texella cokendolpheri is a rare species of arachnid known by the common name cokendolpher cave harvestman .

it may also be called the robber baron cave harvestman .

it is endemic to texas in the united states , where it is only known from a single cave in bexar county .

it is a federally listed endangered species of the united states .

this is " a small eyeless or essentially eyeless " harvestman .

it is pale orange in color .

it is a troglobite that lives its whole life in a subterranean karst cave .

it is known only from robber baron cave in bexar county , a cave which runs underneath a heavily urbanized area .

the cave is owned by the texas cave management association .

this cave is also the only home for the robber baron cave meshweaver ( cicurina baronia ) , a spider .

the worst threat to this and other rare local troglobites is the outright loss of their cave habitat , which is destroyed as the land is consumed for urban development , or during quarrying operations .

caves are also altered and polluted so that they can not support this and other species .

texella cokendolpheri

beetles : rhadine exilis / rhadine infernalis / batrisodes venyivi harvestman : texella cokendolpheri spiders : cicurina baronia / c . madla / c . venii / c . vespera / neoleptoneta microps
texella mulaiki c . j . goodnight and m . l . goodnight , 1942
texella reddelli c . j . goodnight and m . l . goodnight , 1967 – bee creek cave harvestman
the harvestman family phalangodidae . 5 . new records and species of texella goodnight and goodnight ( opiliones : laniatores )
ubick , d . and t . s . briggs . 1992 . the harvestman family phalangodidae . 3 . revision of texella goodnight and goodnight ( opiliones : laniatores ) . texas mem . mus . , speleol . monogr . 3 : 155 - 240 .
ubick , d . and t . s . briggs . 2004 . the harvestman family phalangodidae . 5 . new records and species of texella goodnight and goodnight ( opiliones : laniatores ) . texas memorial museum . speleological monographs , 6 : 101 - 141 .
rhadine exilis and rhadine infernalis were first collected in 1959 and described by barr and lawrence ( 1960 ) as agonum exile and agonum infernale , respectively . barr ( 1974 ) assigned the species to the genus rhadine . batrisodes venyivi was first collected in 1984 and described by chandler ( 1992 ) . texella cokendolpheri was first collected in 1982 and described in ubick and briggs ( 1992 ) . cicurina baronia , cicurina madla , cicurina venii , and cicurina vespera were first collected in 1969 , 1963 , 1980 , and 1965 , respectively . in 1992 , gertsch described these species . neoleptoneta microps was first collected in 1965 and described by gertsch ( 1974 ) as leptoneta microps . the species was reassigned to neoleptoneta following brignoli ( 1977 ) and platnick ( 1986 ) .
summary : we , the u . s . fish and wildlife service ( service ) , determine nine cave - dwelling invertebrates from bexar county , texas , to be endangered species under the authority of the endangered species act of 1973 , as amended ( act ) . rhadine exilis ( no common name ) and rhadine infernalis ( no common name ) are small , essentially eyeless ground beetles . batrisodes venyivi ( helotes mold beetle ) is a small , eyeless beetle . texella cokendolpheri ( robber baron cave harvestman ) is a small , eyeless harvestman ( daddy - longlegs ) . cicurina baronia ( robber baron cave spider ) , cicurina madla ( madla ' s cave spider ) , cicurina venii ( no common name ) , cicurina vespera ( vesper cave spider ) , and neoleptoneta microps ( government canyon cave spider ) are all small , eyeless or essentially eyeless spiders .
known from a single locality , robber baron cave in bexar county , texas . although the cave entrance is protected as a preserve by the texas cave management association ( tcma ) , this cave is relatively large , and the land that overlies the cave is heavily urbanized . the cave has also been historically subject to extensive commercial and recreational use ( veni 1988 ) . in addition , the current gate on the cave is blocking organic material from entering and may be severely limiting food input and impacting the cave fauna ( veni and reddell , pers . comm . 2001 in cockendolpher and reddell , 2004 ) . while no regular biomonitoring occurs in robber baron cave , there are no records of specimens of t . cokendolpheri collected since october 1993 ( usfws 2008 ) .
html public
- / / w3c / / dtd xhtml 1 . 0 strict / / en
urltoken
this species has been referred to by two common names , the robber baron cave harvestman ( usfws 2000 ) and the cokendolpher cave harvestman ( breene et al . 2003 ) . the latter name has been accepted as the official common name ( breene et al . 2003 , usfws 2003 ) .
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
there are nine bexar county , texas invertebrates that were listed as endangered on december 26 , 2000 . the recovery priority number for all bexar county karst invertebrates is 2c , which means that these species face a high degree of threat with a high potential for recovery and there may be conflict between species recovery and economic development .
known only from robber baron cave in the alamo heights karst region , bexar county , texas .
population estimates are unavailable for any of the nine troglobites listed as endangered in bexar county ( usfws , 2000 ) due to lack of adequate techniques , their cryptic behavior , and inaccessibility of habitat ( usfws , 2008 ) . culver et al . ( 2000 ) states that while some troglobites are known from a few specimens , detailed studies suggest that
as a rule
most troglobites
are not numerically rare and thus are not susceptible to the problems of small populations .
however , considering the lack of population estimates and limited study of these species , data are insufficient to indicate whether bexar county karst invertebrates are numerous enough to rule out small population concerns ( usfws , 2008 ) .
based on usfws ( 2008 ) , reduce threats to the species by securing an adequate quantity and quality of caves , including selecting caves or cave clusters that represent the range of the species and potential genetic diversity , then preserving these caves , including their drainage basins and surface communities upon which they rely . maintenance of these cave preserves involves keeping them free from contamination , excessive human visitation , and non - native fire ants by regularly tracking progress and implementing adaptive management to control these and any new threats when necessary . monitoring the population status and threats are also components of recovery . because many aspects of the population dynamics and habitat requirements of the species are poorly understood , recovery is also dependant on incorporating research findings into adaptive management actions .
( < 100 square km ( less than about 40 square miles ) ) known only from robber baron cave in the alamo heights karst region , bexar county , texas .
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 ) .
culver , d . c . , l . l . master , m . c . christman , and h . h . hobbs iii . 2000 . obligate cave fauna of the 48 contiguous united states . conservation biology 14 ( 2 ) : 386 - 401 .
elliott , w . r . 2000 . conservation of the north american cave and karst biota . pages 665 - 689 in wilkens , h . , d . c . culver , and w . f . humphreys ( editors ) . subterranean ecosystems . ecosystems of the world , 30 . elsevier , amsterdam . xiv + 791 pp . corrected version online . available : urltoken
longacre , c . 2000 . endangered and threatened wildlife and plants ; final rule to list nine bexar county , texas invertebrate species as endangered . u . s . fish and wildlife service ( usfws ) . federal register 65 ( 248 ) .
rappaport , c . j . 1998 . department of interior fish and wildlife service 50 cfr part 17 , rin 1018 - af33 , endangered and threatened wildlife and plants ; proposal to list nine bexar county , texas invertebrate species as endangered . federal register , 63 ( 250 ) : 71855 - 71867 .
stanford , r . , and a . shull . 1993 . department of interior fish and wildlife service 50 cfr part 17 ; 90 - day finding on a petition to list nine bexar county , tx , invertebrates . federal register , 58 ( 229 ) : 63328 - 63329 .
u . s . fish and wildlife service ( usfws ) . 1994 . endangered and threatened wildlife and plants ; animal candidate review for listing as endangered or threatened species . federal register 59 ( 219 ) : 58982 - 59028 .
u . s . fish and wildlife service ( usfws ) . 2003 . endangered and threatened wildlife and plants ; designation of critical habitat for seven bexar county , texas , invertebrate species ; final rule . federal register 68 ( 67 ) : 17156 - 17231 .
u . s . fish and wildlife service ( usfws ) . 2008 . draft bexar county karst invertebrate recovery plan . 125 pp .
the small print : trademark , copyright , citation guidelines , restrictions on use , and information disclaimer .
note : all species and ecological community data presented in natureserve explorer at urltoken were updated to be current with natureserve ' s central databases as of november 2016 . note : this report was printed on
trademark notice :
natureserve
, natureserve explorer , the natureserve logo , and all other names of natureserve programs referenced herein are trademarks of natureserve . any other product or company names mentioned herein are the trademarks of their respective owners .
copyright notice : copyright © 2017 natureserve , 4600 n . fairfax dr . , 7th floor , arlington virginia 22203 , u . s . a . all rights reserved . each document delivered from this server or web site may contain other proprietary notices and copyright information relating to that document . the following citation should be used in any published materials which reference the web site .
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 .
data provided by natureserve in collaboration with bruce patterson , wes sechrest , marcelo tognelli , gerardo ceballos , the nature conservancy - migratory bird program , conservation international - cabs , world wildlife fund - us , and environment canada - wildspace .
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 .
no graphics available from this server can be used , copied or distributed separate from the accompanying text . any rights not expressly granted herein are reserved by natureserve . nothing contained herein shall be construed as conferring by implication , estoppel , or otherwise any license or right under any trademark of natureserve . no trademark owned by natureserve may be used in advertising or promotion pertaining to the distribution of documents delivered from this server without specific advance permission from natureserve . except as expressly provided above , nothing contained herein shall be construed as conferring any license or right under any natureserve copyright .
