pancrustacea is a clade , comprising all crustaceans and hexapods .

this grouping is contrary to the atelocerata hypothesis , in which myriapoda and hexapoda are sister taxa , and crustacea are only more distantly related .

as of 2010 , the pancrustacea taxon is considered well accepted .

the clade has also been called tetraconata , referring to the square ommatidia of many of its members .

that name is preferred by some scientists as a means of avoiding confusion with the use of " pan - " to indicate a clade that includes a crown group and all of its stem group representatives .

pancrustacea

linking insects with crustacea : physiology of the pancrustacea : an introduction to the symposium .
morphological , molecular , and hormonal basis of limb regeneration across pancrustacea . - pubmed - ncbi
show trees collapsed to the group level ( pancrustacea ) and order level ( copepoda ) .
phylogenetic relationship of pancrustacea . phylogenetic relationship of 510 species of pancrustacea ( collapsed to major taxa level ) based on 18s rrna sequences . numbers show the posterior probabilities calculated by bayesian analysis .
ebscohost | 110372275 | linking insects with crustacea : physiology of the pancrustacea : an introduction to the symposium .
is the most recent comprehensive analysis on ostracode phylogenetics , and places the ostracoda in the oligostraca clade of pancrustacea .
reconstructed with dna sequence data from three nuclear protein coding genes show that ostracoda are at the base of pancrustacea and might be polyphyletic .
given that pancrustacea and arachnomorpha were ancestrally marine , we would expect the earliest myriapods to be marine also , however definite marine myriapods elude us .
) . the basal divergences within pancrustacea , chelicerata and myriapoda occurred no later than 530 , 578 and 544 myr ago , respectively , and possibly several hundred million years earlier . within pancrustacea , the six traditional crustacean classes had all appeared by 388 myr ago and no earlier than 522 myr ago .
we were able to reliably place several fossils within the pancrustacea , which can be used for calibration points in divergence time studies ( table 5 ) .
the authors would like to thank the speakers in this symposium for reaching outside of either their crustacean or insect expertise to consider evolutionary physiology of the pancrustacea .
+ remipedia appear to be a basal lineage of pancrustacea . the present study also provides considerable data for the application of both population and phylogenetic studies of other crab species .
pancrustacea ( equivalent to
tetraconata
of some authors ) contains all traditional crustaceans and hexapods . work of the last decade has produced the growing consensus that
crustacea
is paraphyetic with respect to hexapoda , but there is no clear consensus of exactly where , within pancrustacea , hexapods go . for now , we follow the morphology - based analysis of
, and these two taxa together are referred to as the ichthyostraca which , together with the mystacocarida and ostracoda , forms the oligostracan branch of the pancrustacea ( e . g .
) either place ostracoda and branchiura as a paraphyletic grade at the base of pancrustacea or else move ostracoda : podocopa to the base of clade 24 , keeping ostracoda : myodocopa + branchiura at the base of pancrustacea . thus , our data consistently indicate that ostracoda and branchiura belong at or near the base of pancrustacea , although their precise phylogenetic positions remain uncertain . we also note that ostracoda is invariably , but inconsistently , split by branchiura , i . e . branchiura is grouped with ostracoda : myodocopa by nucleotides and ostracoda : podocopa by amino acids ( see
the pancrustacea clade itself is composed of several clades , but where the ostracodes fit within these groups has been the subject of considerable research . there has not been much agreement regarding monophyly of the ostracoda [ see
as sister group to the branchiura , and these two taxa together are referred to as the ichthyostraca which , together with the mystacocarida and ostracoda , forms the oligostracan branch of the pancrustacea ( e . g .
online ) , whereas the wag + γ model always failed on that case . this shows that when the phylogenetic question at hand is difficult , such as the short branch at the base of pancrustacea + myriapoda (
sherry l . tamone , jon f . harrison ; linking insects with crustacea : physiology of the pancrustacea : an introduction to the symposium , integrative and comparative biology , volume 55 , issue 5 , 1 november 2015 , pages 765–770 , urltoken
, involved soft tissue characters of the brain and the shape of ommatidia , and exceed the technical limits of this course . indeed , the whole topic is so complex that we can only scratch the surface . beyond possessing the synapomorphies of mandibulata , pancrustacea displays :
for several reasons . first , they represent each of the two major arthropod clades outside of pancrustacea . second , they have much data represented in our ingroup taxa . third , they are relatively short - branch taxa in previous studies ( e . g . ,
- relationships within the pancrustacea : examining the influence of additional malacostracan 18s and 28s rdna - in “crustacea and arthropod relationships” ( koenemann , s . & r . a . jenner , eds . ) taylor & francis , davis , ca . pp . 275 - 294 (
( see the 12 nodes with filled half - circles and also the time - scale below ) . according to these results ( and at the level of two standard deviations ) , pancrustacea separated from both chelicerata and myriapoda no later than 601 myr ago ( in broad agreement with
citation : ribeiro l , tobias - santos v , santos d , antunes f , feltran g , de souza menezes j , et al . ( 2017 ) evolution and multiple roles of the pancrustacea specific transcription factor zelda in insects . plos genet 13 ( 7 ) : e1006868 . urltoken
it turns out that , according to this analysis , the xenocarids are the long sought - after sister group to hexapods . together they form the new clade miracrustacea ( ‘surprising crustacean’ ) . the miracrustaceans are sister to the vericrustaceans , forming a clade that is in turn sister to the oligostraca , rounding out pancrustacea .
this tree further supports a pancrustacean clade consisting of hexapods and crustaceans . in addition , this work supports the mandibulata model of deep evolutionary relationships within arthropoda . pancrustacea is sister to the myriapods , forming the mandibulata clade ; which is then sister to the early - branching chelicerates . this reconstruction strongly refutes the mitochondrial paradoxopoda model .
we find good support for three major pancrustacean clades : a ) oligostraca ( ostracoda , mystacocarida , branchiura , and pentastomida ) , which forms the sister group of the rest of pancrustacea ; b ) multicrustacea ( malacostraca , cirripedia , and copepoda ) ; and c ) allotriocarida ( branchiopoda , remipedia , cephalocarida , and hexapoda ) .
for the full tree ) . malacostraca + ( thecostraca + tantulocarida ) is the sister - group of copepoda . the present tree shows that copepoda is a monophyletic group , with platycopioida ( progymnoplea ) as its most basal clade , followed by calanoida ( gymnoplea ) and podoplea , both also monophyletic . this phylogeny of pancrustacea agrees with regier
2010 ) . this grouping is primarily supported by molecular data ; the synapomorphies given below have all become modified in various subclades . relationships between clades within the pancrustacea remain debated ; nevertheless , certain relationships have been supported by recent molecular analyses . the clade oligostraca unites the bivalved ostracoda with a clade of parasites of vertebrates in the ichthyostraca ; schwenter
broadly , crustaceans and insects can be thought of as aquatic and terrestrial branches of a phylogenetic tree , and thus represent an outstanding clade for understanding structural and physiological aspects to the adaption to a terrestrial lifestyle . throughout this symposium , we sought to use comparative physiology as a topic for the discussion of physiological adaptation of the pancrustacea to their respective habitats . while we could not possibly cover all aspects of comparative physiology of the pancrustacea in a single symposium , the articles contained in this issue represent systems - level investigations into exoskeletons , gas exchange , cardiovascular systems , immunology , excretion , vision , endocrines , and transcriptional responses to stress . here we will highlight some examples of the comparative physiology discussed in this symposium .
the phylum arthropoda consists of four major groups : chelicerata , crustacea , myriapoda , and hexapoda . recent molecular analyses have greatly changed our traditional understanding of arthropod phylogeny and evolution . these studies have rejected the traditional view that the closest relatives to hexapods are myriapods , and instead indicate that hexapods and crustaceans form a common clade , which is now called pancrustacea [
) had little effect on the arrangement of the pancrustacean clades , but the bootstrap values of some nodes increased greatly ( for diplura from 66 % to 91 % , for insecta from 36 % to 97 % ) . in addition , we got similar tree topologies for pancrustacea when removing the onychophoran and myriapods , and using chelicerates as the outgroup ( data not shown ) .
