes ranging from 1–5 individuals per haplotype . at no time was a detected haplotype shared between different species . there were no deletions , insertions , or stop - codons . as expected for fish coi , the nucleotide composition showed a ct bias ( means c = 24 . 0 % , t = 32 . 7 % , a = 25 . 9 % , g = 17 . 3 % ) within nannostomus .
the results indicated that species could be discriminated by the dna barcode approach since the samples of distinct species were represented by a unique haplotype , a single tight cluster of haplotypes , or distinct clusters of haplotypes in neighbor - joining tree ( fig . 2 ) .
deep conspecific divergences shown by 4 species ( n . digrammus , n . trifasciatus , n . unifasciatus and n . eques ) are identified as distinct lineages . bootstrap values based on 1000 replicates are indicated at the branches .
the mean genetic distances ( kimura—2 - parameter distances ) among different nannostomus species ranged from 2 . 2 % to 22 . 5 % ( table 1 ) . the congeneric distance values found in nannostomus were high , with an overall mean of 19 . 0 % ± 1 . 3 % , with values > 22 % between n . britski and its congeners and 2 . 2 % between n . limatus and n . nitidus ( table 1 ) .
the conspecific distance showed less than 2 % divergence in 8 of the 14 species . exceptionally deep sequence divergences were evident between individuals identified as the same morphospecies of n . digrammus ( 9 . 80 % ± 1 . 0 % ) , n . trifasciatus ( 8 . 1 % ± 0 . 7 % ) , n . unifasciatus ( 7 . 1 % ± 0 . 7 % ) , and n . eques ( 4 . % ± 0 . 4 % ) . we constructed a haplotype network based on templeton’s method ( 95 % statistical parsimony ) [ 53 ] , and all four species displayed unconnected networks , which was consistent with the clusters identified through the nj tree . each of these lineages or phylogroups is presented in more detail below .
of the 16 specimens identified as n . eques , 13 unique haplotypes were identified . these haplotypes were assigned to five lineages ( e1—e5 ) . e1 ( n = 6 ) was sampled in rio tapajós , lower rio amazonas and lower rio negro , e2 ( n = 2 ) in upper rio amazonas and upper rio negro : e3 ( n = 5 ) in lower rio amazonas and upper rio negro , e4 ( n = 3 ) in rio guamá , and e5 ( n = 1 ) in rio madeira ( fig . 3a ) . low genetic variation was observed within the lineages ( e1 = 0 . 55 % , e3 = 0 . 26 % , e4 = 0 . 52 % ) , with the exception of e2 ( 2 . 29 % ) ( table 2 ) . the mean genetic distance between the lineages was 4 . 3 % . the smallest genetic distance encompassing different lineages was found between e2 and e3 ( 3 . 4 % ±0 . 7 % ) , while the largest was between e1 and e5 ( 7 . 7 % ± 1 . 1 % ) .
in the network each circle represents one haplotype , and theize of the circles is proportional to the haplotype frequency . color codes represent the lineages found in nannostomus . numbers on the lines represent the mutational steps between haplotypes .
of the 17 specimens identified as n . unifasciatus , 13 unique haplotypes were identified . these haplotypes were assigned to three lineages ( u1—u3 ) ; u1 ( n = 10 ) was sampled in rio purus , rio amazonas , and lower and middle rio negro . u2 ( n = 3 ) in lower and middle rio negro and u3 ( n = 4 ) in middle and upper rio negro ( fig . 3b ) . moderate genetic variation was observed within the u1 , u2 and u3 lineages ( 0 . 71 % , 0 . 57 % and 0 . 93 % , respectively ) . the mean genetic distance between the lineages was 7 . 1 % . the smallest genetic distance comprising different lineages was found between u2 and u3 ( 4 . 6 % ± 0 . 8 % ) , while the greatest was between u1 and u3 ( 12 . 5 % ± 1 . 4 % ) ( table 2 ) .
of the 7 specimens identified as n . digrammus , 6 unique haplotypes were identified . these haplotypes were assigned to lineages ( d1 and d2 ) . d1 ( n = 3 ) was sampled in the tapajós river , and amazonas and takutu river tributaries , while d2 ( n = 3 ) was endemic to the negro river ( fig . 3c ) . moderate genetic variation was seen within the d1 and d2 lineages ( 1 . 50 % and 0 . 80 % , respectively ) ( table 2 ) . the mean genetic distance between the lineages was 16 . 2 % ± 1 . 9 .
of the 12 specimens identified as n . trifasciatus , 9 unique haplotypes were identified . these haplotypes were assigned to 5 lineages ( t1—t5 ) . t1 ( n = 1 ) was sampled in rio preto da eva , t2 ( n = 1 ) in upper rio negro , t3 ( n = 1 ) in upper rio negro , t4 ( n = 3 ) in middle rio negro , and t5 ( n = 6 ) in tributaries of rio takutu , uatumã , and amazonas near macapá city ( fig . 3d ) . low genetic variation was observed within t4 and t5 ( 0 . 11 % and 0 . 33 % , respectively ) ( table 2 ) . the mean genetic distance between the lineages was 8 . 1 % . the smallest genetic distance encompassing different lineages was found between t4 and t5 ( 5 . 0 % ± 1 . 0 % ) , while the largest was between t2 and t4 ( 16 . 2 % ± 1 . 9 % ) .
molecular methodologies developed rapidly in recent years , and currently dna barcoding is considered the most useful tool for species identification . indeed , a great advantage offered by dna barcoding is the possibility of identifying cryptic species , as can be seen in several publications since its launch in 2003 [ 54 ] .
according to the fish barcode of life project database ( urltoken ) , only one nannostomus species had been previously barcoded . in our data set , 14 nannostomus species were dna barcoded , and they were easily identified by this approach , given that all recognized species formed monophyletic clusters ( fig . 2 ) . however , two species ( n . nitidus and n . limatus ) revealed shallow interspecific sequence divergence ( 2 . 2 % ) when compared to other nannostomus species ( table 1 ) . despite this , no evidence of shared sequences among both species were observed , suggesting either recent speciation or the need of synonymization .
