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
eol content is automatically assembled from many different content providers . as a result , from time to time you may find pages on eol that are confusing .
to request an improvement , please leave a comment on the page . thank you !
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


motyxia is a genus of cyanide-producing millipedes ( collectively known as sierra luminous millipedes or motyxias ) that are endemic to the southern sierra nevada , tehachapi , and santa monica mountain ranges of california .

motyxias are blind and produce the poison cyanide , like all members of the polydesmida .

all species have the ability to glow brightly : some of the few known instances of bioluminescence in millipedes .

motyxia

forma motyxia pior f . secca causey & tiemann , 1969 accepted as motyxia pior chamberlin , 1941
during the day , this millipede , motyxia sequoiae looks like another nonglowing critter .
glowing millipede ( motyxia sequoiae ) lights on , lights off . paul merek - youtube
the suggestion that motyxia ' s glow wards off marauding nocturnal predators is supported by the fact that motyxia are blind , so their visual signaling can only be seen by members of other species , such as predators .
analysis of xystocheir bistipita‘s dna revealed that the millipede actually belonged to a different genus , motyxia - the only genus of bioluminescent millipede in the western hemisphere . the identity - challenged millipede was then renamed motyxia bistipita .
this week , we published a study documenting the rediscovery of the millipede xystocheir bistipita , which turns out to be bioluminescent and a species of motyxia , the luminous mountain millipedes ( marek & moore , 2015 ) . a few folks have asked what the distribution looks like with these new data . here’s an updated distribution of the genus including motyxia bistipita ( we transferred the species into motyxia after determining it was more closely related to bioluminescent millipedes in the genus motyxia ) . the distribution of motyxia is overlaid on level iii usgs ecoregions .
at top , millipedes motyxia sequoiae ( left ) and motyxia bistipita ( right ) are seen normal light . in the bottom row , m . sequoiae , which has more predators , glows much stronger than m . bistipita .
motyxia ' s eastern cousins possess bright and conspicuous reds and yellows , apparently also to ward off daytime predators .
analysis of xystocheir bistipita ' s dna revealed that the millipede actually belonged to a different genus , motyxia — the only genus of bioluminescent millipede in the western hemisphere . the identity - challenged millipede was then renamed motyxia bistipita .
this entry was posted in uncategorized and tagged bioluminescent , california , cyanide , millipede , motyxia . bookmark the permalink .
interestingly , motyxia are blind , and so their visual signaling can only be seen by members of other species , such as predators . in addition , motyxia produce hydrogen cyanide , an extremely poisonous gas as a chemical defense . some of our research indicates that motyxia ’s bioluminescence serves as a warning signal to deter nocturnal mammals from eating these highly poisonous millipedes .
the researchers then sequenced several genes in each xystocheir and motyxia species to determine how the millipedes were related to each other .
a prolonged exposure taken in the darkroom , showing the greenish glow of a motyxia millipede . ( photo : paul marek )
their results , combined with x . bistipita ' s unexpected bioluminescence , showed that x . bistipita actually belongs to the motyxia genus . the scientists renamed it motyxia bistipita . ( also see
world ' s leggiest animal found near silicon valley .
)
the study also revealed that m . bistipita evolved before other motyxia —giving the team a theory for how millipedes got their glow .
when scientists discovered that this millipede , formerly named xystocheir bistipita , emits a faint bluish green glow , they renamed it motyxia bistipita .
in a previous article for nsf , i explained that motyxia ' s
glow means no
to predators . that is , the green glow of nocturnal motyxia - - which are exclusively found in the sierra nevada mountains of california - - wards off nocturnal predators . motyxia ' s bright coloring warns predators that when these millipedes feel threatened , they ooze toxins , including hydrogen cyanide , an extremely poisonous gas .
these discoveries provide the bases for a possible explanation of the evolutionary origins of bioluminescence in millipedes . over time , bioluminescence gradually escalated from the faintly glowing species of motyxia that live at low elevations to the brighter and more highly toxic species of motyxia that live at high elevations .
after we sequenced them we were able to place the millipede on an evolutionary tree with other bioluminescent species of motyxia ,
says marek .
after sequencing the bug’s dna , he found that it was in fact related to its luminous cousins , and changed its name to motyxia bistipita .
when you observe bioluminescence , you may wonder about the purpose of this illuminating phenomenon . my research on motyxia indicates that
glow means no !
to predators . that is , motyxia ' s glow warns nocturnal predators that these 60 - legged creatures are armed and dangerous ; any predator that riles a motyxia risks being squirted by toxins , including hydrogen cyanide , an extremely poisonous gas , which the millipede releases when it feels threatened .
at the field study site in giant sequoia national monument in calif . , paul marek assesses predator bite marks on clay models of the millipede motyxia .