is copepoda ( including platycopioida ) a monophyletic group within pancrustacea ? to answer this question , the position of copepoda was interrogated using 18s rrna gene sequences of 305 arthropoda species available from ncbi in addition to our own data from 205 copepod species ; together these data include chelicerata , myriapoda , hexapoda , pentastomida , ostracoda , branchiura , branchiopoda , remipedia , cephalocarida , copepoda , malacostraca , thecostraca and tantulocarida .
the evolutionary success of hexapods is attributed to a combination of features : their segmented body plan and jointed appendages , which were inherited from their arthropod ancestor and ; wings and holometaboly , two features that arose later in insect evolution [ 67 ] . it is interesting to note that the specific pancrustacea gene zld is required for most of these processes , such as embryonic segment formation , wing ( elytra ) patterning , and appendage ( antennae and leg ) formation during beetle development . like several other zinc finger proteins that show tremendous lineage - specific diversity in eukaryotes , zld appears to have specifically arisen within pancrustacea and risen to play an important role as a “master tf” . hence , future studies focusing on how this tf was integrated to the conserved backdrop of developmental tfs and existing grns of arthropods would be of great interest .
the monophyly of copepoda ( including the platycopioida ) within pancrustacea is demonstrated for the first time . previous molecular analyses considered only a few species ( mostly podopleans ) as representatives of copepoda . mallat et al . 47 considered a single taxon , identified no further than “cyclopidae sp . ” ( cyclopoida ) as a representative for copepoda in their ecdysozoan phylogeny , based on 18s and 28s rrna genes . regier et al . 45 , 48 , 49 explored the phylogenetic relationships of arthropoda using 41 kb of nuclear protein - coding genes , but included only three copepod species , namely one derived calanoida and two fresh - water cyclopoida in the analysis . von reumont et al . 50 , presented an analysis of pancrustacea using 454 expressed sequence tags ( est ) from 1 , 886 genes , but included just one harpacticoida species and four species of siphonostomatoida in the analysis .
the sister group relationships of the arachnida are quite well understood . this group includes all the terrestrial chelicerates and has two extant successively more distant marine sister taxa : xiphosura ( horseshoe crabs ) and pycnogonida ( sea spiders ) [ 23 , 36 , 37 ] . in contrast , the exact relationships of the hexapoda within pancrustacea are still unclear , and it is not obvious whether their sister taxon was a marine - , brackish - or freshwater - adapted organism .
within the pancrustacea , hcs first were described only in the malacostracan crustaceans , which include shrimps , lobsters , crabs , woodlice , and related species ( mangum 1983 ; markl 1986 ) . only later did it become obvious that this respiratory protein also occurs in other crustacean classes and in the hexapods ( hagner - holler et al . 2004 ; ertas et al . 2009 ; pick et al . 2009 ; marxen et al . 2014 ; see below ) .
two bivalved arthropod groups have in the past been allied with ostracoda . first , we included two bradoriids , which consistently placed outside pancrustacea in both our concatenated fossil analysis and our site - weighted fossil placement . second , phosphatocopina are bivalved arthropods that were once considered a group of ostracoda until the discovery of soft parts showed major differences , notably the undifferentiated fourth and fifth cephalic appendage ( maxillae in all extant ostracods ) . in our phylogenetic analyses , these species (
recent phylogenetic analyses using molecular data suggest that hexapods are more closely related to crustaceans than to myriapods , a result that conflicts with long–held morphology–based hypotheses . here we contribute additional information to this debate by conducting phylogenetic analyses on two nuclear protein–encoding genes , elongation factor–1α ( ef–1α ) and the largest subunit of rna polymerase ii ( pol ii ) , from an extensive sample of arthropod taxa . results were obtained from two data sets . one data set comprised 1092 nucleotides ( 364 amino acids ) of ef–1α and 372 nucleotides ( 124 amino acids ) of pol ii from 30 arthropods and three lobopods . the other data set contained the same ef––lα fragment and an expanded 1038–nucleotide ( 346–amino–acid ) sample of pol ii from 17 arthropod taxa . results from maximum–parsimony and maximum–likelihood analyses strongly supported the existence of a crustacea + hexapoda clade ( pancrustacea ) over a myriapoda + hexapoda clade ( atelocerata ) . the apparent incompatibility between the molecule–based pancrustacea hypothesis and morphology–based atelocerata hypothesis is discussed .
( barnacles , etc . ) and they , in turn , with copepoda , making the traditional crustacean class maxillopoda paraphyletic . ostracoda ( seed shrimp ) âeither all or a subgroupâis associated with branchiura ( fish lice ) and likely to be basal to all other pancrustaceans . a bayesian statistical ( non - clock ) estimate of divergence times suggests a precambrian origin for pancrustacea ( 600 myr ago or more ) , which precedes the first unambiguous arthropod fossils by over 60 myr . pmid : 15734694
all three conclusions have been partially revealed in previous molecular reconstructions of arthropod or pancrustacean phylogenies 45 , 51 , 52 , but the consequences have never been discussed in detail . it is remarkable that our tree , based on a partial 18s rrna gene alone ( 1 . 7 kb ) , shows almost the same topology as that of regier et al . 48 , which is based on 62 nuclear protein coding genes ( 41 kb ) , although regier et al . 48 . obtained better support values . regarding pancrustacea , both analyses differ only in that regier’s et al . 48 tree does not include tantulocarida , and mystacocarida is sister to ostracoda in the present study but is the sister - group of pentastomida + branchiura in regier et al . 48 . pancrustacea ( hexapoda plus crustacea ) hypothesis is supported here , and , for the first time , the position of tantulocarida is demonstrated to be a sister - group ( outside not inside ) to thecostraca ( as compared to the discussion of petrunina et al . 52 ) .
) in a molecular tree . the tree shows that 1 ) mystacocarida is not a sister to copepoda but to ostracoda and 2 ) mystacocarida + ostracoda are not closely related to copepoda ; therefore , maxillopoda must be rejected as a natural group . its members are distributed into two non - sister major clades in the tree , with the clade consisting of mystacocarida + ostracoda , and pentastomida + branchiura placed at the base of the pancrustacea tree . 3 ) copepoda is sister to a clade comprising thecostraca + tantulocarida and malacostraca .
recent molecular analyses indicate that crustaceans and hexapods form a clade ( pancrustacea or tetraconata ) , but relationships among its constituent lineages , including monophyly of crustaceans , are controversial . our phylogenetic analysis of three protein - coding nuclear genes from 62 arthropods and lobopods ( onychophora and tardigrada ) demonstrates that hexapoda is most closely related to the crustaceans branchiopoda ( fairy shrimp , water fleas , etc . ) and cephalocarida + remipedia , thereby making hexapods terrestrial crustaceans and the traditionally defined crustacea paraphyletic . additional findings are that malacostraca ( crabs , isopods , etc . ) unites with
in the past about 20 years , the view of animal phylogeny has experienced radical changes . one of the most remarkable developments concerns the relative position of the hexapoda ( insects and allies ) , which were considered for a long time to be closely related to the myriapoda . however , molecular phylogenetic studies have demonstrated that hexapods are actually nested within the paraphyletic “crustacea” ( meusemann et al . 2010 ; regier et al . 2010 ; rehm et al . 2011 ; von reumont et al . 2012 ) . together , hexapoda and “crustacea” form a monophyletic taxon referred to as the pancrustacea ( zrzavý and štys 1997 ) .
occurrence of respiratory proteins in pancrustacea . the phylogeny of the “crustacean” classes is derived from regier et al . ( 2010 ) , the tree of the hexapods is based on misof et al . ( 2014 ) . hc denotes taxa in which hemocyanin was found : hb taxa with a respiratory globin . note that not all species of a certain taxon may harbor hc or hb . the dash ( - ) indicates the absence of any respiratory proteins ; the question mark ( ? ) denotes taxa for which there is no or too little information . there was a single example of co - occurrence of hc and hb in malacostraca .
the most notable results of this study are in regard to the internal structure of the pancrustacean clade . instead of a distinct separation between crustaceans and hexapods , we see four resolved clades within pancrustacea . the hexapod clade is upheld and nested completely within crustacea . in addition crustaceans are split into three paraphyletic clades ; vericrustacea , oligostraca , and xenocarida . vericrustacea ( ‘true crustaceans’ ) includes the decapods ( crabs and lobsters ) , mantis shrimp , barnacles , copepods , and branchiopods ; most of the critters commonly thought of as crustaceans . oligostraca is a strange and early branching crustacean group that includes ostracods , tongue worms , and branchiurans . xenocarida ( ‘strange shrimps’ ) includes remipedes and cephalocarids .