) . your comments will be very valuable in improving the overall quality of our databases for the benefit of all users .
designation of critical habitat for nine bexar county , tx , invertebrates final rule . and a 12 - month finding on a petition to revise critical habitat designation by removing unit 13 from designation under the act - not warranted .
designation of critical habitat for nine bexar county invertebrates : proposed rule ; reopening of comment period .
- - natureserve explorer is a source for authoritative conservation information on more than 50 , 000 plants , animals and ecological communtities of the u . s and canada . natureserve explorer provides in - depth information on rare and endangered species , but includes common plants and animals too . natureserve explorer is a product of natureserve in collaboration with the natural heritage network .
fws digital media library - - the u . s . fish and wildlife service ' s national digital library is a searchable collection of selected images , historical artifacts , audio clips , publications , and video .
data and apps idaho fish and wildlife information system bringing information to bear on the management and conservation of fish , wildlife , and plants in idaho .
popular : trivia , history , america , television , tv , usa , geography , cities , . . . more
html public
- / / w3c / / dtd html 4 . 01 transitional / / en
urltoken
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 .
html public
- / / w3c / / dtd html 4 . 01 transitional / / en
action : final rule . - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
these species ( referred to in this final rule as the nine invertebrates ) are known from karst topography ( limestone formations containing caves , sinks , fractures and fissures ) in north and northwest bexar county . threats to the species and their habitat include destruction and / or deterioration of habitat by construction ; filling of caves and karst features and loss of permeable cover ; contamination from septic effluent , sewer leaks , run - off , pesticides , and other sources ; predation by and competition with nonnative fire ants ; and vandalism . this action will implement federal protection provided by the act for these species . we based our decision on the best available information , including that received during public comment on the proposal to list these species .
effective date : the effective date of this rule is december 26 , 2000 .
addresses : the complete file for this rule is available for inspection , by appointment , during normal business hours at the austin ecological services field office , 10711 burnet road , suite 200 , austin , texas 78758 .
for further information contact : alisa shull , supervisory fish and wildlife biologist , austin ecological services field office ( telephone 512 / 490 - 0057 ; facsimile 512 / 490 - 0974 ) .
these nine invertebrates are obligate ( capable of surviving in only one environment ) karst or cave - dwelling species ( troglobites ) of local distribution in karst terrain in bexar county , texas . ` ` karst ' ' is a type of terrain in which the rock is dissolved by water so that much of the drainage occurs into the subsurface rather than as runoff . the subsurface drainage leads to passages or other openings within the underground rock formations . some of the features that develop in karst areas include cave openings , holes in rocks , cracks , fissures , and sinkholes .
habitat required by the nine karst invertebrate species consists of underground , honeycomb limestone that maintains high humidity and stable temperatures . the surface environment of karst areas is also an integral part of the habitat needed by the animals inhabiting the underground areas . openings to the surface allow energy and nutrients , in the form of leaf litter , surface insects , other animals , and animal droppings to enter the underground ecosystem . mammal feces provide a medium for the growth of fungi and , subsequently , localized population blooms of several species of tiny , hopping insects . these insects reproduce rapidly on rich food sources and may become prey for some predatory cave invertebrates ( service 1994 ) . while the life habits of the nine invertebrates are not well known , the species probably prey on the eggs , larvae , or adults of other cave invertebrates .
we funded a status survey ( veni 1994a ; reddell 1993 ) of all nine species through a grant to the texas parks and wildlife department ( tpwd ) under section 6 of the act . researchers obtained landowner permission to study and assess threats to 41 caves in north and northwest bexar county , texas . landowners denied permission to access an additional 36 caves that biologists believed likely to contain species of concern . researchers described all 77 caves , to some extent , before the status survey was conducted and some were already known to contain at least one of the nine invertebrates .
during the status survey , the researchers made a collection of the invertebrate fauna at each cave studied , assessed the condition of the cave environment and threats to the species , and collected geological data . they used this information to prepare two reports . one report discusses the overall karst geography in the san antonio region and the potential geologic and geographic barriers to karst invertebrate migration ( on an evolutionary time scale ) and limits to their distribution ( veni 1994a ) . the other report ( reddell 1993 ) details the fauna of each cave visited during the study and presents information obtained from invertebrate collections .
veni ' s ( 1994a ) report delineates six karst areas ( hereafter referred to as karst regions ) within bexar county . the karst regions he discusses are stone oak , utsa ( university of texas at san antonio ) , helotes , government canyon , culebra anticline , and alamo heights . the boundaries of these karst regions are geological or geographical features that may represent obstructions to troglobite movement ( on a geologic time scale ) which has resulted in the present - day distribution of endemic ( restricted in distribution ) karst invertebrates in the san antonio region .
batrisodes venyivi , the helotes mold beetle , is known from only three caves in the vicinity of helotes , texas , northwest of san antonio . two of these caves are located in the helotes karst region on private property . we do not have reliable information on the collection from the third cave . the collector of the specimen declined to give us a specific site collection record , but we believe it is located on private property .
rhadine exilis is known from 35 caves in north and northwest bexar county . twenty - one are located on department of defense ( dod ) land in the stone oak karst region . the remainder are distributed among the helotes , utsa , and stone oak karst regions , while one location lies in the government canyon region . one of the non - dod sites is located in a county road right - of - way , one is located in a state - owned natural area , and the remainder are located on private property . ongoing efforts by the dod to locate and inventory karst features on camp bullis and to document the karst fauna communities in caves on camp bullis resulted in discovery of 18 of the 35 caves mentioned above ( veni 1994b ; james reddell , pers . comm . 1997 ) .
rhadine infernalis is known from 25 caves . this species occurs in five of the six karst regions - - helotes , utsa , stone oak , culebra anticline , and government canyon . scientists have delineated three subspecies ( rhadine infernalis ewersi , rhadine infernalis infernalis , rhadine infernalis ssp . ) , and described and named two of these in scientific literature ( barr 1960 , barr and lawrence 1960 ) . in a recent report , scientists characterized the third subspecies as distinct , but not named ( reddell 1998 ) . only three caves , all on dod land , contain the subspecies rhadine infernalis ewersi . sixteen caves contain the subspecies rhadine infernalis infernalis and lie in the government canyon , helotes , utsa , and stone oak regions . six caves in the culebra anticline region contain the unnamed subspecies .
cicurina venii is known from only one cave , which is located on private property in the culebra anticline karst region . the species was collected in 1980 and 1983 , but the cave itself was not initially described until 1988 ( reddell 1993 ) . the cave entrance was filled during construction of a home in 1990 . without excavation , it is difficult to determine what effect this incident had on the species ; however , there may still be some nutrient input , from a reported small side passage .
cicurina baronia , the robber baron cave spider , is known only from robber baron cave in the alamo heights karst region . although the cave entrance is owned and operated by the texas cave management association , it is located in a heavily urbanized area .
cicurina madla , madla ' s cave spider , is known from six caves . one cave is within the government canyon karst region in government canyon state natural area , one is on dod land , three are located in the helotes karst region on private property , and one is located on private property in the utsa karst region .
biologists have found cicurina vespera , the vesper cave spider , in two caves . one cave is government canyon bat cave in the government canyon state natural area , and the other is a cave 5 miles northeast of helotes . the location and name of this latter cave have not been revealed to us , but we believe it is located on private property .
neoleptoneta microps is known only from the government canyon karst region , from two caves within government canyon state natural area .
in the course of conducting the 1993 status survey , veni contacted landowners and requested access to as many caves as possible that were believed to be potential habitat for the nine invertebrates . it is possible that these species occur in some of the caves that could not be visited and that new locations of the nine invertebrates will be discovered in the future . although these new discoveries may increase the number of locations where the species are found , they are expected to fall within the same general range and are expected to face the same threats as the known occurrences of these species . the listing of these species is not based on a demonstrable decline in the number of individuals or the number of known locations of each species , but rather on reliable evidence that each species is subject to threats to its continued existence throughout all or a significant portion of its range .


