motobdella montezuma is a species of leech which is only found in montezuma well , central arizona , united states .

it is a nocturnal pelagic predator that feeds almost exclusively on the endemic amphipod hyalella montezuma , which it detects using passive sonar and swallows whole .

motobdella montezuma

phylogenetic relationships of three genera of erpobdellidae ( hirudinoidea ) , with a description of a new genus , motobdella , and species , motobdella sedonensis .
hyalella montezuma , a new species ( crustacea : amphipoda ) from montezuma well , arizona .
govedich , f . r . , d . blinn , p . keim , and r . w . davies . 1998 . phylogenetic relationships of three genera of erpobdellidae ( hirudinoidea ) , with a description of a new genus , motobdella , and species , motobdella sedonensis . canadian journal of zoology 76 : 2164 - 2171 .
leech , glossiphoniidae , helobdella blinni sp . n . , new species , montezuma well
, the well and montezuma castle preserve extraordinary examples of prehistoric sinagua cliff dwelling sites .
- and watkins , r . c . 1977 . hyalella montezuma a new species ( crustecea : amphipoda ) from montezuma well , arizona . - hydrobiologia 52 : 175 - 184 .
montezuma well ( mowe ) became a detached unit of montezuma castle national monument in 1943 to protect the
objects of scientific interest
located there . nps interpretation of montezuma well focuses on three major scientific themes : native american prehistory , water chemistry , and unique biology .
erpobdella montezuma ( hirudinoidea : erpobdellidae ) , a new species of freshwater leech from north america .
wagner vt , blinn dw . ( 1987 ) montezuma well : the living desert oasis . urltoken
yet other aquatic organisms flourish in their place . many of these other aquatic organisms are only found there , including the endemic leech motobdella montezuma . former northern arizona university ( nau ) professor dean blinn identified and documented the life - history of many of these endemic organisms in the 1980s and 1990s including 12 undescribed species of :
proclaimed the montezuma castle national monument on december 8 , 1906 , including 840 acres on and around montezuma well . after passage of the national park service organic act in 1916 , the monument was transferred to the newly formed national park service .
montezuma well is a limestone sinkhole in the verde valley . arsenic seeps naturally from the rocks , which makes the water toxic to most living things . a research team at northern arizona university discovered the montezuma well leech has a unique way of dealing with arsenic .
location of helobdella blinni sp . n . a the northeast side of montezuma well ; and b the swallet where the leeches were collected .
travertine : a mineral deposit of calcium carbonate that forms through precipitation in a hot spring , like montezuma well , or from the calcium - rich water on cave dripstones .
hevly , r . h . 1974 . recent paleoenvironments and geological history of montezuma well . - j . ariz . acad . sci . 9 : 66 - 75 .
blinn , d . w . , dehdashti , b . , runck , c . , and davies , r . w . ( 1990 ) . the importance of prey size and density in an endemic predator - prey couple ( leech erpobdella montezuma - amphipod hyalella montezuma ) . j . anim . ecol . , 59 : 187 - 192 .
- and barry , w . t . 1973 . montezuma well , arizona , as a habitat . - j . ariz . acad . sci . 8 : 7 - 13 .
a leech occupies the top spot in the food chain at montezuma well near camp verde . surprisingly , its body contains the highest level of arsenic ever recorded in a living organism .
govedich , f . r . , blinn , d . w . , keim , p . , and davies , r . w . ( 1998 ) . phylogenetic relationships of three genera of erpobdellidae ( hirudinoidea ) , with a description of a new genus ( motobdella ) and species , m , sedonensis . can . j . zool . , 76 ( 12 ) : 2164 - 2171 .
pelagic : living in or relating to open water , especially surface waters to the middle depths . this would be away from the shore of montezuma well where the leeches feed on the amphipod at night .
endemic : indigenous to , and restricted to , a particular area . the leech and amphipod studied are endemic to montezuma well , meaning they ' re found only in this location and nowhere else on earth .
batchelder , g . l . 1974 . postpluvial ecology of montezuma well , yavapai co . , arizona . - final rep . arizona archaeol . ctr . , nat . park serv . p . 18 .
usa , arizona : yavapai county , montezuma well ( 34 . 6491°n , 111 . 7522°w ( dd ) ) , aquatic system , under rocks , 10 june 2012 , r . k . beresic - perrins .
runck , c . and d . w . blinn . 1990 . population dynamics and secondary production by ranatra montezuma ( heteroptera : nepidae ) . journal of the north american benthological society 9 : 262 - 270 .
) . this location is thermally constant year - round ( 19–24˚c ) and is continuously replenished by two vents located at the well bottom . montezuma well is 0 . 76 ha in area and approximately 20 m deep . most of the shoreline drops off immediately into open water , except at the northeast corner where water drains through a shallow region called the “swallet” and empties into wet beaver creek which is located east of montezuma well ( fig .
usa : az : yavapai co . , montezuma well 34 . 6491°n , 111 . 7522°w ( dd ) , 10 . vi . 2010 , aquatic system , under rocks , rk beresic - perrins , holotype ( usnm )
montezuma well is a quiet but spectacular example of a large , limnocrene ( pool - forming ) spring . it is one of the best - studied springs ecosystems in the world , thanks in large measure to the dedicated research of
blinn , d . w . , davies , r . w . , and dehdashti , b . ( 1987 ) . specialized pelagic feeding by erpobdella montezuma ( hirudinea ) . holarctic ecol . , 10 : 235 - 240 .
fry , d . , b . obit , j . matson and s . m . shuster . 2007 . laboratory survivorship of leeches from montezuma well , arizona . iii undergraduate research expo , northern arizona university , flagstaff . pdf .
( 14 specimens ) usa : az yavapai co . , montezuma well 34 . 6491°n , 111 . 7522°w ( dd ) , 10 . vi . 2010 , aquatic system , under rocks , rk beresic - perrins , paratypes ( usnm )
- , singhal , r . n . and blinn , d . w . 1985 . erpobdella montezuma , a new freshwater leech from north america ( arizona , usa ) . - can . j . zool . 63 : 965 - 969 .
