alasea is a genus of moths in the choreutidae family , containing only one species , alasea corniculata , which is known from costa rica .

the length of the forewings is 4.5 – 5.2 mm for males and 5.2 – 5.7 mm for females .

alasea

alasea is an professional mermaid , model , actress , artist , vocalist and cosplayer .
alasea is located in seattle , washington . this organization primarily operates in the museums and art galleries business / industry within the museums , art galleries and botanical and zoological gardens sector . this organization has been operating for approximately 4 years . alasea is estimated to generate $ 47 , 581 in annual revenues , and employs approximately 1 people at this single location .
hi ! i am alasea , i am a professional mermaid with blue mermaid designs . i also do arts & crafts , singing , modeling , acting , and voice over work .
alasea corniculata was described by rota in 2008 ( see reference to original description below ) from specimens collected in costa rica . the holotype is held by inbio . this is a small moth with forewing length of about 5 mm . it has dark forewings with iridescent specks of scales and a yellow - orange hindwing .
the kingdom of vetulia quickly emerged from the 7th century b . c . as the undisputed military power in a ' vassortivēa . eventually growing immensely wealthy from sea and overland trade with the hevlātian tribes beyond the kutzgrōd mountains , the vetulians began to set their sights on securing control of their borderlands . targeting what would become the modern province of varia and the italorian peninsula , in 574 b . c . a succession of expansionist vetulian kings made use of military alliances with the hevlātian tribes to crush a series of barbarian confederations occupying the eastern lowlands along the kutzgrōd . with the fall of the last bel ' akian confederation stronghold , alasea , in 536 b . c . , vetulian expansion began to secure the benefit of agricultural independence to the growing empire ( immeasurably furthered by the 510 b . c . conquest of the fertile western plains of modern athenia ) .
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i somehow ended up with a copy of an ebook and can ' t find my original book ( i ' m almost 80 % sure it ' s still in storage with a few hundred other books ) , so i ' m rereading the series . i have no map though . no problem right ? i ' ll just go to the author ' s site and find the map there . urltoken the map is so degraded it ' s pretty much unreadable . does anyone know of another copy of the map so i don ' t have to brave my storage unit to try and find the books ? thanks !
hi lori . i found my copy of wit ' ch star if you add me to you permissions list for private message , i ' ll shoot you a scan of the map .
you sir , are a truly good person . thank you so much for this !
let me just see if i can dig up my paperback . i ' ve no idea where they get up and walk off to my books !
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11 , 877 white 3 , 792 hispanic 4 , 800 black 4 , 036 asian 659 native american 1 , 007 hawaiian 2 , 326 other 51 . 4 % white 16 . 4 % hispanic 20 . 8 % black 17 . 5 % asian 2 . 9 % native american 4 . 4 % hawaiian 10 . 1 % other
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the holy pontifical union of astavia ( latin : sancta pontificia ūnio astaviae - astavian high latin : svēti papinsko sávek astáviz ) ( commonly reffered to as astavia , the pontifical union , astavian union , and the h . p . u . ) is a unitary theocratic republic comprised of its main territorial holdings on the efian continent , in addition to various overseas possessions . primarily situated on efia proper , the pontifical union also wields administrative control of the papal states of akkasti and livonia in southern ardas and northern euraso respectively , as well as a military outpost administered by a papal prelaturate on the st . arak islands . bordered to the north by the arctic ocean , and in the east and west by the sassani and halcyon oceans respectively , astavia shares only two land borders with the mid - efian states of auraliae and petruvia . comprised of a land area of 9 , 257 , 161 km2 ( 3 , 574 , 209 sq mi ) , astavia is the largest nation on the efian continent , and in undria by landmass . with 112 . 3 million inhabitants , astavia is also undria’s most populous nation , and is the only country that shares a geographic border with both the arctic and sassani oceans .
astavia is home to one of the world’s oldest civilizations . first settled by nomadic tribes 9 , 000 years before the founding of the ancient city of vetulia , the plains of northern efia eventually became home to numerous diverse polities by the end of the 9th century bc . with the settled population primarily centralized in the lowlands , various powerful kingdoms emerged in the 7th and 8th centuries bc - particularly the kingdom of vetulia , which dominated overland trade routes throughout varangia , as well as river trade along the vestria river . eventually conquering most of the southern polities and securing a political union with the powerful tribal kingdom of hevlatia in 428 bc , the varāngik dominion soon extended its control over the northern mountains through a series of military campaigns , forming one of the largest ancient empires in undria ' s history .
long considered a nation of great geostrategic importance because of its domination over the mineral rich mountain ranges of northern efia , as well as its prominent position on veltari strait and the bay of tytherion , astavia ranks as one of the largest regional trading powers on efia , and has consistently been a leading economic force in undria since industrializing in the 19th century . the country boasts significant reserves of both rare earth minerals and natural gas , as well as a burgeoning industrialized sector focused intensely on arms manufacturing and research . it is also a major diplomatic player , wielding considerable influence over international security issues and energy policy in undria . vaticaina , the country’s capital and 4th largest city , is a leading center of economic and cultural exchange in undria . the seat of the holy see , as well as astavia’s primary governing institutions , vaticania also boasts one of the lowest overall urban crime rates in the world , despite dealing with a rising population living beneath the national poverty line .
the country is a founding member of the undrian regional council , formed in 1943 following the conclusion of the krumaarian succession crisis , and is a leading signatory of the avignon mutual assistance pact . astavia’s longstanding cultural legacy has been further reinforced by the designation of twelve global heritage sites by the urc within its borders . its unique political system , based on the 1884 constitution that introduced significant administrative restructurings of the previous papal state , blends elements of a parliamentary republic with a church - centered theocracy dominated by the country ' s clergy , wherein the supreme pontiff , the head of the ancient catholic church , wields ultimate influence over government activity . varāngiks , the ethnic descendents of the ancient varāngik dominion , constitute a vast majority of the nation’s ethnic makeup , although a small multitude of ethnic and linguistic groups also exist . most astavians identify themselves as practicing catholics in communion with the national church , and high latin ( a northern - efian based blending of varāngik latin and hevlatian ) is the official national language .
( reffering to the fertile plains north of the vassar river in modern aegia ) . the vassaskri were one of several migrant tribes who traveled beyond the vassar and settled along the vestria river . the term ' s association with the region of vassaskri settlement eventually gained it the alternative meaning of
( hence it ' s feminine form ) . the vassaskri were one of several tribes that eventually coalesced into the vetulian kingdom over the 7th century bc , and their development of a written language helped form the basis of a common tongue used by the vetulian royal court . by the 5th century bc , the earliest form of vassaskri - based proto - latin language was widely present in the expanding kingdom , and had become adopted as the chief administrative language for state recordkeeping . the term
gaston d ' ildaro ' s 15th century painting of the assandrian prince , paranus superbus ( center ) , presiding over a rik ' kai council following the collapse of the duonomi bloodline .
in 624 b . c . , the last vassaskrian king , galerion possoli died - leaving behind no direct heir to assume the throne . possoli ' s 12 year old nephew , klio seemed the next reasonable successor , but was quickly challenged by his older sister , alemia . married to the assandrian royal heir , julianos superbus , alemia ' s claim was strengthened not only by the loyalty of her father ' s troops , but also the threat of an assandrian intervention . seizing power in a relatively bloodless palace coup , alemia ascended to the vassaskrian throne the same year - swiftly conducting an alteration of the kingdom ' s laws on primogeniture succession to exclude consideration of gender . by the beginning of the 6th century b . c . , both the vassaskri and assandri lived under the rule of a unified royal dynasty , based in the old assandrian capital of vetulia .
the duonomi , charged originally with protecting the frontier along the vassar river , encountered disaster in 614 b . c . when the steppe army of ok ' loka the cruel smashed their defenses at solsid . crossing the river , ok ' loka ' s army rapidly advanced to the duonomi capital of foggia , laying waste to the countryside and routing local militias organized to halt their advance . eventually capturing foggia following a short siege , ok ' loka ' s army subjected the city to a horrific sacking and brutally massacred most of the duonomi royal family . an assandrian army under the command of crown prince paranus superbus gathered strength and moved south to prevent the loss of any more duonomi lands to the steppe invasion - soundly defeating ok ' loka ' s main force at the battle of zaria in 612 b . c . the assandrians quickly crushed the rapidly dissipating steppe raiders , and regained control of the vassar river plains . exploiting the substantially weakened position of the duonomi family , the assandrians assumed military control of the region - subjecting the duonomi to the status of a protectorate state .
