the proboscis bat ( rhynchonycteris naso ) is a bat species from south and central america .

other common names include long-nosed proboscis bat , sharp-nosed bat , brazilian long-nosed bat and river bat in english , and murciélago narizón in spanish .

it is monotypic within its genus .

this species is in the family emballonuridae , the sac-winged or sheath-tailed bats .

like most bats , it is nocturnal .

it is found from southern mexico to peru , bolivia and brazil , as well as in trinidad .

proboscis bat

information on the proboscis bat is currently being researched and written and will appear here shortly .
proboscis bat ( in dutch : langneusvleermuis ) . cocha blanco , peru , 28 june 2009 . mov
proboscis bats are not threatened in the wild and are hardly ever kept by zoos .
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english : long - nosed bat , sharp - nosed bat ; german : nasenfledermaus ; spanish : murciélago de trompa .
knornschild , m . , c . harview , r . moseley , o . von helversen . 2009 . remaining cryptic during motion - behavioral synchrony in the proboscis bat .
english : white bat , jumbie bat ; german : geisterfledermaus , weisse fledermaus ; spanish : murciélago blanco , murciélago albino .
is a darker shade of brown and lacks the gray - grizzled pattern found in proboscis bats . male proboscis bats also lack wing sacs , which are organs on the propatagial membrane of the wings that store secretions used for mating rituals and are very prominent in
there are no known positive effects of proboscis bats on humans , although in general , insectivores may aid in controlling insect populations .
emballonurids generally give birth to a single offspring each year . an exception is the small proboscis bat that reproduces twice a year . most emballonurid females give birth to their offspring at the beginning of the rainy season .
english : greater white - lined bat , greater two - lined bat ; german : große taschenflügelfledermaus , große sackflügelfledermaus ; spanish : murciélago de listas .
knörnschild m , harview c , moseley r , von helversen o ( 2009 ) remaining cryptic during motion – behavioral synchrony in the proboscis bat , rhynchonycteris naso . acta chiropterologica 11 ( 1 ) : 208 - 211 .
ulanovsky n , moss cf ( 2008 ) what the bat ' s voice tells the bat ' s brain . proc nat acad sci usa 105 : 8491–8498 .
timm , r . m . and m . losilla . ( 2007 ) . orb - weaving spider , argiope savignyi ( araneidae ) , predation on the proboscis bat rhynchonycteris naso ( emballonuridae ) . caribbean journal of science 43 , 282 - 284
tannenbaum br . 1975 . reproductive strategies in the white - lined bat . tannenbaum .
copyright bat conservation international © 2018 . all rights reserved . unless otherwise noted , all images are copyright © merlin d . tuttle and / or © bat conservation international .
knörnschild , m . , harview , c . , moseley , r . and von helversen , o . ( 2009 ) . remaining cryptic during motion – behavioral synchrony in the proboscis bat ( rhynchonycteris naso ) . acta chiropterologica 11 , 208 - 211
) . proboscis bats can be distinguished from these species by their elongated muzzle and evenly - spaced tufts of white to pale gray fur along their forearms . the dorsal side of
emballonurid bats exhibit a variety of different mating systems . similar to other mammalian groups , polygynous mating patterns are most common . however , exceptions are for example the monogamous mating system of cormura brevirostris and possibly also of the greater dog - faced bat , peropteryx kappleri , and some members of the genus taphozous . the proboscis bat is considered to have a promiscuous mating pattern . among emballonurid bats , the greater sac - winged bat , saccopteryx bilineata , is the best studied species .
is the most similar to proboscis bats , neither of these bats hibernate or roost in caves frequently ; however , they both have a similar body mass and a similar number of offspring annually .
nyffeler m , knörnschild m ( 2013 ) bat predation by spiders . plos one 8 : e58120 .
günther l , knörnschild m , nagy m , mayer f ( 2017 ) the evolution of a rare mammalian trait – benefits and costs of male philopatry in proboscis bats . scientific reports 7 : 15632 .
the lesser false vampire bat , megaderma spasma , search in pohara by dr . v . t . ingole
fresh weight and wingspan of bat species reported to be captured by spiders ( arranged in alphabethical order ) .
in this study , we use long - term behavioural observations of individually marked proboscis bats in combination with genetic parentage analyses to assess social dispersion and male mating strategy during day and night . in contrast to the vast majority of other bat species , proboscis bats usually inhabit very exposed and well - lit structures [ 42 ] . thus , we assume that the clumped roosting of mixed sex groups during the day is a derived trait and the result of selection for cryptic behaviour on exposed roost structures . at night , we hypothesize to still observe an ancestral strategy , namely that male proboscis bats establish themselves at preferred sites in their roost where they are territorial or dominant ( see the electronic supplementary material , figure s2 for a sketch of this main hypothesis ) .
this small bat is common throughout central america and into northern south america . unlike many other bat species , they roost in the open , usually on tree trunks leaning over the water . these bats forage for small insects , usually just over water .
washington d . c . bat conservation international 1012 14th street nw , suite 905 washington , d . c . 20005 , usa
data from : from resource to female defence : the impact of roosting ecology on a bat ' s mating strategy . dryad digital repository
citation : nyffeler m , knörnschild m ( 2013 ) bat predation by spiders . plos one 8 ( 3 ) : e58120 . urltoken
speakman jr ( 1991 ) the impact of predation by birds on bat populations in the british isles . mammal rev 21 : 123–142 .
the proboscis bat is a small bat , the forearm ranging from 35 to 41 mm in length and weight averaging about 4 g . the back is grizzled brownish to grayish , with two faint whitish stripes on the lower back and rump and the under side of the animal is pale brownish gray . the pelage is soft and dense . the membranes and ears are dark brown . the prominent ears are narrowly rounded . ( d . l . plumpton , j . knox jones , jr . mammalian species , no 413 , 1992 ) .
goiti u , aihartza j , guiu m , salsamendi e , almenar d , et al . ( 2011 ) geoffroy ' s bat ,
up to 31 male ( range = 20–31 ) and 26 female ( range = 23–26 ) proboscis bats roosted under the eaves of a large house at la selva biological station , costa rica . for details on number and distribution of age classes during the different periods , see
foraging mode and echolocation call frequency of adult bat species reported to be captured in spider webs ( arranged in order of increasing peak frequency ) .
siemers bm , schnitzler h - u ( 2004 ) echolocation signals reflect niche differentiation in five sympatric congeneric bat species . nature 429 : 657–661 .
bats of the family emballonuridae usually give birth to a single offspring per year . exceptions to this rule may be small species like the proboscis bat that reproduce twice each year . most emballonurid bats show a seasonal pattern of reproduction with females giving birth to their offspring at the beginning of the rainy season . sperm storage or delayed embryonic development occurs in some old world members within the family emballonuridae . the mating system varies by species .
knörnschild m , ueberschaer k , helbig m , kalko ekv ( 2011 ) sexually selected infanticide in a polygynous bat . plos one 6 : e25001 .
fernandez aa , knörnschild m ( 2017 ) isolation calls of the bat saccopteryx bilineata encode multiple messages . animal behavior and cognition 4 : 169 - 186 .
knörnschild m , feifel m , kalko ekv ( 2013 ) mother - offspring recognition in the bat carollia perspicillata . animal behaviour 86 : 941 - 948 .
some emballonurids roost in large groups , others gather in smaller groups of about ten to forty , and a few are loners . colonies of african sheath - tailed bats include up to 50 , 000 bats , each of which returns to a precise place in a roosting cave along the kenyan coast . daytime roosts for the sac - winged bat can reach up to sixty individuals . proboscis bat females roost apart from the males when the young are born . different shelters are used by adult male and female gray sac - winged bats during the summer ; most of the other forms seem to remain together throughout the year .
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small bat , forearm 1 . 7–1 . 9 in ( 4 . 3–4 . 7 cm ) , weight 0 . 18–0 . 21 oz ( 5–6 g ) .
lin aq , jin lr , shi lm , sun kp , berquist sw , et al . ( 2011 ) postnatal development in andersen ' s leaf - nosed bat
knörnschild m , feifel m , kalko ekv ( 2014 ) male courtship displays and vocal communication in the polygynous bat carollia perspicillata . behaviour 151 : 781 - 798 .
article by niall o’riain niall is an education specialist from london who came to help with our bat project and was particularly charmed by r . naso and it’s strange nose .
knörnschild m , tschapka m ( 2012 ) predator mobbing behaviour in the greater spear - nosed bat phyllostomus hastatus . chiroptera neotropical 18 ( 2 ) : 1132 - 1135 .
nephilengys cruentata , another bat - catching nephilid ( table 1 ) , weighs 0 . 5–1 . 6 g ( table 2 ) . it is a synanthrophic spider that spins orb webs ( with up to 1 m diameter ) on the corners of walls and ceilings of buildings [ 60 ] – [ 62 ] . nephilengys might be less efficient in bat catching . the only reported incidence where a bat got entangled in a nephilengys web ended with the bat’s successful escape . nevertheless this spider species should not be ruled out as a potential predator of small bats ; it is known to catch and eat various types of vertebrates including small birds [ 25 ] – [ 26 ] , [ 62 ] .