e . montezuma was collected from montezuma well on 11 different days to examine the seasonal diet of the leech . leeches ( 0 . 2 - 0 . 6 g wet wt . ) were collected in nets at different time intervals 2 hours prior and up to 6 hours after sunset on 8 dates and near sunrise on the remaining 3 dates . to sample various specimens and locations , different sampling techniques were used . petite dredges were used to collect specimens in sediment , vertical nets collected from the water column to quantify plankton prey , and net tows were used in the littoral zone to quantify potential prey in the vegetation . ( to make sure the scientists had all the information they needed , different types of collections at different locations were made at varying times in montezuma well . )
boucher , p . , blinn , d . w . and johnson , d . b . 1984 . phytoplankton ecology in an unusually stable environment ( montezuma well , arizona , u . s . a . ) . - hydrobiologia 119 : 149 - 160 .
davies , r . w . , r . n . singhal and d . w . blinn . 1985 . erpobdella montezuma ( hirudinoidea : erpobdellidae ) , a new species of freshwater leech from north america . canadian journal of zoology 63 : 065 - 969 .
beresic - perrins rk , govedich fr , banister k , bain ba , rose d , shuster sm ( 2017 ) helobdella blinni sp . n . ( hirudinida , glossiphoniidae ) a new species inhabiting montezuma well , arizona , usa . zookeys 661 : 137–155 . urltoken
littoral zone : the zone in a body of fresh water where light penetration to the bottom is sufficient for the growth of plants . this is where most of the aquatic plants grow in montezuma well and where the amphipods migrate to in the evening , from the pelagic zone .
in other words , the leeches are coated in a slimy toxic skin . foust said this is probably how they protect their internal organs . other top predators in montezuma well metabolize the toxin and get rid of it . unlike the leech , they have lower concentrations of arsenic than their prey .
davies , r . w . , singhal , r . n . , and blinn , d . w . ( 1985 ) . erpobdella montezuma ( hirudinoidea : erpobdellidae ) , a new species of freshwater leech from north america . can . j . zool . , 63 ( 4 ) : 965 - 969 .
the well was probably first seen by europeans during the 1583 spanish expedition of antonio de espejo , who described an abandoned pueblo ( likely montezuma castle ) and a pond ( likely montezuma well ) . a railroad survey in the mid - 1880s reported abandoned indian villages in the verde valley , but it was edgar mearns , a military physician stationed at fort verde , who in 1884 was the first scientist to see the site . the smithsonian institution took notice of the well , sending cosmos mindeleff in 1892 and jesse w . fewkes in 1895 to investigate . by 1930 , a total of 185 archaeological sites had been identified in the verde valley and surrounding plateau .
beresic - perrins , r . , d . rose , k . banister , f . govedich and s . shuster . 2013 . helobdella blinni ( hirudinoidea : glossiphoniidae ) a new species inhabiting montezuma well , arizona . society for integrative and comparative biology meeting , san francisco , ca usa , january 2013 . abstract . pdf .
montezuma well ' s significance is due , in part , to its unique water chemistry and ecosystem . as we continue to identify endemic species , we help define the well ' s unique environment and distinguish it from all other locations in the region and world . similarly , research provides invaluable insight into the development and evolution of the mowe ecosystem .
blinn , d . w . 2008 . the extreme environment , trophic structure , and ecosystem dynamics of a large , fishless desert spring : montezuma well , arizona . in stevens , l . e . and v . j . meretsky , editors . every last drop : ecology and conservation of aridlands springs in north america . university of arizona press , tucson , in press .
montezuma well—a collapsed carbonate caldron 368 feet across and 55 feet deep—lies at 3 , 618 feet elevation in the verde valley . its waters likely are derived from the southern colorado plateau just to the north , and undoubtedly have a lengthy and circuitous flow path . the springs well up from the bottom of a collapsed limestone sink and discharge through a natural tunnel at a rate of about 1 , 100 gallons per minute into
mowe ' s uniqueness includes a constant water temperature between 69 - 74 o f –cool in the summer and warm in the winter ; ducks love this and return every fall to bathe in their own spa . the well also has elevated levels of dissolved co 2 from the travertine and naturally occurring arsenic . due to these chemical properties , that include a lack of oxygen , fish cannot breathe in the well . there ' s even an anecdote from some of the first owners of montezuma well in the late 1800s , who tried to stock the well for fishing . in the morning all the poor fish were belly up , and the family none the richer .
from an ecosystem standpoint , the well’s vegetation supports a very large population of amphipods that serve as a prey base . by day , montezuma well’s predator assemblage is dominated by the small , red , desert firetail damselfly , as well as the water scorpion and giant water bug . by night , a horde of endemic leeches swarms up from the depths . the larval damselflies and waterbugs are visual predators in the dense pondweed vegetation around the pool’s periphery . leeches spend their days in the ooze at the bottom of the well , swimming to the surface to feed at night . both sets of predators feed on the endemic amphipod , which spends the day in moderately deep open water , avoiding direct sunlight and the predators in the peripheral vegetation . the amphipod moves to the outer margins of the weed beds at night to avoid predation by leeches .
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govedich , fredric r . , dean w . blinn , paul keim , and ronald w . davies
disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
dean w . blinn , ronald w . davis and behrooz dehdashti northern arizona university
the range of prey species taken by freshwater predatory leeches is normally quite high and changes seasonally with prey availability . while leeches sometimes do ingest whole body prey , they more commonly suck out body fluids . when whole bodies are ingested the exoskeletons are normally quickly evacuated .
in collections taken for microscopic analysis of leech gut contents , animals were individually packaged and preserved to stop digestion . ( if the leeches were kept alive , they would continue to digest they food and the scientist wouldn ' t be able to accurately find out what they were eating . ) all visible prey in the dissected gut was counted for each animal .