in 428 b . c . the vetulian king , dominian akrasti , married helena superba , daughter of hevlātian king othorogorus . with othorogorus’ death in 406 b . c . , dominian ascended to the hevlātian throne . the unification of the hevlātian and vetulian kingdoms under the control of dominion in 406 b . c . is considered the end of the monarchal period , and the beginning of the varāngik dominion by most modern historians ( although the degree of cultural assimilation between the two kingdoms leaves some room for debate as to the exact year both states considered themselves one and the same empire ) .
over the next century , the sons of primarch dominian carried on their father’s wars of imperial expansion . in addition to presiding over transkyrior khūzian for 10 years before ascending to the primarch ' s throne , general tarquinias akrasti issued numerous reforms for the dominion’s army . standing armies of professional soldiers were established , military standards for training and equipment were made universal , lines of communication improved , the chain of command was streamlined , and strict codes of conduct were introduced . professional troops quickly became the backbone of the dominion’s military power on the continent . during primarch tarquinias’ rule , the dominion expanded across the sūtgorod mountains towards the northern coastline of the continent . the primarch ' s brother antonais colonized the island of edia in 389 b . c . , and antonias ' son justinarius consolidated the dominion’s control over the far western provinces by pacifying the native dulcedi and botissi tribes . by 320 b . c . most of the northern half of the efian continent had been brought under the control of the dominion .
in 317 b . c . however , king acenatrix of the gutulian confederation ( at the time , a powerful tribal alliance based in what is modern - day etruscia province ) rallied his people to arms after proconsul julius maxentius ( who had been who had been appointed governor or hevlātia ) ordered the destruction of the gutullian border city , averni , in response to repeated gutullian raids on dominion trading caravans .
the gutulian onslaught into dominion territory was unexpected , successfully beating back the inexperienced garrison forces occupying the border cities of northern hevlātia . in 316 b . c . general rutullius maximus was given command of four
( armies ) and traveled north to deal with the barbarian threat . when the dominion reinforcements arrived , the gutullian forces soon found they could not defeat organized armies in open combat - experiencing a decisive defeat at the battle of nos essa . for the next 6 years , the gutullian confederation bled rutullius’ stratiae in small skirmishes in the freezing highlands of the continant’s northern coast . in 309 b . c . , rutullius’ prized , yet severely weakened iii stratia was crushed by a massed gutullian attack at the battle of gliara pass . general rutullius , struck down by an arrow himself during the battle , was relieved of his command .
general tiberius romulus akrasti , the grandson of primarch justinarius akrasti , was then given command of the disheartened dominion force . bolstering the other three stratiae with the survivors of the iii stratia , tiberius set out to starve the gutullians into submission . the arid highlands that the gutullians called home had little land suitable for farming . while the dominion’s navy ( which held uncontested control of the seas ) denied oversea trade to the gutullians , tiberius’ stratiae laid waste to every farm and city they came upon . within two years , tiberius’ armies had pushed the gutullians back to their capital brundisium . starved of food , suffering from disease , and broken in spirit , the people of the city rebelled against king acenatrix , and surrendered the city to the dominion . with the fall of brundisium and the capture of the gutullian leadership , the confederation quickly collapsed . over the next three and a half centuries the emperors of the varāngik dominion held unconditional control over the northern half of the efian continent .
the year 0 a . d . marks the moment the first catholic missionaries set foot on the dominion’s soil . sent by the ailing church in other countries , the missionaries spread far and wide throughout the dominion . the appeal of universal salvation that the missionaries preached took hold over the dominion’s masses ( most of whom had little in the way of “paradise” awaiting them in the next life under the teachings of the polytheistic state religion ) . priests in the cities of the dominion were elevated to bishops , and gained public favor in numerous provinces . as the christian population grew , a church hierarchy began to develop within the dominion . in 12 a . d . , pope gregory i , recognizing the potential for the creation of a christian empire , moved his papal court to the dominion city of avignon .
with the permission of the dominion ' s government , pope gregory began preaching in avignon , and from there coordinated the larger missionary effort . for the most part , the state religion of the dominion had been a polytheistic blend of local religious beliefs and the old vetulian gods - reflecting the empire ' s tendency to incorporate various faiths into the rational of the state religion rather than force conversion . thus the dominion government ' s initial attitudes towards the growing christian population were , more often than not , benevolent .
the city of virakia ' s statue of primarch severus is a popular pilgrimage site and national monument .
primarch austark ii , hoping to cull the growing power of the church and return the dominion to its adherence to the pagan vetulian pantheon , began a persecution of the christians and other religious groups in 60 a . d . - claiming they had tried to set fire to parts of the capital vetulia to destabilize the dominion’s government in the aftermath of the temurid crisis . outrage ensued throughout much of the newly christianized portions of the empire . chaos reigned in several provinces as provincial governors either chose to send troops to put down rebellions staged by their countrymen , or keep their severely depleted forces in the safety of their barracks .
severus ' bond with liara however would be tragically severed through a series of events that eventually culminated in the severan rebellion . in what they had hoped would be a move to protect their family , severus and liara ' s eldest daughter amyra was wed to the primarch ' s nephew , tyrannius austark - at the time , a senator and former military tribune residing in vetulia . as civil strife began to mount in resistance to austark ' s religious pogroms , the primarch began to use close family members of provincial governors trapped in the capital as leverage to ensure their loyalty . when severus ' wife liara and her entourage were refused free movement out of the city after visiting her daughter in the spring of 63 a . d . , severus unsuccessfully pleaded with the primarch to let her return to athenia .
depicts severus ( shown left in gold ) decimating the pagan forces of primarch austark ii .
an 11th century depiction of the
donation of severus ,
which adorns the walls of the sinistrian chapel within the apostolic palace .
upon his deathbed in 76 a . d . , primarch severus , ordered in a letter bearing the seal of the imperial household that custodianship for the dominion be turned over to the church - angering the remnants of the imperial nobility , and signaling the end of the dominion ' s line of primarchs from the aristocratic class . following the deaths of his wife and daughter in the severan rebellion , a grieving severus vowed never to marry again , and committed himself to faithful chastity in memory and honor of the woman he loved totally . much to the distaste of the imperial nobility , not only did severus fail to produce offspring to succeed him , he also openly abandoned his faith in the imperial pantheon - receiving the sacrament of baptism into the church in a ceremony conducted by pope gregory ii in 74 a . d .
long before his death , severus ' imperial administration came to rely heavily on educated clerics , produced by an ever expanding system of church schools - progressively forcing the empire ' s secular nobility to the margins of power . laws were passed elevating catholicism to the imperial faith , though severus himself never outlawed the ancient pagan pantheon . by the year of severus ' death , historians estimate roughly seven out of ten offices in the dominion ' s civil administration were held by ordained members of the catholic clergy , while 8 stratiae out of a standing force of 13 were led by specially appointed cardinal - tribunes .
declining immigration rates prompted the government to enact several reforms in 2008 to simplify access to astavian citizenship . following the 2008
immigration reform act ,
catholics abroad seeking to obtain astavian citizenship may claim a temporary citizenship status , and upon completion of a three year period of law - abiding residence , can be granted full citizenship . an amendment to the act , attached by the astavunitis wing in the npc , stipulated however that this offer of full citizenship was contingent on whether the immigrant renounces allegiance to all other foreign powers , and forfeits all other citizenships .
the astavian government , formed by 1884 constitution , is officially a unitary theocratic republic - composed of an intricate collection of governing bodies , divided conventionally into institutions that exercise executive , legislative , and judicial powers . separated into civilian and ecclesiastically - administered sectors , ultimate
custodianship
of the nation rests in the person of the supreme pontiff , who is empowered to oversee the country ' s national interests , while the day - to - day management of the country ' s legislative work and civil administration are entrusted to an elected legislature and various civilian institutions . though governing power has historically been highly centralized in some form of national government for much of astavia ' s history , since 1940 various powers have been granted gradually to provincial councils to govern on local matters , while county and municipal councils have been extensively involved in local governance and the maintenance of public order since the early 1880s .
under the charter promulgated during the 1884 great constitutional reform , the pope is the pontifical union ' s ex officio head of state and commander - in - chief of the armed forces . in addition to his primary role as bishop of the diocese of vaticaina , the supreme pontiff serves as the primate of varangia ( an honorary title reserved for the pope ) , and the head of the holy see ( and thus , the astavian catholic church ) . as a general term , the holy see represents not only the temporal power of the pope , but also his spiritual and pastoral authority over ecclesiastical and civil matters . the supreme pontiff serves a life term following his election by the college of cardinals during conclave , and may not be removed from office after his ascension to the papacy ( though voluntary resignation is legally permitted ) . the pontiff wields broad administrative powers , including the right to issue papal bulls that carry the full force and effect of national law ( when so stipulated ) , and the power to declare war and offer peace . despite this official concentration of authority , astavian pontiffs have largely delegated most powers to legislative procedure and civilian oversight . pope
, born gregory o ' hare , was elected on the 19th of may , 2014 . prior to his election , cardinal o ' hare served as the
, and has since become a polarizing figure in undria . pope julius ' vigorous focus on serving the poor and marginalized portions of astavian society was seen as the primary initial focal point of his pontificate until international events largely overshadowed most of his domestic work . the
kicked up an international firestorm in mid 2014 , which saw a major falling out between the pontifical union and its southern neighbor , the federal republic of auraliae . among more positive circles , pope julius is regarded as a champion of christian self - determination and a protector of catholic communities around the region . less enamored observers have frequently characterized his military exploits in belmia as that of a
hapless warmonger ,
driven by a misguided
crusading mentality , and a dangerous sense of
catholic supremacism
.