. although every effort is made by google to ensure translation accuracy , errors may occur . bat conservation international does not guarantee or warrant the accuracy or reliability of this tool .
austin bat conservation international 500 n capital of tx hwy . , bldg . 1 austin , tx 78746 , usa + 1 ( 512 ) 327 9721 1800 - 538 - bats
engler s , rose a * , knörnschild m * ( accepted ) isolation call ontogeny in bat pups ( glossophaga soricina ) . * joint senior authors . behaviour 154 : 267–286
there has been no specific study on the role this species plays on the ecosystem , but due to its diet and large population numbers , proboscis bats may contribute to the control of insect populations in its habitat . in panama , where these bats occur , research has shown that insectivorous bats indirectly affect herbivory on plants , by reducing herbivorous
yellow bats in the southern united states sometimes climb into the dead fronds of palm trees ( 5 ) or nestle within clumps of spanish moss ( 6 ) that drape dramatically from tree branches ( 7 ) . because the yellow bats fur matches the color of the dry , faded moss or yellowing fronds , the bat is hidden from view . they cannot protect themselves , however , from the common practice of pruning dead fronds from ornamental palms , which threatens yellow bat habitat in texas and other southern states [ bats , summer 1997 ] .
schoeman cm , jacobs ds ( 2008 ) the relative influence of competition and prey defenses on the phenotypic structure of insectivorous bat ensembles in southern africa . plos one 3 : e3715 .
burgess , h . and lee , n . ( 2004 ) a behavioural study of the silhouette sheath - tailed bat ( coleura seychellensis ) . phelsuma , 12 : 69 - 77 .
strauss m , von helversen o , knörnschild m ( 2010 ) the ontogeny of courtship behaviors in bat pups ( saccopteryx bilineata ) . behaviour 147 ( 6 ) : 661 - 676 .
knörnschild m , behr o , von helversen o ( 2006 ) babbling behavior in the sac - winged bat ( saccopteryx bilineata ) . naturwissenschaften 93 ( 9 ) : 451 - 455 .
knörnschild m , blüml s , steidl p , eckenweber e , nagy m ( 2017 ) bat songs as acoustic beacons - male territorial songs attract dispersing females . scientific reports 7 : 13918 .
smotherman m , knörnschild m , smarsh g , bohn km ( 2016 ) the origins and diversity of bat songs . journal of comparative physiology a 202 ( 8 ) : 535 - 554 .
greiner s , nagy m , mayer f , knörnschild m , hofer h , voigt cc ( 2014 ) sex - biased senescence in a polygynous bat species . ethology 120 : 197 - 205 .
geipel i , kalko ekv , wallmeyer k , knörnschild m ( 2013 ) post - weaning maternal food provisioning in a bat with a complex hunting strategy . animal behaviour 85 : 1435 - 1441 .
deaths of bats in spider webs have been considered to occur very rarely . in two more recent papers , a web - building spider , argiope savignyi , and a theraphosid spider , poecilotheria rufilata , were each reported to predate on a small bat [ 30 ] – [ 31 ] . these authors hypothesized that bat captures and kills due to spiders might be more frequent than previously thought . to test this hypothesis , an extensive global literature survey on bat - catching spiders was conducted , along with an attempt to use web - based sources as well . the insights from this research are reviewed here .
heim o , treitler jt , tschapka m , knörnschild m , jung k ( 2015 ) the importance of landscape elements for bat activity and species richness in agricultural areas . plos one 10 : e0134443 .
knörnschild m , nagy m , metz m , mayer f , von helversen o ( 2010 ) complex vocal imitation during ontogeny in a bat . biology letters 6 ( 2 ) : 156 - 159 .
knörnschild m , von helversen o ( 2008 ) non - mutual vocal mother - pup recognition in the sac - winged bat , saccopteryx bilineata . animal behaviour 76 ( 3 ) : 1001 - 1009 .
joubert , f . ( 2004 ) a preliminary investigation into the general ecology , status and habitat of the seychelles sheath - tailed bat coleura seychellensis ( emballonuridae ) . phelsuma , 12 : 54 - 68 .
nagy m , knörnschild m , günther l , mayer f ( 2014 ) dispersal and mating strategies in the neotropical grey sac - winged bat balantiopteryx plicata . behavioural ecology and sociobiology 68 : 891 - 901 .
knörnschild m , von helversen o , mayer f ( 2007 ) twin siblings sound alike – isolation call variation in the noctule bat , nyctalus noctula . animal behaviour 74 ( 4 ) : 1055 - 1063 .
nagy m , günther l , knörnschild m , mayer f ( 2013 ) female - biased dispersal in a bat with a female - defence mating strategy . molecular ecology 22 ( 6 ) : 1733 - 1745 .
knörnschild m , nagy m , metz m , mayer f , von helversen o ( 2012 ) learned vocal group signatures in the polygynous bat saccopteryx bilineata . animal behaviour 84 ( 4 ) : 671 - 679 .
gerlach , j . & taylor , m . ( 2006 ) habitat use , roost characteristics and diet of the seychelles sheath - tailed bat coleura seychellensis . acta chiropterologica , 8 ( 1 ) : 129 – 139 .
gerlach , j . and taylor , m . ( 2006 ) habitat use , roost characteristics and diet of the seychelles sheath - tailed bat coleura seychellensis . acta chiropterologica , 8 ( 1 ) : 129 - 139 .
fulmer a , knörnschild m ( 2012 ) intracolonial social distance , signalling modality and association choice in the greater sac - winged bat ( saccopteryx bilineata ) . journal of ethology 30 ( 1 ) : 117 - 124 .
carter g , schoeppler d , manthey mk , knörnschild m , denzinger a ( 2015 ) distress calls of a fast - flying bat ( molossus molossus ) provoke inspection flights but not mobbing . plos one 10 : e0136146 .
fernandez aa , fasel n , knörnschild m , richner h ( 2014 ) when bats are boxing : aggressive behaviour and communication in male seba ' s short - tailed fruit bat . animal behaviour 98 : 149 - 156 .
proboscis bats have large home ranges used for foraging and multiple roosting sites . a 10 - month study on 3 different colonies in la pacifica , costa rica , showed a range of 0 . 9 to 1 . 2 hectares , with an average of 1 . 1 hectares . the distance between each roosting site ranges from 25 to 180 m with some rotation of sites ; each site is typically used for 4 to 6 weeks .
some bats simply hang in the open on tree trunks , relying on camouflage to protect them . these proboscis bats ( 10 ) blend well into the multicolored bark on the side of a tree in belize . this species typically roosts only on trees that extend over water , probably as a means of limiting predators . to photograph these bats , bci executive director merlin tuttle had to wade chest - deep into the river ( 11 ) .
an extensive bibliographic search was conducted in order to find any information available on bat - catching spiders . the search was based largely on the thomson - reuters data base ( web of science ) , google scholar , google books , proquest dissertations & theses , and flickr image - hosting website ( hosting more than 6 billion images ) . in addition to this , an internet search for blogger information on this topic was conducted . bloggers who had posted photographs and reports on bat - catching spiders on the internet were contacted to get detailed information on their observations . furthermore , the staff of bat hospitals was contacted to get information on bats rescued from spider webs . finally , an inquiry among fellow arachnologists and chiropterologists was carried out to get access to unpublished reports on this topic . many of these experts had conducted field studies for decades , and their feedback provided valuable information needed to assess how frequent incidences of bat catches by spiders might be . altogether , 52 reports on bat - catching spiders could be gathered ( table 1 ) . twenty - three of these reports have previously been published in scientific journals , books , and a doctoral thesis .
the brazilian long - nosed bat got it ' s name for a reason - look at that snauz . a small bat known to fly in the night skies from southern mexico to central brazil , the brazilian long - nosed bat is only 1 . 25 - 1 . 75 inches ( 3 . 5 - 4 . 5 cm ) long , with a wingspan of just 4 . 75 - 6 . 25 inches ( 12 - 16 cm ) , and it is known for it ' s elongated snout . also different from most other bats , are the tufts of gray hair on it ' s forearms . this helps them blend in with some of the gray , mossy trees of the region .
heckel , g . , c . c . voigt , f . mayer , and o . von helversen .
extra - harem paternity in the white - lined bat saccopteryx bilineata .
behaviour 136 ( 1999 ) : 1173 .