first , waptia is one of the most enigmatic burgess shale arthropods , and we found alternate positions as a sister group either to malacostraca or to multicrustacea as a whole . the possible relationship to malacostraca is supported mainly by eye morphology , which can be homoplastic in pancrustacea ( e . g . , oakley 2003 ) . the ambiguity of its phylogenetic placement makes waptia a poor choice for divergence time constraints . second , the devonian fossil nahecaris has been regarded as a stem - group leptostracan , an idea supported by our concatenated analysis . interestingly , the site - weighed method places nahecaris on the stem lineage of the malacostraca . this seems to occur due to the lack of leptostracan epipod morphology .
cheong et al . ( 2015 ) review the endocrine system of pancrustaceans . all pancrustaceans ( and all arthropods ) have an exoskeleton that must be shed for the animal to grow ; throughout pancrustacea , molting is regulated by ecdysteroids ( and upstream peptides ) . while juvenile hormone has long been known to occur in insects , it was not until 1990 that the crustacean equivalent of a juvenile hormone , methyl farnesoate , was identified ( laufer et al . 1987 ) . emerging evidence suggests that methyl farnesoate modifies the nature of the molt in crustaceans , and also can regulate reproduction , that is , its functions are very similar to those observed for juvenile hormone in insects . cheong et al . ( 2015 ) suggest that juvenile hormone evolved near the origin of insects , and that this hormone is a key player in the diversification and success of that taxon .
the evolution of copepoda as a progenetic malacostracan was proposed by gurney 59 who compared the morphology of the larval “copepodid” stages of copepods to the protozoea of peneid shrimps . this hypothesis was elaborated further by newman 60 , who extended this idea to the origin of maxillopoda from eumalacostraca through progenesis . however , the arguments to support a maxillopodan - malacostracan relationship ( or common origin ) were mostly based on detailed morphological and developmental patterns of copepoda and thecostraca alone 61 , leaving other maxillopodans aside because of a lack of detailed information . although our pancrustacea tree invalidates the progenetic origin of maxillopoda from an eumalacostracan , the progenetic origin of copepoda from a stem line group common to communostraca remains a valid hypothesis . copepoda have developed a terminal moult ( adult copepods do not moult further ) , suppressing the part of development that would be homologous to postlarval development in decapods .
lost limbs can be replaced in insects and crustaceans and then the regeneration . in both arthropod groups , such regeneration requires molting . das ( 2015 ) reviews the current state of knowledge concerning regeneration of limbs with respect to the hormonal , molecular , and morphological mechanisms known to occur in the pancrustacea . she found that insect model systems ( fruitflies and crickets ) are primarily studied to reveal information on the molecular mechanisms involved in successful regeneration . such studies have elaborated the complex profiles of gene expression required for the correct patterning of the lost appendage . because of their size , crustacean model systems ( crabs and lobsters ) have provided the majority of data concerning the hormonal and morphological basis of regenerating limbs . very detailed histological examination of the growing limb describes morphological changes associated with regeneration , whereas repetitive sampling of hemolymph and subsequent analysis of ecdysteroids provide the hormonal changes associated with the regeneration of limbs .
pioneer transcription factors ( tfs ) are considered the first regulators of chromatin accessibility in fruit flies and vertebrates , modulating the expression of a large number of target genes . in fruit flies , zelda resembles a pioneer tf , being essential during early embryogenesis . however , the evolutionary origins and ancestral functions of zelda remain largely unknown . through a number of gene silencing , microscopy and evolutionary analysis , the present work shows that zelda is an innovation of the pancrustacea lineage , governing not only the mzt in the short - germ insect tribolium castaneum , but also posterior segmentation and post - embryonic patterning of imaginal disc derived structures such as wings , legs and antennae . further , zelda regulation of posterior segmentation predates the origin of insects with complete metamorphosis ( holometabolous ) , as supported by gene silencing experiments in the kissing bug rhodnius prolixus . we hypothesize that the emergence of zelda contributed to the evolution of gene regulatory networks and new morphological structures of insects .
author contributions c . w . c . , j . c . r . , j . w . s . , a . z . and j . w . m . designed the project . j . w . s . , j . w . m . , r . w . and j . c . r . designed and carried out taxon sampling and collection . j . c . r . and c . w . c . supervised dna sequencing and editing , with pcr templates generated by j . c . r . , b . b . and others . j . c . r . , a . z . , c . w . c . and j . w . s . decided on the strategy for data analysis and its implementation , with the degen1 coding method developed and implemented by j . c . r . , a . h . and a . z . j . c . r . and a . z . assembled the supplementary information and submitted sequences to genbank . j . w . s . and j . w . m . proposed the names for the new , strongly supported clades in the pancrustacea . c . w . c . wrote the first draft of the manuscript , with major additions by j . c . r . and j . w . s . and additional contributions by j . w . m . and a . z . all authors commented on the manuscript .
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the complete mitochondrial genome of the black mud crab , scylla serrata ( crustacea : brachyura : portunidae ) and its phylogenetic position among ( pan . . . - pubmed - ncbi
warning : the ncbi web site requires javascript to function . more . . .
the complete mitochondrial genome of the black mud crab , scylla serrata ( crustacea : brachyura : portunidae ) and its phylogenetic position among ( pan ) crustaceans .
department of genetics , kasetsart university , bangkok , 10900 , thailand . fscianj @ urltoken
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. this work offers strong evidence for , and opens new , perplexing questions about , the deep evolutionary history of arthropods .
crustacea – crabs , lobsters , shrimp , copepods , ostracods , barnacles , pill bugs , brine shrimp , water fleas , remipedes , ect .
all known arthropods are included in these subphyla . however , the interrelation of the subphyla has long been in contention . up to now , a series of morphological and molecular - based classification paradigms have been employed to parsimoniously resolve the deep phylogeny of arthropods ; with mixed results . two of the competing hypotheses of deep arthropod phylogenetics are the paradoxopoda and mandibulata models .
both mandibulata and paradoxopoda propose a pancrustacean clade consisting of hexapods and crustaceans as closely related sister groups . they differ in their treatment of the myriapods and chelicerates . paradoxopoda has myriapods and chelicerates as closely related sister groups . mandibulata , on the other hand , has myriapods as a sister group to pancrustaceans , and chelicerates as a distant , early - branching clade of arthropods . while mandibulata is supported by taxonomy ( crustaceans , hexapods , and myriapods all have mandibles , where as chelicerates have chelicerae for mouthparts ) , paradoxopoda is primarily supported by recent mitochondrial molecular pylogenetics .
also under contention is the relationship between crustaceans and hexapods within the increasingly well supported pancrustacean clade . it has not yet been settled if the two are distinct sister groups , or if the hexapods are nested within the crustaceans . if the latter is true , researchers have not been able to determine which branch of the crustaceans gave rise to the hexapods .
in the present study , researchers at university of maryland , duke , and the natural history museum of la have dramatically improved on previous molecular approaches for inferring deep arthropod phylogenetics . they did this by increasing the number and diversity of arthropod species included , as well as the type and amount of genetic sequence characters used . whereas previous approaches used mitochondrial genetic sequences , resulting in the morphologically perplexing paradoxopoda model ; the researchers in this study instead used nuclear protein - coding genes . nuclear genes have been increasingly used to resolve deep evolutionary relationships in animals .
the researchers included in their molecular phylogenetic analysis 75 arthropod species . the 75 arthropod taxa were chosen to encompass the broadest spectrum of known arthropod groups . from each of these species , they sequenced 62 nuclear genes ; totaling 41 thousand bases of dna sequence per species . these sequence data sets from the 75 different arthropods were compared to one another , producing this phylogenetic tree :
maximum likelihood phylogenetic tree of arthropods . adapted from regier et al . , 2010 .
this study causes some serious upheavals in the pancrustacean clade ; breaking crustaceans into three paraphyletic clades , placing remipedes as close sister taxa to hexapods , and nesting the hexapods completely within the crustaceans . at the same time it presents new and exciting questions to morphologists and evolutionary - developmental biologists .
further morphological , genetic , and evolutionary study is required in order to understand the full implications of these newly resolved arthropod relationships . however , this research represents a robust step forward towards unraveling arthropoda .