consisting of 12 cardinals is charged with the review of all legislative proposals passed by the npc . the esc may only veto legislation by unanimous agreement , though it is empowered to attach
mandatory amendments
to proposed legislation . such amendments , carrying the support of two thirds of the cardinals on the committee , function as editorial prerequisites for a bill ' s enactment . the amended bill , if not challenged by the national people ' s congress over a period of ten days , becomes law . if the npc challenges the amended bill , it may move to the floor for debate again , followed by a
nullification
vote . a four - fifths vote may override an esc amended bill , rendering it
( a dead law ) . at any time , the npc and esc may appeal to the supreme pontiff to strike down or approve the law . the esc is also responsible for vetting and approving candidates for political office appointments , and national elections . citizens wishing to run as a candidate for one of the nationally recognized parties must be reviewed by the committee , and successfully petition the support of three cardinals ( and not opposed by more than four ) . committee members ( who serve septennial terms ) are exclusively appointed by the church , with six directly appointed by the supreme pontiff , and six elected from within the college of cardinals .
within the church itself , a vast bureaucracy works in coordination with civilian government branches to facilitate effective governance . the curia of vaticaina is comprised of ten congregations , thirteen councils , four secretariats , three tribunals , and one prefecture . among the most active of the major curial institutions are the congregation for the evangelization of the peoples , which oversees the catholic church ' s various missionary activities across undria ( and within the astavian nation itself ) , and the congregation of divine mercy , which functions as the nation ' s primary social aid provider . most astavian social welfare , job training , food stamp , and public assistance programs fall under the administration of the congregation of divine mercy ( typically working with funds provided by the apostolic camera ) , although standards of national healthcare fall under the jurisdiction of the national health council .
rests with the supreme pontiff , who selects a candidate from a list of four . two names are submitted by the chairman of the esc , and two more by the consul ( both the chairman and consul may concur on candidates , and stipulate such in the submitted list ) . in turn , the excelsus advocatus directly appoints the 7 justices of the supreme judicial council and the chief public prosecutor . courts in astavia are largely divided along secular and religious lines . for the most part , the secular courts of the nation bear the brunt of the civil cases and criminal prosecutions that make their way through astavia ' s judicial system on a yearly basis . at the local level , 112
administer justice for both criminal and civil cases . decisions made at the district level may be appealed to one of 17
, which have appellate jurisdiction over specific regions across the country . at the highest civil level , the 7 - judge panel of the
, exercises total authority to reconcile discrepancies ( constitutional or otherwise ) between decisions made by circuit courts .
three established tribunals exercise various forms of judicial power within astavia ' s catholic church . normal judicial appeals on sacramental matters ( such as marriage annulment ) , may work their way up through local church courts to the vetulian rota . some civil cases , if agreed upon by both parties , may also be taken to the vetulian rota provided both parties are practicing catholics in good standing with the church . the high signatura is the supreme appellate and administrative court of the astavian see , wielding jurisdiction over decisions made by the vetulian rota , as well as the administrative decisions of church prelates . additionally , it oversees and regulates the legal framework that innervates the curia ' s administration . the apostolic suprema council ( colloquially referred to as the cloistered council ) deals primarily with matters of conscience and morality . officially , the suprema is the judicial wing of the office of the holy inquisition , which generally falls under the broad categorization of a curial institution , though its administration and operation is largely kept separate from the curia itself .
astavian catholics may also elect ( so long as both the plaintiff and defendant assent ) to have civil cases brought before an ecclesiastical court , officially convened by a bishop , but typically administered by a designated local prelate . decisions by ecclesiastical courts may be appealed to one of 13 canon courts ( one for each province , and one for overseas territories ) , which are each overseen by three appointed canon judges . these courts fall under the supreme jurisdiction of the high signatura , the decisions of which ( since they are carried out on direct papal approval ) , cannot be appealed . while ecclesiastical courts are not empowered to hear criminal cases under normal circumstances , the apostolic suprema council of the ohi wields loose powers to conduct trials and carry out sentencing for certain categories of offenses , including crimes endangering national security , treason , and heresy . like the high signature , the suprema ' s decisions are final and cannot be appealed - rendering it ultimately answerable only to the supreme pontiff . the curate ' s court of review , a sub - court of the ohi , handles crimes allegedly committed by clerics and citizens closely associated with the church . the ohi ' s jurisdictional claim to legitimately investigate , detain , prosecute , and punish clerics and other church officials has been the subject of considerable criticism between catholic nations in undria , particularly in cases where the alleged offending party has been a foreign national who sought refuge in his or her home country .
paf su - 27s flying a patrol run near the astavian - auralian border during the belmian independence crisis .
. in a 2009 administrative audit , the military reported 377 , 000 personnel on active duty , as well as roughly 267 , 000 trained reservists in the home guard . the
, a separate regime security force answering directly to the pope , is estimated to number around 187 , 000 troops , divided amongst 13 infantry divisions , 3 armored divisions , various independent / specialized brigades , and several classified special operations units . the pontifical union also possesses a paramilitary volunteer militia , operating under the direction of the
, of whose 4 . 7 million civilian members , 158 , 000 are uniformed active duty personnel dispersed throughout the country . in wartime however , papal decree may place the order under the overall command of the apostolic guard .
the pontifical union also boasts a vast and fully indigenous arms industry , producing most of its own military equipment , and exporting considerable amounts of hardware to regional allies and organizations . the international arms overwatch committee of the undrian regional council estimated that small arms produced by the pontifical union accounted for at least 31 . 7 % of all traded firearms in 2013 . the same 2013 report concluded that other astavian military hardware accounted for roughly 29 . 2 % of the undrian arms economy .
this page was last modified on 20 january 2018 , at 15 : 51 .
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. . . we tested the best partition model based on protocols intro - duced by rota & wahlberg ( 2012 ) , using bayes factor ( kass & raftery , 1995 ; fan et al . , 2011 ; xie et al . , 2011 ) , partitioning the data first by gene ( the traditional approach , summing up to nine partitions ) and then using tiger ( cummins & mcinerney , 2011 ) . for this , we used tiger v 1 . 02 ( cummins & mcinerney , 2011 ) to calculate the relative rates of evolution of each site in the combined dataset . . . .
. . . lanfear et al . , 2012 ) which has been shown to improve various aspects of phylogenetic inference ( e . g . see poux et al . , 2008 ; rota & wahlberg , 2012 ; leavitt et al . , 2013 ) . partitioned analyses were carried out using partitionfinder 2 . 0 . 0 ( pf ; lanfear et al . , 2012 ) with the raxml option to determine optimal partitioning schemes and best - fit models of substitution for both aa and nt data . . . .
. . . for ml analyses , we used the gtr model of substitution with gamma model of rate heterogeneity ( gtr + g ) and different partition schemes , either gene - based or based on rates of evolution calculated by the program tree independent generation of evolutionary rates ( tiger ) ( cummins & mcinerney , 2011 ) . in gene - based partitions each gene was considered as a separate partition , while in tiger - based partitions , the characters were binned together based on their rate of evolution regardless of gene origin ( used as a partitioning strategy in rota & wahlberg , 2012 ) . tiger partitions for the dataset were derived from the program tiger v1 . 02 that calculates relative rates of evolution of each site in an alignment ( cummins & mcinerney , 2011 ) . . . .
. . . these markers have repeatedly been proven useful in resolving evolutionary relationships of different families of lepidoptera ( e . g . sihvonen et al . 2011 , rota & wahlberg 2012 , rönkä et al . 2016 , õunap et al . 2016 , zenker et al . 2016 ) . pcr was performed in a total volume of 20 μl , with the reaction mixture containing 1x bd advantage 2 pcr buffer , 1u bd advantage 2 polymerase mix ( bd biosciences , san jose , usa ) , 0 . 2 mm dntp ( thermo scientific , pittsburgh , usa ) , 5 pmol of primers and 20 - 80 ng of purified genomic dna . . . .