in contrary , four species ( n . digrammus , n . trifasciatus , n . unifasciatus , and n . eques ) showed deep intraspecific sequence divergence , suggesting the existence of overlooked species within nannostomus , although some lineages were represented by a single individual ( table 2 ) .
the coi delineations of the t1 , t2 , t3 and e5 nannostomus lineages were achieved with only one individual . we are aware that hypothetical intraspecific lineages can be hindered by inadequate sample size in large geographic areas such as the amazon basin . in flathead fishes , for example , the limited number of specimens in a particular lineage and the sparse geographical spread of the samples for some of the proposed lineages restricted the ability to evaluate the extent of genetic diversity across several groups [ 55 ] . thus , we suggest future surveys of these nannostomus lineages to confirm if they do , in fact , represent evolutionary units .
all the lineages found in each of the above mentioned species were well supported by bootstrap values ( > 80 ) in the nj tree , independent of the mutational steps necessary to connect the haplotype networks below the standard statistical parsimony ( figs . 2 and 3 ) . the two lineages observed in n . digrammus ( d1 and d2 ) diverged by 12 . 6 % . the five lineages of n . trifasciatus had a mean divergence of 8 . 1 % , while that of the three lineages of n . unifasciatus was of 7 . 1 % . finally , the five lineages of n . eques diverged by 4 . 3 % ( mean ) . the divergence between the lineages of nannostomus was greater than obtained for other marine and freshwater fish species [ 20 , 56 – 61 ] .
it was evident in this study that the cutoff value of 2 % does not apply to nannostomus species , given the mean congeneric ( ∼19 % ) and conspecific ( ∼3 % ) distances were high . indeed , surveys on north american and neotropical freshwater ichthyofauna have shown that the mean congeneric and conspecific genetic distances are usually > 6 . 8 % and < 0 . 73 % , respectively [ 62 – 63 ] , with the exception of the pampa plain freshwater fishes at , the southernmost distribution range of many neotropical species , where the mean congeneric genetic distance is 1 . 67 % [ 64 ] .
among lebiasinids , only nannostomus has twice been taxonomically revised , in addition to the phylogenetic hypotheses [ 32 – 34 , 36 ] . morphology - based taxonomy has shown that nannostomus cannot be considered to have a phenotypic conservatism . apparently , there are poorly defined nannostomus species complexes based on morphological grounds , including n . beckfordi , n . eques , n marginatus , and n . trifasciatus ( a . l . netto - ferreira , unpublished data ) . recent mitochondrial and nuclear dna data have revealed hypothetical evolutionarily significant units in species of nannostomus . for example , in n . unifasciatus , the dna sequence data of the intron in the s7 ribosomal protein gene revealed two distinct lineages in the rio negro basin [ 43 ] . in n . eques , the mitochondrial dna control region also revealed the existence of two lineages in the rio negro basin [ 44 ] . together , the morphological and genetic studies indicate that species richness in the genus is probably underestimated .
the amazon aquatic ecosystem is rich in diversity and quick access to information on amazonian biodiversity is essential . forest degradation and , fishery over - exploitation enhance the risks of species extinction , and quickly updated information regarding the fishes caught in wild fisheries ( such as the nannostomus species ) is necessary to implement appropriate practices for to their conservation and , management and to prevent exploitation .
pencilfishes are commercial ornamental fish and constitute a source of revenue for the riverine people of the amazon . the discovery of cryptic species becomes very important when they are targets for commercial use . considering the economic importance of the nannostomus species , its dna barcoding contributes to conservation policy in two important ways : by enhancing amazonian biodiversity assessments to prioritize conservation areas ( e . g . , upper rio negro ) , and by providing information about evolutionary histories and phylogenetic diversity ( e . g . , unveiling hidden diversity ) .
dna barcoding of ornamental marine fishes has generated data and provided confidence in species identification , opening new avenues for managing business practices [ 65 ] . of the approximately ∼500 brazilian ornamental fishes allowed to be sold in the ornamental fish trade , 7 are pencilfishes : n . beckfordi , n . digrammus , n . eques , n . espei , n . marginatus , n . trifasciatus , and n . unifasciatus . our study contributes new amazonian fish barcodes , providing for a more comprehensive species identification of the ornamental pencilfishes , in addition to revealing the hidden diversity in the analyzed nannostomus species . it is certain that future species delimitation of nannostomus should be in accordance with the spirit of integrative taxonomy as well and based on congruence across analyses that utilize multiple data sources .
the authors are grateful to mark sabaj and ansp fish collection for the donation of tissue from n . harrisoni ; jorge mori marin of unesp for technical support ; andré luiz netto - ferreira of mpeg for help in identifying the fish species and support in the initial phase of the project ; douglas bastos for preparing the map ; and reinaldo peleja ( ufopa—tapajós river ) and , wilson laurentino ( guyana / lethem—upper takutu ) for making logistical and institutional arrangements for a portion of our field trip .
conceived and designed the experiments : dcb jirp . performed the experiments : dcb . analyzed the data : dcb co jsb jz jirp . contributed reagents / materials / analysis tools : dcb co jsb jz jirp . wrote the paper : dcb jirp co .
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de moraes rlr 1 , bertollo lac 1 , marinho mmf 2 , yano cf 1 , hatanaka t 1 , barby ff 1 , troy wp 3 , cioffi mb 1 .
1 departamento de genética e evolução , universidade federal de são carlos ( ufscar ) , são carlos , brazil .
2 museu de zoologia da universidade de são paulo ( mzusp ) , são paulo , brazil .
3 departamento de ciências biológicas , universidade do estado de mato grosso , campus de tangará da serra , tangará da serra , brazil .
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 .
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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 .