the campbell 's mona monkey , also known as campbell 's guenon and campbell 's monkey , ( cercopithecus campbelli ) , is a species of primate in the family cercopithecidae found in the ivory coast , gambia , ghana , guinea , guinea-bissau , liberia , senegal , and sierra leone .

it was named for henry dundas campbell , in 1838 .

lowe 's mona monkey was previously considered a subspecies of campbell 's mona monkey .

the international union for conservation of nature has rated this species as being of " least concern " because it has a wide range and is able to adapt to degraded habitats .

campbell ' s mona monkey

lowe ' s mona monkey was previously considered a subspecies of campbell ' s mona monkey .
wolf ' s mona monkey was previously considered a subspecies of the crested mona monkey .
campbell ' s mona monkeys , boabeng - fiema monkey sanctuary . # 2 - youtube
campbell ' s mona monkeys , boabeng - fiema monkey sanctuary . vitts in ghana 2010
more campbell ' s mona monkeys , boabeng - fiema monkey sanctuary . vitt family , ghana 2010
wolf ' s mona monkey - cercopithecus wolfi a . meyer , 1891 - details - encyclopedia of life
and arnold et al . ( population differences in wild campbell’s monkeys alarm call use ,
keenan , s . , lemasson , a . , & zuberbu¨hler , k . ( 2013 ) . graded or discrete ? a quantitative analysis of campbell’s monkey alarm calls .
wolf ' s mona monkey is known to associate with several guenon and non - guenon species such as the black crested mangabey , the red - tailed monkey , the angola colobus , allen ' s swamp monkey , and the bonobo . no viable offspring or interspecific mating occurs during its associations with other primates .
when forming associations with other primates it is necessary that there is a difference in diet or feeding height between the species to reduce competition . when in a mixed group , wolf ' s mona monkey will move and forage at a mean height of 17 metres . wolf ' s mona monkey is mainly found in association with the red - tailed monkey ( which forages at 12 m ) and the black crested mangabey ( which forages at 21 . 5 m ) , two species with similar diets to wolf ' s mona monkey . these mixed groups most likely form for predator detection .
spectrographic illustrations of the different loud call types produced by male campbell ' s monkeys in different contexts .
lemasson a , gautier jp , hausberger m ( 2003 ) vocal similarities and social bonds in campbell ' s monkey ( cercopithecus campbelli ) . comptes rendus biologies 326 : 1185–1193 .
wolf ' s mona monkey is also sexually dimorphic in size . males weigh , on average , almost twice as much as females ( 4 . 5 kg and 2 . 5 kg respectively ) .
basic acoustic measurements of the stem of the six different loud calls produced by adult male campbell ' s monkeys .
the wolf ' s mona monkey ( cercopithecus wolfi ) , also called wolf ' s guenon , is a colorful old world monkey in the cercopithecidae family . it is found in central africa , primarily between the democratic republic of the congo and uganda . it lives in primary and secondary lowland rainforest and swamp forest .
zuberbühler k ( 2001 ) predator - specific alarm calls in campbell ' s guenons . behavioral ecology and sociobiology 50 : 414–422 .
campbell ' s monkeys appear to combine the same calls in different ways , using rules of grammar that turn sound into language .
kamara , t . 2001 . saddam ' s oil and taylor ' s timber . urltoken
is most commonly found in the democratic republic of congo and areas in uganda . there are three subspecies of wolf’s monkey :
lemasson , a . , ouattara , k . , bouchet , h . , & zuberbühler , k . ( 2010 ) . speed of call delivery is related to context and caller identity in campbell’s monkey males .
wolters s , zuberbühler k ( 2003 ) mixed - species associations of diana and campbell ' s monkeys : the costs and benefits of a forest phenomenon . behaviour 140 : 371–385 .
, are very colorful . their color is used in intraspecific communication for recognizing individuals , species , and potential mates . wolf ' s mona monkey is dark grey with a red
saddle
on its back . the pelage depends on the subspecies .
lemasson , a . , zuberbühler , k . , & hausberger , m . ( 2005 ) . socially meaningful vocal plasticity in campbell’s monkeys .
ouattara , k . , lemasson , a . & zuberbühler , k . ( 2009a ) . campbell’s monkeys use affixation to alter call meaning .
zuberbühler , k . ( 2002 ) . a syntactic rule in forest monkey communication .
ouattara k , lemasson a , zuberbühler k ( 2009 ) the alarm call system of female campbell ' s monkeys . anim behav 78 : 35–44 .
kuhn . j . ( 2013 ) . do campbell’s monkeys have linguistic morphology ? seminar paper , nyu . available online march 10 , 2014 , from
the diet of wolf ' s mona monkey differs depending on location . although predominantly a frugivore , it may also forage for seeds and insects for protein . since it has no adaptations for leaf eating , its leaf diet mainly consists of young and easily digestible leaves .
lemasson , who is further investigating campbell ' s monkey talk by measuring their reactions to recorded calls , suspects that a dense jungle environment drove the evolution of syntax . since the monkeys had trouble seeing each other , they compensated by talking .
ouattara , k . , lemasson , a . , & zuberbühler , k . ( 2009b ) . campbell’s monkeys concatenate vocalizations into context - specific call sequences .
arnold , k . , keenan , s . , lemasson , a . , & zuberbühler , k . ( 2013 ) . population differences in wild campbell’s monkeys alarm call use . manuscript , university of st andrews .
) . when a mixed group was involved in the association , it always lasted for over an hour . interactions occurred once every seven hours . associations mainly occurred while the bonobos were feeding or resting . wolf ' s mona monkey was found to feed in the trees while the bonobo fed or rested .
arnold k , pohlner y , zuberbühler k ( 2008 ) a forest monkey ' s alarm call series to predator models . behav ecol sociobiol 62 : 549–559 .
lemasson ' s analysis was based on a vast set of recordings , gathered from 10 monkey groups observed for two full years in their african rain forest homes .
zuberbühler k ( 2002 ) a syntactic rule in forest monkey communication . anim behav 63 : 293–299 .
lemasson a , hausberger m , zuberbühler k ( 2005 ) socially meaningful vocal plasticity in adult campbell ' s monkeys ( cercopithecus campbelli ) . j comp psychol 119 : 220–229 .
among cercopithecines , forest guenons such as wolf ' s mona monkey have very developed cheek pouches . these cheek pouches are second only to macaques . the evolution of these cheek pouches in both genera may be a response to the increased potential for interspecific competition in the mixed - species associations which these monkeys frequently form .
arnold , k . , pohlner , y . , & zuberbühler , k . ( 2008 ) . a forest monkey ' s alarm call series to predator models .
citation : ouattara k , lemasson a , zuberbühler k ( 2009 ) campbell ' s monkeys use affixation to alter call meaning . plos one 4 ( 11 ) : e7808 . urltoken
there is little hard data on threatened and declining species in the gambia . there has been a considerable decline in the diversity of large mammalian species , which commenced during the latter part of the nineteenth century . from the species in the gambia in the late 1960s , it was estimated that of the 67 species of mammals listed , 13 had become extinct and a similar number were threatened with extinction ( dpwm 1991 ) . a number of species of mammals still migrate into the gambia including roan antelope ( hippotragus equinus ) and campbell’s mona monkey ( cercopitheocus mona campbell ) , wild dogs ( lycoon pitus ) and lio ( panthera leo ) still infrequently enter the eastern end of the gambia , invariably as vagrants from niokolokoba national park in senegal .
according to lemasson and to jared taglialatela , a chimpanzee communication researcher at clayton state university , it ' s too soon to say whether the monkey talk is proto - human .
lemasson a , hausberger m ( 2004 ) patterns of vocal sharing and social dynamics in a captive group of campbell ' s monkeys ( cercopithecus campbelli campbelli ) . j comp psychol 118 : 347–359 .
lemasson a , blois - heulin c , jubin r , hausberger m ( 2006 ) female social relationships in a captive group of campbell ' s monkeys . american journal of primatology 68 : 1161–1170 .
ouattara , k . , zuberbühler , k . , n’goran , e . k . , gombert , j . - e . , & lemasson , a . ( 2009c ) . the alarm call system of female campbell’s monkeys .
citation :
generating meaning with finite means in campbell’s monkeys .
by karim ouattara , alban lemasson , and klaus zuberbuhler . proceedings of the national academy of sciences , vol . 106 no . 48 , december 7 , 2009 .
primates , this is not the case with the bonobo . no aggressive interactions occurred during the study period . the red - tailed monkey (
karim ouattara , alban lemasson , klaus zuberbühler ( 2009 ) ,
campbell ' s monkeys use affixation to alter call meaning
, plos one 4 ( 11 ) : e7808 , doi : 10 . 1371 / journal . pone . 0007808
here , we studied the alarm calling behavior of free - ranging campbell ' s monkey males in natural and experimental predator situations . we were particularly interested in how males concatenated their repertoire of six call types into call sequences . to this end , we sought to describe the principles governing the organization of sequences in terms of composition and call order and how the different sequences related to external events .
karim ouattaraa , alban lemassona , and klaus zuberbühler ( december 22 , 2009 ) ,
campbell ' s monkeys concatenate vocalizations into context - specific call sequences
, pnas 106 ( 51 ) : 22026–22031 , doi : 10 . 1073 / pnas . 0908118106
riede t , zuberbühler k ( 2003 ) pulse register phonation in diana monkey alarm calls . journal of the acoustical society of america 113 : 2919–2926 .