amphipod : any small , flat - bodied crustacean of the order amphipoda , having one set of limps adapted for swimming and another for hopping . they look like tiny shrimp .
diel : pertaining to a 24 - hour period , a regular daily cycle . the leeches and amphipods follow one diel cycle .
mean : the average ; statistical term for the average of a series of values , calculated by adding together all of the values and dividing by the number of different values .
quantitative : numerical , or capable of being expressed in numbers ; derived from measurements or other numerical values , e . g . , 10 leeches .
blinn , d . w . and dehdashti , b . 1984 . the nocturnal feeding behaviour of erpobdella punctata ( hiudinoidea ) in a near thermally constant environment . - ( abstr . ) 47th annual meeting , am . soc . . limnol . oceangr . , vancouver , b . c . , p . 9 .
- , wagner , v . t . and grim , l . n . 1986 . surface sensilla on the predacious fresh - water leech eprobdella montezuama : possible importance in feeding . - trans . am . micros . soc . 105 : 21 - 30 .
cole , g . a . 1983 . textbook of limonolgy . - 3rd ed . c . v . mosby , co . , pp . 401 .
davies , r . w . 1969 . the production of antisera for detecting species triclad antigens in the gut of predators . - oikos 20 : 248 - 260 .
- and everett , r . p . 1975 . the feeding of four species of freshwater hirudinoidea in southern alberta . - verh . int . verein . limnol . 19 : 2816 - 2827 .
- , wrona , f . j . and everett , r . p . 1978 . a serological study of prey selection oby nephelopsis obscura verrill ( hirundinoidea ) . - can . j . zool . 56 : 587 - 591 .
- , wrona , f . j . and linton , l . 1979 . a serological study of prey selection by helobdella stagnalis ( hidrudinoidea ) . - j . anim . ecol . 48 : 181 - 194 .
- , wrona , f . j . and wikialis , j . 1981 . inter - and intraspecific analysis of the food niches of two sympatric species of erpobdellidae ( hirudinoidea ) in alberta , canada . - , oikos 37 : 105 - 111 .
elliott , j . m . 1973 . the diel pattern , drifting and food of the leech erobdella octoculuta ( l . ) ( hirudinea : erpobdellidae ) in a lake district stream . - j . anim . ecol . 42 : 451 - 461 .
feigenbaum , d . and reeve m . r . 1977 . prey detection in the chaetognatha : response to a vibrating probe and experimental determination of attack distance in large aquaria . - limnol . oceanogr . 22 : 1052 - 1058 . giguere , l . a . and dill , l . m . 1979 . the prey response of chaoborus larvae to acoustic stimuli , and the acoustic characteristics of their prey . - z . tierpsychol . 50 : 113 - 123 .
ivlev , v . s . 1961 . experimental ecology of the feeding fishes . - new haven , conneticut : yale univ . press .
kirk , k . l . 1985 . water flows produced by daphnia and diaptomous : implications for prey selection by mechanosensory predators . - limnol . oceanogr . 30 : 679 - 686 .
nations , j . d . , hevly , r . d . , landye , j . j . and blinn , d . w . 1981 . the paleontology . paleoecology and depositional history of the miocene - pliocene verde formation , yavapi county , arizona . - in : stone , c . and jenny , j . p . ( eds ) geol . soc . digest 13 : 133 - 149 .
wrona , f . j . , davies , r . w . and linton , l . 1979 . analysis of the food niche of glossiphonia complanata ( hirudinoidea : glossiphoniidae ) . - can . j . zool . 57 : 2136 - 2142 .
young , j . o . 1981 . a comparitive study of the food niches of lake - dwelling triclads and leeches . - hydrobiologia 84 : 91 - 102 .
- and ironmonger , j . w . 1979 . the natural diet of eprobdella octoculata ( l . ) ( hirudinea : erpobdellae ) in british lakes . - arch . hydrobiol . 87 : 483 - 503 .
- and ironmonger , l . w . 1980 . alabratory study of the food of three species of leeches occuring in british lakes . - hydrobiologia . 68 : 209 - 215 .
- and proctor , r . m . 1985 . oligochaetes as a food resource of lake dwelling leeches . - j . freshwat . ecol . 3 : 181 - 187 .
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the leech feeds exclusively on a type of shrimp , and the arsenic consumed in the shrimp is then stored in the leech’s body .
“the other thing that’s interesting is the arsenic seems to be moved from the gut of the organism up to the skin or surface of the organism . we don’t know what that mechanism is ,
said richard foust , lead author of the study and former nau chemistry professor .
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rebecca k . beresic - perrins , 1 fredric r . govedich , 2 kelsey banister , 1 bonnie a . bain , 2 devin rose , 1 and stephen m . shuster 1
1 department of biological sciences , northern arizona university , 617 s . beaver st . , po box 5640 flagstaff , az , 86011 - 5640 ,
2 department of biological sciences , southern utah university , 351 w . university blvd . cedar city , ut , 84720 ,
copyright rebecca k . beresic - perrins , fredric r . govedich , kelsey banister , bonnie a . bain , devin rose , stephen m . shuster
this is an open access article distributed under the terms of the creative commons attribution license ( cc by 4 . 0 ) , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
( linnaeus , 1758 ) , all of which inhabit the swallet ( fig .
we documented number of eyes and their placement , color pattern , presence of papillae , number of and structure of gastric caecae , body size , presence of nuchal scute , gonopore placement , egg size and number , and number of offspring using a nikon binocular dissecting microscope . we then deposited the examined materials in the invertebrate zoology collection at the smithsonian institution , national museum of natural history ( usnm ) .
whole dna was extracted from the caudal suckers of the individual leeches using a qiagen dneasy blood & tissue kit ( cat . no . 69504 ) , with each sample incubated overnight in a water bath set at 54°c . using
oxidase subunit i ( coi ) was amplified . the primers were lco1490 5’ - ggtcaacaaatcataaagatattgg - 3’ and hco2198 5’ - taaacttcagggtgaccaaaaaatca - 3’ (
) . the pcr product was purified through the use of the qiaquick pcr purification protocol ( cat . no . 28104 ) , checked for pcr product using gel electrophoresis , and sequenced with an abi prism 3730 sequencer ( applied biosystems ) . we imported the seven “cleanest” sequences and 71 comparative sequences ( table