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gene regulatory networks ( grns ) depend on the coevolution of transcription factors ( tfs ) and their relationship with the cis - regulatory modules ( crms ) they bind [ 1 , 2 ] . in insects , the detailed role of a number of tfs and crms have been well - described , particularly during the embryogenesis of the fruit fly drosophila melanogaster [ 3 ] .
( a ) multiple sequence alignment of zelda proteins , representing major groups of arthropods . ( b ) conserved protein architecture features of zelda proteins . asterisks and x marks represent presence and absence , respectively . question marks denote that the feature is either partially preserved or could be flagged as absent due to sequencing or assembly errors ( e . g . wrong start codons ) . the cladogram was organized according to a previously reported phylogenetic study [ 79 ] . two outgroups without zld orthologs were also included .
previous studies in d . melanogaster using microarrays and chip - seq revealed that dm - zld regulates the transcription of hundreds of genes during early embryogenesis [ 6 , 9 , 29 ] . in d . melanogaster , enhancers bound by dm - zld are characterized by a consensus sequence caggtag ( i . e . tagteam sequence ) , which is overrepresented in early zygotic activated genes , including tfs involved in ap and dv patterning [ 7 , 8 ] . since the tagteam motif identified in d . melanogaster is conserved in a . aegypti [ 30 ] , we investigated whether we could predict tc - zld targets by detecting tagteam motifs in the upstream regions of t . castaneum genes .
( a ) tomtom comparison of d . melanogaster motif similar to the tagteam [ 7 ] obtained by dreme and putative t . castaneum zld dreme motif . ( b ) venn diagram of the d . melanogaster chip - seq mzt regulated genes ( green ) from [ 8 ] and d . melanogaster genes predicted by fimo analysis with the putative dreme motif ( yellow ) . ( c ) venn diagram of the d . melanogaster dm - zld mzt targets ( green ) [ 8 ] and d . melanogaster one - to - one orthologs of putative tc - zld targets .
is maternally provided and progressively confined to the posterior growth - zone during embryonic development .
in all embryos , the anterior region points to the left . ( a ) relative expression of tc - zld in ovary and carcass . ( b ) relative expression of tc - zld 0–3 , 3–6 and 6–9 hours after egg lay . expression values were normalized using the constitutive gene rps3 in both experiments . ( c ) pre - blastoderm stage embryo ( 0–3 hours ) shows tc - zld transcripts uniformly distributed and its respective dapi staining in d . ( e ) at uniform blastoderm stage ( 3–6 hours ) transcripts begin to occupy the germ rudiment , respective dapi in f . ( g ) shortly before posterior invagination , zld expression occurs along the whole germ rudiment , with higher levels at the posterior region where growth zone will form ( 6–9 hours ) . ( h ) shortly before the beginning of germ band elongation zld expression is highly expressed at the gz , although lower levels of mrna can be observed at the along the whole embryonic region . ( i ) approx . 13 hours after egg lay ( ael ) , during the beginning of germ band extension , tc - zld is expressed at the posterior gz ( arrow ) . ( j ) an embryo slightly older than the one in h , highlighting tc - zld expression at ventral serosal cells during serosal window closure . ( k ) approx . 18 - 21hours ael zld is still expressed at the posterior most region ( arrow ) , head lobes ( arrowheads ) and a single gnathal segment ( asterisk ) . ( l ) approx . 39–48 hours ael , during early dorsal closure , expression is observed at the nervous system ( arrow ) .
( a - c ) nuclear dapi staining of t . castaneum embryos between 0–1 hours ( a ) , 3–6 hours ( b ) and 6–9 hours after oviposition ( c ) . ( d , h ) western - blots of embryonic extracts from 0–1 , 0–3 , 3–6 and 6–9 hours after oviposition using an antibody against the transcriptionally active form of rna pol ii , as previously described [ 33 ] . ( d ) or an antibody against the α - tubulin protein as a loading control ( h ) . ( e , f , g ) immunostaining showing nuclear rna pol ii at 3–6 hours ( f ) and 6–9 hours ( g ) , but not at 0–1 hour ( e ) after oviposition . coupling rna pol ii staining with nuclear dapi staining shows that t . castaneum zygotic transcription starts between 3–6 hours after egg laying , when the energids reaches the periphery ( fig 2a - c - dapi and e - g - rna pol ii ) . similar results were obtained by western - blots using the same antibody ( d ) .
it has been shown that injection of dsrna in t . castaneum females leads to reduced expression of a given gene in the females and their offspring , in a phenomenon called parental rnai ( prnai ) effect [ 15 ] . we injected zld dsrna in females and analyzed tc - zld transcriptional levels during embryogenesis by qrt - pcr . after zld prnai , tc - zld mrna levels were reduced in the first two weeks of egg laying , severely impairing larval hatching ( fig 5a–5c ) . importantly , identical knockdown phenotypes during embryogenesis were obtained by using a second , non - overlapping , dsrna construct ( s1 fig ) . further , morphological analyses showed that cellularization was severely disrupted in over 50 % of the zld dsrna embryos ( fig 5d ) , similarly to what was previously reported in d . melanogaster zld mutants [ 6 , 11 ] . the remaining zld prnai embryos were not severely affected during cellularization and developed beyond that stage . finally , we also found that some putative conserved target genes were down - regulated in embryos after zld prnai , such as the early zygotic genes involved in ap patterning , the serosal gene tc - zerknullt [ 36 ] ( fig 5e and 5f ) and the gap gene milli - pattes [ 37 ] ( fig 5g and 5h ) .
parental rnai affects oviposition and cellularization , while embryonic rnai affects embryonic posterior patterning .
( a ) normalized number of dsneo and dszld rnai collected eggs . zld prnai reduces oviposition . ( b ) percentage of hatched larvae obtained from eggs after zld prnai in comparison to the control ( dsneo rnai ) . zld prnai leads to lethality during embryogenesis . ( c ) relative expression of zld in dsneo rnai ( control ) and dszld rnai during the first two egg lays . zld prnai almost completely abolishes zld expression . ( d ) phospho - tyrosine ( red ) and dapi ( blue ) staining in dsneo and zld rnai eggs during cellularization . zld prnai impairs cellularization . ( e , f ) tc - zen expression in the serosa [ 36 ] is reduced after tc - zld prnai ( arrowhead ) when compared to the dsneo control . ( g , h ) expression of the gap gene tc - mlpt [ 37 ] in control ( g ) and zld rnai ( h ) . tc - mlpt loses its anterior expression domain in zld rnai when compared to control . ( i , j ) the expression of the dorsoventral gene tc - cactus [ 38 ] was reduced after tc - zld prnai ( j - arrow ) when compared to the ( i ) control . ( k , l ) the expression of the dorsoventral gene tc - sog was reduced after tc - zld prnai ( j - arrow ) when compared to the ( i ) control . ( m , n ) expression of tc - toll7 was affected in the posterior region of tc - zld rnai embryos ( n ) when compared to dsneo ( m ) . ( o , p ) expression of the segmentation gene tc - eve in control ( o ) and zld rnai ( p ) . tc - eve expression is essential for segmentation via a pair rule circuit in t . castaneum [ 39 ] . after zld rnai the characteristic tc - eve stripe expression in the growth zone is lost . ( q , r ) segment generation from the growth - zone ( gz ) is disrupted , as judged by the analysis of tc - gsb expression after tc - zld prnai ( r ) and dsneo ( q ) . ( s , t ) tc - zld dsrna embryonic injection ( ernai ) into nuclear gfp transgenic line affects segment generation from the posterior gz ( l - asterisk ) , when compared to ngfp embryo injected with dsneo dsrna .
changes in the spatial distribution of transcripts from predicted dorsoventral target genes were also observed after tc - zld rnai . in wild - type ( wt ) embryos , the tf dorsal forms a dynamic transient gradient , which activates tc - cactus ( tc - cact ) and tc - short - gastrulation ( tc - sog ) at the ventral region [ 38 , 39 ] . after tc - zld prnai , the expression of tc - cact and tc - sog is observed in two lateral domains , in contrast to the expression at the single ventral domain in dsneo rnai embryos ( fig 5i , 5j , 5k and 5l ) . these results suggest that tc - zld is required for proper activity of dorsal at the ventral - most region of the embryo .