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horse , ( equus caballus ) , a hoofed , herbivorous mammal of the family equidae . it comprises a single species , …
dog , ( canis lupus familiaris ) , domestic mammal of the family canidae ( order carnivora ) . it is a subspecies…
animal , ( kingdom animalia ) , any of a group of multicellular eukaryotic organisms ( i . e . , as distinct from…
a taxonomic review of copella is presented based on the analysis of the type material of all nominal species and extensive material from south american drainages . six out of ten nominal species are recognized as valid : copella arnoldi , c . callolepis , c . compta , c . eigenmanni , c . nattereri , and c . vilmae . copella carsevennensis is a junior synonym of c . arnoldi , c . nigrofasciata and ‘nannostomus’ stigmasemion are junior synonyms of c . callolepis , c . metae is junior synonym of c . eigenmanni , and c . meinkeni is junior synonym of c . nattereri . species of copella occur in the rio amazonas and orinoco basins , and coastal drainages of guyana , french guiana , surinam , and venezuela . an identification key is provided .
copyright : © 2017 marinho , menezes . 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 .
data availability : all relevant data are within the paper and its supporting information files .
funding : authors are financially supported by fundação de amparo à pesquisa do estado de são paulo , urltoken , 2009 / 15075 - 0 ( mmfm ) and 2011 / 50282 - 7 ( nam ) . visits to ansp and usnm were partially supported by “böhlke travel award
( urltoken ) and smithsonian
short term visitor
( urltoken ) , respectively . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
the genus copella includes small species , reaching a maximum size of 52 . 0 mm sl . they are popular among aquarists due to their peaceful behavior and colorful bodies . the genus was erected by myers [ 4 ] based on the presence of a maxilla triply curved ( “s” shaped ) , more pronounced in males . in the same paper , myers included copeina compta myers , pyrrhulina nattereri steindachner , and copeina callolepis regan in copella and designated copella compta as the type species . although myers [ 4 ] stated that no other species of the “ pyrrhulina - copeina ” group could be identified as copella , géry [ 5 ] found that all the species previously recognized as copeina ( c . arnoldi regan , c . carsevennensis regan , c . eigenmanni regan , c . metae eigenmann , and c . nigrofasciata meinken ) should be transferred to copella , except copeina guttata ( steindachner ) and copeina osgoodi eigenmann . later on , géry [ 6 ] and zarske & géry [ 7 ] described copella vilmae and c . meinkeni , respectively .
the taxonomic problems related to the species of copella range from unclear diagnoses [ 5 ] , type material not represented by the same species [ 1 ] and literature misidentifications [ 7 ] . currently , the genus includes ten nominal species : copella arnoldi , c . carsevennensis , c . callolepis , c . compta , c . eigenmanni , c . metae , c . nattereri , c . nigrofasciata , c . vilmae , and c . meinkeni .
examination of a large amount of material of copella from the amazon and orinoco basins , and coastal drainages of brazil , guyana , french guiana , suriname , and venezuela , as well as type material of all nominal species , made possible the most - comprehensive taxonomic revision of the genus . a dichotomous identification key and distribution maps are also presented .
counts and measurements follow fink & weitzman [ 8 ] , with the addition of depth at dorsal - fin origin , pectoral to pelvic - fin origin , pelvic to anal - fin origin , and anal - fin base length , measured point to point . additional meristic data are the first longitudinal scale row on body , the fourth longitudinal scale row , which is the mid - lateral scale series including the first three small scales posterior to opercle , the longitudinal scale rows between dorsal - fin origin and pelvic - fin origin , and longitudinal scale rows between dorsal - fin origin and anal - fin origin . principal caudal - fin rays include all branched rays plus one unbranched ray in each lobe , following hubbs & lagler [ 9 ] and lundberg & baskin [ 10 ] . teeth and anterior unbranched anal - fin ray counts were taken from clear and stained material ( c & s ) , which was prepared according to taylor & van dyke [ 11 ] . count of maxillary teeth of c . vilmae was made on specimens preserved in alcohol by transparency , because insufficient material was available for c & s preparations . teeth were only counted in adult specimens , since juveniles have fewer teeth that are difficult to detect . meristics of the holotype of ‘ nannostomus’ stigmasemion were not taken due to the poor condition of the specimen .


lycomorpha pholus , the black-and-yellow lichen moth , is a moth in the family erebidae .

it is found in north america from nova scotia to north carolina , west to south dakota and texas .

the habitat consists of short-grass prairie .

the wingspan is 25 – 32 mm .

adults have bluish-black wings , with a yellow , orange or red basal area and black distal area .

the adults are thought to mimic poisonous lycid beetles , such as calopteron species .

the larvae feed on lichen and resemble their host .

lycomorpha pholus

species lycomorpha pholus - black - and - yellow lichen moth - hodges # 8087 - bugguide . net
dyar , h . g . 1897 . the larva of lycomorpha pholus . psyche 8 : 82 - 83
figure 15 . adult black and yellow lichen moth , lycomorpha pholus ( drury ) . photograph by tom murray .
figure 1 . adult black and yellow lichen moth , lycomorpha pholus ( drury ) . photograph by tom murray , university of florida .
figure 13 . adult black and yellow lichen moth , lycomorpha pholus ( drury ) ( pinned specimen ) . photograph by clare scott , university of florida .
lycomorpha pholus is a colorful tiger moth and can be found on flowers during the day throughout the summer . the forewing is 12 - to 15 - mm long .
central north american specimens are subspecies lycomorpha pholus miniata and show red on the wings ; eastern specimens of the nominate race have the red replaced by yellow ( paraphrase of gerald fauske , moths of north dakota ) .
