the tufted puffin ( fratercula cirrhata ) , also known as crested puffin , is a relatively abundant medium-sized pelagic seabird in the auk family ( alcidae ) found throughout the north pacific ocean .

it is one of three species of puffin that make up the genus fratercula and is easily recognizable by its thick red bill and yellow tufts .

tufted puffin

figure 1 . distribution of the tufted puffin in north america and easternmost russia .
the tufted puffin is the largest puffin and is characterized by the long , straw - colored feathers extending back from its crown .
click the range map to learn more about the distribution of tufted puffin in washington .
figure 4 . native american ( tlingit ) dance apron , fringed with tufted puffin bills .
) and may not return to coastal breeding areas for several years . thus , the tufted puffin , even more so than the horned puffin (
the tufted puffin nests mostly in deep burrows that it digs into a cliff by the ocean .
description : the tufted puffin differs from atlantic puffin and horned puffin by its dark body . these three puffins belong to the genus fratercula which means “little brother” . all have the large , typical , laterally compressed beak .
the latest sighting details and map for tufted puffin are only available to our birdguides ultimate or our birdguides pro subscribers .
in the south but not in the north pacific . the tufted puffin and the horned puffin however , are absent from the north atlantic and instead can be found in large colonies in the northern
the tufted puffin is threatened by unsustainable levels hunting , fishing or
by - catch ( coincidental capture in fishing gear ) .
one of the natural predators of the puffin are gulls ( i wrote about puffin predators here ) . this poor little puffin was trying to bring its puffling some sand eels when this gull flew in and took it…mean gull ! you can read the whole story the puffin that got . . .
the oldest recorded tufted puffin was at least 6 years old when it was found in alaska , the same state where it had been banded .
voice : the tufted puffin , as other puffins , is mostly silent outside colonies where it utters some “arrr” sounds , groans and raucous growls .
1 . iucn : red list of threatened species urltoken 13 . washington dept . of fish and wildlife : status report for the tufted puffin urltoken
or swim . in the 1800s and 1900s , the puffin was commonly hunted for its meat and eggs which led to a severe decline in the world ' s puffin population , even completely eliminating some puffin colonies . in the 21st and late 20th century , however , the puffin has enjoyed better protection and
baird , p . h . 1991 . optimal foraging and intraspecific competition in the tufted puffin . condor no . 93 : 503 - 515 . close
wehle , d . h . s . 1980 . the breeding biology of the puffins : tufted puffin ( lunda cirrhata ) , horned puffin ( fratercula corniculata ) , common puffin ( f . arctica ) , and rhinoceros auklet ( cerorhinca monocerata ) . phd thesis , univ . of alaska , fairbanks . close
tufted puffins form long - term pair bonds to take care of the young .
the tufted puffin nests mostly in deep burrows that it digs into cliff edges and slopes . these burrows can be more than 1 . 5 meters ( 5 feet ) deep .
boone , d . l . 1986 . breeding biology and early life history of the tufted puffin fratercula cirrhata . master ' s thesis , oregon state univ . , corvallis . close
the tufted puffin is a seabird of the open waters , islands , and coastal cliffs of the north pacific . it is larger than other puffin species and distinctive in appearance , with a bold white
face - mask
 and golden head plumes in the breeding season .
predators are a natural part of life . and while uber adorable puffins also have predators . each puffin has its own set of predators . what are the puffin predators ? what preys on puffins ? predators of the atlantic puffin include people ; gulls and skuas including the . . .
seabird ecologist , nathalie hamel volunteering her time to assist with puffin surveys in the san juan islands .
diet : the tufted puffin feeds on small fish , squid , krill , shrimps , crabs and marine worms . it performs pursuit - diving at considerable depths , up to 40 - 50 metres .
…southerly pacific distribution is the tufted puffin ( lunda cirrhata ) , which is black with red legs and bill , a white face , and straw - coloured plumes curving backward from behind the eyes .
vermeer , k . 1979 . nesting requirements , food and breeding distribution of rhinoceros auklets , cerorhinca monocerata , and tufted puffin , lunda cirrhata . ardea no . 67 : 101 - 110 . close
in adult breeding plumage , the tufted puffin has glossy black feathers on its body , a white face , a yellow and orange vertically compressed bill , and long yellow tufts of feathers behind each eye .
amaral , m . j . 1977 . a comparative breeding biology of the tufted and horned puffin in the barren islands , alaska . master ' s thesis , univ . of washington , seattle . close
work is being carried out , particularly in north america , to help to save the remaining puffin populations .
the tufted puffin winters in the pacific ocean north and south of the distribution shown and is rarely found in breeding areas during winter . this species also breeds in other parts of asia . see text for details .
a puffin can dive for up to a minute underwater , but most dives usually last 20 to 30 seconds .
the diet of tufted puffins is mostly small fish . they also eat crustaceans , mollusks , and cephalopods .
the tufted puffin can capture and hold multiple small fish crosswise in its bill , routinely 5 to 20 fish at a time , for delivery to chicks at the nest . adults eat their own food while still under water .
predation by introduced red and arctic foxes , norway rats , ground squirrels , and even brown bears has devastated tufted puffin populations on some alaskan islands and coastal areas . eradication of the foxes in particular has helped renew some populations .
tufted puffins are active at their nest colonies during the day and can often be seen sitting upright on rocks .
both parents feed nestling , carrying fish in bill and dropping them on ground in nest or near entrance . tufted puffin may forage farther from colony than horned puffin , and sometimes 1 - 2 days pass between feeding visits to nest . young leave nest about 6 - 7 weeks after hatching . 1 brood per year , or 2 possibly in south .
december 5 , 2008—in iceland ' s remote westman islands , warming weather is threatening a beloved mascot : the atlantic puffin .
tufted puffins are the size of pigeons , but weigh nearly twice as much ( 1 kg , 2 lbs ) .
tufted puffins are highly social , fishing and traveling in groups of 2 - 20 birds and nesting in large colonies .
the tufted puffin winters at sea , well offshore . it performs post - breeding dispersions , solitary or in pairs . they may move s to waters off s japan and s california . they return to the colonies in late april and may .
along the pacific coast of north america , nesting tufted puffins were traditionally preyed upon by indigenous humans for their meat and eggs . tribes such as the tlingit in the pacific northwest fashioned garments from puffin skins and feathers , using their beaks for ceremonial rattles and apron - fringes . this sort of hunting pressure is mostly a thing of the past . these days , tufted puffins struggle with inadvertently ingesting ocean plastic , getting caught and drowning in gill - nets , and protecting their burrows from introduced mammalian predators such as foxes and rats . since the mid - 1990s , tufted puffin populations in oregon and washington have plummeted more than 95 percent .
source / reference article learn how you can use or cite the puffin article in your website content , school work and other projects .
in winter plumage , the tufted puffin has dark body and head , and tufts are absent . the bill is duller and appears smaller and longer , less triangular . the basal ridge is lacking too . eye - ring , legs and feet are much paler .
tufted puffins are medium - sized , stocky seabirds with rounded heads . they are solid brown - black overall , except for distinctive facial coloration . the outer wings are wider than the inner wings , giving the outstretched wing a hand - like appearance . the feet and bill are orange . in breeding plumage , a tufted puffin has a white mask across its face , ending in a tuft of blonde feathers sweeping back from above the ear . the orange beak is very bright and accented with yellow at the base . like other puffins , this species has a large , laterally compressed bill . the tufted puffin can be distinguished from other puffins by its dark belly .
there is not a lot of information on tufted puffin population trends . the north american waterbird conservation plan estimates a continental breeding population of 2 , 750 , 000 - 3 , 000 , 000 birds , rates the species a 9 out of 20 on the continental concern score , and lists it as a species of low concern . tufted puffin is not on the 2014 state of the birds watch list . bycatch in fishing nets killed tens of thousands of tufted puffins each year into the 1980s . the elimination of drift - nets on the high seas has reduced mortality , although bycatch in coastal fishing nets still kills large numbers of puffins . in addition , nesting tufted puffins are highly vulnerable to red and arctic foxes , river otters , brown bears , and other mammals . such predators were once absent from most islands in the northeast pacific , but were introduced in the 1800s and early 1900s . where present , mammalian predators have devastated or eliminated tufted puffins from many islands , but programs to eradicate the introduced species have led to dramatic recovery of puffin populations . back to top
tufted puffin colonies fluctuate in size annually , based in part upon food supply and climatic events , such as el niño . the tufted puffin is one of the most abundant and conspicuous seabirds nesting in the north pacific . many california populations , however , have disappeared or significantly declined . in recent years , potentially serious declines have occurred in washington as well . oil spills and gill - nets are threats to tufted puffins , as is human disturbance . in washington , most breeding colonies are protected as wildlife refuges or are inaccessible to humans . populations of large gulls , which prey on young puffins , have been increasing in washington , which may account for some of the recent decline in numbers .
no author , 1997 .
puffin
arctic studies center
( on - line ) . accessed march 15 , 2000 at urltoken .
they dive and swim underwater , using their wings to paddle and their feet to steer their way through schools of small fish , which they catch in their bills . a puffin can hold a dozen fish crosswise in their bill until they return to the nest to feed the puffin chick .
piatt , john f . and alexander s . kitaysky . 2002 . tufted puffin ( fratercula cirrhata ) , version 2 . 0 . in the birds of north america ( p . g . rodewald , editor ) . cornell lab of ornithology , ithaca , new york , usa .
since mid - october , hundreds of tufted puffins have washed up dead or starving on the pribilof islands in the middle of the bering sea .
nest on land . with more and more inhabitants though , populations began to drastically suffer leading to the extinction of many puffin colonies throughout the northern hemisphere .
byrd , g . v . , j . c . williams and r . walder . 1991 . status and biology of tufted puffin in the aleutian islands after a ban on salmon driftnets . adak , ak : unpubl . rep . u . s . fish wildl . serv . , alaska maritime nwr . close