in order to provide a context to the evolution of bioluminescence , my team and i sequenced the dna of x . bistipita , and positioned it on an evolutionary tree with other species of motyxia and their closest relatives . these and other analyses showed that x . bistipita should now be classified in the genus motyxia along with other glowing millipedes . so in honor of its family ties , i gave x . bistipita a new name : motyxia bistipita .
at the field - study site in giant sequoia national monument in california , paul marek assesses predator bite marks on clay models of the millipede motyxia .
entomologist paul marek rediscovered this faintly glowing millipede in san luis obispo and renamed it m . bistipita to reflect its connection the genus of bioluminescent millipedes called motyxia .
species in the genus motyxia are unique in being the only known bioluminescent millipedes . while all xystodesmids can fluoresce under ultraviolet light , motyxia creates its own glow , visible to the naked eye , in what is thought to be a warning signal to nocturnal predators indicating its noxious chemical defenses ( marek et al . , 2011 ) .
millipede motyxia photographed in natural light ( top ) and solely with its own light from bioluminescence ( bottom ) . the animal is an adult about 25 milimeters long .
supplemental information includes detailed experimental procedures and the phylogeny of motyxia species and close relatives ( figure s1 ) . it can be found with this article online at doi : xxxxxx .
only later in time , as motyxia colonized higher elevations with more predators , was the glow co - opted by evolution as a way to warn predators that the millipedes were toxic .
my research team , which is funded by nsf , explores the evolution of bioluminescence in a genus known as motyxia , the only millipedes in north america that are known to be bioluminescent .
a re - evaluation of the millipede genus motyxia chamberlin , with a re - diagnosis of the tribe xystocheirini and remarks on the bioluminescence ( polydesmida : xystodesmidae ) , rowland m . shelley
( of luminodesmus sequoiae loomis & davenport , 1951 ) causey , n . b . ; tiemann , d . l . ( 1969 ) . a revision of the bioluminescent millipedes of the genus motyxia ( xystodesmidae polydesmida ) . proceedings of the american philosophical society , 113 ( 1 ) : 14 - 33 page ( s ) : 31 ; note : motyxia sequoiae [ details ]
the researchers also collected real glowing millipedes from giant sequoia national monument in california , from the species motyxia sequoiae . these are each about 30 millimeters long and 1 gram in mass on average .
millipedes of the genus motyxia spend the day burrowed under soil and leaves . they come out to forage at night , when they are sought after by predators . ( photo : paul marek )
just like all other millipedes , motyxia are vegetarians , feeding mostly on decaying plant material , but in the course of adapting to a lifestyle primarily underground , they lost the ability to see .
many millipedes also have bright colours , but these would be useless to motyxia species . they spend the daytime buried in the leaf litter , emerging only at night to feed on decaying plants . “night is an excellent time to do millipede things like eating detritus and mating , ” says marek . when they’re active , predators wouldn’t be able to see bright colours anyway . as such , motyxia millipedes are a dull orange , and they publicize their defences by glowing in the dark . “i think that motyxia is better able to exploit this nighttime niche if bioluminescent & toxic , ” says marek .
of all these creepy crawlies , eight species are bioluminescent , meaning they can glow just like fireflies and glowworms . all are part of a genus known as motyxia . [ gallery of glowing creatures ]
interestingly , a few of the species in the glowing genus motyxia can switch their glow on and off . marek and his co - workers measured glowing intensity of species in the genus using darkroom photography and traced the results on an evolutionary tree . they determined that the ability to glow evolved only once in millipedes and is restricted to a set of closely related species , all in the genus motyxia .
one species they examined , xystocheir bistipita , had not been seen since its discovery back in 1967 and was assumed to be non - luminescent . but to their surprise , the bug began glowing in their laboratory , arousing suspicion that this millipede may actually be a mislabeled member of the motyxia group . later genetic analysis by the team revealed that this was indeed the case , so the enigmatic species was renamed motyxia bistipita .
previous research had shown that the ten known species of the motyxia genus glow at different levels of brightness . measuring just how much that brightness varies between species can reveal how luminescence evolved—an important goal of evolutionary biologists .
motyxia , which are the only known bioluminescent millipedes , are found solely in a small region of the sierra nevada mountain range in california . but various types of bioluminescent creatures live throughout the united states . they include :
luminescent millipede and results of the field experiment . a . motyxia sequoiae photographed in natural light , and b . entirely with light from luminescence ; c . mean proportion of millipedes attacked versus luminescence . number of individuals attacked above bars ; d . rodent incisor marks in clay millipede , e . live millipede ( arrows ) with anterior segments 1 – 14 missing . ( phylogeny of motyxia species and close relatives in supplemental information online , figure s1 )
one possible clue to the origins of bioluminescence is provided by a millipede species known as motyxia sequoiae , which inhabits habitats that are normally off - limits to other closely related millipedes . these habitats include exposed areas of the forest floor , open mountain meadows and the trunks of oak trees . so perhaps bioluminescence evolved in motyxia sequoiae to protect these creatures from predators in particularly vulnerable areas , and thereby enable these millipedes to expand their range to these favorable locations .
