exelis pyrolaria , the fine-lined gray moth or plumose gray moth , is a species of moth in the family geometridae .

it is found in north america , where it has been recorded east of the mississippi river and south of illinois and the district of columbia .

the is 22 – 25 mm for males and 19 – 24 mm for females .

adults have been recorded on wing from february to september , with most records from may to august .

the larvae have been recorded feeding on chimaphila umbellate and diospyros virginiana .

exelis pyrolaria

species exelis pyrolaria - fine - lined gray moth - hodges # 6478 - bugguide . net
a revision of the geometrid genus exelis ( lepidoptera ) . frederick h . rindge . 1952 . american museum novitates 1582 .
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upcoming events 2018 bugguide gathering in virginia july 27 - 29 : registration and discussion photos of insects and people from the 2015 gathering in wisconsin , july 10 - 12 photos of insects and people from the 2014 gathering in virginia , june 4 - 7 . photos of insects and people from the 2013 gathering in arizona , july 25 - 28 photos of insects and people from the 2012 gathering in alabama photos of insects and people from the 2011 gathering in iowa photos from the 2010 workshop in grinnell , iowa photos from the 2009 gathering in washington
east of the mississippi river and south of illinois and the district of columbia .
hodges , ronald w . ( ed . ) 1983 . check list of the lepidoptera of america north of mexico .
moth photographers group - map with some distribution data , photos of living and pinned adults .
bold - barcode of life data systems - species account with collection map and photos of pinned adults .
check list of the lepidoptera of america north of mexico . hodges , et al . ( editors ) . 1983 . e . w . classey , london . 284 pp .
disclaimer : dedicated naturalists volunteer their time and resources here to provide this service . we strive to provide accurate information , but we are mostly just amateurs attempting to make sense of a diverse natural world . if you need expert professional advice , contact your local extension office .
contributors own the copyright to and are solely responsible for contributed content . click the contributor ' s name for licensing and usage information . everything else copyright © 2003 - 2018 iowa state university , unless otherwise noted .
confused by a class within a class or an order within an order ? please see our brief essay .
to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
a fine - lined gray moth in anne arundel co . , maryland ( 6 / 9 / 2016 ) . identification verified by roger downer / bamona . photo by timothy reichard . ( mbp list )
a fine - lined gray moth in howard co . , maryland ( 6 / 11 / 2003 ) . photo by larry line . ( mbp list )
use of images featured on maryland biodiversity project is only permitted with express permission of the photographer .
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online list of [ . . . ] the geometridae of the world ( dec 2007 ) , website ( version 01 / 12 / 2007 )
maintained by malcolm j . scoble and axel hausmann . online list of valid and available names of the geometridae of the world , urltoken last update december 2007
disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
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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 .
due to latency between updates made in state , provincial or other natureserve network databases and when they appear on natureserve explorer , for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data . please refer to
distribution data for u . s . states and canadian provinces is known to be incomplete or has not been reviewed for this taxon .
a location where the species occurs or has occurred and where there is potential for persistence or regular recurrence . for most species minimally verification of an adult or larva in association with suitable habitat including larval foodplant . minimum verification standards vary by species from genitalia dissection to decent photographs , but specimens are strongly recommended . it is usually advisable to rear larvae to the adult stage for positive identification .
most of these species are more or less landscape level moths that occupy a variety of wooded habitats and often adjacent shrublands and thickets . in the context of habitat separation , suitable habitat includes marginal habitat and unsuitable means sparsely wooded to treeless places without suitable larval foodplant . for the relatively few included species that are specialized feeders forest or woodland where the foodplant is absent or nearly so can be treated as unsuitable habitats . in particular for the obligate conifer feeders , forest tracts in which suitable ( for that species ) pines , spruces , firs , etc . comprise fewer than 4 canopy trees per hectare may be regarded as unsuitable . see habitat and food comments fields for species - specific information on what constitutes habitat when mapping occurrences .
this figure is arbitrary but a circle of two kilometers radius would define a habitat clearly smaller than most , but well above the smallest ones . it is probably unrealistically low in extensively forested areas . this figure should not be used however if forests are reduced to small woodlots and the landscape is more than 50 % agricultural or otherwise essentially devoid of native tree cover . in such cases the inferred extent is simply the woodlot in which the collection was made . in general with habitats under 1000 hectares assume full occupancy .
use the generic guidelines for the application of occurrence ranks ( 2008 ) . the key for ranking species occurrences using the generic approach provides a step - wise process for implementing this method .
zoological data developed by natureserve and its network of natural heritage programs ( see local programs ) and other contributors and cooperators ( see sources ) .
scoble , m . j . ( ed . ) , m . s . parsons , m . r . honey , l . m . pitkin , and b . r . pitkin . 1999 . geometrid moths of the world : a catalogue . volumes 1 and 2 : 1016 pp . + index 129 pp . csiro publishing , collingwood , victoria , australia .
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natureserve . 2017 . natureserve explorer : an online encyclopedia of life [ web application ] . version 7 . 1 . natureserve , arlington , virginia . available http : / / explorer . natureserve . org . ( accessed :
ridgely , r . s . , t . f . allnutt , t . brooks , d . k . mcnicol , d . w . mehlman , b . e . young , and j . r . zook . 2003 . digital distribution maps of the birds of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
data provided by natureserve in collaboration with robert ridgely , james zook , the nature conservancy - migratory bird program , conservation international - cabs , world wildlife fund - us , and environment canada - wildspace .
patterson , b . d . , g . ceballos , w . sechrest , m . f . tognelli , t . brooks , l . luna , p . ortega , i . salazar , and b . e . young . 2003 . digital distribution maps of the mammals of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
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iucn , conservation international , and natureserve . 2004 . global amphibian assessment . iucn , conservation international , and natureserve , washington , dc and arlington , virginia , usa .
data developed as part of the global amphibian assessment and provided by iucn - world conservation union , conservation international and natureserve .
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aporrectodea is a genus of earthworms in the family lumbricidae .

the genus includes some of the most common earthworms in the palearctic ecozone and in agricultural soils across the temperate regions of the world .

several species are found throughout the world today , having been widely introduced via agriculture .

one of these widespread species , a. trapezoides , was also transported across continents as a popular fishing bait .

some species of this genus are known to be important in the process of soil formation .

among the most familiar species is aporrectodea caliginosa , which has been known as a species complex made up of several very similar taxa whose relationships were not clear .

a phylogenetic analysis confirmed that these taxa are separate species , including a. trapezoides , a. tuberculata , a. longa , and a. nocturna .