reproduction : the peak of laying occurs between late may and mid - june , with some variations of about 1 - 3 weeks according to the breeding range . the laying is usually highly synchronised within the colony . the tufted puffin breeds in small to huge colonies , mainly on steep maritime grassy slopes . this species nests in burrows excavated in sea - facing grassy slopes and flats . like the horned puffin , it also may nest in crevices or cavities in cliffs and boulders .
a puffin can capture and hold multiple small fish crosswise in its bill , routinely 5 to 20 fish at a time , for delivery to chicks at the nest .
wehle , d . h . s . 1982b . food of adult and subadult tufted and horned puffins . murrelet no . 63 : 51 - 58 . close
ceremonial tlingit dance apron , fringed with hundreds of tufted puffin bills that served as rattles ( inset shows detail of puffin bills ) . the woven chilkat blanket panel ( made from goat wool and cedar - bark yarn ) is mounted on buckskin . dates approximately 1840 - 1850 ; collected 1939 , wrangell , ak , by maj . max c . fleischmann . photos and permission to reproduce them here provided by the santa barbara museum of natural history , courtesy of j . timbrook , curator of ethnography .
one . dull white to bluish - white , often spotted with gray and brown . incubation is by both sexes , about 40 - 42 days . young : both parents feed nestling , carrying fish in bill and dropping them on ground in nest or near entrance . tufted puffin may forage farther from colony than horned puffin , and sometimes 1 - 2 days pass between feeding visits to nest . young leave nest about 6 - 7 weeks after hatching . 1 brood per year , or 2 possibly in south .
protection / threats / status : the tufted puffin has large populations throughout the range . severe declines due to introduced predators such as rats , foxes , cats and dogs , are reported over the last 150 years in most of breeding range . human disturbances on breeding islands are important threat too . however , this species is not currently globally threatened .
habitat : the tufted puffin occurs along rocky coasts and offshore islands . it breeds usually on tops of islands or on maritime slopes . unlike the two other puffins , it rarely frequents the coastal cliffs . during winter , it is mostly offshore . it moves into the oceanic area to mid - ocean , beyond the edge of the continental shelf .
wehle , d . h . s . 1982a . color phases in the downy and juvenal plumages of tufted puffins . condor no . 84 : 444 - 445 . close
the tufted puffin is an iconic species of the oregon coast , its likeness gracing untold numbers of t - shirts , coffee mugs , refrigerator magnets and other souvenirs throughout the region . and with good reason : the puffin is an undeniably attractive bird , looking something like a cross between a parrot and a penguin , sharply made up and impeccably dressed . its blonde , back - swept head plumes bespeak an urbane haughtiness ; the contrast of tangerine bill and snow - white mask against its otherwise somber plumage is at once jarring and irresistible .
flight : the tufted puffin shows a stocky black body when flying . it has short , narrow wings , better for underwater pursuit than flight . the flight is laboured , and to take - off from water , it must run some distance over the surface to rise into the air . it is however a powerful flier , able to perform strong wingbeats .
diet : the diet of tufted puffins is mostly small fish which they catch in their beak while swimming . they also eat squids , octopuses , crabs , jellyfish and zooplankton .
our tufted puffins have paired into couples—naia and monty , speedy and val . they have strong personalities and definite likes and dislikes . the noise accompanying their spats sometimes reaches very high levels
tufted puffins have three major predators : snowy owls , bald eagles , and arctic foxes . puffins try to select habitats that will be difficult for foxes to reach , and laying their eggs in burrows makes them inaccessible to the predator . however , foxes seem to prefer the puffin over other birds , making the bird a main target . ( gaston , 1998 , jewett 1953 )
wehle , d . h . s . 1983 . the food , feeding and development of young tufted and horned puffins in alaska . condor no . 85 : 427 - 442 . close
vermeer , k . and l . cullen . 1979 . growth of rhinoceros auklets and tufted puffins , triangle island , british columbia . ardea no . 67 : 22 - 27 . close
puffins are found inhabiting the sea and coastal regions of the northern hemisphere most commonly in the atlantic , pacific and in parts of the arctic circle . the exact location of the puffin is largely dependant on the
the current conservation status and legal protections for tufted puffins include no specific management or recovery requirements for washington’s population . puffins in washington are considered a species of concern by the u . s . fish and wildlife service and have been petitioned for listing under the federal endangered species act , but an evaluation to list the species will not begin until 2016 or 2017 . at the state level , tufted puffins have been a candidate for listing since 1998 and have been identified by audubon washington as a species at high risk from the impacts of climate change . given the rate of recent population decreases , widespread colony abandonment , ongoing threats from multiple factors , and the challenging ocean conditions expected for piscivorous seabirds in the years ahead , tufted puffins are likely to continue declining in washington . if the current 8 . 9 % annual rate of decline continues , the state’s population could become functionally extirpated within about 40 years . for these reasons , it is recommended that the tufted puffin be listed as endangered in washington .
at times a tufted puffin can appear to be walking on water . this occurs when the bird erects itself on the eater , fluffs its feathers , and flaps its wing vigorously to four times in a behavior called wing - flapping . this behavior also occurs when the puffin lies to one side on the water so that one wing is under - water while the other is held in the air vertical to the water . satisfied that one side is clean , the bird turns onto its other side and repeats the wing - flapping . this behavior is often seen in the aquarium’s diving bird habitat in the northern pacific gallery .
puffins can carry 5 - 20 fish back to their nest at a time . the average catch is around 10 fish per trip , but the record in maine is a whopping 62 fish at once ! the puffin ' s beak is specialized to hold lots of fish . the puffin ' s raspy tongue holds fish against spines on the palate , while it opens its beak to catch more fish . ( source : projectpuffin . org )
puffins may dive deeper than 24 m ( 80 ft . ) to catch fish . their bills have specialized sharp hooks that help to hold fish . one puffin was seen holding 62 fish in its bill at one time .
such as skuas and large gulls that compete for the same food that the puffins are hunting . the biggest threat to puffins today though are people and the impact that their increasing activities are having on the puffin ' s natural
gjerdrum , c . 2001 . nestling growth and parental provisioning of tufted puffins ( fratercula cirrhata ) on triangle island , british columbia . master ' s thesis , simon fraser univ . , burnaby , bc . close
baird , p . h . and r . d . jones . 1983 .
tufted puffin ( lunda cirrhata ) .
in the breeding biology and feeding ecology of marine birds in the gulf of alaska . , edited by p . a . baird and p . j . gould , 427 - 469 . natl . oceanic atmos . admin . , ocseap fin . rep . 45 : u . s . dep . comm . close
wehle , d . h . s . 1976 . summer food and feeding ecology of tufted and horned puffins on buldir island , alaska - 1975 . master ' s thesis , univ . of alaska , fairbanks . close
sources : 1 . atlantic puffin information ( date unknown ) available at : [ accessed at : 10 nov 2008 ] 2 . christopher perrins , oxford university press ( 2009 ) the encyclopedia of birds [ accessed at : 01 jan 2009 ] 3 . david burnie , dorling kindersley ( 2008 ) illustrated encyclopedia of animals [ accessed at : 10 nov 2008 ] 4 . david burnie , kingfisher ( 2011 ) the kingfisher animal encyclopedia [ accessed at : 01 jan 2011 ] 5 . dorling kindersley ( 2006 ) dorling kindersley encyclopedia of animals [ accessed at : 10 nov 2008 ] 6 . horned puffin information ( date unknown ) available at : [ accessed at : 10 nov 2008 ] 7 . puffin facts ( date unknown ) available at : [ accessed at : 10 nov 2008 ] 8 . richard mackay , university of california press ( 2009 ) the atlas of endangered species [ accessed at : 01 jan 2009 ] 9 . tom jackson , lorenz books ( 2007 ) the world encyclopedia of animals [ accessed at : 10 nov 2008 ] 10 . tufted puffin information ( date unknown ) available at : [ accessed at : 10 nov 2008 ]
tufted puffins are migratory seabirds of open ocean waters in the winter and coastal islands and rocky cliffs in the summer breeding season . they are the largest and most distinctive in appearance of the three puffin species , and the one that migrates the furthest south in eastern pacific waters to as far as california’s channel islands . they have several other common names—old man of the sea , sea parrot , and the name given to them by the aleuts , toporkie .
tufted puffins protect their young by nesting on offshore islands and in burrows . adults are swift in flight and spend much of their time in the open ocean . puffins may be preyed on by sharks and other large seabirds .
range : the tufted puffin is found in n pacific ocean . this species breeds from s british columbia ( several sites off the washington and oregon coast ) , throughout se alaska and the aleutian islands , kamchatka , the kuril islands and throughout the sea of okhotsk . the southernmost site , the farallon islands shelters a small breeding population , less than 100 pairs . ) it winters at sea along the coasts of kamchatka peninsula and s alaska , through the breeding range .
of all alcids , tufted puffins have the most extensive latitudinal distribution , ranging from japan through the aleutian islands and south from oregon to southern california . few other seabirds call as much of the pacific home as do the puffins .
tufted puffins have a limited range of calls , including a low grumbling noise heard usually from underground in breeding colonies . chicks often peep to indicate that they want food . they use postures and other physical cues to communicate as well .
vermeer , k . , l . cullen and m . porter . 1979a . a provisional explanation of the reproductive failure of tufted puffins lunda cirrhata on triangle island , british columbia . ibis no . 121 : 348 - 354 . close
behaviour : the tufted puffin feeds on small fish , cephalopods and crustaceans all year round . it finds these preys offshore . during the breeding season , it tends to forage in the vicinity of the colony . chicks are exclusively fed with a wide variety of small fish . it pursues the preys underwater and may reach the depth of 40 - 50 metres . it is able to carry up to 20 prey - fish at a time , sometimes more , crosswise in its large beak .
original file name : atlantic puffin ( fratercula arctica ) _ 4677 . jpg resolution : 1881x1263 file size : 810990 bytes date : 2007 : 11 : 09 15 : 05 : 00 upload time : 2007 : 11 : 09 15 : 05 : 36
all appearances aside , tufted puffins are truly remarkable birds . they spend their winters on the open ocean , chasing fish and invertebrates by
flying
hundreds of feet below the waves , yet they dig burrows in soil with their webbed feet