assistant professor of entomology paul marek rediscovered this faintly glowing millipede in san luis obispo and renamed it m . bistipita to reflect its connection the genus of bioluminescent millipedes called motyxia . photo : courtesy of the national geographic society expeditions council
motyxia are extremely common out there ,
marek said .
if you sit there in a moonless night , the ground will look like the starry night sky up above , from all those millipedes glowing in the dark .
according to my explanation for the origins of bioluminescence in millipedes , as their evolution progressed and motyxia colonized higher elevations that support more predators , the millipede repurposed and intensified its glow as a way to warn predators of its greater toxicity .
this is just the tip of the iceberg for the fascinating story of bioluminescence in motyxia ,
marek added .
we still don ' t know a lot of things about their biology and the circumstances under which luminescence evolved .
diminutive in size compared to other millipedes , the species lives at a lower elevation with few predators and was not thought to belong to the genus of millipedes that glow called motyxia — the only genus of bioluminescent millipedes in the western hemisphere .
the location i marked in keene , ca is where i collected the motyxia monica on march 10 , 2013 . the photos were taken the next day at santa barbara city college . my professor verified its bioluminescence and my id for a project of his .
however , my recent research indicates that millipedes may not have always used bioluminescence as a defense mechanism . rather , bioluminescence may have originated in a millipede species named xystocheir bistipita for an entirely different function and slowly evolved into its current defensive function for motyxia .
my research team and i ran an experiment to test whether motyxia ' s coloration warns predators to stay away . our experiment involved positioning 150 glowing clay millipede models and 150 clay non - glowing millipede models in motyxia ' s natural nighttime habitat in california . the results : predators attacked a significantly lower percentage of the glowing vs . non - glowing models ( 18 percent vs . 49 percent . ) the relatively greater ability of the glowing millipede models to repel predators supports the
glow means no !
idea .
my research team , which is funded by the national science foundation ( nsf ) , explores bioluminescence—the biological production of light by natural chemical reactions . specifically we focus on the evolutionary origins of bioluminescence in motyxia , the only millipede genus in california that is bioluminescent .
species such as m . bistipita , which live in low elevations of the sierra nevadas , have to cope with a much hotter , drier climate than other motyxia . their glow is also much dimmer , and they have fewer predators . ( see more pictures of glowing animals . )
but when the members of the motyxia group began to migrate to higher elevations with a greater risk of predation , the bugs repackaged this luminescence system as a warning signal . this was supported by the observation that species containing larger volumes of cyanide were brighter than their slightly less toxic counterparts .
“after we sequenced them we were able to place the millipede on an evolutionary tree with other bioluminescent species in motyxia , ” said marek . “we demonstrated the faint bioluminescence of the low - lying millipedes represented an older trait and the brighter luminescence of their mountain cousins represented a newer trait . ”
causey , n . b . ; tiemann , d . l . ( 1969 ) . a revision of the bioluminescent millipedes of the genus motyxia ( xystodesmidae polydesmida ) . proceedings of the american philosophical society , 113 ( 1 ) : 14 - 33 page ( s ) : 25 [ details ]
causey , n . b . ; tiemann , d . l . ( 1969 ) . a revision of the bioluminescent millipedes of the genus motyxia ( xystodesmidae polydesmida ) . proceedings of the american philosophical society , 113 ( 1 ) : 14 - 33 page ( s ) : 29 [ details ]
causey , n . b . ; tiemann , d . l . ( 1969 ) . a revision of the bioluminescent millipedes of the genus motyxia ( xystodesmidae polydesmida ) . proceedings of the american philosophical society , 113 ( 1 ) : 14 - 33 page ( s ) : 27 [ details ]
after we sequenced them we were able to place the millipede on an evolutionary tree with other bioluminescent species of motyxia ,
said marek .
the faint bioluminescence of the low - lying m . bistipita represented an older trait and the brighter luminescence of their mountain cousins represented a newer trait .
there are around 12 , 000 known species of millipedes , and only the eight motyxia species glow . marek says , “ [ they ] would definitely be on my top 10 for my imaginary “millipede biodiversity global tour” ( along with the shocking pink millipede in thailand & the longest millipede in africa ) . ”
video of my new motyxia cf . tiemanni , this is a rare species of millipede that can actually glow in the dark ! here is a video of them in the light of day , though i will be trying to get a video of them at night while they are glowing , so stay tuned !