meyer , ab ( 1894 ) .
remarks on an african monkey , cercopithecus wolfi
. proceedings of the zoological society of london : 83–84 .
in one study , wolf ' s mona monkeys were found associating with bonobos within 20 metres for an average time of 20 minutes ( although sometimes for over an hour ) . these interactions were mainly initiated by , and departed by , the guenons ; this indicates that the guenons most benefited from these associations . although the
the birth season for wolf ' s mona monkey is from june through december due to rainfall and resource availability . it lives in a single male / multi - female group . it is female philopatric , with males dispersing from the group at sexual maturity . because one male controls several females there is extreme competition for the alpha male position . females , on the other hand , are generally amicable and participate in grooming and allomothering . unlike macaques there are no strong linear dominance hierarchies .
zuberbühler k , noë r , seyfarth rm ( 1997 ) diana monkey long - distance calls : messages for conspecifics and predators . anim behav 53 : 589–604 .
the degree to which the communication system found in campbell ' s monkey is unique or a general feature of primate communication is currently unknown . we suspect the latter , especially for forest primates whose vocal skills must have been under considerable pressure by natural selection due to high levels of predation and low levels of visibility in their dim habitats . the lack of articulatory control , which characterizes nonhuman primates , may have favored the evolution of combinatorial signaling .
thompson , h . s . s . 1993 . status of white - necked picathartes – another reason for the conservation of the peninsula forest , sierra leone . oryx 27 : 155 - 158 .
oates , j . f . , m . abedi - lartey , w . s . mcgraw , t . t . struhsaker and g . h . whitesides . 2000 . extinction of a west african red colobus monkey . conservation biology 14 : 1526 - 1532 .
“wak - oo” calls are given to the same events as “hok - oo” calls ( eagles , other flying animals , diana monkey eagle alarms ) , but not to neighbours .
riede t , zuberbühler k ( 2003 ) the relationship between acoustic structure and semantic information in diana monkey alarm vocalization . journal of the acoustical society of america 114 : 1132–1142 .
though some researchers have ascribed syntax to animals , it ' s never been formally demonstrated – until now .
worldcat is the world ' s largest library catalog , helping you find library materials online . learn more ››
anstey , s . 1991 . wildlife utilization in liberia . wwf and liberian forestry development authority , uk .
comments : c . mona species group . mcallan and bruce ( 1989 ) argued that the original publication of this species should be : the analyst , 24 : 298 - 299 [ publ . 2 july 1838 ] . does not include lowei : see kingdon ( 1997 )
zuberbühler , k . , noë , r . r . , & seyfarth , r . m . ( 1997 ) . diana monkey long - distance calls : messages for conspecifics and predators .
seyfarth , r . m . , cheney , d . l . , & marler , p . ( 1980b ) . vervet monkey alarm calls : semantic communication in a free - ranging primate .
université de rennes 1 , laboratoire d’éthologie animale et humaine , umr 6552 – c . n . r . s .
wolf ' s monkeys occasionally raid local agricultural crops and have a potential for carrying diseases that can be contagious to humans .
seyfarth , r . m . , cheney , d . l . , & marler , p . ( 1980a ) . monkey responses to three different alarm calls : evidence of predator classification and semantic communication .
the study was conducted in the taï national park ( 5°50′n , 7°21w ) , ivory cost , the largest remaining block of intact rainforest in west africa . data were collected between january 2006 and september 2007 on two groups of campbell ' s monkeys ( cercopithecus c . campbelli ) that were fully habituated to the presence of human observers . campbell ' s monkeys routinely form polyspecific groups with other primates , particularly diana monkeys , which whom they spend 77–89 % of their time during feeding , travelling and resting [ 29 ] . campbell ' s monkeys live in small one - male groups with 3–7 adult females with their offspring [ 23 ] . the two study groups have been followed on a regular basis since the early 1990s and all individuals can be recognised individually . we had additional access to four other groups that were partly habituated to human observers . during the study period , we observed one replacement of the single adult male in one habituated group . the new male became quickly habituated to human observers , which effectively increased the sample size of habituated individuals to n = 3 .
wolf ' s monkeys are probably important in seed dispersal of food trees and they may contribute to pollination when they drink nectar .
nowak , r . 1999 . walker ' s primates of the world . baltimore , maryland : the johns hopkins university press .
the third sequence also consisted of a pair of boom calls followed by a k + sequence ( overall median = 12 calls ; range : 9–16 calls ; n = 76 ) , but here the sequence was interspersed with 1–7 h + calls ( fig . 2 ) . this combined sequence was recorded from all four habituated males and two semihabituated groups and always in response to neighboring campbell ' s monkey groups or single stranger males , suggesting that it functioned in territorial defense . to further test this hypothesis , we investigated whether the location of call production varied in systematic ways . we predicted that sequences given in the periphery of a group ' s home range contained significantly more h + calls than sequences given in the center , which turned out to be the case ( fig . 3 ; fisher ' s exact test , p < 0 . 001 ) .
savill , p . s . and j . e . d . fox . 1967 . trees of sierra leone . government printers .
in earlier work , we have shown that campbell ' s monkey males use an affixation rule to increase the size of their call repertoire . adding the suffix “oo” to krak ( k → k + ) or hok ( h → h + ) calls altered these calls ' meanings in predictable ways ( 50 ) . here , we describe regularities at another level , i . e . , in how monkeys combined this repertoire of six call types into nine distinct call sequences ( fig . 2 ) . these call combinations were not random , but the product of a number of principles , which governed how semantic content was obtained .
we develop a formal semantic analysis of the alarm calls used by campbell’s monkeys in the tai forest ( ivory coast ) and on tiwai island ( sierra leone ) —two sites that differ in the main predators that the monkeys are exposed to ( eagles on tiwai vs . eagles and leopards in tai ) . building on data discussed in ouattara et al . ( plos one 4 ( 11 ) : e7808 ,
mulavwa , m . 1991 . notes on the call of mona monkeys ( cercopithecus wolfi ) in the mabali forest : frequency of emission and daily activities . pp . 1 in a ehara , t kimura , o takenaka , m iwamoto , eds . primatology today proceedings of the xiii congress of the international primatological society . amsterdam : elsevier science publishers .
a uniquely valuable compendium of taxonomic and distributional data on the world ' s living and historically extinct mammalian species . contributors and editors alike deserve the thanks of all
davies , g . and b . birkenhager . 1990 . jentink’s duiker in sierra leone : evidence from the freetown peninsula . oryx 24 : 143 - 146 .
mayers , j . , s . anstey and a . peal . 1992 . liberia . pages 214 - 220 in j . a . sayer , c . s . harcourt , and n . m . collins , editors . the conservation atlas of tropical forests : africa . iucn and macmillan publishers , united kingdom .
schel am , tranquilli s , zuberbühler k ( 2009 ) the alarm call system of black - and - white colobus monkeys . j comp psychol 123 : 136–150 .
* brandon keim ' s twitter stream and reportorial outtakes ; wired science on twitter . brandon is currently working on a book about ecosystem and planetary tipping points . *
papworth s , böse as , barker j , zuberbühler k ( 2008 ) male blue monkeys alarm call in response to danger experienced by others . biology letters 4 : 472–475 .
like other primates , wolf ' s monkeys also extensively use grooming for tactile social communication . the use of chemical cues , such as pheromones , is likely , but undocumented .
are these results relevant for understanding the evolution of syntax ? as outlined before , this system pales in contrast to the communicative power of grammar ; using calls in different sequences , even highly predictable sequences , is not the same as grammar . nevertheless , the call sequences observed in this primate were not the results of individuals responding to serial or compound stimuli ( such as “water” and “bird” combined to “waterbird” ) . in campbell ' s monkeys , the different sequences were given to highly discrete external events with very little conceptual similarities .
we carried out long - term observations and predator model experiments to investigate how free - ranging male campbell ' s monkeys of taï national park , ivory coast , communicated about external events . in previous research , we found that males and females produced different alarm calls that , in some cases , were combined into meaningful sequences [ 23 ] , [ 31 ] - [ 34 ] . here , we were interested in how acoustically flexible males were with some of their alarm call types , and how they applied this variation to external events . our study showed that male campbell ' s monkeys produced six different loud alarm calls in response to disturbing or dangerous events . “boom” calls were acoustically and contextually unique , whereas the other five calls shared a number of acoustic features . the most relevant finding was that these five calls consisted of a call stem that differed in terms of the basic frequency contours and could be followed by an optional suffix - like small and inconspicuous vocal unit , which altered the semantic content of the full call in significant and predictable ways .