tufted puffins are northern pacific sea birds that spend a majority of the year over the pacific ocean , but nest along coastlines from lower california to alaska , and across the ocean from japan to the shores of northeastern asia . ( gaston 1998 )
nettleship , d . n . , de juana , e . , garcia , e . f . j . & boesman , p . ( 2018 ) . tufted puffin ( fratercula cirrhata ) . in : del hoyo , j . , elliott , a . , sargatal , j . , christie , d . a . & de juana , e . ( eds . ) . handbook of the birds of the world alive . lynx edicions , barcelona . ( retrieved from urltoken on 7 july 2018 ) .
since 2004 , dr . peter hodum has been studying two closely related “puffins , ” the rhinoceros auklet and tufted puffin , on outer coast and salish sea colonies in washington . at this nature night presentation , dr . hodum will cover the natural history of these special seabirds with fascinating stories and photographs . he will compare and contrast the species ' population trends , diet and reproduction , and will discuss what the emerging patterns mean for the ecological health of regional marine ecosystems and the conservation status of these species .
from boats , tufted puffins are easier to see on the breeding colonies than many other alcids , as they are active during the day and can often be seen standing in front of their nesting burrows . however , although the tufted puffin is one of the most abundant seabirds breeding in washington , it is among the least frequently seen , except from boats . during winter , these puffins are seen only occasionally near land . most of the 29 known nesting colonies in washington are on the outer coast . jagged island and alexander island hold the largest colonies , with 7 , 800 and 4 , 000 breeding birds , respectively . the five known inland colonies are located on protection island , seal and sail rocks , smith island , colville island , and bare island .
breeding colonies of tufted puffins are found on islands and some portions of mainland coastlines throughout the north pacific , from the chukchi peninsula in siberia to the channel islands off southern california . they are mostly pelagic ( live at sea ) during the winter .
dastardly plastic–the evils of the plastic industry are reaching our little puffin populations . a tiny little plastic pebble called a nurdle is finding its way by the millions into the sea . these nurdles are used for packing and shipping — to keep * things * . . .
photo by dailymail when you’ve swooped 300 feet into the cold and murky depths of the atlantic for your dinner , you don’t give it up without a fight . even when you’re a tiny puffin and the thieves outnumbering you are three fiercesome black - headed . . .
despite all the successful efforts by renowned ornithologist steven kress ( and the audubon’s project puffin ) to get ( and keep ) a thriving population of atlantic puffins on the eastern coast of the united states ( eastern egg rock off the coast of maine ) , the . . .
the tufted puffins started washing ashore on st . paul island in mid - october—first a handful , then dozens , then so many that volunteers patrolling to collect dead birds began walking their four - wheelers rather than riding . it was easier than getting off every few feet .
migratory patterns are not well known , but tufted puffins are less likely to be seen near shore in winter than in the breeding season . they are probably the most pelagic of the alcids during their non - breeding season , with many birds wintering 60 - 120 miles offshore .
tufted puffins are protected in some areas of their distribution . although populations are slowly being reduced , at this time the numbers appear to be high enough for sustainability of the species . global climate changes on water temperature and food availability may have a negative impact in the future .
such as gulls . puffins mate for life and the female puffin lays a single white egg that is incubated by both parents until it hatches up to six weeks later in the burrow , covered in medium - brownish feathers . the chick is looked after and fed by both parents who bring it
description : the tufted puffin is a medium - sized , stocky , dark seabird with a rounded head . the breeding adult is all black except for a white face and long golden plumes curling over back of head and neck . the thick bill is large and red - orange , with a bright - orange yellow plate over the base . the nonbreeding adult has a dark gray face with no head plumes or bill plate . the large webbed feet are orange . the eye is ringed with orange . males and females look alike , but males are usually slightly larger than females .
tufted puffins can be found in many coastal habitats adjacent to the washington coast and elsewhere in the northern pacific , with the exception of estuaries . they breed in colonies on islands with steep , grassy slopes or on cliff tops . winter habitat is well offshore , in mid - ocean .
populations have apparently declined significantly in washington state waters , including the salish sea region ( puget sound , strait of juan de fuca , strait of georgia ) , during the 1980s and 1990s . however , quantitative assessments of the extent of the decline and of factors influencing puffin population trends are lacking . to begin to evaluate population trends and potential explanatory factors , we used boat - based surveys to assess occupancy of historic breeding sites . to examine reproductive trends , we monitored the puffin population on tatoosh island , washington since 2005 , and are compared breeding performance to that of sympatric seabird species like the rhinoceros auklet .
with intervening yellow markings and in the same way as their legs and feet , the bill of the puffin is more brightly coloured during the warmer breeding season , becoming duller as they shed for the colder winter months . although both males and females are almost identical in appearance , males tend to be slightly larger in
piatt , j . f . and a . s . kitaysky . 2002 .
horned puffin ( fratercula corniculata ) .
in the birds of north america , no . 611 . , edited by a . poole and f . gill . philadelphia , pa : the birds of north america , inc . close
) conducted the earliest directed studies on tufted puffins and collected wholly new information on breeding biology , behavior , chick growth , diets , and habitat use in the aleutians , gulf of alaska , and british columbia . much incidental information is also available from early general field investigations of seabird colonies ( compiled in
range / habitat : tufted puffins can be found in many coastal habitats adjacent to the washington coast and elsewhere in the northern pacific , with the exception of estuaries . they breed in colonies on islands with steep , grassy slopes or on cliff tops . winter habitat is well offshore , in mid - ocean .
harrison , p . seabirds : an identification guide . boston . houghton mifflin company . 1983 . harrison , p . seabirds of the world : a photographic guide . princeton , nj . princeton university press . 1996 . herron baird , p .
optimal foraging and intraspecific competition in the tufted puffin
. the condor . 93 : 503 - 515 . 1991 . scott , s . l . ( editor ) . field guide to the birds of north america , 2nd ed . washington , d . c . national geographic society . 1987 . gough , g . a . , sauer , j . r . , iliff , m . patuxent bird identification infocenter . version 97 . 1 . patuxent wildlife research center , laurel , md . urltoken 1998 . national wildlife federation : urltoken project puffin . audubon society . urltoken stirling , k .
fratercula cirrhata
( on - line ) , animal diversity web . urltoken fratercula _ cirrhata . html . 2001 .
nesting : tufted puffins probably form long - term pair bonds to help care for the offspring . they nest in burrows at the edges of cliffs , on grassy slopes , or in natural crevices in rocks . the pair spends a great deal of time preparing the nest site , excavating the burrow with their bills and feet .
each female puffin lays one off - white egg , sometimes with faint blue and brown markings , usually between late april and early june . eggs produced later than june are unlikely to produce fledglings . the peak egg - laying period usually lasts about two weeks in each colony . both parents help with incubation , which last betweeen 40 and 53 days .
puffin recordings by william w . h . gunn ©â cornell lab of ornithology macaulay library cornell lab of ornithology 159 sapsucker woods road ithaca new york 14850 united states of america tel : + 1 ( 607 ) 254 - 2404 fax : + 1 ( 607 ) 254 - 2439 email : macaulaylibrary @ urltoken website : www . birds . urltoken / macaulaylibrary
the tufted puffins are monogamous with long - lived pair - bonds . they perform some displays to strengthen the bond , such as the “billing” courtship ceremony . both mates defend the nest - site by using threat displays . they may fight too , but usually , fighting is short . they utter grunts or growls at the nesting colonies .
tufted puffins are primarily offshore feeders . during nesting , and when food is in abundance , they may feed inshore . the diet of chicks is almost entirely small fish , while the adults ' diets are more diverse . adults prey mostly on anchovies and other small fish , but also eat squid , octopus , crabs , zooplankton , and jellyfish .
this dark - bodied puffin is common along the northern pacific coast , nesting on islands offshore , where it may be seen sitting on rocks in an upright posture . although it flies strongly , it must work hard to take off from the water , thrashing along the surface before becoming airborne . the colorful tufts of feathers on the head are present mostly in summer .
seabirds are often used as biological indicators in ecosystem monitoring programs because they are ubiquitous and relatively well studied , and because some species are tightly linked to their habitats , prey resources , and / or climatic and oceanographic conditions . tufted puffins have declined dramatically in california , oregon and washington , and are on the cusp of being listed as endangered in washington .
is not a threatened species , but in some locations its numbers are decreasing . in alaska , they are highly abundant , but the seabird colonies of alaska are protected by federal and state laws . also , a permit is often required to land on islands where puffins are nesting . in the puffin colonies along the coasts of california , oregon , and washington , population size has been declining since the beginning of the century due to decreasing numbers of fish , ocean pollution , and oil spills . as with most species , puffins have fallen victim to the expansion of humans . to attempt to make up for human takeover of land that was once used for puffin nesting , some programs have been set up to restore of former nesting colonies and help reduce the risk to populations by establishing more nesting sites . ( paul 1994 , small 1994 )
tufted puffins are threatened by sea level rise and storm surges which destroy habitats and breeding areas . in some areas of north america warming seas are causing the fish that the puffins feed on to migrate farther north , making it difficult for them to find adequate food . other threats are entrapment in fishing nets , oil spills , pollution , ingestion of plastic , human disturbance of breeding colonies and introduced predators such as rats and foxes .
in 10 years of monitoring , we ' ve only seen six puffins wash in—total ,
said julia parrish , a university of washington professor who coordinates a west coast volunteer bird - monitoring network .
now we ' ve seen nearly 250 in 20 days . and these islands are small dots in the middle of a huge ocean . the entire puffin population is only 6 , 000 birds , and we project half that many may be affected .