lycomorpha pholus drury : arctiidae , lepidoptera ( this observation is from graenicher ; this is the black - and - yellow lichen moth ) asteraceae : solidago canadensis sn ( gr ) insect activities : sn = sucks nectar scientific observers : ( gr ) = s . graenicher
scott , c . 2010 . black and yellow lichen moth ( suggested common name ) lycomorpha pholus ( drury ) ( insecta : lepidoptera : noctuidae : arctiinae : lithosiini ) . university of florida eeny 479 : 1 - 5 , f . 1 - 17 ( pdf )
the black and yellow lichen moth , lycomorpha pholus ( drury ) , is a member of the subfamily arctiinae , tiger moths and woolly bears . the common names of the subfamily refer to the bright wing color patterns found in adults and the long setae covering many of its caterpillars .
yellow brazilian peppertree leaf - galler , calophya latiforceps burckhardt yellow fever mosquito , aedes aegypti ( linnaeus ) yellow flies , diachlorus spp . yellow fly , diachlorus ferrugatus ( fabricius ) yellowjackets and hornets , vespula and dolichovespula spp . yellow lichen moth , black and , lycomorpha pholus ( drury ) yellow mud dauber , black and , sceliphron caementarium ( drury , 1773 ) yellowmargined leaf beetle , microtheca ochroloma stål yellowstriped armyworm , spodoptera ornithogalli ( guenée ) yellowstriped oakworm , anisota peigleri riotte yellow sugarcane aphid , sipha flava ( forbes ) yucca giant - skipper , megathymus yuccae ( boisduval & leconte )
body and wings bluish black . tegulae and bases of forewing and hindwing yellowish orange .
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wingspan 25 - 32 mm ( scott , 2010 ) ; body length about 10 mm .
adult - body and wings bluish - black . basal portion of wings yellow , orange , or red ; distal portion of wings black . amount of yellow / orange / red varies , according to published photos and illustrations .
larva - gray dotted with pale green , sparse long hairs , resemble those of hypoprepia ( forbes , 1960 ; dyar , 1897 ) .
larvae feed on lichens . the strictly diurnal adults are often found on flowers of dogbane (
may take several years to develop , especially in the north . hairy cocoons are attached to rocks or tree trunks near the former food source .
the red color pattern is thought to be mimicry of poisonous lycid beetles , such as calopteron .
see also the presumed model of the mimicry , the lycid beetles of genus calopteron .
covell jr . , c . v . 1984 . a field guide to the moths of eastern north america . houghton mifflin company . p . 61 , plate 11 # 16
drury , d . 1773 . illustrations of natural history 3 : 49 , pl . 28 f . 3
powell , j . a . & p . a . opler 2009 . moths of western north america . university of california press . p . 266 , pl . 46 . 29m
distribution in canada list of provinces ( u . of alberta , using cbif data )
peterson field guides : eastern moths charles v . covell . 1984 . houghton mifflin company .
disclaimer : dedicated naturalists volunteer their time and resources here to provide this service . we strive to provide accurate information , but we are mostly just amateurs attempting to make sense of a diverse natural world . if you need expert professional advice , contact your local extension office .
contributors own the copyright to and are solely responsible for contributed content . click the contributor ' s name for licensing and usage information . everything else copyright © 2003 - 2018 iowa state university , unless otherwise noted .
photographs are the copyrighted property of each photographer listed . contact individual photographers for permission to use for any purpose .
powell , j . a . & p . a . opler , moths of western north america , pl . 46 . 29m ; p . 266 . book review and ordering
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figure 2 . the banded woolly bear caterpillar , pyrrharctia isabella ( smith ) . photograph by whitney crenshaw , colorado state university , urltoken .
tiger moths are found in all of the zoogeographical regions ( six geographical divisions of the world that are used to study the occurrence of land animals ) including the nearctic ( north america ) . however , they are most common in the neotropics ( mexico south of the tropic of cancer , central and south america , and the west indies ) ( scoble 1995 ) . species found in florida include the bella moth , utetheisa bella ( linnaeus ) ; the oleander moth , syntomeida epilais ( walker ) ; and the giant leopard moth , hypercompe scribonia ( stoll ) .
figure 3 . adult bella moth , utetheisa bella ( linnaeus ) , a tiger moth . photograph by don hall , entomology and nematology department , university of florida .
figure 4 . adult oleander moth , syntomeida epilais ( walker ) , a tiger moth . photograph by howard ensign evans , colorado state university , urltoken .
figure 5 . adult giant leopard moth , hypercompe scribonia ( stoll ) , a tiger moth . photograph by usda cooperative extension slide series , urltoken .
some caterpillars in the tiger moth subfamily are considered nuisance species , including the fall webworm , hyphantria cunea ( drury ) . others , such as the caterpillar of the cinnabar moth , tyria jacobaeae ( linnaeus ) , are beneficial . the cinnabar moth caterpillar is used as a successful biological control agent of tansy ragwort in the pacific northwest .
figure 6 . fall webworm larvae , hyphantria cunea ( drury ) , in protective webbing . photograph by g . keith douce , university of georgia , urltoken .
figure 7 . adult cinnabar moth , tyria jacobaeae ( linnaeus ) , nectar feeding on tansy ragwort . photograph by eric coombs , oregon department of agriculture , urltoken .
figure 8 . caterpillar of a cinnabar moth , tyria jacobaeae ( linnaeus ) , feeding on tansy ragwort . photograph by david cappaert , michigan state university , urltoken .
the black and yellow lichen moth is also a member of the tribe lithosiini ( subfamily arctiinae ) , the lichen moths . the common name of the tribe refers to the frequent choice of lichens as food sources by the caterpillars . like other tiger moths , the lichen moths are also most common in the neotropics . they are generally smaller than other tiger moths , and the adults may be either brightly colored or drab .
figure 9 . the painted lichen moth , hypoprepia fucosa hübner , an example of a brightly colored adult lichen moth . photograph by john himmelman , university of alberta .
figure 10 . adult little white lichen moth , clemensia albata packard , an example of a lichen moth with drab coloration . photograph by phil huntley - franck , university of alberta .
the caterpillars in lithosiini also have much fewer setae covering their bodies , making them appear bare in comparison to the woolly bears .