decaying trees with hollow trunks should also be conserved , because they provide a dry and secure shelter for bats that enter and exit through holes in the trunk . some of these bats roost toward the tops of trees so they are able to make the slight drop needed to take flight when leaving their roost . these are generally species with long , narrow wings built for high speed . others , such as this tropical frog - eating bat and sebas short - tailed bat ( 13 ) , have broader wings that permit greater maneuverability and do not require a drop - off for the bat to take flight . these bats can enter a hollow at the base of a tree where many other species cannot .
knörnschild m * , jung k * , nagy m , metz m , kalko ekv ( 2012 ) bat echolocation calls facilitate social communication . * joint first authors . proceedings of the royal society b 279 ( 1748 ) : 4827 - 4835 .
three incidences of bat captures by web - building spiders have been reported from africa ( [ 41 ] – [ 42 ] ; d . schultz , pers . comm . ; fig . 2f ; report # 46–48 ) . only two incidences of bat catches by web - building spiders witnessed in north america have been reported so far ( [ 43 ] – [ 44 ] ; report # 49–50 ) , and these both refer to warm areas in the southern usa . incidences of bat catches by orb - weaving spiders are unknown from the northern part of north america ( b . fenton , pers . comm . ) . likewise , incidences of this type have not been reported in the ukrainian and russian scientific literature ( a . t . bashta , pers . comm . ) . only two incidences of bats being captured in spider webs have been reported from europe ( report # 51–52 ) . in one case , a dead bat was found entangled in the web of an orb - weaving spider on a building site near stuttgart , germany ( german tabloid bild , may 2011 ) . another incidence of a bat caught in a spider web was observed on the isle of wight , south east england ( g . street , pers . comm . ) .
rose a , brändel sd , cvecko p , engler s , hiller h , knörnschild m , tschapka m ( 2017 ) new records of hypopigmentation in two neotropical phyllostomid bat species with different roosting habits ( uroderma bilobatum , glossophaga soricina ) . mammalia . online first .
bat - catching spiders belong to the araneomorph families nephilidae ( golden silk orb - weavers ) , araneidae ( orb - weaver spiders ) , sparassidae ( huntsman spiders ) , and the mygalomorph family theraphosidae ( tarantulas ) . furthermore , an attack attempt by an araneomorph hunting spider of the family pisauridae ( fishing spiders ) was witnessed . seventy - three percent of the known incidences of bat catches were attributable to orb - weaving spiders , 15 % to unidentified web - building spiders , and 12 % to hunting spiders ( table 1 ) .
although bats range in size from the small hog - nosed bat that weighs less than a penny to flying foxes whose wing spans are greater than 5 feet , the majority tend to be small . all species of bat share a fairly similar body form - fur and skin , wings , teeth , reproductive technique , and visual and hearing systems . the skin is normally black or dark grayish brown in color and has small transparent hairs . most bats have small eyes but their auditory technique ( sonar ) is advanced to say the least .
günther l , lopez md , knörnschild m , reid k , nagy m * , mayer m * ( 2016 ) from resource to female defence – the impact of roosting ecology on a bat’s mating strategy . * joint senior authors . royal society open science 3 : 160503 .
proboscis bats are polygynous and although the male to female ratio is usually equal , breeding females mainly copulate with dominant males . the rest of the colony likely follows a hierarchy , where reproductive females rank higher than non - reproductive females and non - dominant males . dominant males forage at the edge of the feeding area and protect their colony from neighboring conspecifics using aerial attacks and audible vocalizations . dominant males are thought to exhibit female - defense over resource - defense , since the males follow the females as they forage , even when they return to their roosting sites at night . mating in proboscis bats is not well - studied , but during an observed copulation , two males approached a female from opposite sides , leading to what appeared to be inaudible vocalizations by the female . one male proceeded to edge over the female’s body and appeared to vocalize inaudibly , followed by copulation . the female and the second male then took off and executed a downward spiraling flight , until nearly reaching the water surface . however , the purpose of this act is not yet known .
until recently very little was known about this rare seychelles bat . colonies roost in caves and are thought to be divided into harem groups that consist of adult females and their young and are led by a single male ( 2 ) . unlike the african sheath - tailed bat c . afra ( 2 ) , the seychelles species hangs upside down from the cave ceiling to roost . females give birth to their offspring during the rainy season that runs from november to december ( 2 ) . these bats feed exclusively on insects ( 3 ) ( 7 ) ( 8 ) .
the neotropical orb - weaving spider eriophora fuliginea has been observed to kill and eat small bats that got entangled in its webs [ 8 ] , [ 37 ] , [ 101 ] . when a bat got caught in a web , the spider immobilized the bat by attack - wrapping and subsequently biting it [ 101 ] . following this , the spiders fed on the dead bats for many hours ( d . e . wilson , pers . comm . ) . the incidences of bats being caught , killed , and eaten in webs of eriophora spp . are without any doubt predation events .
voigt c , behr o , caspers b , von helversen o , knörnschild m , mayer f , nagy m ( 2008 ) songs , scents , and senses : sexual selection in the greater sac - winged bat , saccopteryx bilineata . journal of mammalogy 89 ( 6 ) : 1401 - 1410 .
during these inquiries we got access to several previously unpublished photographs of bat - catching spiders . these photographs were shown to established bat and spider taxonomists for identification of the bats and spiders , respectively . in a few cases photographs of habitats were sent to vegetation specialists for proper habitat classification . nomenclature follows [ 32 ] – [ 33 ] . spiders reported in this paper are divided into two major groups based on foraging mode ( sensu gertsch [ 14 ] ) : “hunting spiders” ( i . e . , spiders that forage without the use of a catching web ) and “web - building spiders” ( i . e . , spiders that forage using a catching web ) . data on spider weight and size as well as bat weight , wingspan , foraging mode and echolocation call frequency were taken from the literature when available . report numbers used in the results , tables and figures refer to the respective detailed report description ( see file s1 ) .
from tree trunks to branches , dead fronds to lush tropical leaves , there is potential bat habitat in much of the vegetation that surrounds us every day . in future issues of this magazine , well look at the many different ways in which bats use caves , rock crevices , and shelters built by other animals .
a third geographic region where bat - catching by web - building spiders was repeatedly witnessed ( seven reports ) is the area of australia ( report # 39–44 ) and papua new guinea ( report # 45 ) . most australian incidences were observed in the coastal areas of new south wales and queensland ( fig . 2j–k ) .
the brazilian long - nosed bat is a slow flier , and therefore isn ' t equipped to hunt in the treetops ( larger , faster birds would end up preying on them ) ; so they have fallen into the habit of feeding on the insects that live on the surface of the ponds , lakes , and rivers over which they live .
throughout the year , breeding proboscis bats can be found in different parts of their distribution , but may cease breeding during the dry season , which is november to march in costa rica and april to september in southeastern brazil . females produce only one offspring per pregnancy , but may have up to two pregnancies annually and can therefore be polyestrus . offspring are quite large at birth and can reach adult size within 2 weeks . weaning occurs after 2 to 4 months , after which , the young bats disperse to nearby colonies . females undergo their first parturition around 18 months of age . overlap between lactation with the first young and gestation of a second offspring has been documented .
proboscis bats prefer tropical lowlands with an elevation of less than 300 m and are almost always found near or over moving water , but rarely near fast - moving water . unlike most nocturnal bats , light does not seem to disturb the colony , individuals roost in well - lit areas , usually around 1 . 8 meters ( 6 feet ) above water . these bats are known to cling upside down in a vertical line on the bark and roots of trees overhanging water , but on occasion , some have been found under bridges , in cave mouths overhanging water , under large curled leaves , such as those of banana plants and under large fabric umbrellas of outdoor brazilian restaurants .
voigt c . c . , g . heckel , and o . von helversen .
conflicts and strategies in the mating system of the sac - winged bat .
in functional and evolutionary ecology of bats , edited by g . f . mccracken , a . zubaid , and t . h . kunz . oxford : oxford university press , 2003 .
a second geographic region where bat - catching by web - building spiders has frequently been reported ( 13 reports ; 2i and l ) is eastern and southeastern asia including locations in china ( report # 26–34 ) , japan ( report # 35 ) , vietnam ( report # 36 ) , malaysia ( report # 37 ) , and sri lanka ( report # 38 ) . here , bat - catching by spiders have been witnessed particularly often in the eastern coastal area of china , specifically in parks and forests of the greater hong kong area ( [ 39 ] – [ 40 ] ; g . ades , pers . comm . ; c . s . k . liu , pers . comm . ; fig . 2l ) .
the seychelles sheath - tailed bat is one of the world’s rarest mammals . these small bats are a reddish - brown colour with paler underparts ( 2 ) . the name ‘sheath - tailed’ refers to a membrane that extends between the hind legs and can be placed over the tail . this provides precise movement in flight between the rainforest trees of their habitat ( 3 ) .