aporrectodea

no one has contributed data records for aporrectodea longa yet . learn how to contribute .
you receive more than 100 of the largest and strongest soil dwelling earthworms - southern worm ( aporrectodea trapezoides ) , blackhead worms ( aporrectodea longa ) and dung worms ( lumbricus rubellus ) .
kari pihlaviita added the finnish common name
peltoliero
to
aporrectodea caliginosa ( savigny 1826 )
.
phylogenetic assessment of the earthworm aporrectodea caliginosa species complex ( oligochaeta : lumbricidae ) based on mitochondrial and nuclear dna sequences .
phylogenetic assessment of the earthworm aporrectodea caliginosa species complex ( oligochaeta : lumbricidae ) based on mitochondrial and nuclear dna . . . - pubmed - ncbi
( of aporrectodea jenensis füller , 1953 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of aporrectodea mediterranea ( örley , 1881 ) ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of aporrectodea bowcrowensis reynolds & clapperton , 1996 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of aporrectodea bowcrowensis reynolds & clapperton , 1996 ) reynolds , j . w . ; clapperton , m . j . ( 1996 ) . new earthworm records for alberta ( oligochaeta : lumbricidae ) including the description of a new canadian species . megadrilogica . 6 ( 8 ) : 73 - 82 . [ details ] available for editors [ request ]
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pérez - losada m 1 , ricoy m , marshall jc , domínguez j .
cibio , centro de investigação em biodiversidade e recursos genéticos , universidade do porto , campus agrário de vairão , 4485 - 661 vairão , portugal . mlosada @ urltoken
( of enterion roseum savigny , 1826 ) savigny , j . c . ( 1826 ) . analyse d’un memoire sur les lombrics par cuvier . mem . acad . sci . inst . france . 5 : 176 - 184 . [ details ]
( of allolobophora alpestris bretscher , 1899 ) bretscher , k . ( 1899 ) . beitrag zur kenntnis der oligochaetenfauna der schweiz . revue suisse de zoologie . 6 : 369 - 426 . , available online at urltoken [ details ]
( of allolobophora mucosa eisen , 1873 ) eisen , g . ( 1873 ) . om skandinaviens oligochaeter . öfv . vet - akad . förh . stockholm . 30 ( 8 ) : 43 - 56 . [ details ]
( of dendrobaena diomedaeus cognetti , 1906 ) cognetti , l . ( 1906 ) . nuovi dati sui lumbricidi dell’europa orientale . bollettino dei musei di zoologia ed anatomia comparata della r . università di torino . 21 ( 257 ) : 1 - 18 . [ details ]
( of lumbricus aquatilis vejdovský , 1875 ) vejdovský , f . ( 1875 ) . beitrage zur oligochaetenfauna böhmens . sitzungsberichte der königlichen böhmischen gesellschaft für wissenschaften praha . 1874 : 191 - 201 . [ details ]
( of eisenia rosea var . glandulosa friend , 1910 ) friend , h . ( 1910 ) . new garden worms . gardener’s chronicle . 329 - 330 . [ details ]
( of lumbricus roseus ( savigny , 1826 ) ) risso , a . ( 1826 - 1827 ) . histoire naturelle des principales productions de l ' europe méridionale et particulièrement de celles des environs de nice et des alpes maritimes . paris , levrault : vol . 1 : xii + 448 pp . , 1 map [ 1826 ] ; vol . 2 : vii + 482 pp . , 8 pls [ november 1827 ] ; vol . 3 : xvi + 480 pp . , 14 pls [ september 1827 ] ; vol . 4 : iv + 439 pp . , 12 pls [ november 1826 ] ; vol . 5 : viii + 400 pp . , 10 pls [ november 1827 ] . . 3 ( xvi ) : 1 - 480 , 14 pls . , available online at urltoken [ details ]
( of helodrilus prashadi stephenson , 1922 ) stephenson , j . ( 1922 ) . some earthworms from kashmir , bombay and other parts of india . records of the indian museum . 24 : 427 - 444 . [ details ]
reynolds , j . w . ; jones , a . g . ( 2005 ) . the earthworms ( oligochaeta : acanthodrilidae , glossoscolecidae , and lumbricidae ) of the falkland islands , south atlantic ocean . megadrilogica . 10 ( 10 ) : 75 - 86 . [ english ; abstr . english , spanish , french ] . [ details ]
( of enterion roseum savigny , 1826 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of helodrilus prashadi stephenson , 1922 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of lumbricus roseus ( savigny , 1826 ) ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of bimastus indicus ( michaelsen , 1907 ) ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of eisenia rosea var . glandulosa friend , 1910 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora prashadi ( stephenson , 1922 ) ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora rosea glandulosa ( friend , 1910 ) ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of helodrilus roseus ( savigny , 1826 ) ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of lumbricus aquatilis vejdovský , 1875 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of nicodrilus roseus savigny , 1826 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of eisenia rosea ( savigny , 1826 ) ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora danieli rosai ribaucourt , 1896 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora rosea f . interposita plisko , 1965 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora rosea macedonica rosa , 1893 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora rosea var . paucipartita tzelepe , 1943 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of eisenia rosea dendrobaenoides omodeo , 1950 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of eisenia rosea var . budensis szüts , 1909 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of dendrobaena diomedaeus cognetti , 1906 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of eisenia jenensis füller , 1953 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of eisenia moderata čekanovskaja , 1959 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of eisenia nobilii cognetti , 1903 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of eiseniona diomedaea cognetti , 1906 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of helodrilus diomedaeus cognetti , 1906 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of helodrilus indicus michaelsen , 1907 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora mucosa eisen , 1873 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora rosea savigny , 1826 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora alpestris bretscher , 1899 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora dairenensis kobayashi , 1940 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora harbinensis kobayashi , 1940 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora hataii kobayashi , 1940 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora jeholensis kobayashi , 1940 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
( of allolobophora mediterranea örley , 1881 ) martin , p . ; martinez - ansemil , e . ; pinder , a . ; timm t . ; wetzel , m . j . ( 2016 ) . world checklist of freshwater oligochaeta species . world wide web electronic publication . , available online at urltoken [ details ]
the worms are carefully collected by hand from the windrows at the mynoke® organic certified worm farm . once collected they are replaced in their bedding and immediately sent to you , ready to be released into their new habitat - your raised beds or garden beds . the numbers of each earthworm species in your order may vary according to availability .
in an ideal pasture and garden system you would have earthworms representing all three earthworm functional groups ( dung , topsoil and deep , see figure below ) so that their activity in the soil can complement each other .
while there are still some pastures , which have no earthworms and would benefit from the addition of all earthworm types , it is common for the deep - burrowing earthworms to be absent from our new zealand pasture systems .
figure : the three functional earthworm groups present in our pasture systems . ( source : schon , nicole 2015 )
the southern worm is dark greyish brown in colour , 40 to 90 mm long and lives in the top 20–30 cm of the soil . the blackhead worms are larger ( 90 to 120 mm ) than the southern worms with a significant dark head and burrow deeper than 1m depending on the soil structure .
as the earthworms burrow they ingest ( swallow ) soil and eat the organic matter . the soil and organic matter passes through the earthworm’s digestive system . the organic matter is transformed into humus , mixed with the soil , and released as earthworm casting also called vermicast . the worms and the bacteria living in their guts produce plant growth stimulators such as auxins and highly effective humic acids , which stimulates root growth and flowering .
for more information on the types of earthworms download the earthworms information sheet by nicole schon from agresearch .
if you have images for this taxon that you would like to share with nbn atlas , please upload using the upload tools .
a pale earthworm that is often easy to identify because of the different shades of colour along its body .
habitat lives in non - permanent horizontal burrows in topsoil . rarely found in leaf litter .
confused by a class within a class or an order within an order ? please see our brief essay .
to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
örley , l . ( 1885 ) . a palaearktikus övben élő terrikolák - nak revisiója és elterjedése . magyar tudományos akadémia . 15 ( 18 ) . [ details ]
reynolds , j . w . ; hänel , c . ( 2005 ) . the earthworms ( oligochaeta : lumbricidae ) of tristan da cunha and nightingale islands , south atlantic ocean . megadrilogica . 10 ( 7 ) : 47 - 56 . [ english ; abstr . english , french , spanish ] . [ details ]
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in the assessment of soil genotoxicity using the comet assay . validation of the comet
lter paper contact test . neutral red retention time ( nrrt ) assay was conducted on copper exposed and
comparison to the earthworms from the reference site . the results of this study con
booth et al . , 2005 ; sanchez - hernandez , 2006 ; nahmani et al . ,