) and then corrected the alignments by hand . we partitioned the data and performed the substitution model test by codon in partitionfinder (
) and included 1 , 000 nonparametric bootstrap replicates . we used mrbayes for bayesian inference analysis with ten million generations with a 25 % burn - in and our support was assessed based on clade posterior probabilities (
) to construct parsimony phylogenies with 100 random additions . we performed the parsimony analysis twice , treating the deletions in the sequences as a 5
state and then as missing data . we performed an uncorrected p - distance analysis to examine nucleotide differences between sequences with 1 , 000 replicates in mega7 . 0 . 18 (
internal and external morphology of helobdella blinni sp . n . a dorsal view of the eyes and extended proboscis b crop and post caecae c testisacs d ventral view of internal eggs which have not been oviposited yet e ventral view of white eggs that have been oviposited f ventral view of attached and detached offspring .
diagram of the external and internal morphology of helobdella blinni sp . n . ( drawn by rebecca beresic - perrins and fredric govedich ) .
length of specimens 11 to 22 mm ( mean + se 16 . 6 + 3 . 2 n = 24 ) and width 3 to 8 mm ( 5 . 7 + 1 . 1 n = 28 ) ( table
) . individual color ranges from translucent with brown spots to dark brown ( fig .
) . no dorsal papillae ; one pair of eyes located at somite ii ( 0 . 07 + 0 . 02 mm diameter , n = 11 ) , distance between eyes 0 . 1 to 0 . 03 mm apart ( n = 13 ) . a scallop - shaped nuchal scute is present on the dorsal side , length 0 . 293 to 0 . 432 mm ( 0 . 335 + 0 . 05 n = 9 ) and width 0 . 27 to 0 . 386 mm ( 0 . 32 + 0 . 04 n = 9 ) . one annulus separates the female and male gonopores . the caudal sucker diameter averages 1 . 6 + 0 . 3 mm ( n = 27 ) . the eggs ( diameter 0 . 5 + 0 . 15 mm , n = 28 ) are laid on the ventral side of the parent in soft - walled transparent cocoons ( 7–11 eggs per cocoon , n = 7 ) . the mouth is located subterminally in the oral sucker ( figs
average oral sucker diameter is 0 . 7 + 0 . 19 mm ( n = 15 ) , proboscis length is 3 . 5 + 1 . 1 mm ( n = 17 ) ( table
) . diffuse salivary glands are located near the anterior of the first pair of crop caecae . there are five pairs of smooth crop caecae and one lobed pair of posteriorly directed post caecae . six pairs of compact testisacs are located in between each of the crop caecae . the intestine contains four pairs of caecae , with the first two pairs anteriorly directed and the other two pairs posteriorly directed . the intestine leads into an unraised anus located two annuli from the caudal sucker ( figure
development and growth . this species breeds year - round with peaks in spring and fall . our specimens had an average of 7 to 11 white eggs ( diameter 0 . 5 + 0 . 15 mm , n = 7 ) fixed to their ventral surface . laboratory collections ( 2007–10 ) of helobdella blinni documented the eggs hatching 1 to 2 weeks after ovipositing ( beresic - perrins 2010 ) . once hatched , the young attach to the ventral surface of the parent , allowing the parent to hunt for food and feed the young , occasionally feeding along with them . prey consists of oligochaetes and other invertebrates . the average number of young per adult is 7 + 3 . 3 ( n = 97 ) ranging from 1 to 14 offspring . the young remain attached to the parent for an additional four to five weeks after hatching . once the juveniles leave the parent , they tend to aggregate together on rocks ( beresic - perrins 2010 ) .
. we include the posterior probabilities and maximum - likelihood branch supports > 50 . the arizona populations of
sp . n . , supported by both the bayesian and parsimony analyses . the results of the uncorrected p - distance analysis revealed a difference of 13 . 3 % ( 233 nucleotides included ) between the two groups ( table
( washington ) which is supported by both bayesian inference and maximum - likelihood .
state in our first analysis and in our second , we treated the deletions as missing data . in the resulting 5
, running the spectrum from grey to pink . additionally , they have a descending pair of post caecae , whereas
differences in brooding season and size between helobdella blinni sp . n . , helobdella stagnalis , and helobdella c . f . helobdella .
are slightly longer ( body length 11–22 mm , 16 . 6 + 3 . 2 , n = 24 ) than
species , both from the same region ( rio de flag and oak creek , arizona populations ) and from europe ( uk sample ) . the sequences yielded a 13 . 3 % –17 . 4 % genetic difference between
although currently classified as helobdella c . f . modesta , the arizona populations from the rio de flag and oak creek may be an additional undescribed species based on our molecular analysis . our next step is to investigate these populations more closely , comparing them to other local populations , including white horse lake and j . d . dam lake , az which also contain helobdella modesta .
higher level relationships of leeches ( annelida : clitellata : euhirudinea ) based on morphology and gene sequences .
contributions to the ecology and biology of the danish freshwater leeches ( hirudinea ) .
beresic - perrins rk . ( 2010 ) the life - history and parental care in an arizona population of helobdella stagnalis ( hirudinea : glossiphoniidae ) . master’s thesis , flagstaff , arizona : northern arizona university .
viaggio del dott . a . borellinella republica argentina e nel paraguay . 21 . hirudine´es . bollettino dei musei di zoologia ed .
a comparison of the life cycle of helobdella stagnalis ( linn . 1758 ) ( hirudinoidea ) in two different geographical areas .
the morphology of ozobranchus margoi ( apathy ) ( hirudinoidea ) , a parasite of marine turtles .
folmerr o , blackr m , hoehr w , lutzr r , vrijenhoekr r . ( 1994 )
dna primers for amplification of mitochondrial cytochrome c oxidase subunit i from diverse metazoan invertebrates .
govedich fr , blinn dw , keim p , davies rw . ( 1998 )
the first record of the genus helobdella ( hirudinoidea : glossiphoniidae ) from australia , with a description of a new species , helobdella papillornata .
a new leech species from north america , helobdella californica nov . sp . ( hirudinea : glossiphoniidae ) .
the golden gate leech helobdella californica nov . sp . ( hirudinea : glossiphoniidae ) : occurrence and dna - based taxonomy of a species restricted to san francisco .
lai y - t , chang c - h , chen j - h . ( 2009 )