as discussed above , tc - zld is highly expressed at the posterior region of the embryo ( fig 3 ) and likely associated with segmentation and regionalization ( s4 table ) . further , several putative tc - zld targets are involved in posterior segmentation , such as caudal ( cdx ) , even - skipped ( eve ) and several hox genes ( e . g . ultrabithorax , abdominal - a and abdominal - b ) . tc - eve , for example , is essential for the establishment of a genetic circuit required for posterior segmentation [ 40 ] . interestingly , tc - zld prnai embryos showed a continuous tc - eve expression domain instead of the typical stripe patterning required for wt segmentation ( fig 5o and 5p ) .
elegant studies on t . castaneum gz patterning showed that cell proliferation is not essential for segment generation , which rather occurs by coordinated cell movement and intercalation [ 19 , 41 ] . we then evaluated whether tc - zld regulates genes involved in cell intercalation , such as toll2 , toll6 and toll8 [ 42 , 43 ] . interestingly , toll7 ( tc004474 ) and toll8 ( tollo : tc004898 ) are among the common zld targets conserved between d . melanogaster and t . castaneum ( s3 table ) . since tc - toll7 is expressed during early segmentation [ 43 ] , we compared its expression in dsneo and tc - zld rnai embryos . while anterior expression of tc - toll7 is apparently unaffected , the striped expression at the posterior region is lost when tc - zld expression is reduced ( fig 5m and 5n ) . further support for the loss of posterior segmentation after tc - zld rnai is also provided by the analysis of expression of the segment - polarity gene , tc - gooseberry ( tc - gsb ) ( fig 5q and 5r ) . in summary , tc - zld regulates the expression of several genes that are critical for early ap ( zen , mlpt ) and dv ( sog , cact ) patterning and , in a second phase , genes required for posterior elongation ( e . g . toll7 , gsb ) .
while prnai diminishes maternal and zygotic expression of tc - zld in t . castaneum [ 15 ] , embryonic dsrna injections ( ernai ) may affect only the zygotic component , since eggs can be injected after the mzt [ 38 , 44 ] . to investigate if tc - zld is specifically required for embryonic posterior patterning , we injected tc - zld dsrna in transgenic embryos expressing nuclear gfp ( ngfp ) , as previously described [ 18 , 19 ] . embryonic injections of tc - zld dsrna after the mzt ( see methods for details ) impaired segment generation from the gz , while dsneo - injected embryos developed like wt ones ( fig 5s and 5t , s1 and s2 movies ) . in addition , expression of the predicted target tc - eve , a key tf involved in gz patterning [ 35 ] , has been largely down - regulated upon tc - zld ernai , as previously observed for zld prnai ( fig 5o and 5p ) . in summary , our results imply that tc - zld is involved not only in the mzt , early patterning and nervous system formation , as described for d . melanogaster [ 6 ] , but also play roles in segment generation from the gz , a structure found only in embryos of short - germ insects like t . castaneum .
( a , c , e ) r . prolixus control ( dsneo embryos ) . ( b , d , f , f’ ) a representative embryo collected from r . prolixus zld prnai . ( f , f’ ) embryo inside the chorion , ventral and dorsal views . ( c , e ) embryo removed from the egg shell . ( g , h ) schematic drawings of control and zld prnai embryos . ( a ) dapi stainings of control ( a , e ) and zld rnai ( b ) embryos . ( d ) asterisk denotes an eye which can be observed at the ventral side ( f ) . ( d ) after dissection , the eye can be identified due to its characteristic red pigmentation and shape . scale bar corresponds to 500μm .
david analysis of d . melanogaster orthologs of the putative tc - zld targets uncovered a cluster of 29 genes involved in imaginal disc development ( s4 table ; go : 0007444 ) . among these genes are several homeodomain tfs , such as distalless ( dll ) , abdominal a ( abd - a ) , abdominal b ( abd - b ) , zen , engrailed ( en ) , caudal ( cad ) , defective proventriculus ( dve ) , mirror ( mirr ) , araucan ( ara - iroquois ) and drop ( dr ) , as well as other tfs such as dachsund ( dac ) , taranis ( tara ) and lim1 , poxn , kn , sob , drm , awh , dp .
as the first step towards the characterization of the post - embryonic role of zld , we analyzed tc - zld expression by qrt - pcr in larvae of third ( l3 ) , fifth ( l5 ) and seventh ( l7 ) stages , and first pupal stage ( p1 ) ( fig 7a–7d ) . interestingly , tc - zld expression increases during successive larval stages and sharply decreases after pupal metamorphosis ( fig 7e ) . this suggests that tc - zld might be required for late larval stages , which take place during larvae - pupae metamorphosis , such as growth and patterning of structures derived from imaginal discs in d . melanogaster ( e . g . antennae , legs , fore - and hindwings ) [ 49 , 50 ] . further , we found that three out of five predicted tc - zld targets , namely dll , wg and lim - 1 , also displayed an increase in expression during late larval and pupal development ( fig 7e ) .
( a - d ) morphology of tribolium castaneum larvae on 3rd ( a - l3 ) , 5th ( b - l5 ) , 7th ( c - l7 ) and early pupal stages ( d - p1 ) . ( e ) relative expression of zld , ubx , wingless ( wg ) , distalless ( dll ) , iroquois ( iro ) , lim - 1 and kruppel - homolog - 1 ( krh - 1 ) at l3 , l5 , l7 and p1 . asterisks represent significant differences between stages ( p < 0 . 05 ) .
to investigate zld´s post - embryonic roles , we injected two non - overlapping tc - zld dsrna constructs into early ( l3 ) and late larval ( l6 ) stages , as previously described [ 50 ] . qrt - pcr confirmed that tc - zld was down - regulated after dsrna injection ( fig 8a ) . tc - zld dsrna injections at early larval stages ( l3 ) led to over 50 % lethality during pupal stages . atypical adult pigmentation in the pupal head and reduction in the wing size were observed after early tc - zld dsrna injection , suggesting that proper tc - zld expression is required for metamorphosis and wing growth ( fig 8b and 8c ) . interestingly , tc - zld dsrna injections at late larval stage ( l6 ) displayed a different phenotype when compared with early larval dsrna injections ( l3 ) . l6 larvae injected with dszld and dsneo reached adulthood at comparable rates ( fig 8d ) . specifically , tc - zld dsrna adults showed a series of morphological alterations in tissues undergoing extensive morphological changes during metamorphosis , such as fore - and hindwings , antennae and legs ( figs 8e , 8f , 9 and 10 ) .
tribolium castaneum larvae were injected during early ( 3rd ) and late ( 6th ) stages as previously described [ 50 , 80 ] . ( a ) relative expression of tc - zld during pupal stages after tc - zld dsrna or neo dsrna injection . qrt - pcr was normalized using tc - rps3 gene , as previously described [ 78 ] . ( b , c ) morphology of late pupae obtained from early larvae injected with zld or neo dsrna . differential pigmentation in the head ( arrow ) and reduced wings ( arrowhead ) were observed in zld dsrna and not in dsneo dsrna pupae . ( d ) number of emerging adult beetles eleven days after larval injection ( dali ) at late ( 6th ) stage . ( e , f ) . adults obtained by late larval injections of neo ( e ) or zld ( f ) dsrna . hindwings are usually not properly folded underneath the forewing ( elytra ) .
( a ) control elytron extracted from a dsneo adult shows a parallel venation pattern and rigid chitinous structure . ( b ) tc - zld rnai elytron displays a disrupted vein pattern and a less resistant structure . ( c , d ) hindwings were dissected and photographed . although the overall morphological pattern is not affected in both wings , zld dsrna wings show severe dehydration after ethanol fixation . ( e ) pd indexes comparison between dsneo and dszld rnai hindwings and forewings , a pd index reflects the shape of the wing based on its dimensions ratio [ 52 ] . the statistical analysis was carried out by unpaired t - test assuming unequal variances ( asterisks refer to p < 0 . 0001 while ns stands for “no significance” ) , indicating that tc - zelda rnai affects hindwing shape . ( f ) tc - zld rnai elytra and hindwing do not show statistically significant differences in length when compared to their respective dsneo rnai controls .