the subsistence hunting of adults and eggs by native americans is traditional and still permitted in most coastal areas of alaska , especially in the bering strait region ; however , the . influence on puffin bird populations is minimal . some fishing methods such as use of gill nets have resulted in puffins being killed as bycatch . pollution ( especially oil spills ) , decreasing availability of food fish , human encroachment , and disturbance of nesting sites are all contributing to a slowly declining population .
tufted puffins spend most of their lives in the open ocean , but during the breeding season mature birds are found on islands and coastal areas in places that have areas suitable for nesting . they prefer such places as the seaward faces of sandy bluffs above beaches , steep grassy slopes , or sometimes in crevices and cracks on cliffs . once chicks fledge and leave the nest , they spend about two juvenile years at sea , coming ashore only after they become adults .
projects to restore natural habitat conditions at current and former nesting islands should also be undertaken or evaluated to conserve tufted puffins in washington . these would include the removal of invasive plants and animals ( see factors affecting continued existence – introduced species ) and the replanting of native vegetation . eradication of european rabbits on destruction island is already under consideration ( s . pearson , pers . comm . ) and control of non - native plants at protection island has been proposed ( usfws 2010 ) .
tufted puffins are active at their nest colonies during the day and can often be seen sitting on rocks in an upright posture . once airborne , they fly well but they need a running start to take flight . they dive and swim underwater , using their wings to paddle and their feet to steer their way through schools of small fish , which they catch in their bills . they can be seen carrying fish crosswise in their bills ( sometimes 12 or more ) , which they take back to their young .
a pigeon - sized puffin , mostly seen sitting upright on a sea cliff , measuring 14 . 5–15 . 5 inches ( 37–39 cm ) . in breeding plumage , stubby body black , face white , down - curved yellowish tufts hang behind eyes , and parrot - like bill enlarged , bright orange - red . in winter , coloured bill plates molt and bill is smaller and duller , face turns dusky , and tufts disappear . immatures are dusky above , light gray below , with small bill . in flight , the large , webbed red feet are conspicuous .
foraging takes place at dawn and dusk . tufted puffins capture and eat prey primarily underwater , in mid - water depths . in the non - breeding season they mostly prey on squid and krill although they will also eat small fishes , octopus , sea jellies , and crabs . it is estimated that 50 - 70 percent of their diet is invertebrates . in the breeding season they forage close inshore on squid , marine worms , krill , and small schooling fish—anchovy , sandlance , capelin , and juvenile pollock . chicks are usually fed a variety of fish species , whatever is available close to the breeding colony . the adult carries 5 - 20 fish to the chick arranged crosswise in its sharp , strong , and heavy beak , a meal called a bill - load .
tufted puffins are medium - sized , compact birds with a large head , short , stiff wings and webbed feet on short legs . plumage is - black above and dark gray to black below . in breeding ( alternate ) plumage adults have a white patch on either side of the face , and long , golden head plumes from just behind their eyes draping down the back . during the breeding season their thick , deep bills are bright red with yellow marking . legs and feet are usually red or orange - red . in non - breeding ( basic ) plumage , the white face patches become gray - brown , the plumes disappear , bill plates are shed , and the bill’s red tip also disappears . juveniles are a brownish - gray and have a shorted bill .
tufted puffins were historically hunted for food . hunting puffins is discouraged nowadays in most places , and forbidden by law in others , but people who do still hunt them try to capture only non - breeding animals . in the past , skins were used to make tough parkas worn feather side in . puffins are also used as tourist attractions for communities near healthy colonies , but visitors must watch the birds from the ocean . because human disturbances may cause puffins to leave their nesting sites , people are often prohibited from landing at nesting sites . puffins are also indicators of a healthy ocean , and show humans when over - fishing is occuring . when there are fewer fish in the ocean , puffins bring a noticeably reduced amount of fish ashore . ( paul 1994 , kessel 1989 )
tufted puffins probably form long - term pair bonds . they nest in burrows at the edges of cliffs , on grassy slopes , or in natural crevices in rocks . the pair spends a great deal of time preparing the nest site , excavating the burrow with their bills and feet . the burrow is 2 - 7 feet long with a nest chamber at the end . this chamber may be lined with grass or feathers , or sometimes nothing at all . digging the nest burrow is a time - consuming job , and the birds most likely do not breed in the season in which they dig the burrow , but wait until the following year . when the pair finally breeds , the female lays one egg , which both parents incubate for 6 - 1 / 2 to 7 - 1 / 2 weeks . both parents care for the young for another 6 - 7 weeks , after which time the fledgling leaves its burrow at night and moves to sea . most young birds are not yet capable of flight at this time , so they walk , or flutter to sea , without parental aid .
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aerc tac . 2003 . aerc tac checklist of bird taxa occurring in western palearctic region , 15th draft . available at : urltoken _ the _ wp15 . xls # .
justification : this species has a very large range , and hence does not approach the thresholds for vulnerable under the range size criterion ( extent of occurrence < 20 , 000 km2 combined with a declining or fluctuating range size , habitat extent / quality , or population size and a small number of locations or severe fragmentation ) . despite the fact that the population trend appears to be decreasing , the decline is not believed to be sufficiently rapid to approach the thresholds for vulnerable under the population trend criterion ( > 30 % decline over ten years or three generations ) . the population size is extremely large , and hence does not approach the thresholds for vulnerable under the population size criterion ( < 10 , 000 mature individuals with a continuing decline estimated to be > 10 % in ten years or three generations , or with a specified population structure ) . for these reasons the species is evaluated as least concern .
the global population is estimated to number > c . 3 , 500 , 000 individuals ( del hoyo
. 1996 ) , while national population sizes have been estimated at < c . 100 breeding pairs and c . 50 - 1 , 000 wintering individuals in japan and c . 10 , 000 - 1 million breeding pairs c . 1 , 000 - 10 , 000 wintering individuals in russia ( brazil 2009 ) .
the population is suspected to be in decline owing to predation by invasive species .
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although puffins spend a majority of the year on the ocean , they build their nests on the shores of islands and coastal regions . they require shores with steep , grassy , sloping land with soil that allows them to burrow . in more rocky areas , puffins build their nests in the rock and on cliff faces . they prefer high places that allow them to swoop down and gain momentum . their stubby wings make it difficult for them to take flight from water or land without help . they prefer secluded areas where some protection is offered by their surroundings . their burrows are typically two to six feet long , and four to six inches in diameter . in highly populated colonies , the burrows of two or three of the animals sometimes run together . ( paul 1994 , gaston 1998 , jewett 1953 )
is similar in size to crows , with an average length of 15 inches , and a 15 inch wingspan . size varies a little from location to location : western pacific animals tend to be a little larger than eastern ones . there is also a difference in size between the sexes as male birds tend to be slightly larger than females .
in the winter , as puffins prepare for spring breeding , their colors become more decorative , presumably to attract mates . during this time they develop a brownish - black body , with some white feathers lining the underside of the wing , a white face and glossy , yellow plumes above and behind eye . the bill is mostly bright red , with yellow and sometimes green markings . when breeding ends in the early summer , puffins lose their plumes , the bright colors of the bill turn to a dull reddish - brown , and the belly is speckled with some pale brown flecks . their legs and feet are red or orange - red throughout the year .
juvenile puffins resemble winter adults , but with a grey - brown breast , white belly , and a shallow , brown bill . ( gaston 1998 , gabrielson 1970 )
during the period prior to egg laying , large groups of puffins congregate off - shore from their nesting colony and engage in intense courtships and frequent copulations . similar behaviors occur on land at the same time , with puffins courting mates through skypointing ( flying straight upwards ) , strutting , and billing ( two birds rubbing their bills together ) .
usually begin breeding in april , although mating activity has been seen as early as march and as late as may in some cases .
once the chicks hatch , their growth rate is variable between colonies and from year to year . the difference is dependent on the feeding conditions of their location . both parents take turns bringing food to the chicks , which happens two to three times daily , and most frequently in the morning and early evening . chicks remain in the burrow and rely on their parents until they are fledged , which usually happens 45 - 55 days after hatching . there is no post - fledging parental care , and the puffling first leaves the nest for the open sea alone and at night . young puffins usually do not return to the colony for almost two years , spending all their time at sea . puffins become sexually mature at the age of three , but most do not mate until they ' re four .
juveniles moult during their first winter at sea , and again the following autumn . adult puffins moult completely following the breeding season , and partially before breeding . ( gaston 1998 , kessel 1989 , paul 1994 )
both parents incubate , protect , and feed their developing young until they are fledged .
puffins are highly social animals ; they live in large colonies , and fish together in flocks of 10 to 25 birds . when faced with an intruder close to their burrow , puffins open their bills and lower their head , and may follow this action with bill grappling or kicking . when conspecifics fight , they spread their wings , open their mouths wide , stomp their feet , and wrestle with their opponents . such displays often draw crowds of birds to watch .
every year puffins over - winter on the ocean . their waterproof feathers and their ability to drink salt water and catch fish make staying long periods on the sea possible . they disperse widely while at sea but return to the colony where they were born , and usually to the same mate , every year for breeding . it is unknown how they do this , but it is speculated that they use sounds , smells , or visual cues from the sky or the ocean to find their way home . ( gaston 1998 , paul 1994 , kessel 1989 )