the majority of bats entangled in spider webs were small species with a wingspan of 10–24 cm and an adult weight of 3–8 g ( table 3 ) . it is noteworthy that adult bat weight can vary by several grams depending on the bat’s feeding status and it is plausible that only the lightest individuals of any given bat species get entangled in spider webs ; moreover , some of the captured bats were juveniles or subadults ( reports # 7 , 15–16 , 20–23 , and 50 ) and thus presumably much lighter than the weights for adults presented in table 3 . some of the entangled bats are among the smallest bats on earth ( e . g . , tylonycteris pachypus weighs only 3 g ; [ 73 ] ) . likewise , the neonatal myotis austroriparius bats found entangled in spider webs in a study in florida ( see [ 43 ] ) weighed only 3 g [ 74 ] . large bats are missing in table 3 because of these bats’ capability to fly right through a web ( see below ) . most of the species listed in table 3 are known to roost in buildings , caves , and tree holes .
although the red bat ( 8 ) of north america prefers to roost in more exposed positions - - usually from a tree branch or the petiole ( stem ) of a leaf - - this species mottled red coat protects it by giving the bat the appearance of a dead leaf . in fact , there are accounts of these bats being found hibernating in piles of leaf litter on the ground . red bats may also resemble rotting fruit as they hang from the branches of peach trees . with the exception of mothers and their offspring , red bats and other bats of the genus lasiurus generally roost solitarily . the mother in this picture is nursing quadruplets while clinging to a grapevine . unlike most other bats , red bats can have more than one pup .
head and body length 2 . 7–3 . 2 in ( 68–82 mm ) ; forearm 2 . 4–3 . 6 in ( 63–93 mm ) ; weight 0 . 6–0 . 8 oz ( 17–24 g ) . large white bat with yellowish ears ; grayish hair bases may be visible ; ears short and rounded ; tail membrane with single brown sac close to the tip of the tail .
based on census observations , we calculated fidelity indices f sensu heckel et al . [ 51 ] . first , based on the constant usage of the same areas of the roost by the same groups of bats , we calculated individual fidelity to a social group ( f group ) . f group was calculated during day and night and corresponds to the proportion a bat was observed in the area of a group in relation to its absolute presence in the roost . second , individual fidelity of adult bats to a certain site in the roost ( f site ) was calculated based on the number of observation events and corresponds to the proportion a bat was present at a certain site in the roost between its first and last day of observation . f site was calculated for morning , afternoon and night separately .
initially discovered in ancient egyptian tombs by the scientists who accompanied napoleon on his campaign . the name tomb bat is basically unjustified , because these animals are not generally associated with tombs . daytime roosts are found in rather open spaces that are occasionally penetrated by daylight , such as underneath roofs or on walls . in colonies , individuals roost as pairs in close association . individuals move quickly sideways or upwards when disturbed .
head and body length 1 . 4–1 . 9 in ( 36–48 mm ) ; forearm 1 . 4–1 . 6 in ( 36–40 mm ) ; weight 0 . 10–0 . 21 oz ( 3–6 g ) . very small bat with two wavy , faint dorsal stripes . upperparts with grizzled , whitish fur . nose projecting beyond mouth . tufts of grayish hair on forearms and wing membrane . wing and tail membrane brown .
there is a variety of different mating customs among the different species of emballonurids . most of these bats are polygamous ( puh - lih - guh - mus ) , meaning that males mate with more than one female during the mating season . yet the chestnut sac - winged bat , and possibly other species , are monogamous ( muh - nah - guh - mus ) , meaning a male and female mate and pair only with each other .
with regard to bat - eating theraphosids photographed in the neotropics and in india , the actual killing of the bats was not witnessed . however , it has been proven by means of observations in captivity that large theraphosids are capable of killing bats . this is shown in a youtube video where a grammostola rosea ( subfamily theraphosinae ) is killing a small bat offered to it in a cage environment ( urltoken ; this report was not included in table 1 because it occurred in captivity ) . the g rammostola rosea shown in this video was identified for us by r . west ( pers . comm . ) . theraphosid venom has been proven to be active on small mammals [ 20 ] . one can therefore assume that theraphosid predation on bats is taking place in the field as well and that the incidences witnessed in the neotropics and in india may have been cases of predation and not scavenging .
some bats feed on insects and whole animals . vampire bats , ranging from mexico south to argentina , feed only on the blood . there are also herbivorous species that feed on fruit and flowers alone . females give birth at roost sites , often while hanging upside down . youngsters are pretty helpless until they can fly , so they tend to mature rapidly like the brown bat which can reach adult size and start flying as young as three weeks old .
while tropical bats are active all year round , others in temperate regions can hibernate in the winter in order to accommodate the diminished food supply . most hibernating bats spend the summer feeding heavily on insects , building up fat to provide energy for sleeping through a winter season . whle hibernating , the bat allows its body temperature to fall to that of the surrounding air , and as its pulse and breathing rates slow down , it fundamentally enters a state of suspended animation .
all five groups of bat - catching spider taxa ( nephilidae , araneidae , theraphosidae , sparassidae , and pisauridae ) are known to be predominantly predaceous on insects [ 14 ] , [ 53 ] , [ 64 ] , [ 69 ] , [ 101 ] . with regard to large - sized theraphosids , sparassids , and pisaurids , their feeding behavior in the field has not been thoroughly investigated and one cannot currently judge whether predation on bats is of significance to them from a feeding ecological point of view .
other orb - weavers reported to have caught bats in their webs are found in the araneid genera argiope and parawixia ( table 1 ) . these lighter spiders , in comparison to nephila spp . and eriophora spp . , spin webs of much smaller size and may be engaged in bat catches less frequently , although it must be noted that argiope spp . have repeatedly been observed killing ( and sometimes eating ) small birds in north america and elsewhere ( e . g . [ 21 ] , [ 29 ] ) .
woodpecker holes and loose bark on tree trunks make convenient hideaways for many bats in north america . here an endangered indiana bat ( 12 ) peeks out from its resting place beneath the bark of a dead but standing trunk , called a snag . unfortunately , a perfectly loosened piece of bark doesnt last for the entire life of the snag , and bats are forced to move often to find new bark crevices . well - designed forest management plans protect snags from being cleared away because of their importance as habitat for bats and other animals .
some emballonurids , such as the greater sac - winged bat , live in year - round stable harems ( hare - um ; group of females associated with one male ) , with one to eight females in an area that is patrolled by a male . male sac - winged bats in the genus saccopteryx defend their harems with energetic flight maneuvers . researchers have found that harem males father an average of 30 percent of the offspring within their harem . the majority of offspring is fathered by other harem males or by males from outside the colony .
our current knowledge of orb - weaver feeding ecology suggests that these spiders depend on flying insects as main prey , whereas bats and also birds occasionally entangled in spider webs might be considered to be by - catch [ 45 ] . it can take a large orb - weaver many hours to consume a bat or bird prey ( [ 26 ] , [ 28 ] , [ 30 ] ; d . e . wilson , pers . comm . ) indicating that spiders might extract a substantial amount of energy while feeding on such a large prey . based on field observations in papua new guinea , robinson & robinson [ 53 ] estimated the average capture rate of nephila pilipes at ∼0 . 19 g wet weight prey killed per spider per day ( which corresponds to ∼0 . 015 g dry weight food ingested per spider per day ) . according to higgins [ 46 ] , the average prey capture rate of the significantly smaller - sized nephila clavipes in texas was ∼1 . 5–2 . 5 times lower than the estimate by robinson & robinson [ 53 ] . thus , the catch of a ∼2 g bat yields a nephila pilipes a potential prey biomass that is about 10 times the average daily prey catch .
the following additional supporting material may be found in the online version of this article : table s1 results from allele frequency calculations with cervus v . 3 . 0 ( kalinovski et al . 2007 ) . figure s2 sketch of the main hypothesis regarding the differences in social dispersion in the roost between day and night . we assume that the clumped roosting of mixed sex groups during the day is a derived trait and the result of selection for cryptic behaviour on exposed roost structures . at night , we hypothesize to still observe an ancestral strategy , namely that male proboscis bats establish themselves at preferred sites in their roost where they are territorial or dominant . the blue bats represent males . the red bats represent females . distances between the individual bats are true to scale , while distances between the sites are not to the scale . figure s3 top view of the roof of ‘cabina 5’ . the extending roof is drawn in black with its grid ( 1 - 36 ) and the five defined sites . table s4–s6 detailed census data from different periods ( postpartum oestrus mating period ‘pemp’ ; seasonal mating period ‘smp’ ; non - mating period ‘nmp’ ) between 2010 and 2014 :
the nature protection trust of the seychelles has monitored the known population of seychelles sheath - tailed bats since 1997 ( 5 ) ( 6 ) . as with other bats , the protection of roosts and foraging habitat are of great importance . existing roosts must be preserved , not only from human disturbance , but also from introduced plant species that smother the cave entrances ( 4 ) . to this end , one roost site had the invasive kudzu vine pueraria phaesolides cut back from its entrance in 2001 and 2004 , and ongoing management aims to completely remove the species from the area ( 4 ) . it is hoped that removal of invasive creepers from existing and abandoned roost sites will help to increase the desperately low numbers of this critically endangered bat to more sustainable levels .