( of motyxia pior f . secca causey & tiemann , 1969 ) hoffman , r . l . ( 1999 ) . checklist of the millipedes of north and middle america . virginia museum of natural history , special publication , 8 : 1 - 584 . martinsville page ( s ) : 377 [ details ]
our analyses of the chemical reaction used by m . bistipita to generate its faint glow suggests that this species might not have originally acquired bioluminescence as a defense mechanism . rather , it might have acquired its faint glow to help adapt to the dry heat of its habitat - - before other motyxia acquired bioluminescence .
every evening , these creatures — which remain hidden underground during the day — emerge and initiate a chemical reaction to produce a green - blue glow , a process called bioluminescence . the eerie night lights of these millipedes highlight nature ' s eccentricities , a fringe benefit as i research the millipede species known as motyxia .
our analyses also showed that the faint bioluminescence of the low - lying cousins of motyxia bistipita - - which i ' ll refer to as simply m . bistipita - - represents an older trait than the brighter bioluminescence of their mountain relatives . in addition , millipede species that live at higher elevations exhibit the brightest bioluminescence .
still , a number of poisonous millipede species active during the day are thought to display bright colors to warn predators that they possess toxins and to steer clear . since motyxia are instead out in the dark , marek and his colleagues reasoned
they use their greenish glow in place of a warning coloration ,
he said .
the researchers then began comparing the genomes of the members of the motyxia genus , including its newest member , in order to examine genetic relationships . after constructing an evolutionary tree , the researchers measured the brightness of their characteristic glow photographically and then investigated their toxicity levels by seeing how much cyanide they possessed in their specialized glands .
so the scientists collected several wild specimens all species of motyxia , plus a control group of non - glowing , related millipedes known as xystocheir . the team then set up experiments in the lab to measure the brightness of each species . ( related :
if you see a glowing millipede , best not to bite it .
)
unlike fireflies , whose gleam emanates from a specialized organ on its abdomen , millipedes in the motyxia genus give off a teal hue . these invertebrates have a special type of protein that allows them to produce light from beneath the tough cuticle that covers their bodies . ( also see
glow - in - the - dark millipede explained .
)
how did bioluminescence evolve ? this question is another focus of our ongoing research on motyxia . by helping to reveal the evolutionary origins of warning colorations — which , by necessity , contribute to some of the most blatant and complex appearances in the living world — we expect to improve our ability to investigate and understand how other complex traits arise in nature .
paul marek , a research associate in the university of arizona ' s department of entomology and center for insect science , and his team now provide the first evidence gained from field experiments of bioluminescence being used as a warning signal . they discovered that the nightly glow of millipedes belonging to the genus motyxia helps the multi - legged invertebrates avoid attacks by predators .
species phylogeny of xystocheirine millipedes showing a single origin of bioluminescence in the most recent common ancestor of motyxia species . tree was estimated by using a partitioned bayesian analysis of the concatenated dataset of five genes . gray branches indicate < 0 . 95 posterior probability . ( scale bar : 0 . 1 expected substitutions per site . ) bioluminescent intensity was back - transformed from log - scale .
when they are disturbed , they ooze toxic cyanide and other foul - tasting chemicals from small pores running along the sides of their bodies as a defense mechanism ,
marek explained .
some millipede species that are active during the day display bright warning colors to announce their defenses to predators , but because motyxia are out when it ' s dark , we hypothesized they use their greenish glow in place of a warning coloration .
the reason why these millipedes glow is a mystery . they are blind , adapted to a lifestyle primarily spent underground , so they cannot use their glow to send messages to other members of their species . also , while deep - sea angler fish dangle glowing lures in front of their mouths to attract prey , motyxia are vegetarians just like all other millipedes , feeding mostly on decaying plant material . so they have no need to pull in victims .
interestingly , they found that the millipedes’ bioluminescence originated in motyxia’s common ancestor and then grew brighter over time . furthermore , species living at lower elevations , such as m . bistipita , glow less than their relatives at higher elevations . since m . bistipita’s habitat is much hotter and drier than bugs living further up the mountains , the researchers reasoned that their glow may not have in fact evolved as a warning signal for predators , but instead to help them deal with the stresses of such a climate .