two new species of helobdella blanchard 1896 ( hirudinida : rhynchobdellida : glossiphoniidae ) from taiwan , with a checklist of hirudinea fauna of the island .
lanave c , preparata g , sacone c , serio g . ( 1984 )
lanfear r , calcott b , ho sy , guindon s . ( 2012 )
life history and production of the leech helobdella stagnalis ( l . ) ( hirudinea ) in a shallow eutrophic reservoir in south wales .
leidy j . ( 1851 ) gen . nov . myzobdella . proceedings of the academy of natural sciences of philadelphia 5 : 243 .
linnaeus c . ( 1758 ) systema naturae per regna tria naturae , secundum classes , ordines , genera , species , cum characteribus , differentiis , synonymis , locis . tomus i . editio decima . holmiae , laurentii salvii .
the breeding , growth and age structure of a population of the leech helobdella stagnalis ( l . ) .
further notes on the leeches ( piscicolidae ) living on fresh - water fishes of north america .
miller ma , pfeiffer w , schwartz t . ( 2010 ) creating the cipres science gateway for inference of large phylogenetic trees . in : proceedings of the gateway computing environments workshop ( gce ) , new orleans , la , 1–8 .
leeches of the united states national museum . proceedings of the u . s .
moser we , klemm dj , richardson dj , wheeler ba , trauth se , daniels ba . ( 2006 )
moser we , richardson dj , hammond ci , lazo - wasem e . ( 2011 )
molecular characterization of helobdella modesta ( verrill , 1872 ) ( hirudinida : glossiphoniidae ) from its type locality , west river and whitneyville lake , new haven county , connecticut , usa .
moser we , fend sv , richardson dj , hammond ci , lazo - wasem ea , govedich fr , gullo bs . ( 2013 )
a new species of helobdella ( hirudinida : glossiphoniidae ) from oregon , usa .
the life history and reproduction of the leech helobdella stagnalis ( hirudinea : glossiphonidae ) in the river ely , south wales .
a new freshwater leech species of helobdella ( annelida : glossiphoniidae ) from central mexico .
oceguera - figueroa a , león - règagnon v , siddall me . ( 2010 )
dna barcoding reveals mexican diversity within the freshwater leech genus helobdella ( annelida : glossiphoniidae ) .
rodriguez fj , oliver jl , marin a , medina jr . ( 1990 )
salas - montiel r , phillips aj , de leon gp , oceguera - figueroa a . ( 2014 )
description of a new leech species of helobdella ( clitellata : glossiphoniidae ) from mexico with a review of mexican congeners and a taxonomic key .
sawyer rt . ( 1972 ) north american freshwater leeches , exclusive of the piscicolidae , with a key to all species . illinois biological monographs , univ . of illinois press , urbana .
phylogeny of leeches ( hirudinea ) based on mitochondrial cytochrome c oxidase subunit i .
phylogeny and revision of the leech genus helobdella ( glossiphoniidae ) based on mitochondrial gene sequences and morphological data and a special consideration of the triserialis complex .
dna - barcoding evidence for widespread introductions of a leech from the south american helobdella triserialis complex .
raxml version 8 : a tool for phylogenetic analysis and post - analysis of large phylogenies .
swofford dl . ( 2003 ) paup * version 4 . phylogenetic analysis using parsimony ( * and other methods ) .
tavare s . ( 1986 ) some probabilistic and statistical problems on the analysis of dna sequences . in : lectures in mathematics in the life sciences , vol . 17 , 57–86 .
the reproductive biology of the leech helobdella stagnalis ( l . ) in utah lake , utah .
vaillant l . ( 1890 ) histoire naturelle des anneles marin et d’eau douce . 3 / 2 , ordre hirudiens ( hirudines ) ou bdelles , paris , 477–542 .
gonimosobdella klemmi n . gen . , n . sp . ( hirudinida : piscicolidae ) from cyprinid fishes in arkansas , illinois , and missouri , usa .
phylogeny of the fish leeches ( oligochaeta , hirudinida , piscicolidae ) based on nuclear and mitochondrial genes and morphology .
. one of the few large limnocrene springs on the colorado plateau , it likely contains the highest concentration of endemic ( unique ) species of any single spring in north america . ecosystem studies in the well have helped us understand the evolution of unique species . located just off interstate 17 near mcguireville , and part of
. the well’s water is relatively constant in temperature , varying from 64 degrees to 77 degrees f , with relatively little dissolved oxygen but a high concentration of dissolved carbon dioxide and relatively high concentration of arsenic . the well has existed at least through the holocene epoch ( 12 , 000 years ) and likely much longer . as an evolutionary microcosm , the pool is essentially free of flood - related disturbance and seasonal impacts . despite its harsh water chemistry , it provides a stable , highly productive environment for the life within it . this , it appears , is a formula for endemism .
, and a new genus and species of leech . in addition to unique species , the well supports several regionally rare aquatic beetles and other insects , including
, a tiny member of the giant waterbug family , and a delicate marsh treader .
concerns over archaeological site pot - hunting and excavation encouraged area residents to propose the area as a national monument .
grand canyon wildlands council , inc . 2002 . a hydrological and biological inventory of springs , seeps and ponds of the arizona strip , final report . arizona water protection fund , phoenix .
springer , a . e . , l . e . stevens , and r . harms . 2006 . inventory and classification of selected national park service springs on the colorado plateau : nps cooperative agreement number ca 1200 - 99 - 009 . national park service , flagstaff .
arizona heritage waters dr . larry stevens museum of northern arizona 3101 north fort valley road flagstaff , az 86001 ( 928 ) 523 - 5211 ext . 204 email dr . stevens
dr . abe springer nau department of geology box 4099 flagstaff , az 86011 ( 928 ) 523 - 7198 email dr . springer
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medicinal leeches became infamous for their utility in bloodletting popularized in the 19 th century , and have seen a recent resurgence in post - operative treatments for flap and replantation surgeries , and in terms of characterization of salivary anticoagulants . notorious throughout the world , the quintessential leech family hirudinidae has been taken for granted to be monophyletic , as has the non - bloodfeeding family haemopidae .