figure 11 . yellow woolly bear caterpillar , spilosoma virginica ( fabricius ) . photograph by david cappaert , michigan state university , urltoken .
figure 12 . lichen moth caterpillar of hypoprepia miniata ( kirby ) feeding on lichens . photograph by jason dombroskie , university of alberta .
the range of the black and yellow lichen moth is restricted to north america . it occurs as far north as ontario and quebec , canada . it is found in the eastern united states from maine to florida and as far west as the rocky mountains ( powell and opler 2009 ) .
adults : both male and female adult black and yellow lichen moths are similar in appearance . their wingspan ranges from 2 . 5 to 3 . 2 cm ( 1 to 1 . 3 inches ) ( covell 1984 ) . the head , thorax ( chest ) , legs , and abdomen are all black with an iridescent blue sheen . the tegula ( shoulders ) , the half of the forewing closest to the body ( basal half ) , and a patch on the front ( costal ) margin of the hind wing next to the body are all orange - yellow .
the orange - patched smoky moth , pyromorpha dimidiata herrich - schäffer , has a similar color pattern . this moth has a range that partially overlaps that of the black and yellow lichen moth . however , it only occurs as far west as missouri ( covell 1984 ) . both moths can be found flying at the same time of the year and both are diurnal ( active during the day ) . the orange - patched smoky moth can be distinguished from the black and yellow lichen moth because the former has no orange - yellow scales on its hind wing . also , its wings appear somewhat translucent , unlike the black and yellow lichen moth .
figure 14 . adult orange - patched smoky moth , pyromorpha dimidiata herrich - schäffer , with wings spread to demonstrate the lack of orange - yellow scales on the hindwing . photograph by jim vargo , moth photographers group .
larvae : the caterpillars of the black and yellow lichen moth are described as having sparse hairs and being similar in appearance to those of the genus hypoprepia ( forbes 1960 ) .
the black and yellow lichen moth flies from july to september in eastern north america ( covell 1984 ) and from late june to august in western north america ( powell and opler 2009 ) . this moth is only active during the day . it is found nectar feeding at flowers of goldenrod ( solidag o spp . ) , apocynum spp . , and ericameria nauseosa ( powell and opler 2009 ) . wagner et al . ( 2008 ) found that multiple generations can occur during the time when the adults are flying , and third instar larvae from later generations will overwinter by entering diapause . this is a period of no growth that can only be broken by receiving an appropriate number of cold hours .
the adults and caterpillars of the subfamily , arctiinae , are often defended from predators , including birds and bats , and from parasitoids , such as some flies and wasps , by chemical compounds . the compounds are often obtained from the hosts on which the caterpillars feed . the compounds are transferred to the adults when the caterpillar pupates . for the black and yellow lichen moth , these chemicals are lichen phenolics , defensive compounds produced by the fungal component of the lichen .
caterpillars of the black and yellow lichen moth have been raised to adults on the lichen physcia millegrana ( wagner et al . 2008 ) .
the color pattern found on adult black and yellow lichen moths occurs on other species in the tiger moth subfamily : members of the genera dycladia and correbia , the orange - patched smoky moth in the family zygaenidae , and in some net - wing beetles in the family lycidae . all of these individuals are regarded as müllerian mimics ( simmons 2009 ) , which means that all of them possess chemicals of some kind that make them distasteful to predators .
figure 16 . adult orange patched smoky moth , pyromorpha dimidiata herrich - schäffer . photograph by robert patterson , moth photographers group .
figure 17 . caenia dimidiata ( fabricius ) , an adult lycid beetle . photograph by tom murray , university of florida .
caterpillars and adult moths from the family zygaenidae , including the orange - patched smoky moth , are chemically defended by hydrocyanic acid that they release when threatened ( powell and opler 2009 ) .
conner we . ( editor ) 2009 . tiger moths and woolly bears : behavior , ecology , and evolution of the arctiidae . oxford university press . new york , new york . 303 pp .
covell cv . 1984 . a field guide to moths of eastern north america . special publication number 12 . virginia museum of natural history . martinsville , virginia . 496 pp .
forbes wtm . 1960 . lepidoptera of new york and neighboring states , part iv : agaristidae through nymphalidae including butterflies . cornell university agriculture experiment station memoir 371 . cornell university , ithaca , ny .
powell ja , opler pa . 2009 . moths of western north america . university of california press . berkeley , california . 369 pp .
scoble mj . 1995 . the lepidoptera form , function and diversity . the natural history museum . oxford university press . new york , new york . 404 pp .
simmons , r . 2009 . adaptive coloration and mimicry . in tiger moths and woolly bears : behavior , ecology , and evolution of the arctiidae . ( conner we , editor ) . oxford university press . new york , new york .
wagner dl , rota j , mccabe tl . 2008 . larva of abablemma ( noctuidae ) with notes on algivory and lichenivory in macrolepidoptera . annals of the entomological society of america 101 : 40 - 52 .
( retired ) , entomology and nematology department , university of florida . ; howard ensign evans and whitney crenshaw , colorado state university ; g . keith douce , university of georgia ; eric coombs , oregon department of agriculture ; david cappaert , michigan state university ;
publication date : july 2010 . latest revision : december 2013 . reviewed : december 2017 .
the basal half of theforewing and basal third of the hindwing is bright orange , with the outer portion of the wings black . this is a diurnal species , and quite unmistakable . wingspan of about 30mm .
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hodges , r . w . et al . , eds . 1983 . check list of the lepidoptera of america north of mexico . e . w . classey limited and the wedge entomological research foundation , london . 284 pp .
following lafontaine and schmidt ( 2010 ) , the traditional arctiidae have been transferred to the family erebidae as a subfamily ( arctiinae ) , with former subfamilies such as lithosiinae now treated as tribes . the circumscription of arctiinae remains virtually identical to recent circumscriptions of arctiidae , but circumscriptions of some taxa within the arctiinae have changed .