. . . animals may use colour signals in activities such as claiming advantage in maleemale combat ( e . g . eagle owl , bubo bubo , penteriani et al . , 2007 ; frog , phyllomedusa boliviana , jansen & k€ ohler , 2008 ) , defence ( e . g . glow - worm , lampyris noctiluca , de cock & matthysen , 2003 ; motyxia millipedes , marek , papaj , yeager , molina , & moore , 2011 ) , prey attraction ( e . g . . . .
here , we document the unexpected discovery of bioluminescence in m . bistipita and infer the evolutionary history of the species in the context of a comprehensive molecular phylogeny of the xystocheirini . based on the tree , and phylogenetic trait mapping , faint light originated in the most recent common ancestor of motyxia and gradually intensified through the evolutionary diversification of the genus . the linear escalation in luminescent intensity as a function of tree depth reverses in some crown group species and may indicate a shift in the functional role of luminescence over time .
to reconstruct the evolutionary tree , marek and his team included every species in the genus motyxia and all its closest nonglowing relatives and tested the relationship between them and m . bistipita . marek and his team measured light emission photographically in a darkroom by placing individual millipedes in a light tube . the researchers also measured toxicity of the bugs by measuring the volume of cyanide glands that line the sides of the millipedes . bugs with larger cyanide glands were also brighter , indicating a functional link between luminescent intensity of displays and toxicity .
the ability to make your own light , known as bioluminescence , has evolved around 40 to 50 times in the history of life . hundreds of animals can do the same thing , from fireflies to squid to deep - sea fish . they use this ability to attract their prey , to recognise their mates , and to hide from predators . motyxia millipedes are part of this extensive club , but they’re unusual in one important respect : they’re blind . they can’t see their own glows ; their light shows are aimed at a different audience .
marek collected 164 motyxia millipedes from california’s giant sequoia national monument and painted half of them to cover up their nightly glow . he then tethered them with a gently knotted string to specific places throughout the forest . marek also built 300 clay millipedes using a bronze cast made by his wife . he covered half of them with the same obscuring paint as the live millipedes , and the other half with a glow - in - the - dark hue . he scattered the fake millipedes throughout the forest just like the real ones , and waited .
if you go down to the woods of california today , you might be in for a big surprise . at night , the forests crawl with sinuous shapes that glow with an eerie greenish - blue colour . they are motyxia millipedes and they shine brightly whenever they’re disturbed . “if you go to the right forest and you let your eyes get adjusted to the night , then you can see them everywhere , ” says paul marek from the university of arizona . some big oak tress can shelter 1 glowing millipede in every square metre . they look like fields of stars .
despite our growing knowledge , much about motyxia remains mysterious . for example , how do these blind creatures find mates ? what triggers their nightly emergence ? with funding from the national science foundation , my team is working to answer these and other questions . this research is part of our larger effort to describe biodiversity and reconstruct the evolutionary histories of arthropods — a group that includes insects , spiders and crustaceans , and accounts for 80 percent of all living species . we contribute our findings to the tree of life , which is a worldwide effort to define the evolutionary histories of animals .
among the largest millipedes are the north american millipede and the giant african millipede . the north american millipede grows 10 cm ( 3 . 9 inches ) long , and is dark reddish - brown or black with a red line on each segment . females lay a single egg in a nest of regurgitated food , and wrap themselves around it to incubate the egg . giant african millipedes are the world ' s largest known millipedes , growing 30 . 5 cms ( 12 inches ) long . they live in rain forests and have 30 to 40 segments . millipedes belonging to the group motyxia live in mountain areas of california and glow greenish - blue at night , which seems to scare off predators .
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edited by may r . berenbaum , university of illinois at urbana–champaign , urbana , il , and approved march 31 , 2015 ( received for review january 5 , 2015 )
bioluminescent millipedes : m . sequoiae ( a and b ) and m . bistipita ( c and d ) . ( upper ) millipedes photographed under incident white light . ( lower ) millipedes photographed with light emitted from the cuticle . ( inset ) a 5× magnification of the granular photocytes of m . bistipita . ( scale bars : 5 . 0 mm . )
we then tested for a directional trend of luminescence through evolutionary diversification by estimating the relationship between light intensity and phylogenetic distance . if the hypothesis of a phylogenetic escalation of brightness is true , we predicted a strong positive correlation between a node’s luminescent intensity and its distance from the root , as opposed to the null hypothesis of no correlation ( one - tailed test , α = 0 . 05 ) . although trends can occur in both directions , in cases where novel predator / herbivore defense traits ( e . g . , plant secondary metabolites ) promote speciation or a shift into a new adaptive zone , than the quantity or intensity of these features will escalate and evolve in a directional and positive manner (