where do bats live ? ask anyone , and the immediate response is likely to be in caves .  more knowledgeable people might also mention trees , buildings , and abandoned mines . in fact , the range of places bats choose to roost is too broad to make into a simple list . from woodpecker holes to tall jungle grasses to old world war ii bunkers , bats housing selections are far more creative and opportunistic than one might imagine . bats take advantage of all manner of natural resources , as well as shelters already built by insects , birds , and other mammals , including humans . as the first installment in a series illustrating the vast diversity of bat roosts , this photo essay focuses on some of the interesting ways in which bats use plants and trees .
the seychelles sheath - tailed bat has been lost from the majority of islands where it was originally found . the precise cause of this dramatic decline is unclear ; it has been speculated that introduction of the predatory barn owl ( tyto alba ) and declines in insect availability resulting from use of pesticides may be involved ( 3 ) . however , the recent discovery of a second roost near marshes on silhouette showed that the bats here feed on different insects than those eaten by roosts that forage in palm woodlands elsewhere ( 4 ) . it has therefore been argued that this variability in the diet demonstrates that food is not a limiting factor for the species , and the decline in the species may rather be attributable to habitat alteration caused by invasive plants obstructing roost entrances ( 4 ) .
in the tropics , huge nephilid orb - webs ( genera nephila and nephilengys ) sometimes block the entrances to bat caves ( [ 61 ] ; c . dietz , pers . comm . ) . such cave entrance inhabiting nephilid populations have been discovered in east and south east asia as well as in the neotropics . so far it is unknown to what extend cave - roosting bats flying back and forth between caves and foraging areas are able to detect and avoid these webs . since it is hypothesized that the bats might navigate by spatial memory while passing through cave entrances [ 121 ] , it is conceivable that some of them may crash into nephilid webs within the cave’s entrance zone , given the fact that in some areas they leave caves at dusk in gigantic swarms . monitoring nephild webs at cave entrances by means of video recording devices could bring an answer to this question .
the social behavior of the greater sac - winged bat can be observed easily from some distance . the mating system has been described as harem - polygynous , but recent studies using molecular genetic techniques proved that harem males father only 30 % of the offspring within their harems . however , harem holders have on average a higher reproductive success than non - harem males and , therefore , males benefit from defending a group of females . the courtship of male s . bilineata includes visual , acoustic , and olfactory displays . besides a large number of social calls that have been described for this species , males emit songs that can last for up to an hour and that include many different syllables . songs of male s . bilineata are partly in the audible range of humans . in addition to singing , males also perform hovering flights during which fragrances are fanned towards roosting females . males possess
the neotropical orb - weaving spider nephila clavipes has been witnessed catching bats quite frequently ( 9 reports ) , but in none of these cases was it seen biting , wrapping or eating a bat . likewise , birds trapped in the webs of this spider species were apparently not consumed [ 22 ] – . only once has a neotropical nephila been observed biting a bird , but without subsequent consumption of the prey [ 22 ] . it has been suggested that nephila clavipes might be unable to deal with large , aggressive prey such as bats and birds [ 24 ] , [ 27 ] . if this latter assumption is true , then the captures of bats in the webs of nephila clavipes would be cases of non - predation deaths ( the bats dying of exhaustion , starvation , dehydration , and / or hyperthermia ) . the two european incidences where bats were killed in spider webs without the spiders feeding on them must be considered to have been cases of non - predation deaths as well .
in recent years , the idea has been proposed that the occasional catch of large , energetically rewarding prey may be essential in order to fulfil the reproductive needs of large orb - weaving spiders ( “rare , large prey” hypothesis ; see [ 122 ] – [ 123 ] ) . while large orb - weavers such as nephila spp . capture predominantly small insects of little energetic value , they derive the bulk of their energy from a few rare , large prey items ( see [ 46 ] , [ 49 ] , [ 53 ] , [ 122 ] ) . in this context “rare , large prey” encompasses large insects ( e . g . , cicadas , moths , coleopterans , orthopterans , and odonates ) as well as small flying vertebrates ( bats and birds ) . in our opinion , the examples of bat - eating orb - weavers reported in this paper are consistent with the
rare , large prey
hypothesis , though one may object to this given the rarity of such events .
at night , a constant observation of the whole social groups was not possible , because the groups split up between different sites . thus , night observations were focused on specific sites within the roost , where only a fraction of the whole group was roosting . these focal site observations were conducted during an overall period of 23 : 02 h in 20 individual nights between september and november 2014 . the focal sites were illuminated with two led - infrared spotlights , recorded with a night vision camera ( bell & howell dnv16hdz full hd rouge ) , and bats were simultaneously monitored on the video camera screen . parallel to this setting a dslr camera was used to capture the coloured bands and allow individual identification of bats at the beginning of each recording session and each time a new bat arrived . in addition to focal site observations , the whole roost was constantly checked for behavioural interactions during an overall period of 24 : 32 h throughout 12 nights in october and november 2014 . during that time , behavioural observations were carried out by constantly scanning the extending roof back and forth with an analogue night scope ( yukon 3 × 50 exelon ) . if an interaction was observed , a photo was taken to determine the involved individuals .
there are 19 reports of neotropical bats being captured in orb - webs of large araneids or nephilinids . these reports refer to incidences witnessed in peru ( report # 7 ) , colombia ( report # 8–10 ) , guatemala ( report # 11 ) , belize ( report # 12–13 ) , costa rica ( report # 14–20 ) , panama ( report # 21–24 ) , or elsewhere in central america ( report # 25 ) and are depicted in fig . 2b–e , g and h . several of these observations were made by researchers stationed at the los amigos biological station in peru , the biological stations la selva and la sirena in costa rica and the smithsonian tropical research station barro colorado island in panama , which are all located in lowland tropical rainforest ( [ 8 ] , [ 30 ] , [ 37 ] – [ 38 ] ; m . knörnschild , unpubl . data ; m . eckenweber , pers . comm . ; m . mallo & c . díez , pers . comm . ; m . nagy , pers . comm . ; h & g . unger , pers . comm . ; m . weber , pers . comm . ) . in the incidences witnessed in costa rica and panama , the spiders had constructed their webs on the outside of or in close proximity to buildings inhabited by bat colonies ( [ 8 ] ; m . knörnschild , unpubl . data ; c . metcalf , pers . comm . ) .


neritimorpha is a taxonomic grouping , an unranked clade of snails , gastropod mollusks .

this grouping includes land snails , sea snails , some deepwater limpets , and also freshwater snails .

this clade used to be known as the superorder neritopsina .

the clade neritimorpha is , based on optimal phylogenetic analysis , deemed monophyletic .

neritimorpha

the neritimorpha are a group of primitive
prosobranch
gilled snails which have a shelly operculum .
the phylogenetic position of neritimorpha based on the mitochondrial genome of nerita melanotragus ( mollusca : gastropoda ) .
gofas , s . 2009 . neritimorpha . accessed on february 7 , 2017 through : molluscabase at urltoken .
the phylogenetic position of neritimorpha based on the mitochondrial genome of nerita melanotragus ( mollusca : gastropoda ) . - pubmed - ncbi
connected with description and systematic evaluation of related neritimorpha ( gastropoda ) . — mitteilungen aus dem geologisch - paläontologischen institut der universität hamburg
2001 . the new family nerrhenidae ( neritimorpha , gastropoda ) from the givetian of germany . — neues jahrbuch für geologie und paläontologie , monatshefte
2003 . relationships of cretaceous neritimorpha ( gastropoda , mollusca ) , with the description of seven new species . — bulletin of the czech geological survey
1999 . notes on the evolution and higher classification of the subclass neritimorpha ( gastropoda ) with the description of some new taxa . — geologica et palaeontologica
neritopsoidea is a taxonomic grouping , a superfamily of sea snails , marine gastropod mollusks in the clade cycloneritimorpha , within the clade neritimorpha , ( according to bouchet & rocroi , 2005 ) , or in the order neritoina within superorder cycloneritimorpha within the subclass neritimorpha , ( according to bandel , 2007 ) .