this study is the first to evaluate molecular evidence from hirudinid and haemopid leeches in a manner that encompasses the global scope of their taxonomic distributions . we evaluated the presumed monophyly of the hirudinidae and assessed previous well - accepted classification schemes . the hirudinidae were found not to be monophyletic , falling instead into two distinct and unrelated clades . members of the non - bloodfeeding family haemopidae were scattered throughout the tree and among traditional hirudinid genera . a combination of nuclear 18s rdna and 28s rdna with mitochondrial 12s rdna and cytochrome c oxidase i were analyzed with parsimony and with bayesian methods .
the family hirudinidae must be refined to include only the clade containing hirudo medicinalis ( european medicinal leech ) and related leeches irrespective of bloodfeeding behavior . a second clade containing macrobdella decora ( north american medicinal leech ) and its relatives may yet be recognized in semiscolecidae in order to avoid paraphyly . the african distribution of species from each of the divergent hirudinid clades suggests that a deep divergence took place in the history of the medicinal leeches hundreds of millions of years ago .
medicinal leech
is a common name that describes bloodfeeding clitellate annelids in the family hirudinidae of the order hirudinida . the use of leeches for bloodletting has been a part of western medicine since galen [
, for physician ( merriam - webster dictionary ) . their utility has also been recorded in several eastern traditions , having been documented in the charaka samhita ( maurya period , roughly 3rd century bce ) as one of five treatments for an imbalance of humors and by wang ch ' ung ( 27 - 100 a . d ) [
centuries , european leech populations were over - harvested and leeches became increasingly scarce in parts of western europe . consequently , various countries , such as italy , hungary , and poland , with seemingly abundant sources , began exporting large numbers in order to satisfy the high demand . as early as 1823 , restrictions were put in place to manage the number of leeches being exported through hannover , germany , and collecting seasons were instituted in russia ; these represent some of the first measures in history meant to conserve an animal species [
] to relieve post - operative venous congestion in patients recovering from tissue flap and replantation surgery . their application in this regard proved so successful that european medicinal leeches were approved by the us food and drug administration in june , 2004 as a medical device due to their mechanically relieving venous congestion and delivering anti - coagulants [
] was only possible in light of the newly available purified hirudin , though it would later be supplanted by widely available and less expensive heparin .
. in fact , within the family hirudinidae , approximately 200 species have been described from all continents , save for antarctica . some of these species are used in medical practices in place of
traditionally , the family hirudinidae included any sanguivorous , swimming , freshwater leech with three jaws ( one dorsal and two ventrolateral ) and a distinctively caecate crop . richardson [
] analyses found the family hirudinidae to be split into two major clades with the terrestrial leeches and the non - bloodfeeding haemopidae falling in between . all taxonomic revisions of the family until now have been performed only with morphological characters [ e . g . , [
] ] . here , we revisit the phylogenetic relationships and systematics of the family hirudinidae while testing the monophyly of the family , and for the first time utilizing an expanded taxon sampling from each continent with representatives of most previously proposed subfamilies .
the combined dataset had a total of 6086 characters ( 18s : 2034 characters , 28s : 2162 characters , 12s : 575 characters , co1 : 1315 characters ) . the parsimony analysis produced 9 equally parsimonious trees with 8266 steps while the harmonic mean of log - likelihood values from two runs of the bayesian ( bi ) analysis averaged - 44555 . 69 . the log - likelihood of the topology produced by the maximum likelihood analysis was - 43311 . 984 .
presumed - monophyletic hirudinidae would require 179 extra steps ( templeton test : z = - 8 . 299 ,
> 0 . 0001 ) . the harmonic mean of log - likelihood values constraining traditional hirudinids to be monophyletic was - 45054 . 72 ( yielding a bayes factor of - 998 . 06 ) . similarly , with this constraint under the likelihood criterion , monophyly of hirudinidae was rejected with treefinder [
- values were highly significant ( shimodiara - hasegawa < 0 . 000001 , approximately unbiased test < 0 . 000001 ) . the harmonic mean of log - likelihood values constraining traditional hirudinids and traditional haemopids together to be a monophyletic group was - 44589 . 01 ( yielding a bayes factor of - 66 . 64 ) . similarly , with this constraint under the likelihood criterion , monophyly of hirudinidae + haemopidae was rejected with treefinder [
- values , while not as profound as in the simple case of constraining hirudinidae to be monophyletic , still were significant at the 5 % level ( shimodiara - hasegawa = 0 . 0195 , approximately unbiased test = 0 . 0164 ) .
maximum parsimony and bayesian inference tree topology based on 18s rdna , 28s rdna , 12s rdna , and coi datasets combined . posterior probabilities are above the node and jackknife values are below the node . branch lengths in orange corresponds to terrestrialism , branch lengths in blue correspond to traditional members of the family hirudinidae , and branch lengths in green correspond to traditional members of the family haemopidae .
varied among analyses and received little support in each of parsimony ( jackknife = 32 ) and bi ( pp = 0 . 54 ) analyses . between the two principal hirudinid clades was a paraphyletic assemblage of terrestrial leeches in the families haemadipsidae and xerobdellidae . the parsimony analysis found the genus
individuals were clustered by locality with caribbean individuals closely related to those from thailand . representatives of the genus
with high support values ( jackknife = 100 ; pp = 1 . 00 ) .
the family hirudinidae , long taken for granted to be monophyletic , is not . hirudinid leeches , characterized as relatively large , vermiform , swimming leeches that feed on blood by making an incision with three armed jaws , fall into two separate clades : one typified by the north american