. . . these markers have repeatedly been used in reconstructing phylogenies of different families of lepidoptera ( e . g . wahlberg & wheat , 2008 ; rota & wahlberg , 2012 ) , including geometridae ( e . g . yamamoto & sota , 2007 ; wahlberg et al . , 2010 ; sihvonen et al . , 2011 ) . . . .
. . . second , dividing nucleotide data into partitions according to their actual rate of evolution instead of some predefined criteria ( e . g . genes or codon positions ) has been shown to improve the results of phylogenetic analysis ( rota & wahlberg , 2012 ) . third , sampling taxa at least pairwise for evolutionarily distant lineages ( tribes in the current case ) helps to reduce the negative effects of ' long - branch attraction ' and thereby improve the result of the phylogenetic analysis ( swofford et al . , 1996 ; bergsten , 2005 ; hedtke et al . , 2006 ) . . . .
. . . in contrast , datasets on the order of tens of genes have been highly useful for resolving relationships at the intrafamilial level , as has been repeatedly shown for e . g . lepidopteran families ( wahlberg et al . 2009wahlberg et al . , 2014 kaila et al . 2011 ; kawahara et al . 2011 ; sihvonen et al . 2011 ; zahiri et al . 2011 zahiri et al . , 2012 zwick et al . 2011 ; regier et al . 2012a regier et al . , 2012b rota and wahlberg 2012 ; sohn et al . 2013 ) . thus , datasets generated with pcr - based methods have been and continue to be very insightful . . . .
. . . like ov , tiger has been applied to selectively delete putatively fast evolving characters from genomic datasets ( e . g . , morgan et al . , 2014 ; xi et al . , 2014 ; katz and grant , 2015 ) . but unlike ov , tiger has also been used to partition characters into rate categories for parametric analyses ( e . g . , rota and wahlberg , 2012 ; heikkilä et al . , 2014 ; nakov et al . , 2014 ) . typically character removal is performed by excluding one or more of the ten rate categories that tiger delimited , with different numbers of characters assigned to each category ( e . g . , greene et al . , 2014 ; morgan et al . , 2014 ) . . . .
. . . this group is an excellent study system for investigating this phenomenon because they have wingspans of about 10–15 mm and they are globally distributed ( table 1 ) . moreover , a robust eight - locus phylogeny of metalmark moths has already been published ( rota & wahlberg , 2012 ) . we significantly improved the existing data set that included 38 species from 13 genera representing mainly the neotropical , nearctic , oriental and palaearctic regions by adding another genus and over 60 species , many of which are from africa and australasia . . . .
. . . we have undersampled brenthia from the oriental region and possibly from the aftrotropics , and if the unsampled species from one or both of these regions form a sister group to the remaining brenthia in the phylogeny , a different area of origin and ranges for some of the other nodes would be estimated as the most likely ( sampling details in appendix s1 ) . sequences partly came from published studies ( rota , 2011 ; rota & wahlberg , 2012 ; rota & miller , 2013 ) and from the barcode of life data systems ( bold ) for the dna - barcode fragment , but were largely obtained de novo for this study . genbank accession numbers and sequence length for each fragment / species can be found in appendix s2 . . . .
. . . the phylogenetic hypothesis for the family is robust and is essentially the same as in rota ( 2011 ) , rota & wahlberg ( 2012 ) , and rota & miller ( 2013 ) ( see appendix s2 ) . out of the 103 nodes in the tree , 68 were strongly supported by both mrbayes and raxml analyses ; another nine nodes received moderate to strong support in both analyses ; 14 nodes had weak , moderate or strong support by at least one of the methods ; and only 12 nodes had no support in either of the analyses . . . .
. . . brandley et al . ( 2005 ) investigated the effect of nine alternative ad hoc partitioning strategies on model fit , topology and node support in bayesian framework and found variation in inferred tree topologies at weakly supported nodes . other studies have noted that partitioning scheme choice affects node support ( ward et al . 2010 ; rota and wahlberg 2012 ; powell et al . 2013 ) , tree topology ( strugnell et al . 2005 ; leavitt et al . 2013 ; tao et al . 2013 ) , and branch - lengths ( poux et al . 2008 ; ho and lanfear 2010 ) . occasionally the choice of partitioning scheme has little or no effect on tree characteristics at all ( cameron et al . 2012 ) . . . .
. . . although these studies demonstrate that partitioning can have important effects on phylogenetic inference , it is difficult to draw any general conclusions from them . the partitioning schemes compared within each study were selected on an ad hoc basis with very different approaches ; some studies compared a small set of very similar partitioning schemes ( e . g . , cameron et al . 2012 ) , and others compared extensive sets of very different partitioning schemes ( e . g . , rota and wahlberg 2012 ; leavitt et al . 2013 ) . furthermore , previous studies used a variety of different phylogenetic methods , software implementations , and tree inference methods . . . .
. . . using the program tiger ( cummins & mcinerney , 2011 ) , all of the sites in the alignment were sorted into 80 bins based on their relative evolutionary rate , with invariable sites being sorted into bin 1 , and the fastest - evolving sites into bin 80 . the bins were then combined into partitions as described by rota & wahlberg ( 2012 ) , resulting in eight partitions ( partition 1 contained the slowest - evolving sites and partition 8 the fastest - evolving sites ) . in some of the analyses , as described later , the fastest partitions were excluded . . . .
. . . the tree shown infig . 5 is the result of the analysis of the dataset ( full _ ex78 ) from which the fastest - evolving sites were excluded using the program tiger ( cummins & mcinerney , 2011 ) , partitioned using a previously described procedure ( rota & wahlberg , 2012 ) , and analyzed with mrbayes v . 3 . 2 ( ronquist et al . , 2012 ) . the topology is identical to that recently presented in regier et al . ( 2013 ) , except that in the latter , the eriocraniidae appear as sister group to all other glossata . . . .
. . . but the influence of increasing the number of partitions on phylogeny was minimal , since most relationships found with nine and 22 partitions were the same , except for the resolution within vespa ( figs . 4c and 5 ) . similar issues with convergence and mixing related to a priori partitioning were reported by rota and wahlberg ( 2012 ) in their phylogenetic study of metalmark moths . . . .
we aim to understand better the phylogenetic relationships of the geometridae in the neotropical region . we already have sequenced ca . 150 genera but we still seek certain taxa for our work . i work…
[ more ]
molecular phylogeny , systematics and the evolution of caterpillar - ant interactions in metalmark butterflies .
the parasitoid wasp family ichneumonidae has been considered as an exception to the latitudinal diversity gradient . however , based on long - term and large - scale primary data , we have shown that the …
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note on taxonomic history , thoraco - abdominal articulation , and current placement of millieriidae ( in . . .
immature stages of metalmark moths from the genus brenthia clemens ( choreutidae ) : morphology and lif . . .
in this paper the immature stages of brenthia monolychna meyrick ( choreutidae : brenthiinae ) , as well as their ultrastructure , are described and figured . this is the first description of a new world brenthiine , in addition , notes on life history for four new world species of brenthia clemens are provided , including mention of their host plants and parasitoids . host plant utilization of the . . . [ show full abstract ]
data partitioning in bayesian analysis : molecular phylogenetics of metalmark moths ( lepidoptera : cho . . .
in this study a multilocus phylogenetic analysis of metalmark moths ( lepidoptera : choreutidae ) focused on resolving the higher - level phylogeny of this group is presented . through the analysis of this dataset , i explore different data - partitioning strategies in bayesian phylogenetic inference , and find that a partitioning strategy can have a large influence on the results of phylogenetic . . . [ show full abstract ]
phylogeny of the dagger moths ( lepidoptera : noctuidae : acronictinae : acronicta ) and the evolution of . . .
highly variable larvae in the genus acronicta prompted a . r . grote to proclaim
there would seem to be no genus which offers a more interesting field to the biologist for exploration
( grote , 1895 ) . along with related genera in the subfamily acronictinae , acronicta underwent a whirlwind of taxonomic revision in its infancy , rife with synonyms and rivalries between lepidopterists . controversies . . . [ show full abstract ]
icanheartheoceancallingme : i love the tails he makes , his use of colours is amazing . | mermaids and mythology < 3 photography and videos | pinterest | mermaid , …


scotinomys is a genus of rodent , the singing mice , in the family cricetidae .

together with baiomys , it forms the tribe baiomyini .

it contains the following species : alston 's brown mouse ( scotinomys teguina ) chiriqui brown mouse ( scotinomys xerampelinus ) they are found in mountainous areas in central america , at altitudes of 1000 m to at least 3500 m .

as their common name indicates , they are notable for their acoustic communication .

they are insectivorous .

the two species show substantial divergence in behaviour and reproduction , with s. xerampelinus generally dominant over s. teguina where the species occur together .