hohmann and surbeck published in 2008 that bonobos sometimes do hunt monkey species . having observed a group of bonobos in salonga national park for five years they witnessed five incidents where bonobos preyed on groups of monkeys . their research indicates it was deliberate hunting , where a group of bonobos would coordinate their actions—contrary to their normal hunting behaviour , which is quite solitary and less purposeful . in three occasions the hunt was successful and infant monkeys were captured , once a redtail monkey and twice a cercopithecus wolfi . the spoils , however , were distributed quite peacefully among the members of the group . [ 17 ] [ 18 ]
laboratoire ethos “ethologie animale et humaine” , u . m . r . 6552 - c . n . r . s . , université de rennes 1 , station biologique , paimpont , france
wolf ' s monkeys are frugivorous , but they supplement their diet heavily with leaves , seeds , and flowers . at salonga national park in the democratic republic of congo , wolf ' s monkeys have been recorded consuming 32 % fruit ( 4 % fleshy and 27 % arils ) , 27 % seeds , 29 % leaves , and 11 % flowers . though not a primary means of sustenance , wolf’s monkeys will occasionally feed on nectars and insects if they are readily available . the principal feeding time for this species is during the early morning and early afternoon .
call sequences to leopards were always composed of krak ( k ) calls , sometimes combined with krak - oo ( k + ) calls ( median = 21 calls ; range : 12–35 ; n = 26 ; fig . 2 ) . in the sequences recorded from the three most habituated males , we found that the level of urgency was associated with a high proportion of k calls in the sequence . sequences with just k calls were given in response to real leopards and leopard models . k + calls were far more common when callers responded to leopard vocalizations or diana monkey leopard alarm calls . compared to real leopards and leopard models , k calls were given significantly less often to leopard vocalizations and diana monkey leopard alarm calls ( fisher ' s exact tests , p < 0 . 001 ; fig . 4 b ) .
when comparing human language to primate communication systems , such as this one , a number of interesting similarities and fundamental differences emerge . first , male campbell ' s monkeys are limited to a relatively small range of messages that they can convey to their audience . this is partly because callers do not take full advantage of the potential of their communicative system . for example , they do not inverse the order of calls ( e . g . , ab to ba ) to generate differences in meaning , and a large number of other possible call combinations are not realized . second , human language is symbolic in the sense that signalers can inform listeners about referents that are not physically present ( 55 ) . in campbell ' s monkeys , we only observed calling in response to real life experiences , and some observations suggested that callers did not attempt to inform others . further research will have to test whether some of the observed contingencies between acoustic morphology and external events were intentionally produced or biproducts of other processes . whatever the outcome , our results demonstrate that the evolution of complex morphology has begun early in primate evolution , long before the emergence of hominids , and hence preceded the evolution of intentional communication .
sayer , j . a . , c . s . harcourt , and n . m . collins . 1992 . the conservation atlas of tropical forests : africa . iucn and simon & schuster , cambridge .
whether their rudimentary syntax echoes the speech of humanity ' s evolutionary ancestors , or represents an emergence of language unrelated to our own , is unclear . either way , they ' re far more sophisticated than we thought .
oates , j . f . , gippoliti , s . & groves , c . p . ( 2008 ) . cercopithecus campbelli . in : iucn 2008 . iucn red list of threatened species . retrieved 4 january 2009 .
guschanski , k . , krause , j . , sawyer , s . , valente , l . m . , bailey , s . , finstermeier , k . , sabin , r . , gilissen , e . , sonet , g . , nagy , z . t . , lenglet , g . , mayer , f . , & savolainen , v . ( 2013 ) . next - generation museomics disentangles one of the largest primate radiations .
species and related genera . wolf ' s monkeys have ischial callosities ( callus - like areas of skin on the buttocks ) . this provides a degree of comfort while sitting on branches and night resting . these callosities are typical of the family
information indicates evolutionary relationships at both the ordinal and the family level . this indispensable reference work belongs in public and academic libraries throughout the world and on the shelf of every biologist who works with mammals .
- - publisher ' s website .
occupies primary and secondary lowland rainforest habitats . wolf ' s monkeys are commonly found in swamp forests and secondary forests along riverbanks . they spend a majority of their time between 15 and 25 meters high in the canopy where they forage and sleep .
grubb , p . , t . s . jones , e . edberg , e . d . starin , j . e . hill . 1998 . mammals of ghana , sierra leone , and the gambia . tenderine press , london , uk .
the gambian forest contributes immensely to the gambia’s economy and the social well being of gambia’s population and provide several environmental services . in additional to maintaining the micro - climatic balance , the stabilisation of the river banks and providing life support systems to many other plants , animals and aquatic life , forests are important to the local communities who depend on them for food , medicines , wood products for construction and energy ( particularly to women who rely on the forests for their subsistence ) . particularly important forest products are for women .
wolf ' s monkeys are one of the species hunted in the bushmeat market . their meat provides food to local inhabitants and a product to trade for other goods . they are also likely to play a role in the regeneration of healthy forests through seed dispersal .
bakarr , m . i . , b . bailey , d . byler , r . ham , s . oliverieri , m . omland . 2001 . from the forest to the sea : biodiversity connections from guinea to togo . conservation international , washington d . c .
kingdon ( 1997 ) included this taxon within his cercopithecus mona superspecies group , groves ( 2001 ) followed the same arrangement whereas grubb et al . ( 2003 ) considered it the nominate subspecies of c . campbelli . the latter treatment was followed for the 2008 red list assessment . groves ( 2005 ) classified c . lowei as a distinct species and that approach is now followed here ( mittermeier et al . 2013 ) , hence the former nominate subspecies is now the species - concept for c . campbelli . this is an updated assessment to reflect the promotion of the nominate subspecies to species - level and the inclusion of information previously contained within the former species - level assessment .
the four habituated males were directly observed in dense forest vegetation for a total focal duration of 43 h ( male 1 , 11 months ; male 2 , 6 months ; male 3 , 15 months ; male 4 , 2 months ) . scan animal samples ( n = 4 , 425 ) were collected every 30 min , during which we recorded ( i ) the presence of other monkey species and ( ii ) the location within the group ' s home range . we also collected ad libitum samples from all habituated and semihabituated groups during ≈2 , 000 h total contact time . this enabled us notably to report on the response of males to their main predators , leopards ( n = 3 ) and crowned eagles ( n = 11 ) .
the baobolon wetland reserve was designated as a ramsar site upon the gambia’s ratification of the ramsar convention in 1996 . a comprehensive study of baobolon along with two additional sites , nuimi national park and the tanji wetland complex , was conducted in 1997 with a view to designating them as ramsar sites .
stuart , s . n . , r . j . adams and m . d . jenkins . 1990 . biodiversity in sub - saharan africa and its islands : conservation , management and sustainable use . occasional papers of the iucn species survival commission no . 6 . iucn , gland , switzerland .
a first sequence , recorded from all four habituated males and one semihabituated male , consisted of k + calls only ( median = 15 calls ; range : 3–25 ; n = 18 ; fig . 2 ) . this sequence was rare and given to any auditory sign of a predator , typically after hearing diana monkey alarm calls ( n = 15 / 18 ) or , more rarely ( n = 3 / 18 ) , after hearing predator vocalizations , but never to any visual signs of a predator .
jenkins , m . and a . hamilton . 1992 . biological diversity . pages 26 - 32 in j . a . sayer , c . s . harcourt , and n . m . collins , editors . the conservation atlas of tropical forests : africa . iucn and macmillan publishers , united kingdom .
vooren , f . and j . sayer . 1992 . côte d’ivoire . pages 133 - 142 in j . a . sayer , c . s . harcourt , and n . m . collins , editors . the conservation atlas of tropical forests : africa . iucn and macmillan publishers , united kingdom .
relative distribution of different call types within predator sequences with varying levels of predator threat . ( a ) eagle , ( b ) leopard . fisher exact test ( * * * , p < 0 . 001 ) were used to compare the relative contribution of obligatory ( black ) vs . optional ( white ) call types in low threat situations ( 1 , mainly diana monkey alarms ; 2 , playback of predator vocalizations ) or high threat circumstances ( 3 , visual predator models ; 4 , real predator encounters ) .