oneida lunulalis , the oak gall snout moth , is a species of snout moth in the genus oneida .

it is found in most of eastern north america , from quebec and ontario to illinois and florida .

the wingspan is about 24 mm .

the forewings are silvery grey or light brown .

the hindwings are grey with a white fringe .

adults are on wing from june to august .

the larvae feed on the leaves and galls on oak .

oneida lunulalis

species oneida lunulalis - orange - tufted oneida - hodges # 5588 - bugguide . net
an orange - tufted oneida moth in prince george ' s co . , maryland . photo by bob patterson . ( mbp list )
similar to oneida grisiella and o . luniferella which occur in the west ( compare images of all 3 species by jim vargo at mpg )
orange - tufted oneida moth in howard co . , maryland ( 8 / 8 / 2005 ) . photo by larry line . ( mbp list )
farther review of the camp wood hills specimen and the three species of oneida , i now think it is o . grisella . what do you think ?
an orange - tufted oneida moth in frederick co . , maryland ( 6 / 8 / 2018 ) . photo by mark etheridge . ( mbp list )
an orange - tufted oneida moth in baltimore co . , maryland ( 7 / 26 / 2014 ) . photo by emily stanley . ( mbp list )
an orange - tufted oneida moth in worcester co . , maryland ( 5 / 28 / 2014 ) . photo by scott housten . ( mbp list )
an orange - tufted oneida moth in calvert co . , maryland ( 6 / 16 / 2006 ) . photo by arlene ripley . ( mbp list )
an orange - tufted oneida moth in howard co . , maryland ( 7 / 14 / 2004 ) . photo by larry line . ( mbp list )
an orange - tufted oneida moth in st . mary ' s co . , maryland ( 7 / 25 / 2017 ) . photo by tyler bell . ( mbp list )
an orange - tufted oneida moth in howard co . , maryland ( 6 / 30 / 2002 ) . determined by john glaser . photo by larry line . ( mbp list )
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terry , it could be but as i read the photos the
tuft
is more orange and the coloration below the tuft rather green and does not have a distinct line below . i tend to favor but i realize it is even further out of range or o . grisella which has been see in the big bend area . not much information on this genus but i think we see more
out of range
moths here from az and nm .
after looking at the three species more carefully , i think it is most likely o . grisella . makes the most since .
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in maryland a moderately common species of oak forests ( glaser , micromoths ) .
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hodges , r . w . et al . , eds . 1983 . check list of the lepidoptera of america north of mexico . e . w . classey limited and the wedge entomological research foundation , london . 284 pp .
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the short-billed canastero ( asthenes baeri ) is a species of bird in the furnariidae family .