2000 ; reinecke and reinecke , 2004 ; martin et al . , 2005 ; zhu et al . ,
2006 ; xiao et al . , 2006a ; manerikar et al . , 20
lled with soil from the farm . earthworms used in this research were adults with
was in the dubrovnikavenue , approx . 1 m from the road while the second site was in
hydrocarbons ( pahs ) in the soils . before the analyses soil samples were dried ,
( 65 % ) and hcl ( min . 36 . 5 % ) in 1 : 3
trometers depending on the metal concentrations in the samples . hg and as in the
microscope slides , and a third layer of 0 . 5 % lmp agarose w
dna ( % tdna ) using the image analysis system , komet 5 , kinetic imaging ltd .
solution and a cover slip . the slides were incubated in a humid chamber and
the red coloration of the cytosol was evident in 50 % of the coelomocytes .
uid in pbs were mixed with 0 . 4 % trypan blue dye and cells
dyed blue were considered non - vital . cell viability was consistently above 95 % in
after exposure to the highest concentration of cadmium ( where it was 80 % ) .
1 . software ( statsoft , inc . ) . two levels were considered signi
1996 ; zang et al . , 2000 ; rajaguru et al . , 2003 ; di marzio et al . ,
2005 ; fourie et al . , 2007 ; li et al . , 20
et al . , 2008 ; prá et al . , 2008 ; yildiz et al . , 2009
pavlica et al . , 2001 ; milowska et al . , 2003 ; villela et al . , 2006
polluted and induced a lesser genotoxic effect ( approx . 1 . 9 - fold over
bohlen , p . j . , 2002 . earthworms . in : lal , r . ( ed . ) , encyclopedia of soil science . marcel
di marzio , w . d . , saenz , m . e . , lemiére , s . , vasseur
espinosa - reyes , g . , ilizaliturri , c . a . , gonzález - mille , d . j . , costilla , r . , díaz - barriga , f . ,
heuser , v . d . , da silva , j . , moriske , h . - j . , dias , j . f
hund - rinke , k . , wiechering , h . , 2001 . earthworm avoidance test for soil assess -
carcinogenic risks of chemicals to humans . iarc , lyon , france . pp . 46 .
assessment of cr ( vi ) using comet assay in earthworm coelomocytes . environ -
. , lapied , e . , van oort , f . , 2003 . effects of heavy metal soil
owojori , o . j . , reinecke , a . j . , voua - otomo , p . , r
rodriguez - castellanos , l . , sanchez - hernandez , j . c . , 2007 . earthworm biomarkers
belgium ) : composition , density and biomass . belgian journal of zoology 130 ,
van schooten , f . j . , maas , l . m . , moon
villela , i . v . , de oliveira , i . m . , da silva , j . , henriques , j . a . p
. , el adlouni , c . , mukhopadhyay , m . j . , viel , g . , nadeau , d . , poirier
. , el adlouni , c . , nadeau , d . , fournier , m . , coderre , d . , poirier , g . g . ,
. . . die bodenfeuchtigkeit wurde auf 70 % der wasserhaltekapazität eingestellt . monroy et al . , 2006 ; klobucar et al . , 2011 ) , und von der bekannt ist , dass ihre resistenz und sensitivität gegenüber bestimmten schadstoffen von denen anderer regenwurmarten abweichen kann ( stenersen et al . , 1992 ; fourie et al . , 2007 ; lukkari et al . , 2005 abbauprozesse allerdings weitgehend stabilisiert und enthält nur geringe mengen an leicht abbaubaren kohlenstoffverbindungen und schnell verfügbaren nährstoffen ( leroy et al . , 2008 ) . ( % ) g kg - 1 tm kas 100 , 0270 , 0 135 , 0 stallmist versuch juni 2014 27 , 1 78 , 2 26 , 0 3 , 0 8 , 5 versuch oktober 2014 19 , 3 61 , 7 24 , 3 2 , 1 9 , 0 versuch juni 2015 21 , 9 69 , 4 29 , 0 2 , 0 7 , 6 versuch oktober 2015 26 , 3 59 , 8 24 , 5 2 , 0 7 , 8 versuch gp mit zuschlagstoffen 2016 24 , 3 n . b . 20 , 3 3 , 0 8 , tabelle 11 : geprüfte faktorkombinationen in den vermeidungstests mit regenwürmern . . . .
. . . two most extensively used methods for the detection of genotoxicity potential of chemicals in the environment are : the single cell gel electrophoresis assay ( known as the comet assay ) and micronucleus test . previous studies have shown that the comet assay is rapid , effective and sensitive method for analyzing and quantifying dna damage levels in the coelomocytes of earthworms exposed to genotoxic compounds , both in vivo and in vitro conditions ( singh et al . , 1988 ; verschaeve and gilles , 1995 ; salagovic et al . , 1996 ; reinecke and reinecke , 2004 ; espinosa - reyes ; liu et al . , 2010 ; klobučar et al . , 2011 ; zhang et al . , 2014 ) . micronucleus assay has emerged as one of the preferred methods for assessing chromosome damage ( chromosome loss and chromosome breakage ) accumulated during lifespan of the cells ( ali and naaz , 2013 ) . . . .
. . . the methods used for determining heavy metal concentrations ( cr , cu , ni , pb , cd , zn , co , and hg ) , arsenic , mineral oil hydrocarbons , and pahs have been previously described in detail [ 20 ] . the concentration of mo was determined in the same way as the heavy metals . . . .
. . . the bibliographic review ( table 1 ) shows that : ( 1 ) pesticides can cause dna damage in earthworms ; two methods can be used to demonstrate dna damage : the micronucleus test and comet assay , the latter being much more sensitive than the former ( casabé et al . 2007 ; klobučar et al . 2011 ) , ( 2 ) pesticides disrupt the activity level of enzymes involved in oxidative stress such as superoxide dismutase , catalase and glutathione - s - transferase ( booth and o ' halloran 2001 ; schreck et al . 2008schreck et al . , 2012wang et al . 2012 ) , ( 3 ) pesticides , in particular organophosphate insecticides , affect the activity of carboxylesterases ( sanchez - hernandez and wheelock 2009 ) and the activity of cholinesterase ( booth and o ' hollaran 2001 ; collange et al . 2010 ; denoyelle et al . 2007 ; gambi et al . 2007 ; hackenberger et al . 2008 ; jordaan et al . 2012 ; olvera - velona et al . 2008 ; schreck et al . 2008 ; venkateswara et al . 2003 ) , ( 4 ) earthworm lysosomal membrane stability , measured using the neutral red retention test , can be altered by pesticides ( booth et al . 2001a , b ; casabé et al . 2007 ; gambi and al . 2007 ; klobučar et al . 2011 ; svendsen et al . 2004 ) and ( 5 ) sub - cellular morphology and histological alterations may be observed following exposure to pesticides ( dittbrenner et al . 2011 ; venkateswara et al . 2003 ) . . . .
. . . the bibliographic review ( table 1 ) shows that : ( 1 ) pesticides can cause dna damage in earthworms ; two methods can be used to demonstrate dna damage : the micronucleus test and comet assay , the latter being much more sensitive than the former ( casabé et al . 2007 ; klobučar et al . 2011 ) , ( 2 ) pesticides disrupt the activity level of enzymes involved in oxidative stress such as superoxide dismutase , catalase and glutathione - s - transferase ( booth and o ' halloran 2001 ; schreck et al . 2008schreck et al . , 2012wang et al . 2012 ) , ( 3 ) pesticides , in particular organophosphate insecticides , affect the activity of carboxylesterases ( sanchez - hernandez and wheelock 2009 ) and the activity of cholinesterase ( booth and o ' hollaran 2001 ; collange et al . 2010 ; denoyelle et al . 2007 ; gambi et al . 2007 ; hackenberger et al . 2008 ; jordaan et al . 2012 ; olvera - velona et al . 2008 ; schreck et al . 2008 ; venkateswara et al . 2003 ) , ( 4 ) earthworm lysosomal membrane stability , measured using the neutral red retention test , can be altered by pesticides ( booth et al . 2001a , b ; casabé et al . 2007 ; gambi and al . 2007 ; klobučar et al . 2011 ; svendsen et al . 2004 ) and ( 5 ) sub - cellular morphology and histological alterations may be observed following exposure to pesticides ( dittbrenner et al . 2011 ; venkateswara et al . 2003 ) . . . .
. . . in particular , the applicability of the comet assay to assess dna damaging potential of environmental contaminants on e . fetida is well known ( klobučar et al . , 2011 and references therein ) . moreover , this species is increasingly employed in many in vivo assessments of genotoxicity and oxidative stress induced by various emerging pollutants , such as triclosan , nanoparticles or polycyclic musks ( chen et al . , 2011 ; li et al . , 2011 ; lin et al . , 2010 ) . . . .
. . . comet assay is able to detect the interaction between xenobiotics and dna as a conse - quence of exposure to complex environmental samples . this assay applied to earthworms ( mostly e . fetida ) has been used in previous studies to investigate the genotoxicity of artificial or natural soils spiked with various chemicals ( reinecke and reinecke , 2004 ; xiao et al . , 2006 ; klobučar et al . , 2011 ) . muangphra et al . ( 2012 ) de - tected no differences in tail dna % , tail length , and tail moment of coelomocytes from earthworms ( pheretima peguana ) exposed to concentrations < lc 50 of gly - based herbicides on filter paper . . . .
. . . the magnitude of lysosomal rupture may induce rep - arable sublethal damage , apoptosis , or necrosis ( hwang et al . , 2004 ) . nrrt has already been established as responsive in earth - worms exposed to metals , such as copper , cadmium , zinc , lead , or - ganic contaminants like polycyclic aromatic hydrocarbons , energetic compounds , and pesticides ( xiao et al . , 2006 ; casabé et al . , 2007 ; klobučar et al . , 2011 ) . it was also previously estab - lished as a useful biomarker of tnt exposure in e . andrei ( robidoux et al . , 2002 ; fuchs et al . , 2011 ) . . . .
s26 - 3 clinical application of psg to rem sleep behavior disorder and periodic leg movements during s . . .
genotoxicity and genetic changes in rtg - 2 fish cell line upon exposure to benzo [ a ] pyrene and ethyl m . . .
genotoxic and reproductive effects of an industrially contaminated soil on the earthworm eisenia fet . . .
polluted soil sampled from a former coking plant in lorraine ( france ) was studied for its genotoxicity and reproductive effects on the eisenia fetida earthworm . genotoxicity was investigated by means of the single - cell gel electrophoresis ( comet ) assay on the coelomocytes of earthworms after 4 and 10 days of exposure to the soil . dna damage and a decline in the number of coelomocytes extruded . . . [ show full abstract ]