before each experiment , the following conditions had to be met : ( i ) the study group was aware of the presence of human observers for at least 30 min ; ( ii ) no alarm calls were produced for at least 30 min ; ( iii ) the predator model ( or playback speaker ) was positioned ahead of the group ' s estimated travel direction ensuring that no associated monkey species could detect it first . one observer ( k . o . ) and one field assistant were necessary for these experiments . the observer walked with the group and recorded the male ' s behavior , while the assistant operated the predator model . for eagle trials , the model or loudspeaker was positioned at an elevation of 2–3 m off the ground . for leopard trials , the model or loudspeaker was positioned on the ground . eagle shrieks were recorded in the study area ; leopard growls were purchased from the national sound archives , london . all acoustic stimuli were broadcast with a sony wmd6c professional walkman connected to nagra dsm speaker - amplifier with the amplitude level adjusted so that the calls sounded natural and could be clearly heard by the group .
not much information is known about the parental investment of wolf ' s monkeys , though it has been observed that infants will ride on the backs of their mothers for the first few months after birth . female young stay in their natal group , male young disperse from their natal group when they become independent .
wilson and reeder ' s mammal species of the world is the classic reference book on the taxonomic classification and distribution of the more than 5400 species of mammals that exist today . the third edition includes detailed information on nomenclature and , for the first time , common names . each concise entry covers type locality , distribution , synonyms , and major reference sources . the systematic arrangement of information indicates evolutionary relationships at both the ordinal and the family level . this indispensable reference work belongs in public and academic libraries throughout the world and on the shelf of every biologist who works with mammals .
- - publisher ' s website .
observations were conducted in 15 - min blocks under a focal animal sampling regime ( 56 ) between 08 : 00 and 17 : 00 gmt . during each observation period , the observer ( k . o . ) recorded all loud calls produced by the focal male . the single adult male is the only one to produce loud calls in the group , so there was no risk of confusion in terms of caller identification . if the male produced a loud call , the observer determined its likely cause among the following : ( i ) predator , presence of a leopard ( panthera pardus ) or crowned eagle ( stephanoaetus coronatus ) ; ( ii ) intergroup , presence of neighboring group ( usually inferred by loud calls of their adult male ) ; ( iii ) tree / branch , crashing sound of falling tree or large branch ; ( iv ) monkey alarm , diana monkey eagle or leopard alarm calls ( 54 ) ; and ( v ) cohesion and travel , male spatially separated far from the group ( < 30–50 m ) or beginning of group travel after spatial separation .
one way of studying language evolution is to compare the communicative abilities of humans and animals . parallels with human language can be found at various levels , both in terms of production and comprehension . particularly relevant are cases of social influences on vocal development ( 1 ) , cases of infant babbling ( 2 ) , and other types of vocal learning ( 3 – 5 ) . in some species , there is evidence for population - wide convergence effects in the form of culturally transmitted dialects [ e . g . , starlings ( 6 ) , whales ( 7 ) , and japanese macaques ( 8 ) ] . in terms of pragmatic use , there is good evidence that call production can be influenced by specific audiences ( 9 , 10 ) . in terms of comprehension , primates and possibly many other species are able to assign meaning to different calls if there is a strong relation between a call ' s acoustic morphology and its eliciting context ( 11 – 14 ) . in some species , there is some evidence for hemispheric specialization when processing conspecific calls [ e . g . , horses ( 15 ) , campbell ' s monkeys ( 16 ) , rhesus macaques ( 17 ) , starlings ( 18 ) , and sea lions ( 19 ) ] .
the second sequence type consisted of a pair of boom calls followed by a k + sequence ( median overall = 10 calls ; range : 4–15 calls : n = 53 ; fig . 2 ) . this combined sequence , recorded from all four habituated males , was mainly given to falling trees or branches with no other noticeable disturbance ( 85 % ) . in the remaining cases ( 15 % ) , the sequence was given in response to fights between other monkey species in the canopy , although this usually led to branches falling as well .
the majority of call sequences to crowned eagles were composed of wak - oo ( w + ) and krak - oo ( k + ) calls , sometimes with the addition of hok ( h ) and hok - oo ( h + ) calls ( median = 25 calls ; range : 15 to 40 ; n = 38 ; fig . 2 ) . by analyzing in more details the sequences produced by the three most habituated males , we found that high levels of urgency were associated with a high proportion of h and h + calls in the sequence . there was a significant difference in the proportion of h and h + calls if the caller spotted a real eagle or encountered the eagle model compared to when he only heard eagle vocalizations or diana monkey eagle alarm calls ( fisher ' s exact test , p < 0 . 001 ; fig . 4 a ) .
this ecoregion also has a diverse fauna ( martin 1991 , happold 1996 , bakkar et al . 1999 ) . there are nearly 1 , 000 vertebrates recorded in taï national park , and the park holds viable populations of the near - endemic pygmy hippopotamus ( hexaprotodon liberiensis , vu ) . in the order artiodactyla , two duikers , jentink’s duiker ( cephalophus jentinki , vu ) and zebra duiker ( cephalophus zebra , vu ) are strictly endemic to this ecoregion . the liberian mongoose ( liberiictis kuhni , en ) is also strictly endemic , and another small carnivore , johnston’s genet ( genetta johnstoni , dd ) , is known from small populations in liberia and côte d’ivoire ( hayman 1958 , schlitter 1974 , taylor 1989 , 1992 ) . miller’s striped mouse ( hybomys planifrons ) is the only other strictly endemic mammal , although more than 15 species of mammal are regarded as near - endemic , with all of these species shared only with the adjacent eastern guinea lowland forest and / or the guinea montane forest ecoregions .
mammals : a list of mammals complied in the late 1960’s indicated that there are 67 species of mammals but may have omitted many of the smaller rodents ( rodentia ) and bats ( chiroptera ) . the most recent assessment ( murphy , 1998 ) , puts the total number of mammals at 99 including marine mammals recorded form gambian waters .
some calls were given to a broad , others to a narrow range of events . crucially , “krak” calls were exclusively given after detecting a leopard , suggesting that it functioned as a leopard alarm call , whereas the “krak - oo” was given to almost any disturbance , suggesting it functioned as a general alert call . similarly , “hok” calls were almost exclusively associated with the presence of a crowned eagle ( either a real eagle attack or in response to another monkey ' s eagle alarm calls ) , while “hok - oo” calls were given to a range of disturbances within the canopy , including the presence of an eagle or a neighbouring group ( whose presence could sometimes be inferred by the vocal behaviour of the females ) . on a few occasions , “hok” and “hok - oo” calls were produced in response to a flying squirrel , whose silhouette somewhat resembles a flying eagle , but never to any other large bird .
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 .
in forest guenons , the single adult male of the group mainly vocalizes in response to predators and other significant external disturbances ( 5 , 48 , 53 ) . females are vocally active in social situations ( 44 – 46 ) and to predators to which they produce a diverse alarm call repertoire that encodes information about predator type and level of threat ( 47 ) . whether campbell ' s monkeys produced these calls to intentionally inform others about the event they have experienced cannot be decided with our data . some observations suggest not . for example , it is puzzling that males produce loud calls in response to the thundering sound of falling trees , as all other group members will have perceived the event as well . the collapse of a large tree provides a significant danger to arboreal animals and fatalities are not unusual . thus , calling males may have the urge to advertise their uninjured state and provide an acoustic focal point for scattered or disoriented group members , rather than attempting to inform others about the danger .
wolf ' s monkeys give birth to one offspring at a time , though twins occur rarely . most births occur from june to december when there is the greatest abundance of food . gestation length is from 160 to 170 days and the young are nursed for 3 months after birth . females produce their first young at 4 to 5 years old .
laboratoire ethos “ethologie animale et humaine” , u . m . r . 6552 - c . n . r . s . , université de rennes 1 , station biologique , paimpont , france , centre suisse de recherches scientifiques , abidjan , côte d ' ivoire , laboratoire de zoologie et de biologie animale , université de cocody , abidjan , côte d ' ivoire
the total staff demand was estimated based on the assumptions that the gfmc is implemented country - wide and that fd’s administrative structure is further streamlined by putting afforestation under divisional management , abolishing the beekeeping section , up - grading training and adaptive research , merging the extension unit with the cf unit , and by transferring management responsibilities to local communities represented by forest committees .