2003 . systematic position of cyrto - neritimorpha within class gastropoda with description of two new genera from siluro - devonian strata of central europe . — bulletin of geosciences
2000 . the new family cortinellidae ( gastropoda , mollusca ) connected to a review of the evolutionary history of the subclass neritimorpha . — neues jahrbuch für geologie und paläontologie , abhandlungen
while less diverse than caenogastropoda and heterobranchia , the clade neritimorpha–which has ~ 2 , 000 extant species–is also ecologically diverse , with members occupying marine , freshwater , and terrestrial habitats ( uribe et al . , 2016 ) . similar to both of these larger clades , slug - and limpet - like forms have also independently evolved within the neritimorpha ( uribe et al . , 2016 ) .
for these reasons some taxa have been abandoned , like the traditional subclass of the prosobranchs ( abandoned as paraphyletic ) that has now been divided into the subclasses patellogastropoda , vetigastropoda , cocculiniformia , neritimorpha , caenogastropoda and the small group of neomphalina .
it is estimated ( zapata et al . , 2014 ; uribe et al . , 2016 ) that neritimorpha originated sometime in the late paleozoic and may share some degree of relationship to the platyceratids ( frýda et al . , 2009 ) , which were important gastropods in the early paleozoic .
uribe , j . e . , d . colgan , l . r . castro , y . kano , and r . zardoya . 2016 . phylogenetic relationships among superfamilies of neritimorpha ( mollusca : gastropoda ) . molecular phylogenetics and evolution 104 , 21 - 31 . doi : urltoken .
some marine genera of neritimorpha are associated with particular marine environments ; for example , smaragdia occurs in association with seagrass , while neritina lives in brackish water environments ( costa et al . , 2001 ) . finding these same taxa in the fossil record therefore is very useful for interpreting past environments .
heutige neritimorpha mit planktotropher larvalentwicklung besitzen eine konvolute , glatte larvalschale , deren windungen innen resorbiert sind . die ältesten bekannten larvalschalen dieses typs sind triassischen alters . gut erhaltene spätpaläozoische neritimorphe gastropoden besitzen larvalschalen mit zwei oder mehr wohl separierten , schnell zunehmenden windungen . diese larvalschalen gleichen planktotrophen larvalschalen von caenogastropoden . dieser larvalschalentyp ist möglicherweise bei neritimorpha und caenogastropoda plesiomorph . permische und oberkarbonische neritimorphe ( naticopsis , trachyspird ) haben glatte larvalschalen ( naticopsidae ) oder larvalschalen mit starken axialrippen ( trachyspiridae , neue familie ) . die stark kon volute niedrig gewundene , runde schalengestalt der modernen neritimorphen entwickelte sich zusammen mit der resorbierung der innenwindung des teleoconchs und protoconchs bzw . beides bedingte einander . die moderne neritimorphenschale mit einem einheitlichen , undifferenzierten innenraum lässt sich von einem naticopsiden vorfahren herleiten , der noch nicht resorbiert war . eine herausgehobene spira , tiefe suturen sowie eine gewundene herausragende larvalschale hätten eine intern resorbierte schale vermutlich anfällig für mechanische verletzung und angriffe von räubern gemacht . die annahme , die aufwindung habe sich bei neritimorphen unabhängig von anderen schnecken entwickelt ist unwahrscheinlich und steht im widerspruch zum fossilbericht . die larvalschale der modernen neritiden entwickelte sich vermutlich aus einer glatten , niedrig gewundenen larvalschale , wie sie hier bei karbonischen naticopsiden dokumentiert wird .
in the bouchet & rocroi taxonomy , clades are used between the rank of class and the rank of superfamily . the clades are unranked . bouchet and rocroi use six main clades : patellogastropoda , vetigastropoda , cocculiniformia , neritimorpha , caenogastropoda , and heterobranchia . the first three of these major clades have no nesting clades within them : the taxonomy goes immediately to the superfamily level . within the caenogastropoda there is one extra clade . in contrast , within the heterobranchia , for some of the nudibranch groups there are six separate clades above the level of superfamily , and in the case of most of the land snails , there are four clades above the level of superfamily .
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bouchet , p . ; rocroi , j . - p . ( ed . ) ( 2005 ) . classification and nomenclator of gastropod families . malacologia : international journal of malacology , 47 ( 1 - 2 ) . conchbooks : hackenheim . isbn 3 - 925919 - 72 - 4 . 397 pp .
part of : malacologia : international journal of malacology . institute of malacology : ann arbor . issn 0076 - 2997 , more
this is a vademecum for about 2 , 400 suprageneric names in recent and fossil gastropods , from the subtribe to the superfamily . for each name full bibliographic reference is given , with date of publication , type genus , nomenclatural availability and validity under the rules of the iczn . further 730 names of higher taxa are listed separately . altogether 611 valid families , 202 of them exclusively fossil , are recognised in the proposed classification .
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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 ]
department of marine ecosystems dynamics , atmosphere and ocean research institute , university of tokyo , tokyo , japan .
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helicina clappi ( helicinidae ) , a terrestrial relative of nerites from florida . picture : robert pilla ( jacksonville shell club ) .
fresh water nerite ( theodoxus danasteri ) . view of the animal and its operculum . picture : © alexander mrkvicka .
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full reference : a . n . golikov and y . i . starobogatov . 1975 . systematics of prosobranch gastropods . malacologia 15 ( 2 ) : 185 - 232
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programa de biología , dir . grupo de investigación en evolución y sistemática molecular , universidad del magdalena , santa marta , colombia . lydaraquelcastro @ urltoken
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from bermuda ( pri 70104 ) . note that the operculum of the shell is
gofas ( 2009 ) recognizes the following gastropod families as members of the heterobranchia ( † indicates extinct families ) . this list was copied from molluscabase on february 7 , 2017 and is reposted here in concordance with the source material’s creative commons by 4 . 0 license . the link associated with each name will direct you to additional information about that family on molluscabase .
family dawsonellidae wenz , 1938 † family deianiridae wenz , 1938 † family helicinidae férussac , 1822 family neritiliidae schepman , 1908 family proserpinellidae h . b . baker , 1923 family proserpinidae gray , 1847 family hydrocenidae troschel , 1857 family naticopsidae waagen , 1880 † family trachyspiridae nützel , frýda , yancey & anderson , 2007 † family neridomidae bandel , 2008 † family neritidae rafinesque , 1815 family parvulatopsidae gründel , keupp & lang , 2015 † family phenacolepadidae pilsbry , 1895 family pileolidae bandel , gründel & maxwell , 2000 † family cortinellidae bandel , 2000 † family delphinulopsidae blodgett , frýda & stanley , 2001 † family globocornidae espinosa & ortea , 2010 family neritopsidae gray , 1847 family plagiothyridae knight , 1956 † family pseudorthonychiidae bandel & frýda , 1999 † family titiscaniidae bergh , 1890 family symmetrocapulidae wenz , 1938 †
family craspedostomatidae wenz , 1938 † family pragoscutulidae frýda , 1998 † family nerrhenidae bandel & heidelberger , 2001 † family oriostomatidae koken , 1896 † family tubinidae knight , 1956 † family palaeotrochidae knight , 1956 † family platyceratidae hall , 1879 † ; learn more at the ordovician atlas .
costa , f . h . a . , r . h . nehm , and c . s . hickman . 2001 . neogene paleontology of the northern dominican republic 22 : the family neritidae ( mollusca : gastropoda ) . bulletins of american paleontology 359 .
frýda , j . , p . r . racheboeuf , b . frýdová , l . ferrová , m . mergl , and s . berkyová . 2009 . platyceratid gastropods — stem group of patellogastropods , neritimorphs or something else ? bulletin of geosciences 84 : 107 - 120 . doi : 10 . 3140 / bull . geosci . 1125 .
zapata , f . , n . g . wilson , m . howison , s . c . s . andrade , k . m . jörger , m . schrödl , f . e . goetz , g . giribet , and c . w . dunn . 2014 . phylogenomic analyses of deep gastropod relationships reject orthogastropoda . prooceeding of the royal society , b , 281 : 20141739 . doi : urltoken . link
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1997 . higher classification and pattern of evolution of the gastropoda . a synthesis of biological and paleontological data . — courier forschungs - institut senckenberg
2002 . reevaluation and classification of carboniferous and permian gastropoda belonging to the caenogastropoda and their relation . — mitteilungen aus dem geologisch - paläontologischen institut der universität hamburg
1994 . the late cretaceous gastropod fauna from ajka ( bakony mountains , hungary ) : a revision . — annalen des naturhistorischen museums wien
2002 . larval shells and shell microstructures of exceptionally well - preserved late carboniferous gastropods from the buckhorn asphalt deposit ( oklahoma , usa ) . — senckenbergiana letheae
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1988 . on the origin and evolution of major gastropod groups , with special reference to the streptoneura . — journal of molluscan studies
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, m . 1990 : mikromorphe mollusken und brachiopoden aus dem oberkarbon des rheinisch - westfälischen steinkohlereviers : systematik , paläoökologie und stratigraphie . — unpublished ph . d dissertation , 141 p . , westfälische wilhelms - universität , münster .