the rusty-bellied brush-furred rat ( lophuromys sikapusi ) is a species of rodent in the family muridae .

it is found in angola , benin , cameroon , central african republic , republic of the congo , democratic republic of the congo , ivory coast , equatorial guinea , gabon , ghana , guinea , kenya , liberia , nigeria , sierra leone , tanzania , togo , and uganda .

its natural habitats are subtropical or tropical moist lowland forests , subtropical or tropical seasonally wet or flooded lowland grassland , arable land , and pastureland .

rusty - bellied brush - furred rat

false rusty - bellied brush - furred rat ( < i > l . lophuromys pseudosikapusi < / i > )
rusty - bellied brush - furred rat - growth and reproduction . . . this type of brush - furred mice produces large pups , but generally small litters . . . the infant rusty - bellied brush - furred mice become active in four to seven days , and their eyes begin to open . . . a young rusty bellied brush furred rat is called a pup , nestling , or pinkie . . .
hutterer ' s brush - furred mouse . . . hutterer ' s brush - furred mouse or hutterer ' s brush - furred rat ( lophuromys huttereri ) is a species of rodent in the family muridae . . .
with reverso you can find the english translation , definition or synonym for rusty bellied brush furred rat [ lophuromys sikapusi ] and thousands of other words . you can complete the translation of rusty bellied brush furred rat [ lophuromys sikapusi ] given by the english - french collins dictionary with other dictionaries such as : wikipedia , lexilogos , larousse dictionary , le robert , oxford , grévisse
millardia . . . it contains the following species sand - colored soft - furred rat ( millardia gleadowi ) miss ryley ' s soft - furred rat ( millardia kathleenae ) kondana soft - furred rat ( millardia kondana . . .
colombian soft - furred spiny - rat . . . the colombian soft - furred spiny - rat ( diplomys caniceps ) , or arboreal soft - furred spiny - rat , is a species of rodent in the echimyidae family . . .
menagesha brush - furred mouse ( < i > l . menageshae < / i > )
chercher mountains brush - furred mouse ( < i > l . chercherensis < / i > )
kilonzo ' s brush - furred mouse ( < i > l . kilonzoi < / i > )
makundi ' s brush - furred mouse ( < i > l . makundii < / i > )
sabuni ' s brush - furred mouse ( < i > l . sabunii < / i > )
stanley ' s brush - furred mouse ( < i > l . stanleyi < / i > )
list of the saga of darren shan characters - others - the dark - furred wolf . . . eventually streak and the dark - furred wolf got into a fight . . . neither of them were seriously injured but afterward the dark - furred wolf never bothered darren again . . .
different types of snakes can be found in temperate forests such as the garter snake or the rat snake . if there is nearby water , various types of toads , lizards , and frogs might be seen around as well . insects of all types love the forest including bumblebees , spiders , and mosquitoes .
our review has revealed three predominant leptospira species and a considerable diversity in reported pathogenic leptospira serogroups in people and animals across the continent . animal hosts , including livestock and invasive and indigenous rodent species , were reported for the majority of species and serogroups detected in human cases . however , there was poor geographical overlap in serogroup reporting between human and animal studies . based on the findings of this review , we suggest that the major animal hosts of human - infecting serovars may vary across africa and that both livestock and rodents may play important roles in human disease transmission . few data were identified that described leptospira spp . diversity in human cases and animal populations from the same country , and few studies attempted to link data on acute human leptospirosis with evidence of leptospira infection in local animal populations . studies on the indian ocean islands of mayotte and madagascar were the exception to this . isolates with unusual patterns of genetic and serological diversity , recently reclassified as a new pathogenic species leptospira mayottensis [ 134 ] , were detected from both human and black rat infections , implicating the black rat as the source of these human infections [ 35 , 86 ] . these studies demonstrate the value of integrated human and animal research to identify sources and transmission routes of human leptospirosis , which can in turn help prioritise investment in disease prevention and control efforts .
leptospira spp . isolated from human patients with acute leptospirosis belonged to five pathogenic leptospira species ( table 5 ) . l . interrogans was the most widespread and common species reported in either human or animal studies in africa . multiple animal hosts were identified for l . interrogans as well as the other common species , l . borgpetersenii and l . kirschneri , from a variety of countries . the widest diversity in leptospira spp . was reported from two kenyan studies of acute human leptospirosis , where isolates belonging to five species were identified ( l . borgpetersenii , l . interrogans , l . kirschneri , l . noguchii and l . santarosai ) [ 38 , 39 ] . however , l . noguchii and l . santarosai were not detected in any other studies . four leptospira species : l . borgpetersenii , l . borgpetersenii - like , l . interrogans and l . kirschneri ; were identified in two human studies on mayotte , as well as by a concurrent study of black rats performed during the same period [ 34 , 35 , 86 ] . divergent leptospira spp . described as l . borgpetersenii - like and l . borgpetersenii group b were detected in human and animal studies respectively in mayotte , and in a study of indigenous small mammals in madagascar [ 34 , 35 , 86 , 125 ] . sequencing and alignment of the atypical isolates from rat kidneys in mayotte [ 86 ] showed perfect identity with isolates derived from people [ 35 ] .
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justification : listed as least concern in view of its wide distribution , tolerance of a degree of habitat modification , presumed large population , and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category .
this species is largely distributed in west africa and central africa . it is found from sierra leone in west africa to tanzania and kenya in east africa and southward to northern angola . the eastern range limits are not well resolved . in tanzania its distribution is patchy , and it is found around high mountains in the appropriate altitudinal and climatic habitat . at mount nimba ( guinea ) it occurs up to 1 , 600 m asl .
this species inhabits dense , moist grasslands , secondary growth , agricultural fields , abandoned farmlands , swamps and grassy plantations where there is abundant low cover . in primary and secondary forests , it occurs only in patches of grassland and herbaceous cover , most especially in open areas . it is a terrestrial and nocturnal , crepuscular species . this species has both an insectivorous and omnivorous diet .
there are no conservation measures in place ; in vieew of its wide range it is probably present in some protected areas .
this errata assessment has been created because the map was accidentally left out of the version published previously .
( errata version published in 2017 ) . the iucn red list of threatened species 2016 : e . t12356a115104961 .
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family : muridae ( mice , rats , voles , gerbils , hamsters , etc . )
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most of us know that a forest is a
large , densely wooded area
full of trees and plants not to mention a whole world of insects and animals . however , if we were to ask the question as to what types of animals dwelt in the forest , we might be hard pressed to give a correct answer .
a relatively temperate forest full of deciduous trees is quite different from a rain forest and houses different types of animals . typical mammals found in this type of forest include the black bear , white - tailed deer , raccoon ,
, grey squirrel , chipmunk , and other small rodents . many different types of birds are also found in the temperate , deciduous forest including the cardinal , turkey , eagle , and