we are grateful to “office ivoirien des parcs et réserves” ( oipr ) for permission to conduct research in the taï national park . our special gratitude goes to the field assistants of the taï monkey project for all their invaluable help during data collection and for their stamina and support during the political turmoil of the previous years . we are equally grateful to n ' goran kouakou elizer , gombert jean emile , tecumseh fitch , catherine blois - heulin , laurence henry , isabelle charrier , julia fischer and martine hausberger for various contributions , including hélène bouchet , pascaline legouar , véronique biquand and muriel guernion for their help with the statistics .
impressive as that was , however , it was still relatively one - dimensional , not much different from verbalizations heard in many animal species , from other non - human primates to songbirds . the team ' s latest findings , published monday in the proceedings of the national academy of sciences , describe something far more complicated : syntax , or principles of word sequence and sentence structure .
wilson and reeder ' s mammal species of the world is the classic reference book on the taxonomic classification and distribution of the more than 5400 species of mammals that exist today . the third edition includes detailed information on nomenclature and , for the first time , common names . each concise entry covers type locality , distribution , synonyms , and major reference sources . the systematic arrangement of
) . when these birds are spotted by wolf ' s monkeys , they will sound an alarm call and retreat to the ground . though less common , leopards also pose a threat to this species . more recently , humans have become a major predator of this species for the bush meat market . in addition , their primary habitat is being destroyed at an extremely rapid pace for lumber .
forest resources are a source of biological diversity in themselves . the forest resources are also important for the conservation of biological diversity and for overall environmental protection . a good amount of vegetative cover for the land provides a good measure against soil erosion and other forms of land degradation . the gambia’s biodiversity resources are under increasing pressures and there is an alarming decline in both faunal and floral specie numbers .
although private forest both natural and plantations are foreseen in the forest policy and legislation , to date only one private gmelina plantation of about 100 ha exits which was even established before the new policy was formulated . at the moment the fd’s highest priority is to bring as much as possible of forest reserves under community management . considering the increasing wood demand , however the fd should pay equal attention to promotion and supporting private woodlots .
harcourt , c . , g . davies , j . waugh , j . oates , n . coulthard , n . burgess , p . wood and p . palmer . 1992 . sierra leone . pages 244 - 250 in j . a . sayer , c . s . harcourt , and n . m . collins , editors . the conservation atlas of tropical forests : africa . iucn and macmillan publishers , united kingdom .
bakarr , m . i . , b . bailey , m . omland , n . myers , l . hannah , c . g . mittermeier and r . a . mittermeier . 1999 . guinean forests . pages 239 – 253 in r . a . mittermeier , n . myers , p . r . gil and c . g . mittermeier . hotspots : earth’s biologically richest and most endangered terrestrial ecoregions . toppan printing company , japan .
being an arboreal quadruped , wolf ' s monkeys have forelimbs and hindlimbs that are fairly equal in length giving it an intermembral index number close to 100 . the head and body length of males varies from 445 to 511 mm with an average of 485 mm . the length of the tail in males ranges from 695 to 822 mm with an average of 779 mm . there has not been enough data collected from females to adaquetely determine these measurements .
we thank the office ivoirien des parcs et réserves for permission to conduct research in the taï national park , the field assistants of the taï monkey project for all their invaluable help during data collection , and n . elizer , g . emile , c . blois - heulin , v . biquand , h . bouchet , j . fischer , m . hausberger , and the gis ( groupement d ' intérêt scientifique ) “cerveau , comportement , société” for support and discussions . this work was supported by the european commission ( sixth framework programme , what it means to be human ) , european science foundation eurocore programmes ( origin of man language and languages ) , the wissenschaftskolleg zu berlin , the french ministry of foreign affairs ( egide ) , and the centre suisse de recherches scientifiques in abidjan .
the wildlife conservation et , 1977 , has defined protected areas as any area of land set aside by the government for purposes of preserving and managing the habitat and ecology , including any forest park or local sanctuary . the current protected area system in the gambia comprises six national parks and nature reserves under the mandate of the department of parks and wildlife management , dpwm covering a total land area of 39 , 772 ha i . e . about 3 . 7 % of the gambia’s land area .
plantation establishment , using gmelina aborea was the main preoccupation of the department of forestry in the years between 1953 and 1985 . a total of about 13000 hectares of monoculture of gmelina aborea and very small amount of tectona grandis were planted mainly in the western division . the high cost of plantation establishment and fire protection as well as to conserve biodiversity caused the department to reconsider it’s forest management priorities and potentials and since 1985 , a policy decision was reached to re - orient forestry department to focus more on natural forest management .
perelman , p . , johnson , w . e . , roos , c . , seuánez , h . n . , horvath , j . e . , moreira , m . a . m . , kessing , b . , ponitus , j . , roelke , m . , rumpler , y . , schneider , m . p . c . , silva , a . , o brien , s . j . , & pecon - slattery , j . ( 2011 ) . a molecular phylogeny of living primates .
the gambia has designated 6 protected areas for the conservation of wildlife resources . in addition to fulfilling that function , the protected areas also contain significant amounts of plant species , particularly the rare plant species . the total area under this form of land use currently stands at around 3 . 4 % of the gambia’s total land area . following the ratification of the international convention on biological diversity in 1994 , the government of the gambia under the auspices of its department of parks and wildlife management elaborated the biodiversity strategy and action plan ( bsap ) . the bsap provides a coherent framework for the management of the gambia’s biological resources on a sustainable basis as well as to ensure the fair and equitable sharing of the benefits arising . one of the policy objectives of the bsap is to increase the area under wildlife to about 5 % of the total land area of the gambia . furthermore , the policy also aims to involve local communities in the management of biological resources . the implementation of such a policy is likely to promote to the better management and enhancement of forest - based biodiversity .
the herpetofauna is also diverse ( welch 1982 ) , and contains a large number of endemic species . in the amphibians there are 13 strictly endemic species and a number of others shared with the eastern guinea lowland forest ecoregion . the strict endemics include the rare frog merlin’s clawed frog ( pseudhymenochirus merlini ) known only from guinea and sierra leone ( chabanaud 1920 , menzies 1967 ) , and the freetown long - fingered frog ( cardioglossa aureoli ) , which is only known from the mountains close to freetown in sierra leone . other notable endemics include the tai river frog ( phrynobatrachus taiensis ) , liberian long - fingered frog ( cardioglossa liberiensis ) and ivory coast toad ( bufo danielae ) ( schiøtz 1964 , 1967 , wcmc 1994 , harcourt et al . 1992 , vooren and sayer 1992 ) . the reptile fauna is less rich in endemics , with three strictly endemic species and thirteen shared only with other ecoregions in the upper guinea forest block . the strict endemics are los archipelago worm lizard ( cynisca leonine ) , benson’s mabuya ( mabuya bensonii ) and liberia worm snake ( typhlops leucostictus ) .
lemasson ' s team previously described the monkeys ' use of calls with specific meanings in a paper published in november . it detailed the monkeys ' basic sound structures and their uses :
hok
for eagle ,
krak
for leopard ,
krak - oo
for general disturbance ,
hok - oo
and
wak - oo
for general disturbance in forest canopies . a sixth call ,
boom ,
was used in non - predatory contexts , such as when calling a group together for travel or arguing with neighboring groups .
to leopards , males sometimes produced pure k sequences ( n = 9 / 26 ) , but if k + calls were added , then typically toward the end of the sequence . we compared the number of k and k + calls in the first and second half of the sequences and found a significant difference ( fisher ' s exact test , p < 0 . 001 ) . to crowned eagles , most sequences began with a series of h ( 16 of 38 sequences ) and typically ended with a series of k + ( 36 of 38 sequences ) . this specific order probably reflected the decrease in urgency or threat perceived by the caller . if w + or h + were produced , then they appeared more or less randomly throughout the sequence without any detectable patterns , while h and k + followed the ordering outlined before . there was a significant difference between the number of h and k + at the beginning and at the end of sequences ( fisher ' s exact test , p < 0 . 001 ; n = 38 ) , which had the effect that the distribution patterns differed significantly between the four call types ( χ 2 test x 2 = 311 . 3 ; dl = 21 ; p < 0 . 001 ) .