scotinomys

kari pihlaviita added the finnish common name
guatemalanruskohiiru
to
scotinomys teguina ( alston , 1877 )
.
reproduction , growth and development in two contiguously allopatric rodent species , genus scotinomys . misc . publ . mus . zool . univ . mich
citation : campbell p , pasch b , warren al , phelps sm ( 2014 ) vocal ontogeny in neotropical singing mice ( scotinomys ) . plos one 9 ( 12 ) : e113628 . urltoken
neotropical singing mice ( scotinomys ) are diurnal , insectivorous rodents that inhabit the cloud - cloaked mountains of middle america . males commonly emit elaborate vocalizations to attract mates & repel rivals . please visit urltoken for more information on current research
dimitri v . blondel , jorge pino , steven m . phelps ; space use and social structure of long - tailed singing mice ( scotinomys xerampelinus ) , journal of mammalogy , volume 90 , issue 3 , 2 june 2009 , pages 715–723 , urltoken
comparisons of mean home - range sizes of scotinomys xerampelinus across seasons for a , c ) 85 % minimum convex polygon and b , d ) 100 % minimum convex polygon . data shown are means ± 1 se . all comparisons are not statistically significant ( p > 0 . 05 ) .
home ranges for individual scotinomys xerampelinus located ( captured or fixed via telemetry ) on the study grid ≥4 times for ( a ) 2003 , 85 % mcp home ranges , ( b ) 2003 , 100 % mcp home ranges , ( c ) 2004 , 85 % mcp home ranges , and ( d ) 2004 , 100 % mcp home ranges . home ranges are calculated using the arithmetic mean method . home ranges for males ( solid lines ) and females ( dashed lines ) are shown ; 10 - m scale lines are indicated .
here , we characterize the vocal development of two sister species of neotropical mice in which vocal communication plays a major role in adult social behavior . commonly referred to as singing mice , scotinomys teguina and s . xerampelinus are small ( 10–15 g ) diurnal , insectivorous muroid rodents . both species are restricted to cool , montane habitats in middle america ; s . teguina ranges from southern méxico to panamá at 1000–2930 m , whereas s . xerampelinus occurs only in costa rica and panamá where it replaces s . teguina at altitudes between 2200 and 2900 m [ 31 ] .
comparison of expected versus observed areas of home - range overlap of scotinomys xerampelinus . analyses are based on 85 % minimum convex polygons calculated with the arithmetic mean method . comparisons of expected and observed values were according to batzli and henttonen ( 1993 ) . obs . overlap = observed home - range overlap on the study grid . exp . overlap = expected area of home - range overlap based on random placement of home ranges on the study grid . means are given ± 1 se . ♂♂ = males overlapped by males ; ♂♀ = males overlapped by females ; ♀♀ = females overlapped by females ; ♀♂ = females overlapped by males .
mean ± 1 se expected ( random ) and observed areas of home - range overlap of scotinomys xerampelinus . analyses are based on 85 % minimum convex polygons ; data from 2003 and 2004 were pooled . observed = observed average area of home - range overlap on the study grid . expected = average area of home - range overlap predicted from random placement of home ranges on the study grid . an asterisk ( * ) indicates p < 0 . 05 , wilcoxon sign rank comparisons of observed and expected areas of overlap . n = 10 males and 10 females . ♂♂ = males overlapped by males ; ♂♀ = males overlapped by females ; ♀♀ = females overlapped by females ; ♀♂ = females overlapped by males .
during the 1st field season , both livetrapping ( “trapping localities” ) and radiotracking ( “fixes” ) data were used to characterize individual home ranges . these data were collected over a period of 18 days , from 19 august to 5 september 2003 . because of technical problems , including receiver failure , we were unable to collect telemetry data during the 2nd field season , which continued over a period of 35 days , from 24 may to 27 june 2004 . thus , only livetrapping data were used for the 2nd field season . this difference in data sets should not have affected our statistical comparisons of home - range attributes , because our method of overlap analysis generated both expected and observed values from the same data set ( 2003 : livetrapping and radiotelemetry ; 2004 : livetrapping only ) . although our data collection was limited to may , june , august , and september , the mountainous regions of western panama have consistently wet weather throughout the year and scotinomys reproduces aseasonally ( hooper and carleton 1976 ) , suggesting that space use by members of the study population should not vary greatly over the course of the year .
study population attributes . —in 2003 , 24 adults ( 9 males and 15 females , sex ratio = 0 . 6 : 1 ) were trapped on the study grid . in 2004 , 20 adults ( 9 males and 11 females , sex ratio = 0 . 8 : 1 ) were trapped on the study grid . the density of residents ( i . e . , animals captured ≥4 times on the study grid ) in 2003 , when vegetation was overgrown , was 62 mice / ha ; in 2004 , when vegetation was grazed , density was 28 mice / ha . females were found to be pregnant or lactating or both from june to september 2003 and may to august 2004 ; this represents the entire study period as well as some pre - and poststudy trapping of animals on the site . this finding is consistent with the data of hooper and carleton ' s ( 1976 ) indicating that reproduction by scotinomys is aseasonal . the ratio of scrotal males to perforate , pregnant , or lactating females was 9 to 13 ( 0 . 7 : 1 ) in 2003 and 8 to 10 ( 0 . 8 : 1 ) in 2004 .
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baiomyini . until diagnosed by thomas , species associated here had been affiliated with akodon ( e . g . , bangs , 1902 ) and later the genus with akodontines ( vorontsov , 1959 ) . common ancestry with baiomys inferred on the basis of shared morphological traits ( carleton , 1980 ; carleton et al . , 1975 ; hooper , 1960 ; hooper and musser , 1964 b ) and phylogenetic evaluation of mitochondrial and nuclear genes ( d’elía , 2003 ; engel et al . , 1998 ) , a relationship weakened by karyological banding data ( rogers and heske , 1984 ) . also see comments under baiomys . revised by hooper ( 1972 ) ; for other aspects of biology and systematics , see carleton et al . ( 1975 ) , hooper and carleton ( 1976 ) , and rogers and heske ( 1984 ) .
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i thought you might be interested in this item at urltoken title : mammal species of the world : a taxonomic and geographic reference author : don e wilson ; deeann m reeder publisher : baltimore : johns hopkins university press , ©2005 . isbn / issn : 0801882214 9780801882210 0801882389 9780801882388 0801882397 9780801882395 oclc : 57557352
wilson and reeder ' s mammal species of the world is the classic reference book on the taxonomic classification and distribution of the more than 5400 species of mammals that exist today . the third edition includes detailed information on nomenclature and , for the first time , common names . each concise entry covers type locality , distribution , synonyms , and major reference sources . the systematic arrangement of
information indicates evolutionary relationships at both the ordinal and the family level . this indispensable reference work belongs in public and academic libraries throughout the world and on the shelf of every biologist who works with mammals .
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this indispensable reference work belongs in public and academic libraries throughout the world and on the shelf of every biologist who works with mammals .
a uniquely valuable compendium of taxonomic and distributional data on the world ' s living and historically extinct mammalian species . contributors and editors alike deserve the thanks of all
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wilson and reeder ' s mammal species of the world is the classic reference book on the taxonomic classification and distribution of the more than 5400 species of mammals that exist today . the third edition includes detailed information on nomenclature and , for the first time , common names . each concise entry covers type locality , distribution , synonyms , and major reference sources . the systematic arrangement of information indicates evolutionary relationships at both the ordinal and the family level . this indispensable reference work belongs in public and academic libraries throughout the world and on the shelf of every biologist who works with mammals .