it is found in argentina , bolivia , brazil , paraguay , and uruguay .

its natural habitat is subtropical or tropical dry shrubland .

three subspecies are recognized : a. b. chacoensis brodkorb , 1938 - bolivia and paraguay a. b. baeri ( berlepsch , 1906 ) - argentina , bolivia , brazil , paraguay , and uruguay a. b. neiffi ( contreras , 1980 ) - argentina

short - billed canastero

remsen , j . v . , jr ( 2018 ) . short - billed canastero ( asthenes baeri ) . 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 9 july 2018 ) .
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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
sacc . 2005 and updates . a classification of the bird species of south america . available at : urltoken .
justification : this species has an extremely 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 has not been quantified , but it is not believed to 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 size has not been quantified , but this species is described as ' fairly common ' ( stotz et al . ( 1996 ) . trend justification : this population is suspected to be in decline owing to ongoing habitat degradation ( del hoyo et al . 2003 ) .
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this species has an extremely 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 has not been quantified , but it is not believed to 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 .
recommended citation birdlife international ( 2018 ) species factsheet : asthenes baeri . downloaded from urltoken on 09 / 07 / 2018 . recommended citation for factsheets for more than one species : birdlife international ( 2018 ) iucn red list for birds . downloaded from urltoken on 09 / 07 / 2018 .
voice , nest structure , plumage and tail morphology all indicate that this species is part of a group formed by a . dorbignyi and a . berlepschi . races chacoensis and neiffi perhaps no more than extremes in clinal variation , and species may be better treated as monotypic ; reassessment needed . three subspecies tentatively recognized .
brodkorb , 1938 – extreme sc bolivia ( sc santa cruz ) and nw paraguay .
( berlepsch , 1906 ) – s bolivia ( e tarija ) , w paraguay , n & c argentina ( salta , w formosa and w corrientes s to e mendoza , la pampa , ne río negro and s buenos aires ) , extreme se brazil ( sw rio grande do sul ) and w uruguay .
( contreras , 1980 ) – w argentina ( nw & c mendoza , w córdoba , n & c san luis ) .
song a few introductory notes followed by long , fast trill of mechanical , dry notes ; terminal trill . . .
recorded items include ants , orthoptera ( including acrididae ) , diptera , coleoptera ( including cerambycidae ) , hymenoptera , dermaptera . . . .
season during austral spring - summer ; eggs in oct–jan and nestlings in oct . presumably monogamous . nest an oval mass c . 24–35 × . . .
not globally threatened . uncommon to fairly common . occurs in several protected areas . habitat occupied by this species is subject to at least moderate disturbance and . . .
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recent phylogenetic work supports subdivision into 5–6 clades ( herein tribes ) , depending on inclusion or exclusion of xenopini # r .
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2 . savanna - > 2 . 2 . savanna - moist suitability : suitable season : resident major importance : no 3 . shrubland - > 3 . 5 . shrubland - subtropical / tropical dry suitability : suitable season : resident major importance : yes
iucn . 2016 . the iucn red list of threatened species . version 2016 - 3 . available at : urltoken . ( accessed : 07 december 2016 ) .
stotz , d . f . , fitzpatrick , j . w . , parker , t . a . and moskovits , d . k . 1996 . neotropical birds : ecology and conservation . university of chicago press , chicago .
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upiga is a genus of moths of the crambidae family .

the genus has also been placed in the family pyralidae .

it contains only one species , upiga virescens , the senita moth , which is found in north america , where it has been recorded from arizona and california .

senita cactus flowers are pollinated nearly exclusively senita moth adults .

the larvae feed on lophocereus schottii .

they bore into the top of developing fruit , where they consume seeds and fruit tissue .

upiga

mla style :
upiga .
acronym finder . 2018 . urltoken 9 jul . 2018 urltoken
chicago style : acronym finder . s . v .
upiga .
retrieved july 9 2018 from urltoken
apa style : upiga . ( n . d . ) acronym finder . ( 2018 ) . retrieved july 9 2018 from urltoken
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[ globiz ] global information system on pyraloidea globiz ; globales informtationssystem zünslerfalter ; note this information is not automatically synchronized with globiz and can sometimes be lagging behind .
[ nacl ] ; hodges , 1983 check list of the lepidoptera of america north of mexico check list lep . am . n of mexico
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