( a ) relative expression of tc - dll at pupal stages after neo or zld dsrna injections . ( b - d ) antennae extracted from control injected individuals ( b ) display 11 segments , including two proximal ( s = scape , p = pedicel ) , six intermediate ( flagellum = dots ) and three distal modified segments that form the club ( cl ) . ( c , d ) injection of zld dsrna caused fusion of the club segments in mild phenotypes ( c ) , while in stronger phenotypes ( d ) the antennae fail to segment , developing as an antennal rudiment . ( e , f ) legs of t . castaneum display six podomeres : coxa ( cx ) , trochanter ( tr ) , femur ( fe ) , tibia ( ti ) , tarsus ( t ) and pretarsus ( pt ) . ( e ) mesothoracic legs of neo dsrna injected individuals show five tarsal segments , or tarsomeres . ( f ) zld dsrna beetles lacks tarsal segments resulting in a reduced tarsus , while other leg structures do not display large morphological defects .
the most visible effect of tc - zld dsrna beetles was a failure of the forewings ( elytra ) to enclose the hindwings , leading to the exposure of the dorsal abdomen ( fig 8e and 8f ) . elytra , which are highly modified beetle forewings , have been proposed to be an important beetle innovation , being required for protection against mechanical stresses , dehydration , and predation [ 51 ] . in line with this hypothesis , tc - zld dsrna adults with exposed abdomens started to die a few days after metamorphosis , probably due to dehydration .
interestingly , all tc - zld dsrna beetles that showed this ‘opened wing’ phenotypes ( fig 8f ) also displayed defects in the legs and antennae . antennae and legs share similar developmental grns , involving the so called serial homologs . distaless ( dll ) , one of the putative tc - zld targets , is essential for appendage segmentation [ 53 ] . tc - dll is also expressed during late embryogenesis on the distal part of the leg and , as its name suggests , disruption of its function leads to the absence of distal leg and antennae segmentation [ 54 ] . interestingly , we found that tc - zld rnai resulted in a significant decrease of tc - dll mrna levels ( fig 10a ) , indicating that this gene is indeed downstream of tc - zld .
insect antennae possess three primary segments : scape , pedicel and flagellum . in t . castaneum , the adult antennae display eleven segments , out of which nine form the flagellum . the three most distal flagellar segments are enlarged and form the club , while the six intermediate flagella are called the funicle ( fig 10b ) [ 55 ] . in mild phenotypes , dszld caused a joint malformation in distalmost flagellar segments resulting in a fusion of the club ( fig 10c ) . on the other hand , strong tc - zld dsrna phenotypes resulted in fusion of the scape and pedicel , leading to severe loss of flagellar joints and formation of a single truncated segment ( fig 10d ) . in contrast , no differences in scape and pedicel were observed .
t . castaneum legs originate during late embryogenesis and can be recognized as a small outgrowth of the body wall , the limb bud . in the adult stage , there are three pairs of segmented legs with six segments : coxa , femur , trochanter , tibia , tarsus and pretarsus [ 56 ] . in t . castaneum the tarsus is subdivided in smaller segments ( i . e . the tarsomeres ) , five in prothoracic and mesothoracic legs and four in metathoracic legs . tarsal segmentation occurs during beetle metamorphosis and this subdivision of the tarsus evolved in the common ancestor of insects , since the tarsus is not subdivided in non - insect hexapods [ 57 ] . after tc - zld rnai , tarsal segments were absent or fused , resulting in leg shortening ; in dsneo insects , legs were identical to that of wt animals ( fig 10e and 10f ) . this indicates that some of the tc - zld targets are involved in segment development or joint formation . interestingly , a large domain of dll expression is observed in beetle leg tarsus [ 58 ] , and tc - dll knockdown in beetles also generated legs with tarsomere deletion [ 58 , 59 ] .


dentifusus is a genus of sea snails , marine gastropod mollusks in the family fasciolariidae , the spindle snails , the tulip snails and their allies .

dentifusus

dentifusus rosenberg , g . & g . j . vermeij , 2003 type species : dentifusus deynzeri vermeij , g . j . & g . rosenberg , 2003
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subfamily : fasciolariinae - genus : dentifusus g . rosenberg & g . j . vermeij , 2003 ( db : 1 sp , 1 img )
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, invalid : junior homonym of fusus helbling , 1779 . see nomenclature note below . )
» genus clavella swainson , 1835 accepted as clavilithes swainson , 1840 † ( junior homonym of clavella oken , 1815 . has been renamed clavilithes . )
» genus fredenia cadée & a . w . janssen , 1994 † accepted as latirulus cossmann , 1889
» genus fusus bruguière , 1789 accepted as fusinus rafinesque , 1815 ( invalid : junior homonym of fusus helbling , 1779 . see nomenclatural note below . )
» genus sinistralia h . adams & a . adams , 1853 accepted as fusinus rafinesque , 1815
» genus chasca clench & aguayo , 1941 accepted as chascax r . b . watson , 1873 accepted as hemipolygona rovereto , 1899 ( spelling error )
» genus chascax r . b . watson , 1873 accepted as hemipolygona rovereto , 1899 ( junior homonym of chascax ritgen , 1829 [ reptilia ] ; hemipolygona is a replacement name . )
» genus fusilatirus mcginty , 1955 accepted as dolicholatirus bellardi , 1884 ( synonym of dolicholatirus . )
snyder m . a . ( 2003 ) . catalogue of the marine gastropod family fasciolariidae . academy of natural sciences . of philadelphia , special publication . 21iii + 1–431 . , available online at urltoken [ details ]
couto d . , bouchet p . , kantor yu . i . , simone l . r . l . & giribet g . ( 2016 ) . a multilocus molecular phylogeny of fasciolariidae ( neogastropoda : buccinoidea ) . molecular phylogenetics and evolution . 99 : 309 - 322 . , available online at urltoken [ details ]
bouchet p . , rocroi j . p . , hausdorf b . , kaim a . , kano y . , nützel a . , parkhaev p . , schrödl m . & strong e . e . ( 2017 ) . revised classification , nomenclator and typification of gastropod and monoplacophoran families . malacologia . 61 ( 1 - 2 ) : 1 - 526 . [ details ] available for editors [ request ]
microfulgur finlay , h . j . & j . marwick , 1937 type species : microfulgur longirostris marshall , 1937
fasciolaria lamarck , j . b . p . a . de , 1799 type species : fasciolaria tulipa tulipa linnaeus , c . , 1758
africolaria snyder , m . a . , g . j . vermeij & w . g . lyons , 2012 type species : africolaria rutila watson , r . b . , 1882
aurantilaria snyder , m . a . , g . j . vermeij & w . g . lyons , 2012 type species : fasciolaria aurantiaca lesson , r . p . , 1842
australaria snyder , m . a . , g . j . vermeij & w . g . lyons , 2012 type species : australaria australasia perry , g . , 1811
cinctura hollister , s . c . , 1957 type species : cinctura hunteria hunteria perry , g . , 1811
conradconfusus snyder , m . a . , 2002 type species : conradconfusus parilis conrad , t . a . , 1832
filifusus snyder , m . a . , g . j . vermeij & w . g . lyons , 2012 type species : filifusus filamentosus röding , p . f . , 1798
granolaria snyder , m . a . , g . j . vermeij & w . g . lyons , 2012 type species : granolaria salmo wood , w . , 1828
kilburnia snyder , m . a . , g . j . vermeij & w . g . lyons , 2012 type species : kilburnia heynemanni dunker , r . w . , 1870
lugubrilaria snyder , m . a . , g . j . vermeij & w . g . lyons , 2012 type species : lugubrilaria lugubris adams , a . & l . a . reeve in reeve , l . a . , 1847
microcolus cotton , b . c . & f . k . godfrey , 1932 type species : microcolus dunkeri jonas , j . h . , 1844
pleuroploca fischer , 1884 type species : pleuroploca trapezium linnaeus , c . , 1758
pleuroploca ( pleia ) finlay , h . j . , 1930 type species : pleuroploca ( pleia ) decipiens tate , r .