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editor : brenton g . cooper , texas christian university , united states of america
copyright : © 2014 campbell 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 .
data availability : the authors confirm that all data underlying the findings are fully available without restriction . all relevant data are within the paper and its supporting information files .
funding : the research was funded by grants from the national science foundation to smp ( ios 0548404 , career 0845455 ) . pc was supported by a national institutes of health nrsa fellowship from the national institute on deafness and other communication disorders ( f32 dc008269 ) . bp was supported by a national science foundation ddig ( 0909769 ) . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
offspring - to - parent signaling is among the most fundamental forms of communication [ 1 ] . for nocturnal mammals with altricial young ( e . g . , rodents , bats ) , acoustic signals produced by neonates ( isolation calls ) are integral to maternal localization and retrieval [ 2 ] , [ 3 ] , and can promote maternal behaviors such as grooming and nursing . thus , across distantly related mammalian orders , both the intended receiver and the function of isolation calls seem to be conserved , and are unique to the period of maternal dependence . while vocal communication remains important to the adult social behavior of many species , the receivers and functions of acoustic signals are diverse and context - dependent , ranging from alarm calling , territorial advertisement and mate attraction , to courtship songs that encode individual identity [ 4 ] , [ 5 ] , [ 6 ] . the functional disconnect between neonate and adult vocal behavior raises several interesting questions . for example , are the acoustic properties of isolation calls unique or are elements retained as the animal matures and the social context and function of vocal behavior changes ? conversely , what is the developmental time course of adult vocal communication ? are adult vocalizations produced de novo post - weaning or can the trajectory of vocal development be traced back to isolation calls ?
breeding pairs were housed in polycarbonate cages bedded with wood chips . the mice were provided with sphagnum moss and a pvc tunnel for nesting ; water and food ( kitten chow , seeds , dried beans and peanuts ) were given ad lib . pups were housed with both parents until weaning and litter sizes were not manipulated . a total of 30 s . teguina from 27 litters were recorded for this study ; 16 of these were the offspring of wild - caught parents , captured in cartago , costa rica , and 14 were the progeny of either first generation or wild - caught individuals from boquete , panamá . s . xerampelinus ( n = 15 pups , 9 litters ) were the offspring of first , second or third generation lab - reared individuals , derived from mice captured in parque internacional la amistad , panamá . all mice were housed in the same room . all animal protocols were approved by the institutional animal care and use committee at university of florida ( no . e436 ) .
mice were recorded between 9 : 00 am and 6 : 00 pm in a room acoustically isolated from the rest of the colony . the recording chamber was a polycarbonate cage bedded with clean wood chips inside a 42×42×39 cm opaque plexiglas cube lined with anechoic foam . one side of the cube was left open to allow behavioral observations and a microphone was fitted through a small opening in the ceiling , 25 cm above the cage floor . calls were sampled at a rate of 195 khz , 16 bits with an aco pacific microphone and tucker - davis hardware . file sizes were set to 15 s . fifteen seconds is > 2× the average duration of adult advertisement songs [ 29 ] , and reliably captured pup calling bouts in preliminary recording trials .
species , sex ( f / m ) and population means ( sd ) for litter size and developmental landmarks in singing mice .
we use “isolation call” to refer to any vocalization produced by a pre - weaning pup that does not comprise the rapidly articulated series of fm sweeps characteristic of the adult advertisement song . we delineated the end of the isolation calling period for each individual as the last day on which an isolation call was produced . we define a note as a continuous sound , and a calling bout as a series of three or more notes occurring with an inter - note interval of less than 2 s . notes were classified manually according to quantitative differences in duration , bandwidth , number of changes in direction of frequency ( 0 , 1 or ≧2 ) , and the presence / absence of nonlinear elements ( e . g . , deterministic chaos and bifurcations ; [ 37 ] ) .
all measurements were made in raven pro 1 . 3 ( bioacoustics research program 2008 ) . we analyzed a maximum of three files / individual / day and took the average of all variables . when more than three files were available we used the first three good quality files ( i . e . , high signal : noise ) from that recording session . in each 15 s file we measured overall maximum , minimum and dominant frequency , and bandwidth . when harmonics were present , maximum and minimum frequency were always measured from the fundamental frequency ( i . e . , the lowest frequency in the harmonic series ) . we manually counted the total number of notes in each file , the number of notes in the first calling bout , and the number of notes of each type . in addition , note duration , bandwidth , and maximum , minimum and dominant frequency were measured for each of three notes in the first calling bout in the file . to avoid bias in note choice we measured bout duration , divided by three , and took measurements from notes that fell in the middle of each third ( i . e . , for a 6 s bout we measured the notes closest to 1 . 5 , 3 . 5 and 5 . 5 s ) . internote interval was measured from end of each focal note to the beginning of the next note in the file .
principle components analysis ( pca ) was used to visualize the distribution of age - and species - specific vocal parameters in acoustic space . because the same mice were recorded at multiple ages , call data for a given individual were statistically non - independent . we controlled for non - independence in several ways . to test for age - specific differences within and between species , we binned recordings into age classes ( postnatal days 1–3 , 4–6 , 7–9 , 10–12 , 13–15 , 30 + ) , and analyzed vocalizations from a single recording session for each individual within each age class . for the three - day age classes , we retained data for all individuals that were recorded in the middle of each age range ( e . g . , day 2 for days 1–3 ) , and otherwise picked one representative day / individual / age class at random . adult songs were analyzed from a single recording session / individual ( mean age , sd : 40 . 2 , 6 . 1 days ) .
repeated measures analysis of variance ( anova ) was used to test the effect of age on vocal development over time . in this case , only individuals that were represented in each age class were included in the analysis . to test for correlations between body mass and frequency measures from birth to maturity , we randomly assigned each individual to a focal age , such that total age range was maximized and each individual was used only once in the analysis .
we used the coefficient of variation ( defined as the ratio of the standard deviation to the mean ) to describe change in vocal stereotypy across age classes . within each age class we took the average of the coefficient of variation for seven descriptors of frequency and timing ( maximum , minimum and dominant frequency , number of notes / bout , bout duration , note duration for a randomly selected note in the middle third of a bout , and internote interval to the next note ) from three different call files recorded from the same individual during a maximum time interval of three days ( birth to 15 days ) or 5 days ( > 30 days ) . in most cases , we were able to use calls captured during the same recording session . we used one - way anovas to test for species and sex differences in the coefficient of variation within each age class . the α value for all statistical analyses was set to 0 . 05 , with correction for multiple testing as appropriate .
in most individuals of both species , vocal behavior was developmentally disjunct with a non - vocal period separating the last recorded isolation call and the first production of the adult advertisement song ( table 1 ) . of the 30 s . teguina pups recorded in this study , 27 produced isolation calls when removed from the nest . the three pups that never called were males derived from the boquete population and were all from single pup litters . all 23 s . teguina that were recorded from birth to maturity produced adult songs , including the three males that did not call at earlier stages . of the 15 s . xerampelinus pups used in the study , one female from a single pup litter did not produce isolation calls . of the 11 individuals ( 5 females , 6 males ) recorded to maturity , three females and one male were never observed singing as adults .
unexpectedly , the largest differences in the duration of isolation calling and the timing of first adult song production were within s . teguina . on average , pups derived from the cartago population stopped producing isolation calls almost a week later than those from boquete ( mean , sd : 13 . 0 , 2 . 3 days vs . 7 . 5 , 1 . 7 days ; f 1 , 23 35 . 5 , p < 0 . 0001 ) and produced their first adult song over a week later than boquete mice ( 35 . 5 , 6 . 2 days vs . 26 . 7 , 9 . 7 days ; f 1 , 21 6 . 7 , p = 0 . 02 ) , whereas the duration of the non - vocal period did not differ between populations ( 22 . 1 , 6 . 5 days vs . 21 . 5 , 9 . 0 days ; f 1 , 18 0 . 03 , p = 0 . 9 ) ( table 1 ) . we asked whether these population differences in vocal development might be a secondary consequence of different rates of growth and development , or an artifact of social or sex - specific biases in our sample .
boquete litter sizes were smaller ( f 1 , 26 21 . 6 , p < 0 . 0001 ) and boquete pups ' eyes opened earlier ( f 1 , 39 167 . 0 , p < 0 . 0001 ) ( table 1 ) . therefore , between - population differences in vocal behavior could be explained by overall slower development in cartago mice due to proportionally less maternal resource allocation / pup . however , there was no effect of population on body mass for any age class ( all p ≧0 . 2 ) . likewise , the end of isolation calling was three days prior to when boquete pups opened their eyes , whereas these events were approximately coincident in cartago pups ( table 1 ) . we also considered that there might be an effect of litter size on motivation to vocalize ; pups in this study that did not vocalize had no siblings and a similar relationship between small litter size and low isolation call production was reported in microtus [ 22 ] . we could not test this hypothesis statistically since only one cartago single - pup litter was included in the study . however , this female produced isolation calls , albeit for a period less than the population mean ( 8 vs . 13 . 0 days ) . finally , the boquete sample was strongly female - biased for all pup age classes . therefore , if female s . teguina stop producing isolation calls earlier and start producing adult song later than males , population effects on vocal behavior might be an artifact of sex differences . within the cartago sample , however , there was no sex difference in the duration of the isolation calling period or age at first adult song production ( both p > 0 . 1 ) .
these results , together with population differentiation in acoustic properties of vocalizations ( [ 29 ] ; described below ) , suggest that this bimodal pattern of vocal behavior within s . teguina reflects population differences in nature , which persist across several generations of lab - rearing in a common environment . to avoid confounding intraspecific variation with interspecific divergence , we carried out between - species analyses both with and without the boquete samples . we report results for the full dataset unless exclusion of boquete mice changed significance .