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distribution data for u . s . states and canadian provinces is known to be incomplete or has not been reviewed for this taxon .
a location where the species occurs , or has occurred , where there is potential fro persistence or regular recurrence . minimally a place where at least one specimen has been collected and verified by an expert in seemingly appropriate habitat . habitats are actually very poorly understood but seem to be xeric to mesic forest or woodland in most cases . exact habitat probaly would be best defined by the lichens upon which larvae feed if these are ever actually known .
it will sometimes be possible to map boundaries based on plant communities , e . g . occurrences for crambidia sp . 1 probably coincide with more open portions of pitch pine lowlands . occurrences for crambidia sp . 3 seem likely to coincide with xeric to ultraxeric oak or oak - pine woodlands in at least some cases .
if the occurrence seems to be linked to a specific type of woodland , consider all collections within this woodland as one occurrence regardless of distance .
these are small , weak flying moths so two kilometers should be more than enough to justify separate eos across obviously unsuitable habitats . however in practice it will often be difficult to determine whether wooded habitats are suitable or not and so usually in relatively uniform forested regions one should apply the suitable habitat distance .
given that it will seldom be possible to really define the local habitat and probably never possible to know the larval foodplant , and that some occurrences seem to be small it seems best to use a very conservative ie with this group . in most cases additional sampling will reveal a much larger occurrence but this cannot be assumed .
as far as known these specs can be used for any north american species , but reasonable judgments must be made as to likely suitable habitats . for the lichen feeders obviously areas with no lichens are not suitable . if these specs do not seem applicable consider customizing specs for the species involved .
use the generic guidelines for the application of occurrence ranks ( 2008 ) . the key for ranking species occurrences using the generic approach provides a step - wise process for implementing this method .
zoological data developed by natureserve and its network of natural heritage programs ( see local programs ) and other contributors and cooperators ( see sources ) .
lafontaine , j . d . and b . c . schmidt . 2010 . annotated check list of the noctuoidea ( insecta , lepidoptera ) of north america north of mexico . zookeys 40 : 1 - 239 .
lafontaine , j . d . and m . fibiger . 2006 . revised higher classification of the noctuoidea ( lepidoptera ) . canadian entomologist 138 : 610 - 635 .
schmidt , b . c . and p . a . opler . 2008 . revised checklist of the tiger moths of the continental united states and canada . zootaxa 1677 : 1 - 23 .
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ridgely , r . s . , t . f . allnutt , t . brooks , d . k . mcnicol , d . w . mehlman , b . e . young , and j . r . zook . 2003 . digital distribution maps of the birds of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
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photograph : © 2008 rkd peterson date of photograph : 19 august 2008 location : montana , gallatin co .
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quebec west to extreme southern alberta , south to texas . the only alberta record is from milk river ( canadian national collection of insects , ottawa ) .
hilty , j . editor . 2015 . insect visitors of illinois wildflowers . world wide web electronic publication . illinoiswildflowers . info , version ( 09 / 2015 ) see : abbreviations for insect activities , abbreviations for scientific observers , references for behavioral observations
the wingspan is 25–32 mm . adults have bluish - black wings , with a yellow , orange or red basal area and black distal area . adults are on wing from july to september . they fly during the day and are attracted to flowers , including goldenrod . the adults are thought to mimicry poisonous lycid beetles , such as calopteron species .
and resemble their host . the species may take several years to develop , depending on the location . pupation takes place in hairy cocoons attached to rocks or tree trunks near the host .
comments : following lafontaine and schmidt ( 2010 ) , the traditional arctiidae have been transferred to the family erebidae as a subfamily ( arctiinae ) , with former subfamilies such as lithosiinae now treated as tribes . the circumscription of arctiinae remains virtually identical to recent circumscriptions of arctiidae , but circumscriptions of some taxa within the arctiinae have changed .
imperial moth , eacles imperialis imperialis ( drury , 1773 ) imported cabbageworm , pieris rapae ( linnaeus ) imported fire ant , red , solenopsis invicta buren indianmeal moth , plodia interpunctella ( hübner ) insidious flower bug , orius insidiosus say io moth , automeris io ( fabricius ) ips engraver beetles , ips spp .
jadera bugs , jadera spp . jamaican field cricket , gryllus assimilis ( fabricius ) japanese beetle , popillia japonica newman japanese orange fly , bactrocera tsuneonis ( miyake ) japanese weevil , twobanded , pseudocneorhinus bifasciatus roelofs jewelwing , ebony , calopteryx maculata ( beauvois , 1807 ) jumping gall wasp , neuroterus saltatorius edwards jumping spider , dimorphic jumper , maevia inclemens ( walckenaer , 1837 ) jumping spider , gray wall , menemerus bivittatus ( dufour ) jumping spider , pantropical , plexippus paykulli ( audouin ) jumping spider , regal , phidippus regius c . l . koch jumping spiders , habronattus , habronattus ( pickard - cambridge )
katydid , broad - tipped conehead , neoconocephalus triops ( l . ) katydid , oblong - winged , amblycorypha oblongifolia ( de geer ) khapra beetle , trogoderma granarium everts ked , deer , lipoptena mazamae rondani kermes scale , northern red - oak , allokermes kingii ( cockerell ) killers , bee , mallophora spp . killer , cicada , sphecius hogardii ( latreille ) killer , cicada , sphecius speciosus ( drury )
nantucket pine tip moth , rhyacionia frustrana ( comstock ) native subterranean termite , reticulitermes hageni banks native subterranean termite , reticulitermes virginicus ( banks ) nematode , burrowing , radopholus similis ( cobb , 1893 ) thorne , 1949 nematode , citrus , tylenchulus semipenetrans ( cobb ) nematode , grasshopper , mermis nigrescens dujardin nematode , lance , hoplolaimus galeatus ( cobb ) thorne nematode , amaryllis lesion , pratylenchus hippeastri inserra et al . , 2006 nematode , mole cricket , steinernema scapterisci nguyen & smart nematode , red ring , bursaphelenchus cocophilus ( cobb ) baujard nematode , reniform . rotylenchulus reniformis linford & oliveira nematode , spiral , helicotylenchus spp . nematode , sting , belonolaimus longicaudatus rau nematode , stubby - root , nanidorus minor ( colbran ) nematode , stubby - root , trichodorus obtusus cobb nematodes , awl , dolichodorus spp . nematodes , dagger , xiphinema spp . nematodes , entomopathogenic families steinernematidae and heterorhabditidae nematodes , free - living marine , nematoda nematodes , soil - inhabiting , nematoda nematodes of thrips , entomopathogenic , thripinema spp . net - winged beetle , banded , calopteron discrepans ( newman ) northern red - oak kermes scale , allokermes kingii ( cockerell ) no - see - ums , culicoides spp .