forest animals vary from area to area . forest animals in the southern hemisphere are quite different than those in the northern . even forests on the west and east coasts of the united states vary in habitants . the different temperatures , climate changes , and tree coverings attract different types of animals , but this is the general overview of animals living in a temperate , deciduous forest .
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the golden spiny mouse ( acomys russatus ) gets its name from the reddish - orange spiny fur that covers it body from head to tail . this coarse , inflexible fur is thought to protect it from predation . [ 2 ] aside from the golden fur that covers its head and upper parts , its flanks are yellow and its underside is pale . it has gray legs with pale feet and black soles . it is also described as having a small , but distinct white spot under each eye . [ 3 ] it is often found in the wild missing a part or all of its tail because it is able to shed this as a defense mechanism . however , it is not known how this is done , how often it can occur , or under what conditions . [ 4 ] it lives an average of three years in the wild . it is omnivorous and feeds on seeds , desert plants , snails , and insects . living in desert regions , it is a xeric animal that obtains water from the plants that it eats and produces very concentrated urine in order to conserve water . [ 2 ] a . russatus is naturally nocturnal , but adapts to being diurnal when it shares a habitat with a . cahirinus .
the golden spiny mouse thrives in dry , hot deserts and is common in egypt and much of the middle east , including jordan , israel , oman , saudi arabia , and yemen . it is limited to these areas because its metabolic rate is much slower than predicted for an animal of its size , [ 5 ] so it is unable to maintain a viable body temperature in colder environments where the temperature falls below 18 °c [ 6 ] and can live in temperatures as high as 42 . 5 °c . a . russatus does not have a thermal neutral zone , but shows lowest oxygen consumption at 30 °c . [ 5 ] however , one population was found in the high summits of the mountains in southern sinai where the temperature falls well below this threshold and there is often snow in the winter . [ 6 ] a . russatus is also very good at conserving the little water it gets from its diet of insects and plants . it does this by producing extremely concentrated urine with urea content up to 4800 mm and chloride concentration up to 1500 mm . this means that it could survive drinking sea water , which is very rare in mammals . [ 5 ]
acomys russatus is an opportunistic breeder that reaches sexual maturity at approximately 58 days of age . there is no mating season for spiny mice , but rather they continue to breed throughout the year . each female has an average of 3 . 5 litters per year with about 34 days between litters . [ 7 ] they are viviparous and each gestation period lasts 5–6 weeks and produces 1–5 offspring . [ 2 ] weaning of each litter takes two weeks . [ 7 ] when rainfall decreases , the salt concentration in their food sources increases , which negatively impacts reproduction in both males and females . [ 8 ] [ 9 ]
golden spiny mouse ( acomys russatus ) . arkive . org . retrieved on 2012 - 12 - 28 .
hoath , r . a . ( 2004 ) field guide to the mammals of egypt . the american university in cairo press , cairo and new york isbn 9774248090
shargal , eyal ; rath - wolfson , lea ; kronfeld , noga ; dayan , tamar ( 1999 ) .
shkolnik , amiram ; borut , arieh ( 1969 ) .
temperature and water relations in two species of spiny mice ( acomys )
.
haim , a . ; borut , a . ( 2009 ) .
size and activity of the cold resistant population of the golden spiny mouse
.
spiny mouse ( acomys cahirinus ) longevity , ageing , and life history . urltoken ( 2012 - 09 - 17 ) . retrieved on 2012 - 12 - 28 .
shanas , u . ; haim , a . ( 2004 ) .
diet salinity and vasopressin as reproduction modulators in the desert - dwelling golden spiny mouse ( acomys russatus )
.
wube , tilaye ; haim , abraham ; fares , fuad ( 2009 ) .
effect of increased dietary salinity on the reproductive status and energy intake of xeric and mesic populations of the spiny mouse ,
zisapel , nava ; eilon barnea ; ido izhaki ; anis yossi & abraham haim ( 1999 ) .
daily scheduling of the golden spiny mouse under photoperiodic and social cues
.
levy , ofir ; dayan , tamar ; kronfeld - schor , noga ( 2007 ) .
the relationship between the golden spiny mouse circadian system and its diurnal activity : an experimental field enclosures and laboratory study
.
kronfeld - schor , n ; t . dayan ; m . e . jones ; i . kremer ; y . mandelik ; m . wollberg ; y . yassur & d . d . gaton ( 2001 ) .
retinal structure and foraging microhabitat use of the golden spiny mouse (
this page was last edited on 8 march 2018 , at 01 : 34 .
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the central african republic is located in central africa and covers mostly savannah ( treeless expanses of open grassland ) in the north , and dense congo jungle to the south along the ubangi and sangha rivers . a wide range of vegetation can be found within these different zones such as scrubby drought resistant shrubs and trees to the larger , more leafy forests in the congo . this difference in vegetative zones gives the republic a much larger range of animal species to fill the two different habitats .
, leopards , baboons , antelope , buffalo , and other species of insects , amphibians and reptiles . in the dense congo jungle you can find the smaller forest elephants ,
, leopards , and other primates . located in the rivers you cannot forget the crocodiles , hippos and large variety of fish species .
a special population of
bili
apes are also established within the congo region ; this particular troop of chimps are larger then their cousins and also build their nests on the ground instead of in the trees . this behavior is unusual as most chimps are afraid of larger predators , but the bili chimps have been seeing feasting on a leopard carcass . no big cat hunting has been observed however , so they may have scavenged the cat .
the black rhino also inhabits the area , one of the last niches this rare species can be found . along with the rhino several other species are endangered such as the
, apa , bongo , cheetah , chimpanzee and many other species . endangered plants can also be found in the area such as neoschumannia kamerunensis and nesogordonia papaverifera .
the central african republic homes many different species , endangered and none alike . its rich jungles and open plains are home to many different critters , all of which need to be protected for future generations .
the animals displayed on this page are grouped in their scientific order . view also countries of the world ordered by :
plos neglected tropical diseases is the top open access tropical medicine journal , featuring an international editorial board and increased support for developing country authors .
correction : epidemiology of leptospirosis in africa : a systematic review of a neglected zoonosis and a paradigm for ' one health ' in africa . plos neglected tropical diseases 10 ( 3 ) : e0004552 .
leptospirosis is an important but neglected bacterial zoonosis that has been largely overlooked in africa . in this systematic review , we aimed to summarise and compare current knowledge of : ( 1 ) the geographic distribution , prevalence , incidence and diversity of acute human leptospirosis in africa ; and ( 2 ) the geographic distribution , host range , prevalence and diversity of leptospira spp . infection in animal hosts in africa .
following preferred reporting items for systematic reviews and meta - analyses ( prisma ) guidelines , we searched for studies that described ( 1 ) acute human leptospirosis and ( 2 ) pathogenic leptospira spp . infection in animals . we performed a literature search using eight international and regional databases for english and non - english articles published between january 1930 to october 2014 that met out pre - defined inclusion criteria and strict case definitions .