the western heather vole , phenacomys intermedius , is a small vole found in western north america .

until recently , the eastern heather vole , phenacomys ungava , was considered to be a subspecies .

these animals are similar in appearance to the meadow vole .

they have short ears and a short thin tail which is paler underneath .

their long soft fur is brownish with silver grey underparts .

they are 14 cm long with a 3.5 cm tail and weigh about 40 g. they are found in alpine meadows , open shrubby areas , dry forests with shrubs below to provide cover and tundra regions , usually near water , in british columbia , the yukon and the western united states .

in summer , they live in underground burrows and , in winter , they tunnel under the snow .

they store food for later use year-round .

they feed on plant leaves and berries in summer and plant bark and buds in winter , also seeds and fungi .

predators include owls , hawks and carnivorous mammals .

the female vole has 2 or 3 litters of 2 to 9 young in a nest made from grasses .

they are active year-round , and are crepuscular .

the population of this animal has been reduced in some parts of its range because of clearcutting of forests .

western heather vole

western heather vole ( family cricetidae ) . phenacomys intermedius intermedius - male ; skin ; skull .
there are no known ways that the heather vole positively benefits humans at this time .
( heather vole ) is found on the western mainland of the u . s . as far south as new mexico . in canada the vole is found across northern canada from labrador to the yukon territory . ( banfield 1974 ) ( fitzgerald 1994 )
the only adverse affect known at this time is the heather vole has been known to carry the virus listeriosis ( banfield 1974 ) .
heather voles are difficult to trap , and leave virtually no sign of their presence . the meadow vole ( microtus pennsylvanicus ) will out compete heather voles where their ranges overlap . consequently , grassy habitat , which is preferred by meadow voles , should not be allowed to impinge on heather vole habitat ( u . s . forest service 2000 ) .
the heather vole lives in stands of spruce , lodgepole pine , aspen , and grassy meadows in montane forest , subalpine , and alpine tundra ( fitzgerald 1994 ) . the vole usually stays close to water . the heather vole has been seen traveling 200 yards from its nest . the understory of their habitat may contain shrubs such as blueberry , dwarf birch , and soapberry . the heather vole has been found as high as 12 , 100 feet elevation in colorado . ( banfield 1974 ; fitzgerald 1994 ; armstrong 1972 )
the heather vole usually leads a solitary life except during breeding season . in canada , it was reported that in the winter family groups huddle together in communal nests ( banfield 1994 ) . the female heather vole takes care of the young and has a territory that she defends from other voles .
the range of the eastern heather vole includes much of canada , and extreme northeastern minnesota . minnesota represents the southern edge of this species ' range . though fairly widespread , the heather vole is most likely limited to small , semi - isolated populations . the first documentation of a heather vole in minnesota was in lake county in 1940 , when a single specimen was identified . since then , less than 20 individuals have been found in st . louis , lake , and cook counties . because of the small number of locations that have been found despite targeted surveys , the heather vole has been classified as a species of special concern .
the adult heather vole is 4 1 / 2 to 6 inches long , weighing a little over an ounce . it is gray with a brown to dark - brown cast to the fur , white face , and silvery belly .
etnier , d . a . 1989 . small mammals of the boundary waters canoe area , with a second minnesota record for the heather vole , phenacomys intermedius . the canadian field - naturalist 103 ( 3 ) : 353 - 357 .
kinsella , j . m . 1966 . helminths of microtinae in western montana . m . a . thesis . university of montana , missoula . 61 pp .
the heather vole does not hibernate . in the winter , the vole builds its nest on the surface of the ground under the snow . these nests are better insulated and have thicker walls made of lichen and twigs , and the nest is lined with grasses . winter nests can be found in places of cover , such as under shrubs and logs . the vole in winter and summer months has an area it uses as a toilet . the toilet area is in a tunnel that extends from the nest .
this species is known from the western united states and canada , but with a highly fragmented distribution ( at least seven separate population units ) . one lies completely within british columbia , a second extends from southwest british columbia into central oregon , a third lies completely in western washington , a fourth extends from central british columbia to central oregon and idaho , a fifth lies within central california , a sixth occurs along the nevada - utah border , and a seventh occurs in eastern utah and western colorado and extends slightly into arizona and new mexico . it generally occurs above 750 m asl in the western united states and to above 3 , 000 m asl in the rockies and california .
jannett , f . j . , jr . , and r . j . oehlenschlager . 1997 . range extension and unusual occurrences of the heather vole , phenacomys intermedius , in minnesota . the canadian field - naturalist 111 ( 3 ) : 459 - 461 .
the heather vole fur is long and soft . the color varies geographically . generally , the dorsal colors are brown to grayish , while the ventral fur is gray . the feet are white to gray and the ears can have orange hairs . the tail is slightly longer than the hindfeet ( 31 - 34 mm ) and has dark fur on top and is pale on the ventral side . the total length of the animal is between 130 and 140 mm . the species closely resembles the meadow vole ; skull characteristics are often needed to help tell the species apart . the skull characteristic used is that the cheek teeth have deep lingual angles in the heather vole . ( banfield 1974 ; fitzgerald 1994 )
the heather vole searches for green vegetation , bark of trees and shrubs , twigs , seeds , berries , and fungi . their diet consists of leaves and fruits of willows , myrtle blueberry , snowberry , bog birch , kinnikinnnik ( bearberry ) in the summer months . in the winter , spring , and fall they tend to focus more on the bark and buds of willow , birch , and blueberry . the heather vole stores food for use in the summer and winter . the food reserves are found close to the burrow entrances . ( banfield 1974 ; fitzgerald 1994 ) .
george , s . b . 1999 . eastern heather vole ( phenacomys ungava ) . pages 618 - 619 in d . e . wilson and s . ruff , editors . the smithsonian book of north american mammals . smithsonian institution press in association with the american society of mammalogists , washington .
geographic map of western heather vole ( phenacomys intermedius ) sampling localities ( a–r , appendix i ) in the pacific northwest . also identified is the geographic extent of each of the 3 major clades as defined in fig . 2 . the inset shows the entire contemporary distribution of p . intermedius in dark outline ( redrawn from locality data in hall [ 1981 ] , nagorsen [ 2005 ] , and verts and carraway [ 1998 ] ) , approximate margins of the cordilleran and laurentide ice sheets ( ∼14 , 000 years before present ) as the shaded area north of the dotted line ( siegert 2001 ) , and major alpine glaciers as shaded areas south of the ice sheets ( porter et al . 1983 ) . late - pleistocene and holocene fossil localities outside the modern heather vole range are indicated by black circles .
the minnesota biological survey targeted this species in 2003 . despite intense trapping activities , they were only able to locate two individuals . more research is needed to determine the extent of heather vole distribution in minnesota and the ecological requirements necessary to ensure a viable population ( u . s . forest service 2000 ) .
the heather vole is a small , short tailed rodent with a total length of 135 - 155 mm ( 5 . 3 - 6 . 1 in . ) , and a weight of 30 - 45 g ( 1 . 1 - 1 . 6 oz . ) . the species resembles other voles ( microtus spp . ) and bog lemmings ( synamtomys spp . ) . it has fur that is brown on the back , usually more yellowish or tan than other voles or lemmings , and white or gray underneath . heather vole ' s feet are white or pale gray ( mcallister and hoffman 1988 ; george 1999 ) . their tail is bicolor and shorter than that of other vole species , but longer than that of bog lemmings . heather voles are best distinguished from other species by their teeth . unlike other species , the inner reentrant angles on the lower molars are much deeper than the outside angles ( george 1999 ) . additionally , the upper incisors lack deep grooves that are present in bog lemmings .
this species has a wide distribution in western north america , there are at least hundreds of known locations where it occurs . density estimates range from 0 . 5 to 10 per ha in different habitats in different areas . irregular population fluctuations are typical .
we have shown that the combination of major climatic fluctuations and topographical variation found within a regional montane system has had an important impact on the genetic variation of an alpine mammal , the western heather vole , in the pacific northwest . p . intermedius supports some of our predictions for high - elevation species , such as postglacial decline in population size and differentiation of populations restricted to isolated mountain ranges . however , in some ways p . intermedius showed similar patterns to those of low - elevation species ; for example , the separation of refugial populations into distinct geographical regions and the asynchronous northward expansion by divergent lineages into areas previously glaciated by continental ice sheets . studying the extreme environments and restricted ranges of alpine organisms is useful for understanding the tolerances of temperate mammals to strong environmental heterogeneity .
throughout its range , the heather vole may inhabit coniferous forests with a heath understory , wet meadows , rocky hillsides , forest edge , or deciduous shrubby habitats ( george 1999 ) . proximity to water , boulders , coarse woody debris , and plants of the ericaceous family are important to its survival ( u . s . forest service 2000 ) . in minnesota , one heather vole was trapped in a poorly drained black spruce and balsam fir woodland with groundcover of mosses , ferns , horsetail ( equisetum spp . ) and ground pine ( lycopodium spp . ) ( etnier 1989 ) . other minnesota specimens have been documented in the following habitats : 1 ) a boulder field with a black spruce ( picea mariana ) and balsam fir ( abies balsame a ) overstory and a groundcover of lichens , ferns , and mosses ; 2 ) a clearcut with boulders covered in lichens , ferns , and mosses ( jannett and oehlenschlager 1997 ) ; and 3 ) a large , semi - open black spruce sphagnum bog .
western heather voles live in mountains , near or above the timberline . where high - elevation forests have been clear - cut , they have been seen at lower elevations . like eastern heather voles , they feed on a variety of vegetation , including leaves , seeds , berries , and the bark of willow and other shrubs . they build summer nests underground and winter nests at ground level , next to a bush , rock , or stump . the nests are made of twigs , leaves , and grass . as many as three litters of 2 - 9 young are produced a year . males are aggressive toward each other during the mating season , but several voles may nest together for warmth in winter . these voles do not do well in captivity ( if they survive trapping ) , which makes it hard to study them .