) . because the ecological phenomenon of defense - trait escalation underlying our hypothesis predicts that the sign of a relationship should be positive , we formulated a one - tailed hypothesis ( the two - tailed probability is also shown in parentheses ) . using the comparative framework afforded by the phylogeny , we also tested the null hypothesis with nonluminescent xystocheirini , predicting conformity with the null expectation . first , we estimated phylogenetic conservatism of luminescent trait values under a brownian motion model of character evolution with pagel’s λ (
) , where λ = 0 indicates a lack of fit between the data and the tree . we found that phylogeny strongly influences our observed luminescent data with a pagel’s λ = 0 . 718 ( not significantly different from 1 , with a bayes factor = 0 . 656 ; compare with λ = 0 , bayes factor = 3 . 53 ) . we then used the branch length scaling factor κ to differentiate between a gradual vs . punctuational mode of continuous trait evolution on the tree , where κ = 0 indicates a punctuational model with change associated with speciation events ( i . e . , clumped at nodes ) , as opposed to a gradual change model ( κ = 1 ) where character evolution is proportional to estimated branch lengths of the phylogram (
) . when the models are compared by using a bayes factor test , the empirical κ = 0 . 544 is not significantly different from κ = 1 ( bayes factor = 1 . 59 ; compare with κ = 0 , bayes factor = 3 . 49 ) . therefore , a gradual model of evolution better fits our observed luminescent trait values , given the phylogeny . we then assessed whether bioluminescence underwent a directional change on the phylogeny by testing a correlation between luminescent intensity and a node’s path length from the root ( calculated from branch lengths ) . using phylogenetically independent contrasts , we estimated the phenotype of the internal nodes of the tree and compared these values with their root - to - node distance using the pearson product moment rho (
) . we conducted the same analyses and tested for a correlation between toxicity ( estimated from cyanide gland area ) and phylogenetic distance . the tests were then repeated with the nonluminescent xystocheirine taxa . in
) . lastly , to test whether the observed correlation could have been due to the random association of traits with species , we generated a probability distribution by repeating the processes of node state reconstruction and correlation tests of light intensity ( or gland area ) and path length from the root for 10 , 000 datasets after shuffling the luminescent intensity data among the tips . we then evaluated whether our empirical pearson
was within the range of permuted values representative of the null distribution of random assignments between traits and species . if < 5 % of our permuted values were greater than our empirical value (
= 0 . 712 ) , then we rejected the null hypothesis that luminescent intensity or gland area is negatively related or unrelated to taxon assignment . because the proportion of instances in the randomized distribution where
= 0 . 045 ( significant as a one - tailed test α = 0 . 05 , but not as a two - tailed test ,
tests of phylogenetic escalation of bioluminescent intensity and cyanide gland area . ( a ) in luminescent millipedes , a positive linear relationship exists between a taxon’s luminescent intensity and its phylogenetic distance from the root . ( c ) a similar relationship is observed between its toxicity and phylogenetic distance . ( b and d ) a weak negative or no evidence of a correlation is evident between luminescence and toxicity with phylogenetic distance in nonluminescent taxa . raw luminescent data points are shown for clarity of illustration . open circles denote luminescent taxa , and dots indicate nonluminescent taxa . regression lines : phylogenetically corrected values ( dotted ) , uncorrected raw values ( solid ) .
we inferred the evolution of continuous traits on the tree using a least - squares parsimony method of estimating ancestral states where the cost of a state change is equivalent to the squared difference in values of the states , ( x – y ) 2 . to correct for the phylogenetic nonindependence of species , we used the pdap module in mesquite to produce a set of independent contrasts for statistical analysis ( 43 ) . if light and toxicity evolved in a directional manner , we expected a positive correlation between the trait value at a node and the distance of the node from the root of the tree . we calculated the nodal values of contrasts and the node’s path length from the root and tested for a correlation using the pearson product - moment correlation coefficient r .
the results of a stepwise multiple regression with bioluminescent intensity as the dependent y - variable and root - to - tip distance ( rt ) , cyanide gland area ( cn ) , and elevation ( el ) as independent predictor variables—and with a minimum akaike information criterion as a stopping rule to select the best model—indicate that cyanide gland area contributes most to the multiple regression equation of y = 18 . 000x rt + 0 . 722x cn + 0 . 0004x el - 3 . 690 , with an r 2 = 0 . 75 . to evaluate millipede body width as a possible confounding factor , because we expect it to covary with cyanide gland area as a result of isometric growth , we conducted a second multiple regression including width as a fourth predictor variable and found that cyanide gland remains the most significant contributor to the regression model .
two anonymous reviewers , charity hall , and jackson means provided improvements to earlier drafts . we appreciate the careful work of tim mccoy and elizabeth francis with dissections . tsutomu tanabe , deren ross , pavel stoev , michael jorgensen , derek hennen , avery lane , brent hendrixson and rob marek helped collect specimens for the project . this work was supported by national science foundation grant deb1410911 and national geographic society grant ec0564 - 12 .
author contributions : p . e . m . and w . m . designed research , performed research , analyzed data , and wrote the paper .
data deposition : the sequences reported in this paper have been deposited in the genbank database ( accession nos . kr135885 – kr136081 ) .