we identified 97 studies that described acute human leptospirosis ( n = 46 ) or animal leptospira infection ( n = 51 ) in 26 african countries . the prevalence of acute human leptospirosis ranged from 2 3 % to 19 8 % ( n = 11 ) in hospital patients with febrile illness . incidence estimates were largely restricted to the indian ocean islands ( 3 to 101 cases per 100 , 000 per year ( n = 6 ) ) . data from tanzania indicate that human disease incidence is also high in mainland africa ( 75 to 102 cases per 100 , 000 per year ) . three major species ( leptospira borgpetersenii , l . interrogans and l . kirschneri ) are predominant in reports from africa and isolates from a diverse range of serogroups have been reported in human and animal infections . cattle appear to be important hosts of a large number of leptospira serogroups in africa , but few data are available to allow comparison of leptospira infection in linked human and animal populations . we advocate a ‘one health’ approach to promote multidisciplinary research efforts to improve understanding of the animal to human transmission of leptospirosis on the african continent .
leptospirosis is an important bacterial zoonosis that affects people and animals worldwide . it is common in tropical areas where people and animals live in close contact , but the disease has been widely neglected in africa . in this study we aimed to demonstrate the extent of leptospirosis in africa and describe the diversity of the causative agent leptospira spp . in human and animal infections across the continent . through a systematic literature review , we identified 97 studies from 26 african countries that described human disease or animal infection and met inclusion criteria . leptospirosis was the cause of illness in 2 3 % to 19 8 % of hospital patients with a fever . where population - level data were available , leptospirosis was estimated to affect 3 to 102 people per 100 , 000 every year . a variety of animal hosts of leptospira spp . were identified . cattle were reported as carriers of a variety of serological types of leptospira spp . infection . the role of cattle and many other different animal hosts in human disease transmission remains unclear . our review demonstrates that leptospirosis is a substantial cause of human illness in africa , and we recommend integration of human and animal studies in the future to help us understand the epidemiology of leptospirosis on this continent .
citation : allan kj , biggs hm , halliday jeb , kazwala rr , maro vp , cleaveland s , et al . ( 2015 ) epidemiology of leptospirosis in africa : a systematic review of a neglected zoonosis and a paradigm for ‘one health’ in africa . plos negl trop dis 9 ( 9 ) : e0003899 . urltoken
copyright : © 2015 allan et al . this is an open access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited
funding : this research was supported by the joint us national institutes of health ( nih : urltoken ) – national science foundation ( nsf : urltoken ) ecology of infectious disease program ( r01tw009237 ) and the rcuk , dfid and uk biotechnology and biological sciences research council ( bbsrc : urltoken ) ( grant numbers bb / j010367 / 1 , bb / l018926 , bb / l017679 , bb / l018845 ) . kja is supported by the wellcome trust ( urltoken ) ( 096400 / z / 11 / z ) . rrk also received support from the wellcome trust under the afrique one initiative ( wt 087535ma ) . hmb received support from the nih interdisciplinary research training program in aids ( niaid - ai007392 ) . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
endemic zoonotic diseases affect impoverished and developing communities worldwide but are frequently overshadowed in public and clinician awareness by high profile infections such as malaria and hiv / aids [ 1 , 2 ] . in africa , zoonotic infections are both directly responsible for human illness and death and indirectly impact human well - being as a result of reduced livestock productivity and food security [ 3 – 5 ] . however , bacterial zoonoses including leptospirosis remain under - diagnosed and under - reported in africa , and as a result are overlooked as public health priorities [ 1 , 2 , 6 ] .
leptospirosis is one of the most common and widespread zoonotic infections in the world and is recognised as a neglected disease by the world health organisation ( who ) [ 7 ] . human leptospirosis is caused by infection with pathogenic strains of leptospira spp . bacteria [ 8 , 9 ] . more than 250 pathogenic leptospira serovars are known to exist worldwide , which are classified into 25 serogroups on the basis of their serological phenotype [ 10 , 11 ] . recent species determination by dna homology has identified 13 pathogenic leptospira spp . , and seven of these ( l . interrogans , l . borgpetersenii , l . santarosai , l . noguchii , l . weilli , l . kirschneri and l . alexanderi ) are considered as the foremost agents of human and animal disease [ 10 , 12 ] . both serological and dna - based classification systems are currently in use for clinical diagnosis and in understanding the pathogenesis and epidemiology of the disease [ 11 , 13 , 14 ] .
a wide range of animals can carry pathogenic leptospira bacteria and act as a source of infection [ 8 , 11 ] . leptospira serovars often demonstrate a degree of animal host preference and some common relationships between serovars and their hosts are reported [ 9 , 15 ] . following infection , the bacteria colonise the renal tubules and urogenital tract and are shed in the urine of infected animals . animal species may be asymptomatic carriers of infection ( maintenance hosts ) or develop clinical disease ( accidental hosts ) depending on the infecting serovar [ 11 , 16 ] . in food producing animals , cattle and pigs are relatively susceptible to clinical infection resulting in production losses including reduced milk yield , reproductive failure and abortions [ 16 , 17 ] .
in people , disease occurs through direct or indirect contact with infected urine from an animal host [ 8 , 9 , 15 ] . good knowledge of leptospira serovars circulating in local animal populations is important to determine sources and transmission routes for human infection [ 8 ] . in the early stages , human leptospirosis manifests most commonly as a non - specific febrile illness that is hard to distinguish from other aetiologies of febrile disease particularly in tropical areas [ 11 , 18 , 19 ] . infection can result in severe secondary sequelae including renal failure and pulmonary haemorrhagic syndrome , and a case fatality ratio of up to 50 % has been reported in complicated cases [ 15 , 19 ] .
leptospirosis is particularly common in the tropical areas where people and animals live in close contact , and warm and humid conditions favour environmental survival and transmission of the pathogen [ 8 , 9 ] . in south - east asia and south america , leptospirosis is recognised as an important cause of renal failure and febrile disease [ 18 – 22 ] . however , despite its global importance , large gaps persist in our knowledge of the burden and epidemiology of leptospirosis in africa . reports from the who leptospirosis epidemiology reference group ( lerg ) indicate that leptospirosis incidence may be high in africa , but also highlight the lack of available data [ 7 , 23 ] . although reported seroprevalence data demonstrates widespread exposure to leptospira spp . in humans and animals in africa , [ 24 ] little is known about the extent of human disease or the epidemiology of leptospira infection in different animal species in africa .
to tackle these gaps in current understanding and awareness of human and animal leptospira infection in africa , we performed a systematic review of peer - reviewed and grey literature following the preferred reporting items for systematic reviews and meta - analyses ( prisma ) guidelines [ 25 ] . our aims were summarise and compare : ( 1 ) current knowledge of the geographic distribution , prevalence , incidence and diversity of acute human leptospirosis in africa ; and ( 2 ) the geographic distribution , host range , prevalence and diversity of leptospira spp . infection in animal hosts in africa .
a detailed protocol for this study can be found in the supplementary material ( s1 file ) . following the prisma guidelines and checklist ( s1 checklist ) references for this review were identified through searches of eight international and regional databases ( table 1 ) using the search string ‘leptospirosis’ or ‘ leptospira ’ and ‘africa * ’ for articles published between january 1930 and october 2014 inclusively . additional articles for inclusion were identified by bibliography hand searches of relevant articles [ 26 ] .