postglacial routes for range expansion of western heather voles into northern areas previously occupied by continental ice sheets appear to have occurred along an interior axis rather than a coastal or cascade mountain axis . examination of our data shows minimal northward expansion of washington clade populations in comparison to northern and interior clade populations . in fact , only populations belonging to the northern and interior clade reoccupied areas previously overlaid by continental ice sheets , including the north cascades . this spatial distribution suggests an interior dispersal route along or near the rocky mountain axis . the earlier retreat of the laurentide ice sheet , as opposed to the cordilleran ice sheet , during late - pleistocene deglaciation ( siegert 2001 ) could have played a role in allowing northward expansion by interior refugial populations and the prevention of major northward expansion by the more - coastal washington refugiai populations . this pattern of asynchronous northward expansion by divergent lineages has been found in several low - elevation boreal mammals ( arbogast 1999 ; arbogast et al . 2001 ; demboski et al . 1999 ) and in another high - elevation arvicoline with similar habitat associations , the water vole ( microtus richardsoni — carstens et al . 2005 ) . the american pika ( ochotona princeps ) , however , displays a more symmetrical northward expansion between cascade and rocky mountain lineages despite occupying similar high - elevation mountains in western north america ( galbreath et al . 2009 ) . this demonstrates that climate - mediated movements did not necessarily occur in all assemblages of mammals but were likely more individualistic and dependent on each species’ ecological breadth ( grayson 2006 ; lawlor 1998 ) .
thompson , richard w . , western resource dev . corp . , boulder , co . , 1996 , wildlife baseline report for the montana [ montanore ] project , lincoln and sanders counties , montana . in application for a hard rock operating permit and proposed plan of operation , montanore project , lincoln and sanders counties , montana . vol . 5 . stroiazzo , john . noranda minerals corp . , libby , mt . revised september 1996 .
the interaction between climate - mediated shifts in geographic distribution during the pleistocene and variation in topography associated with regional occurrence of mountains has played a strong role in the phylogeographic structure of western heather voles in the pacific northwest . the initial southward range expansion of phenacomys along the pacific coastal region into the conterminous united states ( ∼1 . 8 mya ) from its arctic origins in alaska and northeastern siberia occurred during a warm interglacial period ( repenning 1990 ) . repeated intervals of glaciation through the remainder of the pleistocene could have resulted in range expansions and contractions and subsequent secondary contact between diverging lineages in lowland refugia or in recolonized high - elevation regions . however , the geographic distinctiveness of the 3 contemporary lineages suggests that barriers were persistent throughout these repeated cycles of climate change and did not allow secondary contact to occur .
bayesian gene tree based on 325 base pairs of the mitochondrial control region of haplotypes of phenacomys intermedius sampled from the northern and western portions of the species’ range . the phylogeny inferred with maximum - likelihood methods yielded an essentially identical topology . numbers above nodes represent maximum - likelihood bootstrap percentages > 80 and those below represent bayesian posterior probabilities > 0 . 85 . sampling localities ( a–r ) and haplotype numbers ( 1–58 ) , as shown in appendix i , are indicated at branch tips .
heather voles are active year round . the voles have different nesting areas depending on the time of the year . in the summer months , they live in burrow systems . these burrows are usually around 20 cm deep . the entrance is usually hidden by some sort of vegetation or leaf litter . the nest is made of grasses , lichen , and other types of vegetation .
heather voles have several estrus periods per year . most of the breeding occurs between may and september . however , the voles located at higher elevations have a shorter season than the voles at lower elevations . heather voles have a gestation period of 19 to 24 days . the litter size varies between 2 and 9 , but variation is large because of the differences in litter sizes between adult and juvenile females . adult females produce larger litters ( average 5 . 9 ) , while the juvenile females produce smaller litters ( average 3 . 8 ) their first season . three litters per year is believed to be the maximum produced . the voles are born blind and deaf . they finally wander out of the nest at about 3 weeks of age . males don ' t reach sexual maturity until the next spring , while females reach sexual maturity 4 - 6 weeks after birth . ( banfield 1974 ; fitzgerald 1994 )
sampling . —our analysis is based on 89 samples of p . intermedius from 18 localities representing the northern and western portions of the geographic range , with a special emphasis on isolated montane populations of the pacific northwest ( appendix i ; fig . 1 ) . the samples consisted of 36 frozen tissues from the liver of specimens collected between 1999 and 2008 , which were deposited in the burke museum , university of washington ( uwbm ) , and the museum of vertebrate zoology , university of california ( mvz ) . the remaining 53 samples were obtained from skins of museum study specimens collected between 1925 and 2000 , which were deposited in the uwbm ; the slater museum of natural history , university of puget sound ( psm ) ; and the department of fisheries and wildlife , oregon state university ( osufw ; appendix i ) . research was conducted in adherence with guidelines of the american society of mammalogists ( gannon et al . 2007 ) .
results from our bayesian skyline plots reveal population declines in all 3 major lineages of heather voles since the beginning of the holocene . this pattern is also shared by pikas ( galbreath et al . 2009 ) and likely reflects the reduced amount of habitat in restricted mountain environments following postglacial upslope movements from more expansive lowland refugia . this pattern contrasts with those of low - elevation temperate mammals , which commonly show signatures of postglacial demographic expansion resulting from geographic expansion into large areas of recently available habitat ( lessa et al . 2003 ; runck and cook 2005 ) . this pattern also might help explain why some of the more southerly interior populations , such as those that existed in the great basin desert , are no longer extant . nevertheless , some of our demographic analyses indicate that the northern and interior clade experienced rapid population growth . although these results conflict with the outcome of the bayesian skyline plots , they are consistent with the observation that this clade is the only major lineage to experience major geographic expansion since the end of the pleistocene and the only case of substantial long - distance haplotype sharing ( northern british columbia and montana ; fig . 2 ) .
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there is uncertainty about the taxonomic status of phenacomys ungava . in recent decades , most authors have regarded ungava as a subspecies of p . intermedius . musser and carleton ( in wilson and reeder 1993 , 2005 ) noted the present validity of earlier statements that the relationship between intermedius and ungava needs further detailed study ; nevertheless , they listed p . ungava as a separate species . jones et al . ( 1997 ) , baker et al . ( 2003 ) , and george ( in wilson and ruff 1999 ) also recognized ungava as a distinct species .
justification : listed as least concern because it is very widespread , its populations are stable , there are no major threats , and it occurs in protected areas throughout its range .
it occurs from sea level to above the treeline ; open coniferous forest with heath , shrub understorey ; shrub areas on forest edge ; mossy meadows in forests ; alpine tundra with cover . it nests on the ground under snow ( winter ) or in burrows ( summer ) . it is solitary in summer except during the breeding season . family groups may occupy communal nests in winter . gestation lasts 19 - 24 days . young are born mid - june to early september , though the season possibly is more restricted at high elevations . litter size averages 3 - 4 for young - of - year , 4 - 6 for older females ( which may produce two litters per year ) ( mcallister and hoffman 1988 ) . in winter , these voles feed on bark and buds of shrubs and heaths . in summer , they feed primarily on green vegetation , berries , and seeds . stores food winter and summer . they are active throughout the year .
there are many protected occurrences of this species and no known large - scale threats .
it is protected in wilderness areas and national parks and provincial parks of british columbia , alberta , washington , oregon , montana , utah , and elsewhere .
this errata assessment has been created because the map was accidentally left out of the version published previously .
( errata version published in 2017 ) . the iucn red list of threatened species 2016 : e . t42636a115197827 .
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during the breeding season males fight among each other . the voles are active at dusk . their predators include martens , weasels , owls , and hawks . ( banfield 1974 ; fitzgerald 1994 )
the population size of this species is not well known . scientists have found the species difficult to trap and the mortality rate during livetrapping is high . ( fitzgerald 1994 )
brian putz ( author ) , university of michigan - ann arbor , phil myers ( editor ) , museum of zoology , university of michigan - ann arbor .
living in the nearctic biogeographic province , the northern part of the new world . this includes greenland , the canadian arctic islands , and all of the north american as far south as the highlands of central mexico .
having body symmetry such that the animal can be divided in one plane into two mirror - image halves . animals with bilateral symmetry have dorsal and ventral sides , as well as anterior and posterior ends . synapomorphy of the bilateria .
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature . endothermy is a synapomorphy of the mammalia , although it may have arisen in a ( now extinct ) synapsid ancestor ; the fossil record does not distinguish these possibilities . convergent in birds .
forest biomes are dominated by trees , otherwise forest biomes can vary widely in amount of precipitation and seasonality .
this terrestrial biome includes summits of high mountains , either without vegetation or covered by low , tundra - like vegetation .
the area in which the animal is naturally found , the region in which it is endemic .
coniferous or boreal forest , located in a band across northern north america , europe , and asia . this terrestrial biome also occurs at high elevations . long , cold winters and short , wet summers . few species of trees are present ; these are primarily conifers that grow in dense stands with little undergrowth . some deciduous trees also may be present .
a terrestrial biome with low , shrubby or mat - like vegetation found at extremely high latitudes or elevations , near the limit of plant growth . soils usually subject to permafrost . plant diversity is typically low and the growing season is short .
u . s . fish and wildlife service homepage
( on - line ) . accessed december 12 , 1999 at urltoken .
to cite this page : putz , b . 2000 .
phenacomys intermedius
( on - line ) , animal diversity web . accessed july 07 , 2018 at urltoken
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this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