mesquite : a modular system for evolutionary analysis , version 2 . 73 . available at urltoken
pdap package of mesquite , version 1 . 15 . available at urltoken . accessed january 24 , 2014
bayestraits , version 2 . 0 ( beta ) . available at urltoken . accessed march 1 , 2014
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researchers used field data from 2012 to 2015 to study mortality and allele frequency changes in the sea star pisaster ochraceus during a mass mortality event in northcentral california , and found that surviving adult and juvenile sea stars experienced 81 % mortality and allele shifts , according to the authors .
a survey of more than 4 , 600 american adults conducted in 1995 - 1996 and in 2011 - 2014 suggests that among individuals of low socioeconomic status , negative affect increased significantly between the two survey waves , whereas life satisfaction and psychological well - being decreased .
a study of cognitive ability in norwegian males born from 1962 to 1991 suggests that environmental factors rather than changing genetic composition of families likely account for most of the change in norwegian population iq .
the mathematical tools behind recent gerrymandering cases have brought a modicum of precision into the political arena—but this rigor hasn’t always been enough to spur policy changes .
but why do the californian ones glow ? marek knows the answer . with hundreds of millipedes , some clay , and a bit of paint , he has shown that they light up to ward off predators . you might expect that the light shows would make the millipedes easier to find and eat . in fact , it deters hungry mouths .
the next morning , he found that rodents like grasshopper mice , had savaged around a third of the millipedes . the dull ones took the brunt of the attacks – they had between two and four times as many bite marks as the glowing ones . nearly half of the dull clay millipedes bore the wounds of a rodent attack compared to just 22 percent of the glowing models . similarly , rodents had attacked around 18 percent of the painted live millipedes but only 4 percent of the glowing ones .
the experiment showed that the millipedes’ glow repels predators , and the models proved that it’s the light , rather than the smell or taste of the animals , that puts off attackers . the glow sends a clear message : “don’t eat me . i’m dangerous . ” and they are – the millipedes create cyanide in their bodies and secrete the poison through pores along their flanks . they make for an unpleasant and possibly lethal mouthful .
if doesn’t matter that an animal is poisonous if its predators have to bite it to find that out . the predator would get a mouthful of poison , but the prey would incur a serious wound . this is why many poisonous animals advertise their toxic payloads with bright colours .
now , marek wants to find out more about how the millipedes got their lights . by analysing the genes of all 8 species , he found that bioluminescence has evolved only once in this group . while many animals glow by harnessing luminous bacteria , the millipedes rely on their own light - producing protein . what that protein is , and how it’s related to those of other glowing animals , is still a mystery .
reference : marekt , papaj , yeager , molina & moore . 2011 . bioluminescent aposematism in millipedes . current biology . current biology ; citation tbc .
some california millipedes first evolved bioluminescence to cope with harsh desert living , according to scientists who may have cracked the mystery of glowing millipedes .
watch : when the lights go out , these millipedes emit a green glow .
in california ' s sierra nevada mountains ( map ) live a group of blind millipedes that glow in the dark—and now scientists have cracked the mystery of how their brightness evolved , a new study says .
these
species glow like a neon light . it ' s enough light to read something with if you get close enough ,
said rowland shelley , an emeritus scientist at the north carolina state museum of natural sciences who wasn ' t involved in the study .
scientists already knew that these creatures ' radiance serves as a warning to predators .
now , paul marek and wendy moore entomologists at virginia tech and the university of arizona , respectively , have discovered that the animals ' bioluminescence first evolved for another reason entirely : to help them cope with california ' s hot , dry environment .
i thought it was going to be just another boring day in the lab ,
said marek , who has received funding from national geographic ' s expeditions council .
but then he made a surprising discovery : a faint glow emanating from one supposedly non - luminescent millipede species : xystocheir bistipita , according to the study , published may 4 in proceedings of the national academy of sciences .
based on the research , the team thinks these millipedes ' low - watt glow isn’t a warning signal , but rather the result of their bodies ' responses to heat stress .
for instance , millipedes have trouble metabolizing oxygen when it ' s really hot , which creates chemical byproducts such as peroxide . the bioluminescent proteins help neutralize these byproducts and prevent harm to the millipede .
as evidence , the researchers found that the millipedes with the brightest glow also contained the most cyanide in their bodies . ( also see photos :
cyanide millipede , huge spider among new species .
)
the paper and work is very nice , and is a rare combination of real fieldwork with modest molecular methods ,
said peter vrsansky , a scientist at the slovak academy of sciences who was not involved in the new research .
for his part , marek calls the study
a surprising , unique evolutionary story .