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the arizona state university hasbrouck insect collection ( asuhic ) is one of the vital southwest arthropod collections in america north of mexico , providing important biological information . the principal objective of the catalog is to give a complete list of the lepidopterous insects held in the asuhic . furthermore , it will be an online catalog of the lepidoptera of arizona . the preliminary lepidoptera checklist is presented , consisting of 1983 species and 175 subspecies of 55 families in approximately 60 , 000 holdings at the asuhic . this article follows the recent classification and nomenclature ( hodges rw . 1983 . check list of the lepidoptera of america north of mexico . london , uk : e . w . classey ltd . and the wedge entomological research foundation ; moth photographers group ( mpg ) . 2014 . urltoken ) .
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copyright © 2014 national science museum of korea ( nsmk ) and korea national arboretum ( kna ) . production and hosting by elsevier b . v .
copyright © 2018 elsevier b . v . or its licensors or contributors . sciencedirect ® is a registered trademark of elsevier b . v .
to 2 inches ] ) with the body as if standing at attention . although most are drab browns and grays , some species are colourful . the scarlet lichen
) has black wings with yellow markings . the larvae of the footman moth have long , sparse hairs arising from wartlike tubercles and feed on lichen .
tiger moth , , ( family arctiidae ) , any of about 11 , 000 species of moths ( order lepidoptera ) , the common name of which is derived from that of one of its most common genera , grammia , which have dark wings with red or orange spots and white stripes , sometimes displayed in striking geometric patterns . …
lepidopteran , ( order lepidoptera ) , any of about 180 , 000 species of butterflies , moths , and skippers . this order of insects is second in size only to coleoptera , the beetles . because of their day - flying habits and bright colours , the butterflies are more familiar than the chiefly night - flying and…
moth , ( order lepidoptera ) , any of about 160 , 000 species of overwhelmingly nocturnal flying insects that , along with the butterflies and skippers , constitute the order lepidoptera . moths vary greatly in size , ranging in wingspan from about 4 mm ( 0 . 16 inch ) to nearly 30 cm ( about 1 foot ) . highly…
insect , ( class insecta or hexapoda ) , any member of the largest class of the phylum arthropoda , which is itself the largest of the animal phyla . insects have segmented bodies , jointed legs , and external skeletons ( exoskeletons ) . insects are distinguished from other arthropods by their body , which is…
arthropod , any member of the phylum arthropoda , the largest phylum in the animal kingdom , which includes such familiar forms as lobsters , crabs , spiders , mites , insects , centipedes , and millipedes . about 84 percent of all known species of animals are members of this phylum . arthropods are…
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5’ tall . very prickly leaves . leaves not wooly underneath . 2” tall flower heads .
just as i left the house , i noticed a small butterfly nectaring on the common milkweed along the driveway . at first , i took it for one of the ubiquitous little glassywings , but when i took a second look , i realized the shape was all wrong . closer inspection revealed that it was a banded hairstreak , a small , woodland species that i had never seen in the yard before ! the caterpillars of this species feed on the leaves of various oak and hickory species , both of which are common in the woods around the yard .
along the edge of the woods , i came across this tiny american toad crossing the path . this time of year , numerous toads and frogs can be found hopping through the woods as they leave the ponds and pools in which they were born .
in a meadow at the top of the property , i noticed an interesting insect sitting on a dried grass blade . this insect turned out to be this beautiful black - and - yellow lichen moth (
as i continued along the path , i found a small yellow and black beetle sitting on a gray birch leaf . this little beetle is a fourteen - spotted lady beetle (
as i reached the end of the property , i turned over a few rocks to look for salamanders , spiders , or whatever i could find . under one of the larger rocks , i found this northern ringneck snake . this is a fairly common species in the region , but one that is infrequently encountered because it hides under rocks and fallen logs .
beetles ( order coleoptera ) are another group of insects that i am just beginning to learn . with over 350 , 000 described species in the world , beetles make up an incredibly large portion ( about 40 % ) of the insect species on the planet . in north america alone , there are about 24 , 000 known species . this huge diversity creates an incredible identification challenge .
however , some beetles are relatively simple to identify and can tell us a lot about the environment in which they live . here are a few examples of beetles i have found recently :
this first beetle belongs to one of the most well - recognized groups of beetles , although many people would not know it from a view like this , but rather from the bright yellow flashes it emits on warm summer nights . this beetle in the genus
belongs to a group of beetles known as lepturinae , the flower longhorns . members of this subfamily often have skinny bodies , long antennae , and ( as the name suggests ) they are often associated with flowers . species in this group of beetles become extremely common as summer flowers begin blooming . i found this particular beetle on the flower of a swamp rose (
) that was filled with various beetles , bees , flies , and wasps .
is commonly found on flowers . every year , i begin seeing these when the wild hydrangea starts blooming . in fact , i found this individual on a flower head of a wild hydrangea . it is thought that the appearance and flight of this genus mimic that of bees in the genus
, the invasive japanese beetle . this scarab beetle causes severe damage as it eats the leaves of anything it can find . the japanese beetle is a true generalist , so it will eat the leaves , flowers , and fruits of many shrubs and perennials ( native and non - native ) , but it seems to particularily enjoy plants in the rose family .