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the interaction of historical climate changes with topographical variation can produce both intermittent lowland biogeographic refugia and high - altitude isolation of montane biota . the pacific northwest region of north america possesses a heterogeneous landscape that transects major mountain ranges and intervening lowland valleys and plateaus . throughout the glacial cycles of the pleistocene , between 2 . 58 million years ago ( mya— gibbard et al . 2010 ) and 11 , 700 years ago ( walker et al . 2009 ) , the pacific northwest was exposed to a succession of about 20 glacial periods characterized by expansion and retraction of continental ice sheets and alpine glaciers . however , the timing of peak expansion of alpine glaciers varied across major mountain ranges ( porter et al . 1983 ; thackray 2008 ) resulting in dynamic changes in the location of lowland ice - free zones that served as biotic refugia . furthermore , periods of climate warming forced the redistribution of lowland biota into alpine zones that were isolated from one another by the topography of the pacific northwest .
our understanding of the responses of north american plants and animals to quaternary climate history has been enhanced by studies of genetic variation of populations across geographic space ( avise 2009 ) . the genetic signatures of populations that resulted from historical vicariance and dispersal events developed through the interplay of 2 important evolutionary forces : genetic drift and mutation . when populations become isolated and small , genetic drift acts to reduce genetic diversity and thereby to promote differentiation among populations . by comparing both the genetic diversity of contemporary populations and the genealogical relationships among populations over geographic space , investigations of phylogeography can provide clues as to where and when isolation events and dispersal routes occurred .
climate - mediated impacts on the structuring of genetic variation of mammal populations in the pacific northwest have been investigated in several low - elevation species ( arbogast and kenagy 2001 ; miller et al . 2006 ; yang and kenagy 2009 ; zheng et al . 2003 ) . one common pattern that has emerged from these studies is the apparent diversification of lineages during full glacial periods due to climate - driven shifts of populations into disjunct refugia . another pattern is the major latitudinal shift in distribution and accompanying population expansion that follows the retraction of continental ice sheets ( lessa et al . 2003 ) . however , mammals currently restricted to high - altitude environments experienced major elevation shifts due to these climate changes , which has led to different predictions on the outcomes of their genetic structuring and demographic changes . the expansion of alpine glaciers during cold climatic periods will force downslope shifts in populations from disjunct mountain ranges and possibly allow them to merge in common lowland refugia with greater habitat availability , which will lead to increases in population size . moreover , upslope shift of populations into restricted mountaintops during warm periods will lead to smaller population sizes and facilitate significant lineage divergences due to founder effects ( wright 1942 ) . recent examples of the response of alpine specialists to these environmental scenarios demonstrate patterns that are consistent with these predictions ( carstens et al . 2005 ; galbreath et al . 2009 ; knowles 2001 ; spaeth et al . 2009 ) .
to assess the impacts of late - quaternary climate change and geological features on the historical biogeography of p . intermedius across montane systems of the pacific northwest , we obtained samples throughout several montane regions from northern british columbia , canada , southward to california . our objective was to incorporate phylogenetic inference analyses , relaxed molecular - clock dating computations , and historical demographic estimations into a phylogeographic framework to test the hypotheses that contemporary isolated populations were forced into a series of disjunct refugia separated by topographical barriers during full glacial conditions in the pleistocene rather than into a common refugium ; dispersal into northerly areas that were previously occupied by continental ice sheets occurred from multiple refugial sources ; postglacial shifts of elevation into isolated mountain ranges resulted in lineage divergence ; and major lineages are experiencing declines in population since the end of the pleistocene due to their upslope range shift and reoccupation of restricted montane regions .
laboratory methods . —we extracted whole genomic dna from fresh liver or spleen tissue using the prescribed protocol of dneasy tissue kit ( qiagen , valencia , california ) . for samples from museum study skins genomic dna was extracted following a protocol developed by mullen and hoekstra ( 2008 ) , which included an ethanol wash every 3 h for 24 h to remove salts and polymerase chain reaction ( pcr ) inhibitors that may have been used in preserving museum skins . for these samples we also undertook several steps to avoid contamination . first , we used a new sterilized razor blade for the removal of snippets from each specimen . second , we performed ethanol washes in a separate room from where pcr amplifications were performed to avoid contamination from pcr amplicons . next , we divided our samples into smaller subsets and staggered the dates of extractions and pcr amplifications for each subset . we also checked for contamination by checking for pcr products from negative - control extractions and negative - control pcr master mixes with gel electrophoresis . finally , we repeated dna extractions for 5 random individuals using skin snippets from another part of the study specimen and compared their sequences with original samples .
we amplified a 325 - base pair ( bp ) sequence of the mitochondrial control region using the pcr with primers rtvl16000 ( 5′ - gtcaacacccaaagctgaca - 3′ ) and rtvlntlr ( 5′ - gttggtttcacggaggatgg - 3′— miller et al . 2006 ) . pcr mixtures totaled 25 μl and followed slightly different protocols for frozen tissue and museum study skin samples . for the frozen tissue samples each reaction mixture contained 14 . 9 ul of nuclease - free h 2 o , 2 . 5 μl of 10x pcr buffer , 1 μl of 25 mm mgcl 2 , 0 . 2 mm of each deoxynucleoside triphosphate , 2 . 5 μl of each 1 x primer , 0 . 125 μl of jumpstart taq dna polymerase ( sigma , st . louis , missouri ) , and 1 μl of genomic dna . we performed pcr amplifications in a geneamp pcr system 9700 thermocycler ( applied biosystems , foster city , california ) . the cycle conditions included a denaturing step at 94°c ( 10 min ) , followed by 30 cycles ( 1 min at 94°c , 30 s at 50°c , and 1 min at 72°c ) with a final extension period of 5 min at 72°c . because dna from study skins was presumed to be somewhat degraded , we modified the pcr master mix to improve pcr amplification by adding 2 . 5 μl of 10x bovine serum albumin , 2 μl of 25 mm mgcl 2 , 0 . 5 μl of jumpstart taq dna polymerase , and 9 . 9 μl of genomic dna , and by not adding the 14 . 9 μl of nuclease - free h 2 o . we also increased the number of amplification cycles to 40 during pcr . we treated all pcr products with exosapit ( usb corp . , cleveland , ohio ) to remove unincorporated nucleotides and primers . sequencing reactions for pcr products of each sample were performed separately for each forward and reverse primer . sequencing reaction mixtures of 10 μl for each sample included 1 μl of pcr product , 7 . 5 μl of nuclease - free h 2 o , 0 . 5 ( μl of the big dye terminator cycle sequencing ready reaction mix ( applied biosystems ) , 0 . 5 μl of the big dye 5x sequencing buffer , and 0 . 5 μl of primer . samples were run on an abi 3100 genetic analyzer ( applied biosystems ) for sequencing of nucleotides . sequences were aligned using sequencher 4 . 6 ( gene codes corp . , ann arbor , michigan ) and deposited in genbank under accession numbers gq903588 – gq903676 .
molecular diversity and demographic analyses . —we computed haplotype ( h ) and mean nucleotide ( π ) diversities ( nei 1987 ) for each clade using arlequin ( excoffier et al . 2005 ) for comparisons of genetic diversity . we also examined the demographic history of each major clade using fu ' s f s test for neutrality ( fu 1997 ) . large and negative f s values suggest an excess of rare alleles , which can be an indication of population growth or selection . we also investigated patterns of historical demographic expansion by examining the distribution of pairwise differences ( mismatch distributions ) for each clade ( rogers and harpending 1992 ) using arlequin . in general , populations undergoing recent and sudden expansion exhibit a poisson - shaped mismatch distribution , and populations in equilibrium tend to have ragged distributions ( slatkin and hudson 1991 ) . we assessed the statistical significance of these distributions with sum of squared distances and harpending ' s raggedness index ( harpending 1994 ) .
we also explored demographic history of each major lineage using bayesian skyline plots ( drummond et al . 2005 ) in beast . this coalescent - based inference method uses a markov chain monte carlo sampling procedure with gene sequence data to estimate a posterior distribution of effective population size through time . we used divergence date estimates of time to the most recent common ancestor from our divergence analysis as root age priors for each of the major lineages so that we could model changes in effective population size against a timescale of years rather than expected substitutions per site . we also used the hasegawa - kishino - yano substitution model with a gamma distribution and proportion of invariant sites , along with a relaxed clock model with an uncorrelated lognormal prior . we forced coalescent intervals into 5 groups for each lineage because test runs for higher group numbers did not converge as well . we used default settings for the remaining model parameter priors . we performed 5 runs with different random seeds , sampled every 10 , 000 generations , and discarded 10 % of the samples as burn - in for each major lineage . we varied the length of runs for each lineage with the northern and interior clade set at 100 million generations , the washington clade set at 80 million generations , and the oregon and california clade set at 40 million generations . effective sample size values for all parameters in each run exceeded 200 . we combined results for each lineage in logcombiner and produced skyline plots in tracer .
phylogenetic inferences and estimates of divergence dates . — both the maximum - likelihood and bayesian phylogenetic inferences reveal 3 major clades ( fig . 2 ) , each associated with distinct geographic areas : a northern and interior clade represented by populations in the northern rockies , wallowa mountains , cascades of northern washington and southern british columbia , and the northern interior mountains of british columbia ( fig . 1 , localities a–j ) ; a washington clade represented by populations in the southern washington cascades and the olympic mountains ( k–n ) ; and an oregon and california clade represented by populations in the oregon cascades and the sierra nevada of california ( o–r ) .
within each major clade we found 1 well - supported subclade associated with a geographically isolated mountain range , as follows . within the northern and interior clade the isolated wallowa mountains of eastern oregon ( j ) contain a lineage with high bayesian nodal support ( fig . 2 ) . within the washington clade the lineage in the isolated olympic mountains ( n ) shows strong nodal support . within the oregon and california clade the disjunct population in the sierra nevada ( r ) shows strong nodal support .