, or the sierra luminous millipedes , is a genus of cyanide - producing millipedes in the order polydesmida endemic to the southern sierra nevada , tehachapi , and santa monica mountain ranges of california ( refs . 1 , 2 ) . members of this genus are blind and produce cyanide ( like
in the order polydesmida ) . one of the most remarkable features of these millipedes is their ability to glow at a peak wavelength of
, which are the only known bioluminescent species in the millipede class diplopoda ( about 12 , 000 known species , ref . 5 ) , spare
, are restricted to three counties in california ( los angeles , kern , and tulare ) . all 8 species of
the darkest ( 4 ) . light is emitted from the exoskeleton of the millipede continuously ( the light intensifies when the millipede is handled , ref . 3 ) . emission of light is uniform across the exoskeleton , and all the appendages ( legs , antennae ) and body rings emit light . the internal organs and viscera do not emit light . luminescence is generated by a biochemical process in the millipede ' s exoskeleton ( 3 ) . the light originates by way of a photoprotein , which differs from the photogenic molecule luciferase in firefly beetles ( 6 ) .
another animal that produces light from a photoprotein is the jellyfish aequorea victoria , which is notable for green fluorescent protein ( gfp ) and widely used as a biomarker in molecular biology ( 7 ) .
in size ( 6 , 8 ) . however , the structure of the luminescent photoprotein remains uncertain , and its homology to molecules of closely related arthropods is unknown .
. various functions were suggested : a nighttime aposematic warning signal , that it had no function at all , or that it inadvertently attracted predators ( 1 , 2 ) . a field study tested the hypothesis that bioluminescence acts as a warning signal . based on the results of the field experiment conducted in california , in a spot where
are native , researchers found that bioluminescence strongly deterred nocturnal mammalian predators ( 4 ) .
medium ( 3 cm in length ) ,
flat - backed
polydesmidan millipedes . typically orange - pink in color ( except
fluorescent ( millipedes in the tribe xystocheirini display some of the brightest fluorescence of the u . s . xystodesmidae species )
species occur in live oak and giant sequoia forests , and notably also in meadows . the presence of xystodesmid millipedes in meadows is atypical for the family . most species are observed under canopies of broad - leaf deciduous forests . all
species are exclusively nocturnal . during the day , individuals are burrowed beneath the soil . at night , they emerge ( by an unknown mechanism potentially not related to light since they ' re blind ) and feed on decaying vegetation . individuals of the species
have been observed climbing on tree trunks , possibly consuming algae , lichens , and other cryptogams adhering to the bark .
species are geographically restricted to 3 counties in california : los angeles , kern and tulare . species predominately occur in the santa monica , tehachapi , and southern sierra nevada mountains . the northernmost population of the genus is of the species
in sequoia national park near crystal cave ( 2 ) . the southernmost population is of the species
, collected in 1944 , was recorded from sherman oaks in los angeles county , california . a dubious historical record indicates a riverside county locality ; however , recent collections in the area have not confirmed this .
chamberlin , with a re - diagnosis of the tribe xystocheirini and remarks on the bioluminescence ( polydesmida : xystodesmidae ) .
p . e . marek , d . r . papaj , j . yeager , s . molina , w . moore ( 2011 ) . bioluminescent aposematism in millipedes . current biology 21 : r680–r681 .
p . sierwald & j . e . bond ( 2007 ) . current status of the myriapod class diplopoda ( millipedes ) : taxonomic diversity and phylogeny .
v . r . viviani ( 2002 ) . the origin , diversity , and structure function relationships of insect luciferases .
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marek , p . , papaj , d . , yeager , j . , molina , s . , & moore , w . ( 2011 ) . bioluminescent aposematism in millipedes . current biology , 21 ( 18 ) , r680–1 ( link )
finding glow - in - the - dark - millipedes . npr , science friday video
checklist of the millipeds of north and middle america richard l . hoffman . 1999 . virginia museum of natural history special publications .
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millipede , which scientists have found uses the glow to warn predators of their toxic oozing cyanide .
mysterious glowing millipedes apparently use their radiance to warn nighttime predators to stay away , scientists now find .
more than 12 , 000 species of millipede are currently known to science , but researchers suggest the vast majority remain undiscovered . the real number of species could actually be as high as 100 , 000 or so .
california is the only place on the planet where you can see glow - in - the - dark millipedes ,
said researcher paul marek , an evolutionary biologist at the university of arizona .
the santa monica mountains , the tehachapi mountains and the southern sierra nevada mountains , all of which are in southern california .
[ see photos of glowing millipede ]
they spend the day burrowed beneath the soil and leaf material , but even though they are blind , they somehow sense when night falls , and come to the surface to forage and mate and to go about their millipede business ,
marek said . it currently remains completely unknown how the millipedes detect the fall of night .