there was no difference between s . xerampelinus and s . teguina in the duration of the isolation calling period ( f 1 , 37 2 . 4 , p = 0 . 1 ) , the duration of the non - vocal period ( f 1 , 25 0 . 9 , p = 0 . 4 ) , or the age at which the first adult song was recorded ( f 1 , 28 0 . 4 , p = 0 . 6 ) ( table 1 ) . with the boquete samples excluded , however , the isolation calling period was significantly longer in s . teguina ( f 1 , 30 9 . 4 , p = 0 . 005 ) .
adult s . teguina sing longer songs and are typically more vocal than adult s . xerampelinus , and males of both species sing longer songs and sing more often than females [ 32 ] , [ 35 ] . therefore , we asked whether species and sex differences in vocal production are established at earlier stages . s . teguina pups that produced isolation calls did so on a significantly higher proportion of days than s . xerampelinus pups ( f 1 , 36 12 . 0 , p = 0 . 001 ) ( table 1 ) . there was no species difference in bout length for any pup age class ( all p ≧0 . 3 ) , but 1–3 day old s . teguina pups from cartago tended to produce more notes / 15 s than age - matched s . xerampelinus ( f 1 , 22 4 . 0 , p = 0 . 06 ) .
within s . xerampelinus , there was no sex difference in pup vocal production ( proportion of days with isolation calls , bout length , note rate , all p ≧0 . 1 ) , but male pups stopped producing isolation calls significantly later than females ( f 1 , 13 13 . 9 , p = 0 . 003 ; table 1 ) . the small sample size for adult s . xerampelinus females that sang precluded tests for sex differences in the duration of the non - vocal period , or the first production of adult song . while there was no effect of sex on the proportion of days that s . teguina pups produced isolation calls , 4–6 and 7–9 day old s . teguina males produced more notes / 15 s ( f 1 , 18 4 . 5 , p = 0 . 05 and f 1 , 18 5 . 6 , p = 0 . 004 , respectively ) and produced longer bouts ( f 1 , 18 4 . 4 , p = 0 . 051 and f 1 , 18 8 . 9 , p = 0 . 008 , respectively ) .
we identified six distinct note types . representative spectrograms of isolation calling bouts are shown in figure 2 ; the five most common note types are shown in figure 3 .
notes a - e are indicated to the right of the first note of that type in each spectrogram .
change in the proportional abundances of common notes types during vocal development in singing mice .
representative examples of notes a–e ( dominant frequency only ; see fig . 2 for harmonics ) are shown on the x - axis . black ( s . teguina ) and white ( s . xerampelinus ) bars are species means for notes in each age class . error bars are + 1 se . sample sizes by age class are 20 ( 1–3 and 30 + days ) , 19 ( 4–6 and 7–9 days ) and 13 ( 10–12 days ) for s . teguina , and 9 ( 1–3 days ) , 11 ( 4–6 days ) , 7 ( 7–9 days ) , 4 ( 10–12 days ) and 8 ( 30 + days ) for s . xerampelinus . species differences in the proportional abundances of note types within age class were tested with one - way anova , * p < 0 . 05 , * * p < 0 . 01 .
a . long fm downward sweep with harmonics and ≧10 khz bandwidth . note a is the main component of the adult advertisement song of both species .
b . short fm downward sweep with < 10 khz bandwidth and no harmonics .
c . notes with nonlinear components , typically an fm downward sweep combined with one or more complex elements , all with multiple harmonics and deterministic chaos .
d . complex fm warble with harmonics and two or more directional changes in frequency ≧2 khz .
e . complex fm with harmonics and one directional change in frequency ≧2 khz .
we used pca to visualize age , species and population differences in frequency and timing . variables and loadings for the first two components are provided in table s2 , mean scores are plotted in figure 4 , species means for selected variables are in table s1 . all measures of frequency loaded strongly on the first axis , which accounted for 34 . 5 % of the total variance . this axis provided strong separation between s . teguina and s . xerampelinus , and between younger pup age classes within species , with positive scores for higher frequencies . there was a clear reduction in frequency with age in both species , with the largest change over time in s . xerampelinus , and a striking convergence in 10–12 day old pups of both species . this axis also provided minor separation between age - matched boquete - and cartago - derived s . teguina , with higher frequencies in boquete pups , but lower frequencies in boquete adults . the second axis ( 19 . 6 % total variance ) defined differences in bandwidth and note duration with positive scores reflecting larger bandwidth and longer notes . s . xerampelinus age classes were largely indistinguishable along this axis , whereas there was strong separation between boquete and cartago s . teguina , with cartago pups falling closer to age - matched s . xerampelinus than to boquete conspecifics . strikingly , while bandwidth and note duration in cartago mice increased from 4–6 day old pups to adults , boquete mice exhibited the opposite pattern , with a sharp decrease between 7–9 day old pups and adults . the third axis ( 8 . 2 % total variance , data not shown ) described among individual differences in timing and did not provide separation between species , populations or age classes .
plot of mean scores from the first ( pc1 ) and second ( pc2 ) principle components axes for age classes in s . xerampelinus ( white ) , and in s . teguina split by population ( cartago , black ; boquete , gray ) . age classes are indicated next to each point . pc1 explains 35 % of the total variance with higher scores corresponding to higher frequency . pc2 explains 20 % of the total variance with higher scores corresponding to larger bandwidth and longer notes . error bars are + / −1 se .
black ( s . teguina ) and white ( s . xerampelinus ) circles are species means for maximum ( max , dashes ) , dominant ( dom , dots ) , and minimum ( min , dots and dashes ) frequency in each age class . error bars are + / −1 se . sample size for s . teguina in the 13–15 day age class is 9 ; no calls were recorded for s . xerampelinus in this age class . see figure 3 caption for all other sample sizes .
interspecific differences in frequency across vocal development were generally larger than population differences within s . teguina . with boquete mice excluded , s . xerampelinus pup dominant and maximum frequencies were marginally higher at days 4–6 ( p = 0 . 03 ) and 7–9 ( p = 0 . 02 ) , respectively , but overall patterns were unchanged .
there were several marginal age - specific differences between s . teguina males and females ( table s3 ) , but these were not consistent either within or across age groups . this general lack of sex differences in spectral measures of pup isolation calls is consistent with the sexually monomorphic spectral features of adult song in both species . however , we cannot exclude the possibility of detecting sex differences with larger sample sizes , particularly for s . xerampelinus .
in both species , there was a significant negative relationship between body mass and all whole call frequency measures ( fig . s1 ) . in s . teguina ( n = 24 ) , the effect of body mass on dominant frequency was marginal ( f 1 , 23 4 . 7 , p = 0 . 04 ) , but highly significant for minimum frequency ( f 1 , 23 37 . 1 , p < 0 . 0001 ) and maximum frequency ( f 1 , 23 16 . 3 , p = 0 . 0006 ) . similarly , in s . xerampelinus ( n = 13 ) , the negative effect of body mass was weaker for dominant frequency ( f 1 , 12 6 . 3 , p = 0 . 03 ) than for minimum or maximum frequency ( f 1 , 12 15 . 5 , p = 0 . 002 and f 1 , 12 16 . 0 , p = 0 . 002 , respectively ) .
the advertisement songs of adult singing mice are highly stereotyped . therefore , we were interested in whether stereotypy increases with age as pups acquire greater motor control , or whether stereotypy is a unique feature of the adult song . for each age class , we used the coefficient of variation ( cv ) to summarize the within - individual repeatability of seven descriptors of frequency and timing . species means for each age class are shown in figure 6 . because the composition of pup vocal repertoires was heterogeneous relative to that of adults , our analysis was biased toward detecting lower stereotypy ( higher cv ) in pups vs . adults . nonetheless , if increase in stereotypy were an important component of pup vocal development , we would expect to observe a reduction in intra - individual variation across pup age classes .
black ( s . teguina ) and white ( s . xerampelinus ) cicles are species means for the coefficient of variation in each age class . error bars are + / −1 se . sample sizes by age class are 15 ( 1–3 days ) , 16 ( 4–6 days ) , 14 ( 7–9 days ) , 11 ( 10–12 days ) , 6 ( 13–15 days ) and 18 ( 30 + days ) for s . teguina , and 7 ( 1–3 days ) , 8 ( 4–6 days ) , 5 ( 7–9 days ) and 6 ( 30 + days ) for s . xerampelinus . there were insufficient data for s . xerampelinus in the 10–12 age class and no calls were recorded for the 13–15 age class .
contrary to this expectation , intra - individual variation increased from birth to the end of the isolation calling period in both species . however , the reduction in stereotypy with age was minor in s . teguina but pronounced in s . xerampelinus . the stereotypical properties of the adult song , evidenced by low cv , were present from the first song recorded in young adult mice ( fig . 6 , 30 + days ) . in the subset of individuals for which we had a second series of adult songs , captured at least 10 days after the first songs we recorded , there was no evidence for an increase in stereotypy with age ( n = 7 , paired t - test , p = 0 . 3 ) . there were no species differences in adult vocal stereotypy .