the estimation of time to the most recent common ancestor ( fig . 3 ; table 1 ) indicates that the tribe phenacomyini diverged from its sister lineage in the subfamily arvicolinae ∼5 . 37 mya . the estimated mean time to the most recent common ancestor for the split of the genera phenacomys and arborimus was ∼2 . 45 mya . this analysis estimated an early - pleistocene origin of ∼1 . 8 mya for the oregon and california clade and an early - pleistocene divergence of ∼1 . 54 mya between the washington clade and northern and interior clade . sublineage diversification of isolated mountain populations ( wallowa , olympic , and sierra nevada ) occurred in the middle pleistocene within each of the 3 major clades .
bayesian tree with divergence dates from the relaxed molecular - clock analysis of haplotypes of phenacomys intermedias . circled numbers refer to estimated divergence dates , and gray horizontal bars indicate 95 % credibility intervals ( table 1 ) . specimen numbers of phenacomys indicated by letters a–j are provided in appendix i , and outgroup sequence numbers and fossil calibration dates , indicated by f1 and f2 , are provided in the “materials and methods . ” all nodes possessed posterior probability support > 0 . 95 except for those indicated by gray stars .
estimates of divergence dates for key splits in phenacomys as shown in fig . 3 .
mean dates estimated in millions of years before present ( mya ) with 95 % higher posterior density in parentheses .
molecular diversity and demographic analyses . —all 3 clades had high haplotype diversity and low nucleotide diversity ( table 2 ) , and no haplotype was shared among the 3 regions occupied by the major clades . only 4 haplotypes were found at > 1 sampling locality . the significantly negative fu ' s f s for the northern and interior clade suggests that this clade experienced recent rapid expansion . this is corroborated by the mismatch distributions , which were unimodal for this clade and did not possess a statistically significant harpending ' s raggedness index or sum of squared deviation . the multimodal mismatch distributions and nonsignificant fu ' s f s values for the other 2 major clades suggest population stability ; however , this conflicts with the nonsignificant harpending ' s raggedness index and sum of squared deviation of the mismatch distribution for both clades . this lack of correspondence could be due to the more conservative test based on pairwise sequence differences ( mismatch distribution ) as opposed to the fu ' s f s ( ramos - onsins and rozas 2002 ) .
haplotype distribution , molecular diversity , and demographic parameters for 3 major clades of phenacomys intermedius .
the median estimates from the bayesian skyline plots indicate that the populations of all 3 major lineages have rapidly declined following a long period of relative stability . this decline occurred after the last glacial maximum and at the beginning of the warming period in the early holocene ( fig . 4 ) .
demographic history of the 3 major clades of phenacomys intermedius . a ) mismatch distributions are represented by solid lines for observed distributions of pairwise nucleotide differences between sequences and dashed lines for expected distribution under a sudden expansion model ( rogers and harpending 1992 ) . b ) bayesian skyline plots , in which central line indicates the median value for the log effective population size ( effective population size times number of generations ) , and the upper and lower lines represent 95 % range of posterior density . dashed vertical line represents the pleistocene - holocene boundary ( 11 , 700 years before present ) , following walker et al . ( 2009 ) .
within the smaller geographic scale of each major clade we also found significant substructuring of populations of p . intermedius associated with shifts of elevation into isolated montane systems . the results from our divergence dating analyses indicate that the initial divergence of these sub - lineages began during the middle pleistocene . as with the case for the 3 major clades , these sublineages persisted in distinct refugia despite repeated range expansions and contractions during glacial cycles associated with the remainder of the pleistocene . barriers that prevented mixing of lineages might have resulted from various prominent geographic features . the divergence between the olympic mountain and washington cascade populations possibly was reinforced by repeated advances of the puget lobe of the cordilleran ice sheet during full glacial periods ( easterbrook 1986 ) . the divergence between populations in the wallowa mountains and the rest of the populations in the northern and interior clade likely was due to the presence of the snake river , whereas the divergence between populations in the oregon cascades and the sierra nevada could have been reinforced mostly by postglacial changes of vegetation in intervening regions from cool subalpine parkland to present - day mesic forests ( briles et al . 2005 ; daniels et al . 2005 ; hakala and adam 2004 ; west 2004 ) . furthermore , lineage divergence also appears to be dependent on sufficient time for genetic drift to sort shared ancestral variation . none of the populations that recolonized northerly areas , which were covered by continental ice sheets , were shown to be divergent .
this project was funded by the burke museum mammal program and a genome training grant fellowship from the department of genome sciences at the university of washington . we thank the museum of vertebrate zoology at the university of california , berkeley ; the slater museum of natural history at the university of puget sound ; and the department of fisheries and wildlife at oregon state university for providing materials for this research . we thank j . bradley , c . welch , c . himes , d . yang , j . whorley , s . vignieri , and x . zheng for collecting specimens used in this study . we also thank l . hauser for comments on the manuscript . lastly , we thank b . lim and 2 anonymous reviewers for their comments and suggestions on previous versions .
formal ratification of the quaternary system / period and the pleistocene series / epoch with a base at 2 . 58 ma
north america and adjacent oceans during the last deglaciation . vol . k - 3 , the geology of north america
late quaternary environments of the united states . vol . i , the late pleistocene
list of 89 specimens of phenacomys intermedias from 18 localities used in our analyses . museum acronyms are defined in the “materials and methods . ” mtdna = mitochondrial dna . bc = british columbia ; ab = alberta ; wa = washington ; mt = montana ; or = oregon ; ca = california .
letters a–j indicate individuals used for divergence dating as shown in fig . 3 .
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merriam , c . h . , 1889 . description of a new genus phenacomys and four new species of arvicolinae , p . 32 . north american fauna , 2 : 27 - 45 .
mammal species of the world ( opens in a new window ) . mammalian species , american society of mammalogists ' species account ( opens in a new window ) .
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unlike females , males only breed in their second year . estimates based on the amount of tooth wear suggested that these animals live up to 4 years in the wild [ 0627 ] . without more detailed studies maximum longevity is classified as unknown .
[ 0036 ] savage et al . ( 2004 ) , the predominance of quarter - power scaling in biology
[ 0455 ] virginia hayssen et al . ( 1993 ) , asdell ' s patterns of mammalian reproduction : a compendium of species - specific data
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u . s . forest service . 2000 . population viability assessment in forest plan revision . questions for animal population viability assessment panel : phenacomys ungava . united states forest service , region 9 , duluth , minnesota .
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most common in subalpine spruce / fir forest with evergreen shrub ground cover , also in timberline krummholz , alpine tundra . sometimes in montane yellowpine / douglas - fir forests with bearberry / twinflower understory ( hoffmann and pattie 1968 ) .
using personal observations and reviewing literature that summarize the breeding , overwintering , or migratory habitat requirements of each species ( dobkin 1992 , hart et al . 1998 , hutto and young 1999 , maxell 2000 , foresman 2012 , adams 2003 , and werner et al . 2004 ) ;
calculating the percentage of observations associated with each ecological system relative to the percent of montana covered by each ecological system to get a measure of
observations versus availability of habitat
.
species that breed in montana were only evaluated for breeding habitat use , species that only overwinter in montana were only evaluated for overwintering habitat use , and species that only migrate through montana were only evaluated for migratory habitat use . in general , species were listed as associated with an ecological system if structural characteristics of used habitat documented in the literature were present in the ecological system or large numbers of point observations were associated with the ecological system . however , species were not listed as associated with an ecological system if there was no support in the literature for use of structural characteristics in an ecological system ,
point observations were associated with that system . common versus occasional association with an ecological system was assigned based on the degree to which the structural characteristics of an ecological system matched the preferred structural habitat characteristics for each species as represented in scientific literature . the percentage of observations associated with each ecological system relative to the percent of montana covered by each ecological system was also used to guide assignment of common versus occasional association . if you have any questions or comments on species associations with ecological systems , please contact the montana natural heritage program ' s senior zoologist .
species associations with ecological systems should be used to generate potential lists of species that may occupy broader landscapes for the purposes of landscape - level planning . these potential lists of species should not be used in place of documented occurrences of species ( this information can be requested at :
) or systematic surveys for species and evaluations of habitat at a local site level by trained biologists . users of this information should be aware that the land cover data used to generate species associations is based on imagery from the late 1990s and early 2000s and was only intended to be used at broader landscape scales . land cover mapping accuracy is particularly problematic when the systems occur as small patches or where the land cover types have been altered over the past decade . thus , particular caution should be used when using the associations in assessments of smaller areas ( e . g . , evaluations of public land survey sections ) . finally , although a species may be associated with a particular ecological system within its known geographic range , portions of that ecological system may occur outside of the species ' known geographic range .
adams , r . a . 2003 . bats of the rocky mountain west ; natural history , ecology , and conservation . boulder , co : university press of colorado . 289 p .
dobkin , d . s . 1992 . neotropical migrant land birds in the northern rockies and great plains . usda forest service , northern region . publication no . r1 - 93 - 34 . missoula , mt .
foresman , k . r . 2012 . mammals of montana . second edition . mountain press publishing , missoula , montana . 429 pp .
hart , m . m . , w . a . williams , p . c . thornton , k . p . mclaughlin , c . m . tobalske , b . a . maxell , d . p . hendricks , c . r . peterson , and r . l . redmond . 1998 . montana atlas of terrestrial vertebrates . montana cooperative wildlife research unit , university of montana , missoula , mt . 1302 p .