pustulatirus vermeij , g . j . & m . a . snyder , 2006 type species : pustulatirus mediamericanus hertlein , l . g . & a . m . strong , 1951
tarantinaea monterosato , t . a . de m . di , 1917 type species : tarantinaea lignaria linnaeus , c . , 1758
triplofusus olsson , a . a . & a . harbison , 1953 type species : triplofusus giganteus kiener , l . c . , 1840
turrilatirus vermeij , g . j . & m . a . snyder , 2006 type species : turrilatirus turritus gmelin , j . f . , 1791
fusinus rafinesque , c . s . , 1815 type species : fusinus ( fusinus ) colus linnaeus , c . , 1758
fusinus ( fusinus ) rafinesque , c . s . , 1815 type species : fusinus ( fusinus ) colus linnaeus , c . , 1758
fusinus ( fusinus ) ansatus caboblanquensis ( var . ) † ? ( 3 )
fusinus ( aptyxis ) troschel , f . h . , 1868 type species : fusinus ( aptyxis ) syracusanus linnaeus , c . , 1758
fusinus ( barbarofusus ) grabau , a . w . & shimer , 1909 type species : fusinus ( barbarofusus ) barbarensis trask , 1855
fusinus ( heilprinia ) grabau , a . w . , 1904 type species : fusinus ( heilprinia ) caloosaensis heilprin , a . , 1887
amiantofusus fraussen , k . , yu . i . kantor & r . hadorn , 2007 type species : amiantofusus amiantus dall , w . h . , 1889
chryseofusus hadorn , r . & k . fraussen , 2003 type species : chryseofusus chrysodomoides schepman , m . m . , 1911
cyrtulus hinds , r . b . , 1843 type species : cyrtulus serotinus hinds , r . b . , 1844
glaphyrina finlay , h . j . , 1926 type species : glaphyrina vulpicolor sowerby , g . b . iii , 1880
granulifusus kuroda , t . & t . habe , 1954 type species : granulifusus niponicus niponicus smith , e . a . , 1879
harasewychia petuch , e . j . , 1987 type species : harasewychia harasewychi petuch , e . j . , 1987
harfordia dall , w . h . , 1921 type species : harfordia harfordii stearns , r . e . c . , 1871
marmarofusus snyder , m . a . & w . g . lyons , 2014 type species : marmarofusus nicobaricus röding , p . f . , 1798
ollaphon iredale , t . , 1929 type species : ollaphon molorthus hedley , c . & w . l . may , 1908
sinistralia adams , h . g . & a . adams , 1853 type species : sinistralia maroccensis gmelin , j . f . , 1791
trophonofusus kuroda , t . & t . habe , 1971 type species : trophonofusus muricatoides yokoyama , m . , 1920
viridifusus snyder , m . a . , g . j . vermeij & w . g . lyons type species : viridifusus buxeus reeve , l . a . , 1847
peristernia mörch , o . a . l . , 1852 type species : peristernia nassatula lamarck , j . b . p . a . de , 1822
peristernia ( nodopelagia ) hedley , c . , 1915 type species : peristernia ( nodopelagia ) brazieri angas , g . f . , 1877
bullockus lyons , w . g . & m . a . snyder , 2008 type species : unknowngenustype
dolicholatirus bellardi , l . , 1884 type species : dolicholatirus lanceus gmelin , j . f . , 1791
dolicholatirus ( fractolatirus ) iredale , t . , 1936 type species : dolicholatirus ( fractolatirus ) normalis iredale , t . , 1936
hemipolygona rovereto , g . , 1899 type species : hemipolygona armatus adams , a . , 1855
latirolagena harris , g . d . , 1897 type species : latirolagena smaragdula linnaeus , c . , 1758
benimakia habe , t . , 1958 type species : benimakia rhodostomus dunker , r . w . , 1860
latirus montfort , p . d . de , 1810 type species : latirus aurantiacus montfort , p . d . de , 1810
latirus ( latirulus ) cossmann , a . e . m . , 1889 type species : latirus ( latirulus ) subaffinis orbigny , a . v . m . d . d ' , 1850
latirus ( polygona ) schumacher , h . c . f . , 1817 type species : latirus ( polygona ) fusiformis schumacher , h . c . f . , 1817
latirus ( pseudolatirus ) bellardi , l . , 1884 type species : latirus ( pseudolatirus ) bilineata hörnes , m . , 1853
crassibougia stahlschmidt , p . & k . fraussen , 2012 type species : niso terebellum dillwyn , l . w . , 1817
cryptofusus beu , a . g . , 2011 type species : cryptofusus cryptocarinatus dell , r . k . , 1956
fusolatirus kuroda , t . & t . habe in kuroda , t . , t . habe & k . oyama , 1971 type species : peristernia pilsbryi kuroda , t . & t . habe , 1952
lamellilatirus lyons , w . g . & m . a . snyder , 2013 type species : lamellilatirus ceramidus dall , w . h . , 1889
leucozonia gray , j . e . , 1847 type species : leucozonia nassa nassa gmelin , j . f . , 1791
lightbournus lyons , w . g . & m . a . snyder , 2008 type species : unknowngenustype
nodolatirus bouchet , ph . & m . a . snyder , 2013 type species : unknowngenustype
opeatostoma berry , s . s . , 1958 type species : opeatostoma pseudodon burrow , e . j . , 1815
taron hutton , f . w . , 1883 type species : taron dubius hutton , f . w . , 1878
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subfamily : fasciolariinae - genus : cyrtulus r . b . hinds , 1843 ? ( db : 2 sp , 1 img )
subfamily : fasciolariinae - genus : fascinus c . hedley , 1903 ( db : 1 sp , 1 img )
subfamily : fasciolariinae - genus : fasciolaria j . b . p . a . lamarck , 1799 ( syn : terebraespina , fascolaria - db : 47 sp , 35 img )


trochidrobia is a genus of very small freshwater snails , aquatic gastropod mollusks in the family tateidae .

the genus trochidrobia is endemic to the lake eyre supergroup , australia .

trochidrobia

the species contained in trochidrobia are similar in shell and opercular characters to posticobia . however , trochidrobia species have more depressed shells , and lack the peripheral keel typical of some specimens of posticobia .
facts summary : trochidrobia is a genus of snails of concern and found in the following area ( s ) : australia .
wikipedia article copyright notice : this article is licensed under the gnu free documentation license . it uses material from the wikipedia article
trochidrobia
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glenn , c . r . 2006 .
earth ' s endangered creatures - trochidrobia spp . facts
( online ) - licensed article from wikipedia : the free encyclopedia . accessed
ponder , w . f . 1996 . trochidrobia punicea . in : iucn 2010 . iucn red list of threatened species . version 2010 . 4 . available at : www . iucnredlist . org > .
shell with short spire and open umbilicus ; reaches 2 . 1 mm in maximum diameter . this species and trochidrobia punicea are virtually identical in shell characters but differ anatomically . they are sympatric in only a few springs .
the small umbilicus and relatively high spire distinguish this species from other species of trochidrobia . it differs from the sympatric t . minuta in its larger size ( up to 1 . 7 mm maximum diameter ) , wider umbilicus and taller spire .
all of the trochidrobia species are very similar in shell morphology with minor differences in size and protoconch sculpture . the main differences are anatomical - particularly the coiling pattern of the oviduct in the female genital system . there are also geographical clusters of species and species associations ( see ponder , hershler and jenkins , 1989 ) .
original name : trochidrobia smithi ponder , hershler and jenkins , 1989 . ponder , w . f . , hershler , r . & jenkins , b . ( 1989 ) . an endemic radiation of hydrobiid snails from artesian springs in northern south australia : their taxonomy , physiology , distribution and anatomy . malacologia 31 : 1 - 140 .
original name : trochidrobia inflata ponder , hershler and jenkins , 1989 . ponder , w . f . , hershler , r . & jenkins , b . ( 1989 ) . an endemic radiation of hydrobiid snails from artesian springs in northern south australia : their taxonomy , physiology , distribution and anatomy . malacologia 31 : 1 - 140 .
ponder w . f . , hershler r . & jenkins b . ( 1989 ) . an endemic radiation of hydrobiid snails from artesian springs in northern south australia : their taxonomy , physiology , distribution and anatomy . malacologia . 31 ( 1 ) : 1 - 140 . page ( s ) : 59 [ details ]
ponder , hershler & jenkins , 1989 . accessed through : world register of marine species at : urltoken ; = 818613 on 2018 - 07 - 09
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dyer , e . , soulsby , a . - m . , whitton , f . , mcguinness , s . , de silva , r . , milligan , h . t . , kasthala , g . , herdson , r . , thorley , j . , mcmillan , k . & collins , a .