there was no effect of sex on stereotypy for any age class in either species . there was , however a species difference in age - matched pups , with higher repeatability in s . teguina . this difference was significant for age classes 1–3 ( f 1 , 21 8 . 01 , p = 0 . 01 ) and 7–9 ( f 1 , 18 22 . 01 , p = 0 . 0002 ) .
the survival of altricial neonates that are displaced from their nest depends on rapid maternal retrieval . therefore , selection should favor isolation calls that are localizable and readily detected by adult females . in general , frequency modulated sounds are easier to localize than constant frequency sounds : directional changes in frequency enhance detectability [ 38 ] , [ 39 ] , and nonlinearities ( e . g . , noise or biphonation ) are thought to prevent receiver habituation [ 37 ] , [ 40 ] . in keeping with these signal design rules , most singing mouse pup vocalizations are frequency modulated and , while the unidirectional fm sweep is the most common note type in both pup isolation calls and adult advertisement song , the vocal repertoire of pups includes notes with multiple directional changes in frequency and notes with nonlinear elements , which are rare or absent in adult songs .
singing mice are unusual among muroid rodents in that long distance acoustic communication is an important modulator of adult social interactions . this motivated our focus on the ontogeny of the spectral and stereotypic properties of the adult advertisement song . however , like many species of social mammals , including other rodents , adult singing mice vocalize during close - range interactions [ 32 ] . interestingly , complex fm warbles that resemble an elaborated version of notes d and e in young pups are the most common note type produced by adult s . teguina in close - range social interactions ( warren , campbell , pasch and phelps unpublished data ) . this suggests that the distinct vocal repertoires used by adults in long - distance vs . close - range communication arise from the same source during early development . similar patterns of vocal development , in which the heterogeneous repertoire of dependent young is modified and used in discrete contexts by adults , are reported in shrews , echolocating bats , and lab mice [ 10 ] , [ 11 ] , [ 12 ] , [ 13 ] .
female s . teguina produced their first advertisement song ( 34 . 3±8 . 2 d ) coincident with the onset of sexual receptivity indicated by vulval opening ( 33 . 8 d ; range 28–39 d ; [ 31 ] ) . this pattern suggests that steroid hormones associated with the estrous cycle activate female vocalizations . androgen manipulations indicate that dht , which cannot be aromatized to estrogen , is sufficient to activate male s . teguina vocalizations , suggesting that estrogens are not necessary for song production . interestingly , the minority of males who continued to sing following castration were those castrates with the highest levels of circulating testosterone , indicating that extra - gonadal androgens may influence song output [ 33 ] . periovulatory androgen release from the adrenals is associated with female sexual behavior in several mammalian species [ 43 ] , and may activate female vocal behavior [ 44 ] .
from an ultimate perspective , the non - vocal period corresponds to the span of time when pups of both species become ambulatory , are weaned , and initiate dispersal [ 31 ] . initially , silence reflects emancipation from immobility and the ability to actively suckle by day 12 [ 31 ] . subsequent silence in subadults likely facilitates competitor avoidance as individuals disperse in search of unoccupied habitat . because adult vocalizations advertise signaler presence to potential rivals , remaining silent reduces the probability of escalation of costly antagonistic encounters [ 35 ] . indeed , immigrant s . teguina males ( who were smaller and younger ) counter - sang less in response to playback of a conspecific male song compared to larger and older residents ( pasch and phelps , unpublished data ) . together , the data suggest that the emergence of adult vocalizations represents a compromise between proximate mechanisms that promote advertisement of the onset of sexual maturity , and selection that suppresses advertisement until the social environment is opportune .
the capacity for vocal learning is rare in non - human mammals . traditionally , rodents are classified as vocal non - learners , and recent studies using approaches such as deafening and cross - fostering indicate that the courtship songs of adult male lab mice require no auditory input for normal development and are invariant relative to the vocal environment in which animals are reared [ 45 ] , [ 46 ] , [ 47 ] . however , patterns of forebrain activation associated with lab mouse courtship song production , pitch convergence in songs of co - housed males from vocally distinct strains , and song degradation following deafening , suggest that the basic neural substrates and the capacity for vocal imitation ( plasticity ) may be present in some rodents ( [ 48 ] ; see also ref . 12 ) . arriaga and jarvis [ 49 ] proposed that these conflicting results could be reconciled if vocal learning is treated as a continuum , placing taxa like mice with a modest capacity for vocal plasticity at one end of the spectrum , and taxa like humans and song birds with the capacity to learn and subsequently modify complex vocal output at the other .
taken together , these observations suggest that singing mouse advertisement songs do not require extensive learning . testing this hypothesis awaits manipulation of early acoustic environments or auditory function . finally , given the evidence for pitch convergence in the close - range courtship songs of adult male lab mice [ 48 ] , it will be of particular interest to determine whether the close - range vocal repertoires of adult singing mice are similarly sensitive to auditory feedback .
the results of this study demonstrate that vocal development in sister species of singing mice follows the same basic trajectory as that described in other mammalian orders ( i . e . , chiroptera and soricomorpha ; [ 10 ] , [ 11 ] ) , and in laboratory mice [ 12 ] . while the acoustic structure , mechanistic basis , and social context of singing mouse vocalizations all undergo major shifts in the transition from isolation calls to adult advertisement songs , the basic elements of adult song are present from birth . likewise , spectral components of isolation calls that are rare or absent in adult long distance signals are common in adult close - range interactions . this suggests that the acoustic signals of need produced by altricial mammalian neonates provide the raw material for diverse forms of adult vocal behavior and communication .
the relationship between body mass and maximum ( black triangles ) , dominant ( open diamonds ) , and minimum ( gray squares ) frequency in a ) s . teguina , and b ) s . xerampelinus .
age - specific means ( sd ) for selected measures of frequency , timing , and vocal behavior in s . teguina ( st ) and s . xerampelinus ( sx ) .
principle component ( pc ) axis loadings for 25 acoustic variables measured from the isolation calls and adult songs of singing mice .
we are very grateful to amanda arner and molly phillips for their assistance with data collection ; we thank andreas george for his help with animal care . an earlier version of the manuscript was improved by comments from monica guerra .
conceived and designed the experiments : pc bp smp . performed the experiments : pc bp alw . analyzed the data : pc alw . contributed reagents / materials / analysis tools : smp . wrote the paper : pc bp .
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eagle rescue 101 . . . . the basics . . . . thumbs and all . . . águila de rescate 101 / sauvetage aigle 101
frontal view , showing head and feet , july 15 , 2010 . n ordonez - garza
distribution : intermediate elevations of middle america from e oaxaca , méxico , to w panamá .
this species occurs in intermediate elevations from eastern oaxaca , mxico to western panam ( musser and carleton 2005 ) . it occurs from 900 to 2 , 900 m ( reid 1997 ) .
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urltoken
musser , guy g . , and michael d . carleton / wilson , don e . , and deeann m . reeder , eds .
mammal species of the world : a taxonomic and geographic reference , 3rd ed . , vol . 2
mammal species of the world : a taxonomic and geographic reference , 2nd ed . , 3rd printing
with contributions by bernadette n . graham , adam p . potter , and mariana m . upmeyer
comments : level of differentiation and status of populations north ( teguina ) and south ( irazu ) of the nicaraguan depression merit reconsideration . hooper ( 1972 ) recognized four subspecies
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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the spatial distribution of individuals may be strongly influenced by social processes and thus patterns of space use are often used to infer the mating and social systems of a population ( gaulin and fitzgerald 1988 ; shier and randall 2004 ; steinmann et al . 2005 ) . in particular , sex - specific differences in home - range use can yield insights into the mating system and can improve understanding of the behavior of little - studied species ( reviewed in clutton - brock 1989 , 1991 ; gaulin and fitzgerald 1988 ; heske and ostfeld 1990 ; ostfeld 1985 ; reichard 2003 ; schradin and pillay 2005 ; shier and randall 2004 ; steinmann et al . 2005 ) . for example , in polygynous mating systems , home ranges of males tend to overlap less than those of females , and a given female typically overlaps with only 1 male . in contrast , in promiscuous mating systems , home - range use by males and females is expected to be more equitable , with each individual ' s home range overlapping those of several members of the opposite sex . finally , socially monogamous systems should be characterized by exclusive ( nonoverlapping ) home ranges , each of which is shared by a male–female pair ( clutton - brock 1989 ; ophir et al . 2008 ; ostfeld 1985 ; shier and randall 2004 ) .