abstracts and titles were compiled in endnote ( thomson reuters , philadelphia , pa , usa ) and reviewed independently by two researchers ( kja , hmb ) to determine whether each article met pre - determined abstract inclusion and exclusion criteria ( s1 file ) . a third researcher ( jebh ) served as a tiebreaker for any discordant decisions . citations were included if they presented data on human or animal leptospira spp . infection from any country within the united nations ( un ) definition of africa [ 27 ] . we excluded abstracts that did not refer to original human or animal leptospirosis research data , or did not describe naturally occurring cases of leptospirosis in human or animal populations . we included case reports but excluded reports of returned travellers because of potential uncertainty around the specific location where infection was acquired .
case definitions for study inclusion : acute human leptospirosis and confirmed animal carrier hosts .
two reviewers ( kja , hmb ) independently extracted pre - determined qualitative and quantitative data from each included article . data on infection prevalence and incidence for comparable studies ( i . e . similar study inclusion criteria and diagnostic methodologies ) were compiled , and ranges were presented by study type ( human studies ) , location or host species ( animal studies ) if three or more citations reporting comparable data were identified . data on serological and genetic typing of leptospiral isolates from people and animals were compiled and summarised by country and by animal species . additional data on serogroup and genetic species of reported serovars was obtained from the leptospirosis library , maintained by the leptospirosis reference centre , royal tropical institute ( kit ) , netherlands [ 29 ] .
the risk of bias in included studies such as selection or reporting bias was assessed following the cochrane guidelines for systematic reviews of medical interventions [ 30 ] . full text study validity and methodological quality was assessed by comparison to pre - determined case definition criteria to control for heterogeneity in study design and diagnostic methodology ( table 2 ) . studies classified as high - risk for bias were not included in quantitative analysis of leptospirosis prevalence and incidence data .
our searches yielded 681 unique articles from a total of 1201 abstracts identified by database searches . data can be accessed through :
after abstract and full text review , 95 citations published between 1956 and 2014 were eligible for inclusion . hand searches identified two additional articles that met inclusion criteria but were not identified in the original database search . reasons for full - text exclusion are detailed in
. in total we included 97 articles that described human or animal studies conducted in 26 ( 44 . 8 % ) of 58 countries included in the un macro - geographical definition of the african continent (
] . major risks of bias identified in eligible studies were selection bias , attrition bias in studies that relied on paired serology ( mat ) for confirmatory diagnosis , and reporting bias , as descriptions of diagnostic methodology and results were often incomplete .
acute human leptospirosis was reported in 46 eligible studies from 18 african countries ( fig 2 ) [ 31 – 76 ] . south africa was the most frequently represented country with a total of six articles [ 43 , 47 , 54 , 57 , 65 , 71 ] , followed by egypt [ 45 , 55 , 56 , 58 , 59 ] and kenya [ 31 , 37 – 39 , 42 ] with five included articles . twenty - one articles described acute human leptospirosis in hospital or health centre - based cohort studies ( table 3 ) . five articles described data from passive population - based surveillance [ 35 , 41 , 64 , 70 , 73 ] , and two articles described active case - finding in the setting of an outbreak of acute febrile illness [ 31 , 72 ] . non - specific febrile illness was the most common clinical criteria described for cohort or surveillance study inclusion . jaundice was stated as a primary inclusion criterion in three hospital - based cohort studies [ 44 , 61 , 66 ] . haemoglobinuria was stated as the only inclusion criterion in one study conducted in the democratic republic of the congo ( drc ) [ 40 ] .
summary of eligible cohort and surveillance studies reporting human acute leptospirosis in africa , 1930–2014 .
the majority of studies ( n = 41 / 46 ) used microscopic agglutination test ( mat ) as a primary method to diagnose human cases of acute leptospirosis . igm enzyme linked immunosorbent assay ( elisa ) testing was the only diagnostic method used in three studies [ 31 , 40 , 63 ] , but was more commonly used as part of a multi - faceted diagnostic approach ( n = 6 / 46 ) [ 44 , 45 , 58 , 64 , 68 , 73 ] . fifteen ( 32 6 % ) of 46 eligible human studies demonstrated leptospirosis infection by blood culture in combination with serological diagnostics [ 34 , 35 , 37 – 39 , 41 , 44 , 54 – 56 , 58 , 59 , 64 , 69 ] , and nine ( 19 5 % ) studies also used pcr detection as well as culture and serology [ 34 , 35 , 41 , 56 , 58 , 59 , 64 , 70 , 73 ] . genetic targets for diagnostic pcr assays included lbf1 , [ 34 , 35 ] lipl32 [ 34 , 35 ] , rrs [ 34 , 35 , 70 ] , and liga [ 58 , 59 ] . no culture - independent genetic typing of leptospira spp . was reported in any included human studies .
leptospirosis prevalence varied by study design and inclusion criteria ( table 3 ) . in hospital - based prospective cohort studies in mainland africa that enrolled patients with non - specific febrile illness and used mat serology for diagnosis of acute leptospirosis with or without adjunct diagnostics , prevalence ranged from 2 3 % to 19 8 % ( n = 11 ; number of patients : median = 166 ; range = 39–2441 ) [ 33 , 36 – 39 , 42 , 44 , 45 , 55 , 58 , 68 ] . a hospital - based prospective cohort study of febrile patients in mayotte that diagnosed acute leptospirosis by pcr and culture without serology reported a prevalence of 13 7 % ( number of patients = 2523 ) [ 34 ] . in hospital - based cohort studies that used jaundice as the main study enrolment criterion , prevalence of acute leptospirosis ranged from 2 0 % to 16 1 % ( n = 3 ; number of patients : median = 102 ; range = 99–392 ) [ 44 – 46 ] . acute leptospirosis was also reported in one patient ( 2 3 % ) of 38 with haemoglobinuria [ 40 ] , three patients ( 25 0 % ) of 12 involved in an outbreak of acute febrile disease in a pastoralist community in northern kenya [ 31 ] , and eight patients ( 9 8 % ) of 82 involved in an outbreak of acute pulmonary disease ( pneumonia ) in a mining camp in drc [ 72 ] .
incidence estimates were calculated in five population - based surveillance studies [ 35 , 41 , 64 , 70 , 73 ] and two hospital - based prospective cohort studies [ 63 , 74 ] . the only estimate of incidence from mainland africa came from northern tanzania , where regional incidence of 75 to 102 cases per 100 , 000 people per year was reported . this estimate was obtained by combining data on leptospirosis prevalence from hospital - based surveillance of febrile disease with multipliers derived from a population - based health - care utilisation survey [ 74 ] . for the indian ocean islands , incidence estimates were available for the seychelles where the average annual incidence was estimated as 60 to101 cases per 100 , 000 [ 63 , 70 ] ; réunion where the average annual incidence reported in three studies using a variety of data sources ranged from 3 1 to 12 0 cases per 100 , 000 [ 41 , 64 , 73 ] and mayotte , where the average annual incidence calculated from cases identified through four years of active hospital - based surveillance between 2007 and 2010 was reported as 25 cases per 100 , 000 [ 35 ] .
sixteen case reports describing acute leptospirosis in a total of 34 individuals were considered eligible for study inclusion . a wide range of clinical manifestations were reported including febrile illness , jaundice , meningitis , and acute respiratory distress syndrome . case reports described confirmed or probable acute leptospirosis in patients in south africa ( n = 6 ) [ 43 , 47 , 54 , 57 , 65 , 71 ] , gabon ( n = 3 ) [ 48 , 62 , 76 ] , morocco ( n = 3 ) [ 50 , 52 , 53 ] , algeria ( n = 1 ) [ 32 ] , mali ( n = 1 ) [ 51 ] , réunion ( n = 1 ) [ 75 ] , and senegal ( n = 1 ) [ 60 ] . with the exception of réunion and senegal , case reports were the only eligible data on acute human leptospirosis from these countries .