has been assessed as least concern due to the abundant and widespread distribution throughout the spring system . although its extent of occurrence is estimated as less than or equal to 20 , 000 km
, with habitat that is susceptible to a range of threats which result in water drawdown , it is unlikely that the global population is being negatively impacted .
this species is found throughout the southern and middle artesian spring groups , south of william creek between maree and oodnadatta in central south australia , australia ( ponder 1996 ) . its extent of occurrence is estimated as approximately 29 , 000 km 2 .
this species can be extremely abundant , but has a localised distribution ( ponder 1996 ) . at the dead boy and bopeechee spring groups in northern south australia , niejalke and richards ( 1998 ) recorded a mean population density of up to 18 . 7 individuals per cm² near to the spring outflow .
this species inhabits springs and can be found in the spring head and outflows , amongst small sedges at the edges , living sympatrically with four other hydrobiid species ( ponder 1996 ) . this species can occupy a variety of microhabitats and has been found to be particularly abundant in shallow , firm - bottomed outflows ( ponder et al . 1989 ) . this species has an adult size of approximately 1 . 65 mm ( niejalke and richards 1998 ) .
artesian springs are found in south australia and western queensland . there also used to be artesian springs in the north - west of new south wales , but these are now mostly extinct , presumably as a result of water extraction from the basin by the pastoral industry ( ponder et al . 1989 ) . in south australia , each spring is typically separated by arid agricultural land ( ponder 1996 ) which provides a marked discontinuity of habitat , like that of an island surrounded by sea . this fragmented distribution makes this species especially vulnerable to threat processes ( ponder 1996 ) , and it has been found that the life history characteristics of this species make it more sensitive to changes in prevailing environmental conditions ( niejalke and richards 1998 ) , such as those brought on by climate change . although locally very common , this species is dependent on a continuing supply of clean water , and yet it is currently threatened by the over - extraction of artesian water for agricultural , industrial and municipal uses ( wilmer et al . 2005 ) .
this species was previously assessed as near threatened in 1996 ( version 2 . 3 ) . the unique flora and fauna of the artesian spring systems have been listed as an ‘endangered community’ under the australian environmental protection & biodiversity conservation act of 1999 ( wilmer et al . 2005 ) . management of the springs in which this species is found - via an area - based management plan - is the only option to ensure its long - term survival . continued local and regional impacts on artesian water must be carefully controlled to prevent further drawdown . habitat damage due to livestock and other pastoral activities should be minimized ( ponder 1996 ) .
1 . land / water protection - > 1 . 2 . resource & habitat protection
11 . climate change & severe weather - > 11 . 3 . temperature extremes
2 . agriculture & aquaculture - > 2 . 3 . livestock farming & ranching - > 2 . 3 . 4 . scale unknown / unrecorded
7 . natural system modifications - > 7 . 2 . dams & water management / use - > 7 . 2 . 5 . abstraction of ground water ( domestic use )
7 . natural system modifications - > 7 . 2 . dams & water management / use - > 7 . 2 . 6 . abstraction of ground water ( commercial use )
7 . natural system modifications - > 7 . 2 . dams & water management / use - > 7 . 2 . 7 . abstraction of ground water ( agricultural use )
iucn . 2011 . iucn red list of threatened species ( ver . 2011 . 2 ) . available at : urltoken . ( accessed : 10 november 2011 ) .
niejalke , d . p . and richards , j . 1998 . population biology of some spring snails . proceedings to the second mound spring researchers forum and spring management workshop , adelaide .
ponder , w . f . , hershler , r . and jenkins , b . 1989 . an endemic radiation of hydrobiid snails from artesian springs in northern south australia : their taxonomy , physiology , distribution and anatomy . malacologia 31 ( 1 ) : 1 - 140 .
wilmer , j . w . , hughes , j . m . , ma , j . and wilcox , c . 2005 . characterization of microsatellite loci in the endemic mound spring snail fonscochlea accepta and cross species amplification in four other hydrobiid snails . molecular ecology notes 5 : 205 - 207 .
ponder , hershler & jenkins , 1989 . accessed through : world register of marine species at : urltoken ; = 821643 on 2018 - 07 - 09
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shell small ( up to about 2mm in diameter ) , trochiform to depressed - trochiform , umbilicate , smooth , with only sculpture weak axial growth lines . aperture oval , peristome thin , no external varix ; outer lip simple , with thin edge . periostracum smooth , thin . operculum oval , nucleus subcentral , thin , simple . penis with swollen basal portion and tapering distal portion that is usually pigmented ; coiled anti - clockwise when at rest .
original reference : ponder , w . f . , hershler , r . & jenkins , b . ( 1989 ) . an endemic radiation of hydrobiid snails from artesian springs in northern south australia : their taxonomy , physiology , distribution and anatomy . malacologia 31 : 1 - 140 .
type locality : blanche cup spring , west of lake eyre south , south australia .
abundant in most of the springs in which they occur , living in a variety of microhabitats and appear to be particularly abundant in shallow , firm - bottomed outflows . egg capsules are spherical , cemented in umbilicus of shell with mucus ( known only in t . punicea ) .
beesley , p . l . , ross , g . j . b . & wells , a . , eds . ( 1998 ) . mollusca : the southern synthesis . parts a & b . melbourne , csiro publishing .
murphy , n . p . , guzik , m . t . , cooper , s . j . , & austin , a . d . ( 2015 ) . desert spring refugia : museums of diversity or evolutionary cradles ? . zoologica scripta 44 : 693 - 701 .
ponder , w . f . ( 2004 ) . endemic aquatic macroinvertebrates of artesian springs of the great artesian basin—progress and future directions . records of the south australian museum monograph series 7 : 101 - 110 .
ponder , w . f . , eggler , p . e . & colgan , d . j . ( 1995 ) . genetic differentiation of aquatic snails ( gastropoda : hydrobiidae ) from artesian springs in arid australia . biological journal of the linnean society 56 : 553 - 596 .
ponder , w . f . , hershler , r . & jenkins , b . ( 1989 ) . an endemic radiation of hydrobiidae from artesian springs in northern south australia : their taxonomy , physiology , distribution and anatomy . malacologia 31 : 1 - 140 .
to cite this resource : ponder , w . f . , hallan , a . , shea , m . and clark , s . a . 2016 . australian freshwater molluscs . urltoken
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urn : lsid : biodiversity . org . au : afd . taxon : 6f5353eb - 12fc - 44d1 - bd1a - 9bbb5bce1ed3
urn : lsid : biodiversity . org . au : afd . taxon : f5e1c410 - 6f90 - 4506 - ab6a - 975a2285b361
urn : lsid : biodiversity . org . au : afd . name : 389788
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creatures with albinism and leucism are beautiful and rare animals . they have all the characteristics of others of their species except they are white in color . the lack of melanin generally results in the animal looking bleached all over , appearing white or pink . it happens in many animals ranging from squirrels to whitetail deer . here are ten incredible and rare , white - colored creatures that you ' ll probably never see in real life .
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the tasmanian devil is endemic to australia . although this species is called tiger ( named for its stripes ) and wolf ( due to its canid - like appearance ) , it is not a member of the cat or wolf family . it is a member of the marsupial family . other members of this family include kangaroos and koala bears .
the last known tasmanian tiger died in a zoo in hobart , tasmania in 1936 , but there have been hundreds of unconfirmed sightings , and a reserve has been set up in southwestern tasmania in the hopes that possible surviving individuals can have adequate habitat .
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shallow water in lower parts of artesian spring outflow living together with species of fonscochlea .
additional information on the biology and ecology of members of this family can be found in fauna of australia , vol . 5b , p . 752 - 755 .
found in the middle springs , south - western springs and the northern springs of the northern south australian artesian spring complex .
shallow water in lower parts of artesian spring outflow living together with t . minuta and species of fonscochlea .
freeling springs complex , northern part of lake eyre supergroup , northern south australia .
urn : lsid : biodiversity . org . au : afd . taxon : 8f476ec3 - a43f - 4b37 - b6d5 - 05d42d628231
urn : lsid : biodiversity . org . au : afd . taxon : f61be664 - 8697 - 46a0 - b732 - b0feb668f7d8
urn : lsid : biodiversity . org . au : afd . name : 278919
nicholas p . murphy , martin f . breed , michelle t . guzik , steven j . b . cooper and andrew d . austin
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