2004 . the evolution of conch ontogeny in mesozoic open sea gastropods . — palaeontologia polonica
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2006 . usefulness of the opercular nucleus for inferring early development in neritimorph gastropods . — journal of morphology
2002 . major adaptive radiation in ner - itopsine gastropods estimated from 28s rrna sequences and fossil records . — proceedings of the royal society of london b
1933 . the gastropods of the st . louis , missouri , pennsylvanian outlier : vi . the neritidae . — journal of paleontology
1934 . the gastropods of the st . louis , missouri , pennsylvanian outlier : vii . the euomphalidae and platyceratidae . — journal of paleontology
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2004 . a late permian chinese gastropod species , possibly larval , in the middle pennsylvanian of new mexico . — journal of paleontology
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1994 . early carboniferous mollusca from the tamworth belt , new south wales , australia . — records of the australian museum
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classification of the gastropod molluscs in the three traditional subclasses of the prosobranchs ( marine snails ) , opisthobranchs ( sea slugs and sea hares among others ) and pulmonates ( land snails and land slugs ) has ruled from the first third of the 20th century until very recently , and still remains in most zoology textbooks .
modern phylogenetic evaluation techniques , based on morphological , embryological , molecular and combined characters , have allowed to demonstrate that the phylogenies of these groups are not that simple and there has been a very significant change in different groups when considering evolutionary independent lines ( monophyletic ) .
the concepts of opisthobranchs and pulmonates ( until now infraclases of the heterobranchia , joined under the term euthyneura based on the structure of their nervous system ) have been replaced by the following groupings with category of subterclass : acteonimorpha ( grouping the traditional acteonoidea and rissoelloidea ) , tectipleura ( grouping the traditional euopisthobranchia and pulmonata ) and ringipleura ( grouping nudipleura and ringiculimorpha ) .
the bases of this new classification are in the enormous taxonomic work by bouchet & rocroi ( 2005 ) , recently updated by bouchet et al . ( 2017 ) and followed by world register of marine species ( worms ) .
the taxonomic synopsis of each species descriptive sheets at opk - opisthobranchs are basically according to these ideas .
except for heterobranchia , the previous subclasses correspond to groups of former “prosobranchs” , preferably marine snails , of separated genders , with pallial cavity and typical gills . the groups marked with the dagger ( † ) are fossil taxa .
burmeister , 1837 : includes the gastropoda that have a gill origin different from that of the other gastropods ( former prosobranchs ) , and comprises species marine , freshwater and terrestrial snails .
includes shelled species of marine and freshwater snails , of still uncertain relationships , which had traditionally been considered prosobranchs or opisthobranchs . until recently it also included the superfamilies acteonoidea , pyramidelloidea and ringiculoidea ( for some time they have not been considered opisthobranchs ) but now these superfamilies are considered again within the euthyneura and therefore no longer belong to this group , actually formed by :
: includes a few groups of the fossil register whose adscription to the other groups is not clear .
, includes those that were traditionally considered opisthobranchs ( slugs almost exclusively marine , with or without shell ) and pulmonates ( terrestrial and marine slugs and snails ) .
, of pelagic habits , they can swim thanks to body extensions that they use as swimming fins .
suborder gymnosomata , without a shell , carnivorous , feeds on euthecosomata and pseudothecosomata .
superorder eupulmonata , this superorder and the following groups includes species of snails and slugs , preferably terrestrial or freshwater , hermaphroditic and whose pallial cavity has become a lung without gills whose epithelium exchanges gases with the outside .
superorder [ unassigned ] euthyneura includes those species that could not be clearly assigned to other groups .
jörger et al . ( 2010 ) carried out a phylogenetic analysis of the small group of acochlidia to investigate their origin and their relationships with the other heterobranchia groups . after a short time , schrödl et al . ( 2011 ) analyzed the phylogeny of the euthyneura based on molecular analysis and obtained results that substantially modified the established taxonomic and phylogenetic schemes . these works were the trigger - taking advantage of the great advances in molecular techniques and their popularization - that started the long journey of discovering the relationships among these gastropods in the last decade .
due to the complexity of the problem , we must emphasize that there are still doubts in many points of the phylogenetic cladogram , more molecular analysis are necessary to confirm or discard previous statements . the latest published works , such as bouchet et al . ( 2017 ) try to synthesize and consolidate all the researcher’s discoveries in this complex matter , of which the “last say” has not been said… yet .
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neritopsoidea was placed in the order neritoida , the superorder neritopsina and the subclass orthogastropoda according to the taxonomy of the gastropoda by ponder & lindberg , 1997 .
† naticopsidae - bandel ( 2007 ) recognizes natisopsinae ( in neritopsidae by bouchet & rocrois 2005 ) at family level .
cuvier , g . 1795 . second mémoire sur l ' organisation et les rapports des animaux à sang blanc , dans lequel on traite de la structure des mollusques et de leur division en ordre , lu à la société d ' histoire naturelle de paris , le 11 prairial an troisième [ 30 may 1795 ] . magazin encyclopédique , ou journal des sciences , des lettres et des arts , 1795 [ 1 . année ] 2 : 433 - 449 . , available online at urltoken page ( s ) : 448 [ details ]
nomenclature the taxon gastropoda was first introduced by cuvier ( 1795 : 448 , as french vernacular name
les gastéropodes
) , alongside . . .
nomenclature the taxon gastropoda was first introduced by cuvier ( 1795 : 448 , as french vernacular name
les gastéropodes
) , alongside with
les céphalopodes
and
les acéphales
, proposed as three orders of his
classe des mollusques
. the name was first latinized by duméril ( 1805 : 160 ) . [ details ]
bouchet p . & rocroi j . - p . ( 2005 ) . classification and nomenclator of gastropod families . malacologia . 47 ( 1 - 2 ) : 1 - 397 isbn 3 - 925919 - 72 - 4 . [ details ]
ponder w . f . & lindberg d . r . 1997 . towards a phylogeny of gastropod molluscs : an analysis using morphological characters . zoological journal of the linnean society , 119 : 83 - 265 . [ details ]
zapata , f . ; wilson , n . g . ; howison , m . ; andrade , s . c . s . ; jorger , k . m . ; schrodl , m . ; goetz , f . e . ; giribet , g . ; dunn , c . w . ( 2014 ) . phylogenomic analyses of deep gastropod relationships reject orthogastropoda . proceedings of the royal society b : biological sciences . 281 ( 1794 ) : 20141739 - 20141739 . , available online at urltoken [ details ]
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freshwater snails are gastropod mollusks which live in freshwater . there are many different families . they are found throughout the world in various habitats , ranging from ephemeral pools to the largest lakes , and from small seeps and springs to major rivers . the great majority of freshwater gastropods have a shell , with very few exceptions . some groups of snails that live in freshwater respire using gills , whereas other groups need to reach the surface to breathe air . most feed on algae , but many are detritivors and some are filter feeders .
( some of the highlighted families consist entirely of freshwater species , but some of them also contain , or even mainly consist of , marine species . )
basommatophora : chilinidae , latiidae , acroloxidae , lymnaeidae . planorbidae ► and physidae - all these six families together form the clade hygrophila
( marine gastropods ( siphonarioidea , sacoglossa , amphiboloidea , pyramidelloidea ) are not depicted within panpulmonata for simplification . some of these highlighted families consist entirely of freshwater species , but some of them also contain , or even mainly consist of , marine species . )
, largely confined to the tropics , also the rivers of europe , family includes the marine
nerites
.
the caenogastropoda are a large group of gilled operculate snails , which are largely marine . in freshwater habitats there are ten major families of caenogastropods , as well as several other families of lesser importance :
, an exclusively freshwater family that is largely tropical and includes the large
apple snails
kept in aquaria .
, medium to large snails , live - bearing , commonly referred to as
mystery snails
. worldwide except south america , and everywhere confined to fresh waters .
, family native to rivers draining to the mediterranean , also middle east , and some south pacific islands .
native to south and central america . formerly included with the pleuroceridae by many authors .
- about 100 species in south asia , diverse in african lakes , and sri lanka .
, abundant and diverse in eastern north america , largely high - spired snails of small to large size .
snails of northwestern north america . formerly included with the pleuroceridae . about 50 species .
, high - spired parthenogenic snails of the tropics , includes those referred to as
trumpet snails
in aquaria .
, small amphibious snails scattered worldwide , most diverse in eastern and southeast asia .
, small low - spired snails referred to as
valve snails
. 71 species .
basommatophorans are pulmonate or air - breathing aquatic snails , characterized by having their eyes located at the base of their tentacles , rather than at the tips , as in the true land snails stylommatophora . the majority of basommatophorans have shells that are thin , translucent , and relatively colorless , and all five freshwater basommatophoran families lack an operculum .
, small to medium - sized snails confined to temperate and cold south america .
these are the dextral ( right - handed ) pond snails . about 100 species .