pachypsylla is a genus of psyllids .

each of its four species lay eggs on the leaves of the celtis occidentalis tree .

upon hatching , the young psyllids become encased in a " gall " which the young leaf parts grow in response to the infestation .

species of pachypsylla include pachypsylla celtidisgemma ( hackberry bud gall maker ) , pachypsylla celtidismamma ( hackberry nipplegall maker ) , pachypsylla celtidisvesiculum ( hackberry blistergall psyllid ) and pachypsylla venusta ( petiolegall psyllid ) .

pachypsylla

figure 4 . petiole gall psyllid nymph , pachypsylla venusta ( osten - sacken ) . photograph by jerry f . butler , university of florida .
what made you want to look up pachypsylla ? please tell us where you read or heard it ( including the quote , if possible ) .
pachypsylla venusta is a gall - forming psyllid ( insecta : hemiptera ) specializing on hackberry trees , which are widely distributed in the united states .
figure 2 . adult hackberry petiole gall psyllid , pachypsylla venusta ( osten - sacken ) . photograph by jerry f . butler , university of florida .
genome size of pachypsylla venusta ( hemiptera : psyllidae ) and the ploidy of its bacteriocyte , the symbiotic host cell that harbors intracellular mu . . . - pubmed - ncbi
genome size of pachypsylla venusta ( hemiptera : psyllidae ) and the ploidy of its bacteriocyte , the symbiotic host cell that harbors intracellular mutualistic bacteria with the smallest cellular genome .
figure 3 . petiole gall psyllid nymph , pachypsylla venusta ( osten - sacken ) , with gall showing individual compartments . photograph by jerry f . butler , university of florida .
figure 7 . hackberry , celtis occidentalis l . ( celtidaceae ) , is a host for the hackberry petiole gall psyllid , pachypsylla venusta ( osten - sacken ) . photograph by don hall , university of florida .
figure 8 . sugarberry , celtis laevigata willd . ( celtidaceae ) , is a host for the hackberry petiole gall psyllid , pachypsylla venusta ( osten - sacken ) . photograph by don hall , university of florida .
figure 6 . apical view of abdomen of petiole gall psyllid , pachypsylla venusta ( osten - sacken ) , showing abdominal cutting teeth ( scanning electron micrograph ) . photograph by harvey l . cromroy , university of florida .
figure 5 . dorsal view of tip of abdomen of petiole gall psyllid , pachypsylla venusta ( osten - sacken ) , showing abdominal cutting teeth ( scanning electron micrograph ) . photograph by harvey l . cromroy , university of florida .
biosystematics of hackberry psyllids ( pachypsylla ) and the evolution of gall and lerp formation in psyllids yang , m . - m . & c . mitter . 1993 . the ecology and evolution of gall - forming insects . united states dept . of agriculture .
the genome of the bacterial symbiont carsonella from pachypsylla venusta has been sequenced and represents one of the most extreme cases of genome reduction ever identified . at only 160 kb in size , this bacterial genome lacks many genes thought to be essential for cellular life , making this system an important model for elucidating the genomic mechanisms of host - symbiont interactions .
yang m - m , mitter c . 1994 . biosystematics of hackberry psyllids ( pachypsylla ) and the evolution of gall and lerp formation in psyllids ( homoptera : psylloidea ) : a preliminary report . in : price pw , mattson wj , baranchikov yn . ( editors ) the ecology and evolution of gall - forming insects . u . s . d . a . forest service ( north central forest experiment station ) general technical report nc - 174 . st . paul , minnesota . pp . 172 - 185 .
infested hackberry trees do not seem to be harmed by these galls , but their abundance makes hackberry leaves look pretty ugly .
in september and october , people who have hackberry trees , or live in neighborhoods where there are hackberry trees , often notice tiny greyish bugs that congregate on their homes , on window screens , front doors and siding . these insects are attracted to lights at night and , at 1 / 10
long , are tiny enough to pass through ordinary window screen . people describe these bugs as gnats , flies or fleas . the photo above hackberry gall psyllid on the tip of a pencil !
these insects are adult hackberry gall psyllids ( pronounced , sill - ids ) . another name is
hackberry nipple gall maker
. under magnification , they look like miniature cicadas ( what people in nebraska commonly call
locusts
) , which makes perfect sense , because they are in same order ( homoptera ) as cicadas , leafhoppers and aphids .
in the fall , these insects are looking for cracks and crevices to squeeze into so they can hibernate without succumbing to lethal temperatures . normally , they overwinter under the bark of trees , but psyllids don ' t distinguish between
good
and
bad
overwintering locations so they also squeeze into cracks and crevices around windows , doors and siding . those that come inside are likely to die .
hackberry psyllids are not harmful to people or pets and will not attack house plants , stored products or furnishings . they are a temporary nuisance . as temperatures get colder , their activity will decrease and hibernation will set in .
hackberry psyllids are so annoying that people sometimes ask about spraying hackberry trees to control them . during the summer , psyllids are protected inside the gall ( photo right ) from insecticides sprayed on the leaves so foliar treatments won ' t be effective then . unfortunately , by the time psyllids emerge and start to move out of the trees , it is probably too late to achieve effective chemical control by spraying the trees .
a more effective preventative approach would be to treat trees in the spring to kill newly hatched nymphs before the onset of gall formation . but , because egg laying occurs over a period of several weeks beginning when new leaves unfold from the bud , several foliar insecticide applications would be needed . the labor involved makes this approach impractical , especially with large trees .
treating hackberry trees with a systemic insecticide to kill psyllids when they feed would be ideal , but this proactive approach means planning ahead . check out systemic insecticides at your home and garden store . one fairly new systemic product , bayer advanced garden tree & shrub control , contains imidacloprid which provides year - long control . it controls sucking insects , like aphids , psyllids , lacebugs and scale insects . it is even pretty easy to use . after determining the amount needed ( based on the diameter of the tree ) , just mix the liquid insecticide in water and pour around the base of the tree . when using any insecticide product , be sure to read and follow all label directions .
spraying with insecticides is not recommended . people who are really
bugged
by this problem and just have to do something can try hosing down their siding with water . this is likely to drown many of the psyllids , but , as long as temperatures are warm , more may show up the next day . a fine mesh window screen ( 18 mesh ) may be small enough to prevent entry through open windows . for those insects that get inside , sucking them up with a vacuum cleaner is very effective .
again , once the psyllids get indoors they will die in your home - even if you do absolutely nothing .
this article was written by dr . barb ogg , phd , extension educator emeritus and it appeared in the nebline newsletter . the information was updated november 2015 by soni cochran , extension associate .
nebraska extension in lancaster county is your on - line educational resource . the information on this web site is valid for residents of southeastern nebraska . it may or may not apply in your area . if you live outside southeastern nebraska , visit your local extension office
nebraska extension in lancaster county 444 cherrycreek road , suite a , lincoln ne 68528 402 - 441 - 7180 | lancaster @ urltoken office hours are 8 a . m . - 4 : 30 p . m . , monday through friday with the exception of designated holidays . learn more about us
in lancaster county , the 4 - h youth development program is a partnership between nebraska extension and the lancaster county government .
html public
- / / w3c / / dtd xhtml 1 . 0 transitional / / en
urltoken
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
select your preferred way to display the comments and click ' save settings ' to activate your changes .
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 .
but are smaller and with wing pattern usually more diffuse . tuthill notes that the sudden narrowing midway of the ventral valve of the female genital segment is the most reliable way to distinguish the two species
the psyllids of america north of mexico : ( psyllidae : homoptera ) ( subfamilies psyllinae and triozinae ) tuthill , l . d . . 1943 . iowa state college journal of science 17 : 443 - 660 .
html public
- / / w3c / / dtd xhtml 1 . 0 strict / / en
urltoken
as its name implies , the hackberry petiole gall psyllid forms woody galls on the leaf petioles of its hackberry ( celtis spp . ) hosts . our native florida hackberry , celtis laevigata willd . , is called sugarberry . the petiole gall psyllid is usually not sufficiently abundant to cause serious damage to its host , but gall infested leaves are unsightly during late fall and winter .
figure 1 . petiole gall on sugarberry , celtis laevigata . photograph by jerry f . butler , university of florida .
the petiole gall psyllid is found throughout the range of its four hackberry hosts - from connecticut to idaho , southwest to arizona and southeast throughout florida .
adult petiole gall psyllids are fairly large for psyllids ( 5 . 0 to 6 . 0 mm to tip of folded wings ) and resemble small cicadas . they are dark colored with tawny markings . the dorsum of the thorax is longitudinally striped . the forewings are whitish with black spots . the full - grown nymph has a green head with brown markings and a green thorax with four light reddish - brown longitudinal stripes . the wing pads of nymphs are brown . the abdomen is green with dark brown transverse bands .
last instar nymphs emerge from the galls in spring and molt to the adult stage as new leaves are opening on the host trees . the apex of the abdomen of the last instar nymph is armed with heavily sclerotized teeth which the nymph uses to cut its way out of the woody gall by wagging the tip of the abdomen .
subspherical galls form around the young nymphs . galls are polythalamous ( having several developing individuals in separate compartments ) with each compartment lined with wax . i have found as many as 13 nymphs per gall . nymphs develop throughout the summer and overwinter as last instars . as in the other species of gall - forming hackberry psyllids , many petiole gall psyllid nymphs are parasitized by hymenoptera larvae .
infested leaves die in the fall instead of undergoing abscission and do not fall from the trees . heavily infested trees are recognizable during the winter by the presence of the dead leaves after uninfested leaves have fallen from the tree .
because significant damage to the tree does not result from infestation , control of hackberry petiole gall psyllids is not recommended .
carsonella ruddii , the endosymbiont of the hackberry petiole gall psyllid , is remarkable for having the smallest known bacterial genome with only about 160 , 000 base pairs - only about 1 / 3 the size of the next smallest genome which is found in an aphid endosymbiont , buchnera sp . ( nakabachi et al . 2006 ) .
there are three hymenopterous parasitoids listed from the hackberry petiole gall psyllid ( krombein et al . 1979 ) .
two trees in the family celtidaceae are hosts for the hackberry petiole gall psyllid . these are hackberry , celtis occidentalis l . , and sugarberry , celtis laevigata willd .
baumann , p . 2006 . diversity of prokaryote - insect associations within the sternorrhyncha ( psyllids , whiteflies , aphids , mealybugs ) . in bourtzis k , miller ta ( editors ) . insect symbiosis . crc press , boca raton , florida . vol . 2 : pp . 1 - 24 .
johnson wt . 1988 . insects that feed on trees and shrubs . 2nd edition . cornell university press . ithaca , new york . pp . 452 - 453 .
krombein kv , hurd jr . pd , smith dr , burks bd . 1979 . catalog of hymenoptera in america north of mexico . volume 1 . symphyta and apocrita ( parasitica ) . smithsonian institution press . washington , d . c . 1198 pp .
nakabachi a , yamashita a , toh h , ishikawa h , dunbar he , moran na , hattori m . 2006 . the 160 - kilobase genome of the bacterial endosymbiont carsonella . science 314 ( issue 5797 ) : 267 .
tuthill ld . 1943 . the psyllids of america north of mexico ( psyllidae : homoptera ) . iowa state college journal of science . pp . 534 - 535 .
photographs : jerry f . butler and harvey l . cromroy , university of florida
love words ? you must — there are over 200 , 000 words in our free online dictionary , but you are looking for one that’s only in the merriam - webster unabridged dictionary .
the story of an imaginary word that managed to sneak past our editors and enter the dictionary .
take this quiz and discover 12 words for things you didn ' t know had words .
like many closely - related hemipterans such as whiteflies , aphids , and mealybugs , psyllids have a nutritionally imbalanced diet consisting primarily of plant sap . to compensate for the paucity of essential amino acids and other required nutrients in their diets , these insects have evolved ancient and intimate symbiotic relationships with intracellular bacteria that are capable of synthesizing these compounds .
a complete genome from this gall - forming psyllid will also provide a valuable resource for investigating plant - insect interactions and gall - induction .
please cite the following publication when using this dataset : sloan , d . b . , nakabachi , a . , richards , s . , qu , j . , murali , s . c . , gibbs , r . a . , & moran , n . a . ( 2014 ) . parallel histories of horizontal gene transfer facilitated extreme reduction of endosymbiont genomes in sap - feeding insects . molecular biology and evolution , 31 ( 4 ) , 857 - 871 . urltoken
data were generated by the baylor college of medicine ' s i5k pilot project .
sequence generation for assembly . for this project we are generating fairly high coverage in a number of different insert sized libraries . the assembly strategy is based around a seed allpaths assembly ( the broad allpaths assembler ) followed by seed assembly improvement using homegrown tools , atlas - link and atlas - gapfill , which can significantly improve the results . thus we generate sequence data to enable the allpaths assembly . as of nov 2011 this is : - 40x genome coverage in 180bp insert library ( 100bp reads forward and reverse ) ; and 40x 3kb insert data . to enable better scaffolding and local gap filling we additionally generate 500bp , 1kb , 2kb , and 8kb insert sizes at > 20x coverage .
please cite the use of our resources : doi : 10 . 1093 / nar / gku983
warning : the ncbi web site requires javascript to function . more . . .
advanced science institute , riken , 2 - 1 hirosawa , wako , saitama 351 - 0198 , japan . bachi @ urltoken