naturally occurring leptospira spp . infection in animal hosts was reported by 51 eligible citations describing studies performed in 17 african countries ( fig 2 ) [ 77 – 127 ] . south africa [ 84 , 100 , 101 , 104 , 117 , 120 – 122 ] and zimbabwe [ 83 , 93 – 99 ] were the most frequently represented countries with a total of eight included articles per country , followed by tanzania with seven articles [ 106 , 110 – 112 , 114 – 116 ] . wild animal surveys were most commonly described ( n = 21 / 51 ) followed by strain typing of leptospira spp . previously isolated from naturally infected animal hosts ( n = 13 / 51 ) , livestock disease outbreaks ( n = 7 / 51 ) and abattoir surveys ( n = 7 / 51 ) . four citations ( n = 4 / 51 ) reported human leptospirosis outbreaks as the inciting cause for investigations into animal carrier status [ 86 , 109 , 117 , 123 ] .
culture and isolation was the most common detection method for leptospira infection in animal studies ( n = 43 / 51 ) . pcr assays were used to demonstrate leptospira spp . infection in 13 ( 25 5 % ) out of 51 studies [ 85 , 86 , 92 , 103 , 105 , 107 , 115 , 118 , 120 , 123 – 126 ] . in three studies , culture and pcr were used in combination to determine infection status [ 92 , 115 , 118 ] . as with human studies , a variety of genetic targets were used in pcr assays to detect pathogenic leptospiral dna , including lipl32 / hap1 , [ 85 , 86 , 118 ] , secy , [ 103 ] , rrl [ 105 ] , and rrs [ 107 , 115 , 120 ] . pcr was predominantly used to demonstrate leptospira spp . infection in rodents and wild animal species . only one study in réunion also used pcr assays to demonstrate infection in domestic animals [ 85 ] .
leptospira infection prevalence varied widely by target animal species and diagnostic methodology ( s1 table ) . studies that used pcr diagnosis reported higher infection prevalence than studies that relied on leptospira culture and isolation . overall leptospira infection prevalence reported in black rats tested by pcr ranged from 11 0 % to 65 8 % ( n = 6 ; number of animals : median = 79 , range = 33–141 ) [ 85 , 86 , 92 , 103 , 118 , 124 ] . in two studies where black rats were tested by both pcr and culture , prevalence was higher by pcr ( 11 0 % , n = 100 ; and 28 . 7 % , n = 94 ) than by culture ( 4 0 % and 3 2 % ) in egypt [ 92 ] , and madagascar respectively [ 118 ] . a similar relationship was observed in brown rats , house mice and asian house shrews tested in madagascar [ 118 ] . cattle and brown rats were the most common species tested by culture . prevalence in brown rats ranged from 2 7 % to 8 5 % by culture ( n = 3 ; number of animals : median = 256 , range = 130–919 ) [ 82 , 108 , 117 ] but was considerably higher in three studies that used pcr to detect infection ( 10 0 % to 4 7 % ; number of animals : median = 11 , range = 10–96 ) [ 103 , 118 , 124 ] . in four abattoir - based surveillance studies of cattle from egypt , nigeria and zimbabwe [ 87 , 89 , 93 , 127 ] , renal leptospira spp . carrier status was detected by culture in 1 1 % to 10 4 % of sampled animals ( number of animals : median = 480 , range = 74–625 ) , compared to 18 2 % ( number of animals = 77 ) in a single pcr - based study from mayotte [ 85 ] .
this systematic review is the first to synthesize and compile data on the epidemiology of acute human leptospirosis and pathogenic leptospira spp . infection in animals in africa . leptospirosis remains amongst the neglected tropical diseases and is frequently overlooked in research priorities for african countries [ 1 ] . yet , through this systematic review we have revealed a wealth of scientific evidence for acute human infection demonstrating that acute leptospirosis is an important cause of febrile illness in hospital patients across the african continent . few studies providing population - level data on leptospirosis incidence in africa were identified but available estimates indicate that the disease incidence is high in both island and mainland populations . in reports of human disease and animal infection , three predominant species , leptospira borgpetersenii , l . interrogans and l . kirschneri , and a variety of leptospira serogroups were diagnosed . leptospira infection was reported in a wide range of domestic and wild animal species from across africa but studies linking data on animal infections with studies of acute human disease were rare .
acute leptospirosis was diagnosed in up to 19 8 % of inpatients with non - specific febrile illness in hospital - based cohort studies conducted in several countries identified by this review . in sub - saharan africa , recent studies have highlighted that clinical over - diagnosis of malaria may conceal other aetiologies of febrile illness [ 20 , 128 ] . consistent with findings in other resource - limited tropical settings ( e . g . south america [ 15 , 129 ] and south - east asia [ 130 – 132 ] ) , the evidence synthesised here demonstrates that acute leptospirosis infection is geographically widespread across the continent and should be considered as an important differential diagnosis for non - specific febrile illness in africa .
addressing the neglect of leptospirosis in africa will be a major challenging for the future of leptospirosis research . systematic review studies such as this can help to raise awareness of the human health threat of leptospirosis in africa among researchers and policy makers . for medical clinicians , the non - specific presenting signs of acute leptospirosis in patients poses a substantial diagnostic challenge in developing countries where laboratory capacity rarely exists to diagnose the infection [ 18 – 20 ] . hence , increasing clinician awareness and the development of treatment guidelines for the management of febrile patients should be a priority in resource - limited settings [ 138 ] . integration of risk factor analysis in human cohort studies of febrile disease is also strongly advocated and would be a valuable next step in identifying groups at high risk of infection , and defining important animal to human transmission routes .
knowledge of reservoir or carrier animal hosts is considered essential to understanding the epidemiology , transmission and control of leptospirosis in each setting [ 9 , 11 ] , yet our review has revealed that the linkages between leptospira infections in people and animals are rarely addressed in the existing literature . human and animal leptospira infections are inextricably linked , and the multi - host epidemiology of leptospirosis means that there may be many potential sources of infection in a given setting . in the future , greater emphasis should be placed on performing multidisciplinary human and animal leptospirosis studies in the same geographical settings . connecting investigations of animal reservoir populations with confirmed human cases would improve our understanding of the role that different animal species play in the transmission of pathogenic leptospira serovars in a variety of geographic and environmental settings [ 8 ] [ 139 ] . using an integrated ‘one health’ approach to explore the relationship between human and animal leptospira infection in areas where human disease is identified would also provide invaluable evidence to quantify the direct and indirect impacts of leptospirosis on human and animal populations in africa [ 140 , 141 ] .
control measures to prevent human leptospirosis often focus on rodent hosts of the disease . however , this review reveals that livestock are also important hosts of leptospira infection in africa , and may play a more substantial role in human disease transmission than is widely recognised . the clinical and sub - clinical productivity impacts of leptospira infection in domestic animal populations in africa are poorly understood . around the world , several leptospira serovars are considered to be of economic importance and cause production losses in a variety of livestock farming species including cattle , sheep , goats and pigs [ 17 , 100 , 142 , 143 ] . more than 300 million of the world’s poorest people live in africa , and at least 60 % of these are in some part dependent on livestock for their livelihood [ 144 ] . therefore , we consider that evaluating the impact of leptospira infection on livestock health and productivity is also an important priority for prospective research in africa . in the future , control of leptospira infection in livestock species may have considerable scope to directly and indirectly improve human health and well - being in africa , through reduced human leptospirosis transmission and increased productivity in livestock that subsistence farming communities [ 8 , 142 , 143 , 145 ] .