this gorgeous , metallic beetle is often confused with the previous species , but is shinier and more colorful . unlike the japanese beetle , the dogbane beetle (
) is a native species that is found on plants in the dogbane family . i found this one on indian - hemp (
this final beetle was one of the more excitings finds of the day . while in the field during a late afternoon walk with the dog , my brother spotted a large beetle on a flower of timothy grass (
most people have seen a light during a warm , summer night with numerous moths flying around it . the reason for this is often thought to be related to the idea that moths use the moon as a point of reference while flying around or migrating . to moths and other insects , a bright artificial light may look like the moon , so they use that as a guideline rather than the real moon . as a result , the moths get disoriented and flutter around the light .
this disorientation and movement towards the light ( not technically
attraction
) allows close observation of many insects species that would be difficult to find otherwise . for the past week or so , i have been going out to the porch light after dark and checking to see what shows up . over the course of about one week , i have recorded about 60 species of moths near the light , as well as numerous species of other insects including mayflies , wasps , beetles , and stink bugs .
) . this species is easily recognized by the two light - colored bands that break up the dark areas of the wing .
) is a fairly plain species . this is a very light - colored individual .
moths come in many shapes , sizes , and colors . this grape leaf - folder moth (
sp . ) is a relatively small moth , with a wingspan of about 2 cm and a distinctive shape and pattern .
, is one of many species of moths that i had never encountered before .
) . the beautiful patterns and colors on this moth made it stand out from many of the brown and gray species at the light .
this leuconycta diphteroides is rather dull , but if you look carefully , the wings have very interesting patterns .
. this one is much brighter and more colorful , so the patterns really stand out .
most of the previously mentioned moths are relatively large and easy to see as they sit on the edge of the house . however , some moths require a bit of searching because their small size makes them difficult to find . one example of such a moth is this diamondback moth (
has a wingspan of about one centimeter . although hard to imagine , this species is a well - documented transoceanic migrant !
) is another small moth that blended in with the wall on which it was perched . this species has distinct
ridges
on the top of its wings .
most common small moth at the porch light this time of year . this species is also commonly found in the woods and meadows near the house .
plenty of non - moth insects come to the light as well . this stinkbug ,
, is a species that i have only ever seen around lights at night .
is only one of several wasps that i see at the porch light each evening .
mayflies , like this one , are frequently found at the porch light . a few weeks ago , during the bioblitz at jacobsburg state park , lots of mayflies were found on the illuminated walls of the park office .
this saturday , my mom , brother , and i traveled to the area around ricketts glen state park . our first stop was state game lands # 57 , which is located directly north of the state park . upon arriving , we parked in a gravel lot situated next to this rocky stream . the stream was bordered with plants such as meadowsweet and silky dogwood .
the muddy parking lot was a perfect location for
puddling
. puddling is a behavior exhibited by many butterflies in order to collect minerals . certain butterflies , like this red - spotted purple , are commonly found at puddles and muddy areas .
we found this white admiral in the same parking lot . this white admiral actually belongs to the same species as the previous butterfly . the two subspecies are different in appearance , but are almost identical in structure . the white admiral is the more northern subspecies of the species .
from the parking lot , we walked along a grassy road that meanders along the creek . this partially wooded area was filled with dragonflies including this female american emerald .
chalk - fronted corporals were another extremely common dragonfly along this road . over the course of a few hours , we saw over one hundred of these skittish insects .
after a little while , the road became muddy , and we found ourselves in a sphagnum bog . the waterlogged
moss provides an excellent place for certain unusual plants to grow . my mom spotted this round - leaved sundew , a carnivorous plant , growing along the edge of the trail . sundew plants trap prey in the sticky
dew
on the leaves . the plant then secretes enzymes that dissolve the insect and allow the plant to receive essential nutrients .
the bog , which is very similar to habitats found farther north , is home to
northern
butterflies like this common ringlet .
leatherleaf is another plant that is common in the bog on state game lands # 57 .
one of the most well - known bog plants is the cranberry . at the time we explored the bog , the flowers were just beginning to open .
the bog and the road , we walked along a section of forest that was recently logged . the logging created perfect habitat for many bird species . while hiking through this area , we heard several least flycatchers , yellow - rumped warblers , and white - throated sparrows .
the logged area also opened up habitat for unusual dragonfly species . i photographed this maine snaketail after it caught a small insect , and then perched to eat .
after exploring the game lands , we headed into the state park . our first stop was at
the hayfields ,
a large expanse of grasses and highbush blueberries . from the top of the hill , we saw a large swath of purple flowers in the wetland below .
as we walked down to the wet area , we realized that the purple flowers were blue flag irises .
from the hayfields we travelled to the main attraction of ricketts glen , the waterfalls . along the wooded trail , we found numerous lichen - covered trees and moss - covered rocks .
the damp , rocky forest provided perfect habitat for plants such as this hobblebush .
the woods at ricketts glen are a wonderful place for wildflowers . we found several spring flowers that were going to seed . trilliums were some of the more common wildflowers along with starflower , indian cucumber - root , and northern wood - sorrel .
after walking for a while , we finally reached the first waterfall . the sight of the falling water , green hobblebush , and textured bark of the hemlocks was a splendid sight !
the falls trail at ricketts glen travels down along one creek then climbs up along another providing great views of over twenty waterfalls .
while hiking along the stream , we found this mountain maple growing along the stream .
unlike many other species in the genus , mountain maple has flowers that grown on a cluster .
the most abundant wildflower along the waterfalls was northern wood - sorrel , which has flowers ranging from white to pink .
i was born and raised in eastern pennsylvania , where i explored the outdoors and fell in love with traditional music from an early age . i am currently studying chemistry at harvard university .