, left - handed ( sinistral )
pouch snails
, native to europe , asia , north america .
in the developed world , people encounter freshwater snails most commonly in aquaria along with tropical fish . species available vary in different parts of the world . in the united states , commonly available species include ramshorn snails such as planorbella duryi , apple snails such as pomacea bridgesii , the high - spired thiarid malaysian trumpet snail melanoides tuberculata , and several neritina species .
& bouchet p . ( 2008 ) .
global diversity of gastropods ( gastropoda ; mollusca ) in freshwater
.
strong e . e . , colgan d . j . , healy j . m . , lydeard c . , ponder w . f . & glaubrecht m . ( 2011 ) .
phylogeny of the gastropod superfamily cerithioidea using morphology and molecules
.
dillon r . t . ( 2006 ) . chapter 21 . freshwater gastropoda . pages 251 - 259 . in : sturm c . f . , pearce t . a . & valdés a . ( eds . ) ( 2006 ) . the mollusks : a guide to their study , collection , and preservation . american malacological society , 445 pp . isbn 978 - 1 - 58112 - 930 - 4 .
jörger k . m . , stöger i . , kano y . , fukuda h . , knebelsberger t . & schrödl m . ( 2010 ) .
on the origin of acochlidia and other enigmatic euthyneuran gastropods , with implications for the systematics of heterobranchia
.
banarescu p . ( 1990 ) . zoogeography of fresh waters , vol . 1 , general distribution and dispersal of freshwater animals . aula - verlag , weisbaden .
schrödl m . & neusser t . p . ( 2010 ) .
towards a phylogeny and evolution of acochlidia ( mollusca : gastropoda : opisthobranchia )
.
entre la política y economía : resultados preliminares de las primeras temporadas del proyecto arqueológico motul de san josé .
xiii simposio de investigaciones arqueológicas en guatemala , 1999 ( edited by j . p . laporte , h . escobedo , b . arroyo and a . c . de suasnávar )
chandler a . c . & read c p . ( 1961 ) . introduction to parasitology . john wiley and sons , new york . 822 pp .
adams a . m . ( 2006 ) . foodborne trematodes . in : ortega i . r . ( ed . ) ( 2006 ) . foodborne parasites . isbn 0 - 387 - 30068 - 6 . page 178 .
adema c . m . , bayne c . j . , bridger j . m . , knight m . , loker e . s . , yoshino t . p . & zhang s . - m . ( 2012 ) .
will all scientists working on snails and the diseases they transmit please stand up ?
.
j . mouthon , typology of molluscs of flowing water ; biotypological organisations ; socioecological groupings ; annls limnol . volume 17 , number 2 , 1981
haynes a . ( 2000 ) .
the distribution of freshwater gastropods on four vanuatu islands : espiritu santo , pentecost , éfate and tanna ( south pacific )
.
vermeij j . & wesselingh f . p . ( 2002 ) .
neogastropod molluscs from the miocene of western amazonia , with comments on marine to freshwater transitions in molluscs
.
this article is issued from wikipedia - version of the 11 / 7 / 2016 . the text is available under the creative commons attribution / share alike but additional terms may apply for the media files .
according to the taxonomy of the gastropoda ( ponder & lindberg , 1997 ) neritopsina is a gastropod superorder in the subclass orthogastropoda . the superfamily palaeotrochoidea is contained within neritopsina but its order placement is undetermined .
. it is one of the 6 highest clades in gastropoda . it contains the clades
^ bouchet p . & rocroi j . - p . ( ed . ) ; frýda j . , hausdorf b . , ponder w . , valdes a . & warén a . 2005 . classification and nomenclator of gastropod families . malacologia : international journal of malacology , 47 ( 1 - 2 ) . conchbooks : hackenheim , germany . isbn 3 - 925919 - 72 - 4 . issn 0076 - 2997 . 397 pp . urltoken
— the taxonomy of the gastropoda as it was revised by philippe bouchet and jean pierre rocroi is currently the most up to date overall system for classifying gastropod mollusks . ( gastropods are a taxonomic class of animals which consists of snails… …
— la taxonomía actual de las gastropoda es la revisión de bouchet rocroi ( 2005 ) . [ 1 ] la sistemática según bouchet rocroi está reconocido como sistemática de referencia [ 2 ] [ 1 ] …
— die gegenwärtig aktuelle taxonomie der schnecken ist die revision von bouchet rocroi ( 2005 ) . [ 1 ] die systematik von bouchet rocroi ist als referenzsystem anerkannt . [ 2 ] [ 3 ] [ 4 ] die taxonomie von bouchet rocroi ( 2005 ) ist ein schritt weiter in… …
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get taxonomy of the gastropoda bouchet & rocroi , 2005 essential facts below . view
the taxonomy of the gastropoda as it was revised in 2005 by philippe bouchet and jean - pierre rocroi is a system for the scientific classification of gastropod mollusks . ( gastropods are a taxonomic class of animals which consists of snails and slugs of every kind , from the land , from freshwater , and from saltwater . ) the paper setting out this taxonomy was published in the journal malacologia . the system encompasses both living and extinct groups , as well as some fossils whose classification as gastropods is uncertain .
in some parts of the taxonomy , instead of
clade
, bouchet and rocroi labelled groupings of taxa as a
group
or an
informal group
. a clade must by definition contain only one lineage , and it was considered to be the case that these
informal groups
may either contain more than one lineage , or only contain part of a lineage . further research will eventually resolve these questions . since the publication of this taxonomic system in 2005 , various proposals for changes have been published by other authors , for more information see changes in the taxonomy of gastropods since 2005 .
in 2017 this taxonomy was superseded by a revised taxonomy
revised classification , nomenclator and typification of gastropod and monoplacophoran families
by philippe bouchet & jean - pierre rocroi , bernhard hausdorf , andrzej kaim , yasunori kano , alexander nützel , pavel parkhaev , michael schrödl and ellen e . strong in malacologia , 2017 , 61 ( 1 - 2 ) : 1 - 526 . . [ 1 ]
systems of classification such as this one are primarily of value to malacologists ( people who study mollusks ) and other biologists . biological classification schemes are not merely a convenience , they are an attempt to show the actual phylogeny ( the evolutionary relatedness ) within a group of organisms . thus a taxonomy such as this one can be seen as an attempt to elucidate part of the tree of life , a phylogenetic tree .
the bouchet & rocroi 2005 system of gastropod taxonomy was laid out in a book - length paper entitled
classification and nomenclator of gastropod families
, which was published in the journal malacologia [ 2 ] and which was written in collaboration with j . frýda , b . hausdorf , w . ponder , á . valdés and a . warén . this system supersedes the system of ponder and lindberg from 1997 . subsequent revisions by other authors have been made since the publication of this paper .
in this taxonomy 611 valid families are recognized . of these , 202 families are exclusively fossil , and this is indicated here with a dagger + . superfamily names are standardized so that they all end in the suffix
- oidea
, also commonly used for superorders and subclasses , replacing the
- acea
ending found especially in the older literature . classification of families into subfamilies however is often not well resolved , and should be regarded as the best possible hypothesis .
the publication bouchet & rocroi ( 2005 ) [ 2 ] also includes a nomenclator of about 2400 suprageneric taxa ( taxa above the level of genus ) of gastropods , from the subtribe to the superfamily . a full bibliographic reference is provided for each taxon , giving the name of the authority , the original publication , the date of that publication , the type genus for the taxon , its nomenclatural status , and its validity under the rules of the iczn .
since the publication of this taxonomic system in 2005 , various changes have been published by other authors , for more information see changes in the taxonomy of gastropods since 2005 .
here the information is displayed in the form of a cladogram ( an evolutionary tree of descent . ) it is worth bearing in mind however that this taxonomy is provisional : many of the taxa are still only known as
groups
or
informal groups
, and these are very likely to be changed as more information becomes available .
this cladogram is based on the following information . the list format used below makes clear which taxa are informal groups rather than clades :
in the following , more detailed list , indentation is used only for the ranks of superfamily and family . the clade names are not indented , but their hierarchy is indicated by the size of the font used . a clearer sense of the hierarchy of the clades can be drawn from the list immediately above this one .
+ family procerithiidae - if the genus argyropeza is placed in the procerithiidae , then this family is not all fossil .
contains the clades cephalaspidea , thecosomata , gymnosomata , aplysiomorpha , sacoglossa , umbraculida , nudipleura and the groups acochlidiacea and cylindrobullida .
family aegiretidae - aegiretidae is incorrect subsequent spelling of this family in bouchet & rocroi ( 2005 ) . correct is aegiridae .
revised classification , nomenclator and typification of gastropod and monoplacophoran families ; malacologia 61 ( 1 - 2 ) : 1 - 526 . 2017
guido t . poppe ; sheila p . tagaro ( 2006 - 02 - 23 ) .
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