mesaxonians are a clade of ungulates whose weight is distributed on the third toe on all legs through the plane symmetry of their feet .

for a while it was often seen to only contain the order perissodactyla ( which includes the equines , rhinos and tapirs ) .

recent work in morphological cladistics and ancient dna suggests that several extinct lineages , like the desmostylia and some of the south american ungulates of meridiungulata ( both groups seen as afrotherian relatives ) are related to the perissodactyls .

mesaxonia

the mesaxonia , as defined here , include the horses , rhinos , elephants , and dugongs , among others . the group shares a number of foot , skull , and tooth characteristics which support its monophyly . however , no official name has been published for this group , and so we have borrowed the term mesaxonia , which is still used by some authors to refer to the perissodactyla .
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fisher ms , tassy p ( 1993 ) the interrelation between proboscidea , sirenia , hyracoidea , and mesaxonia : the morphological evidence . in : szalay fs , novacek mj , mckenna mc , editors . mammal phylogeny : placentals . new york : springer - verlag . pp . 217–234 .
fischer ms , tassy p ( 1993 ) the interrelations between proboscidea , sirenia , hyracoidea , and mesaxonia : the morphological evidence . in : szalay fs , novacek mj , mckenna mc , editors . mammal phylogeny : placentals . new york : springer - verlag . pp . 217–243 .
[ mesaxonia ] include the perissodactyla and some related fossil types . the middle toe may be the only one ( e . g . in present day horses ) , or may be the largest of three , as in some ancestral horses and in the hind - foot of the rhinoceros .
mesaxonians are a group of mammals that form the , not widely accepted , superorder mesaxonia . the superorder includes the perissodactylids , hyracoids , proboscidians , and sirenians , all part of the infraclass eutheria . however , this group is not yet published , it ` s a theory based on the number of digits , teeth , and skull characteristics . . . .
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anthracobunidae is an eocene family of large mammals from south asia that is commonly considered to be part of the radiation that gave rise to elephants ( proboscideans ) and sea cows ( sirenians ) . we describe a new collection of anthracobunid fossils from middle eocene rocks of indo - pakistan that more than doubles the number of known anthracobunid fossils and challenges their putative relationships , instead implying that they are stem perissodactyls . cranial , dental , and postcranial elements allow a revision of species and the recognition of a new anthracobunid genus . analyses of stable isotopes and long bone geometry together suggest that most anthracobunids fed on land , but spent a considerable amount of time near water . this new evidence expands our understanding of stem perissodactyl diversity and sheds new light on perissodactyl origins .
citation : cooper ln , seiffert er , clementz m , madar si , bajpai s , hussain st , et al . ( 2014 ) anthracobunids from the middle eocene of india and pakistan are stem perissodactyls . plos one 9 ( 10 ) : e109232 . urltoken
editor : andrew a . farke , raymond m . alf museum of paleontology , united states of america
copyright : © 2014 cooper 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 .
data availability : the authors confirm that all data underlying the findings are fully available without restriction . all relevant data are within the paper and its supporting information files .
funding : the national science foundation is acknowledged for support to m . t . c . ( ear - 0847413 ) , e . r . s . ( bcs - 0819186 and bcs - 0416164 ) , and j . g . m . t . ( ear - 0207370 ) . also , s . b . acknowledges financial support received from the department of science and technology , government of india for field work in the eocene sections of gujarat . the funders had no role in the study design , data collection and analysis , decision to publish , or preparation of the manuscript .
anthracobunidae is a family of large mammals that is only known from the middle eocene of south asia . historically , anthracobunids have played a prominent role in debates surrounding the origin of tethytheria , the group that includes the living and extinct members of the placental mammalian orders sirenia ( sea cows ) and proboscidea ( elephants ) . anthracobunids are often considered to have branched off near the base of tethytheria [ 1 ] – [ 8 ] , sharing most recent common ancestry with either proboscidea [ 1 ] , [ 9 ] – [ 13 ] , sirenia [ 14 ] , [ 15 ] , or the extinct desmostylia . however , until now , no cranial parts , only some partial dentitions and two postcranial elements ( a scapula associated with a maxilla [ 11 ] and an isolated astragalus [ 13 ] ) , were known for the group .
the paucity of fossil evidence has left anthracobunidae as a phylogenetically and adaptively enigmatic group [ 2 ] , [ 8 ] , [ 16 ] – [ 20 ] , with little known about their diets , habitats , and time and place of origin . in light of this , anthracobunid genera have also been aligned with two other , non - tethytherian clades , including artiodactyla [ 21 ] – [ 23 ] and perissodactyla [ 24 ] , as well as phenacodontids [ 25 ] , [ 26 ] , one of the extinct families of basal ungulates . the convergent evolution of “ungulate” - like features in paenungulata ( containing tethytheria ) and laurasiatheria ( containing artiodactyla and perissodactyla ) revealed by molecular data [ 27 ] requires that anthracobunidae be re - evaluated within this new phylogenetic context .
here we describe new anthracobunid fossils from the kuldana and subathu formations of the early - middle eocene of pakistan and india [ 28 ] – [ 32 ] . these new findings more than double the number of anthracobunid fossils , and include cranial and postcranial material . we test the phylogenetic relationships of anthracobunids using a taxonomically rich morphological data set and investigate anthracobunid diet and habitat . our results have important implications for character transformations near the origins of perissodactyla , the habitats of stem perissodactyls , and the early biogeography of paenungulata and perissodactyla .
fossils were collected in punjab province , pakistan , under a collaborative agreement of howard university and the geological survey of pakistan ( h - gsp ) . localities where fossils were collected , with topographical coordinates , were described in thewissen et al . [ 33 ] . specimens were prepared in the u . s . a . at northeast ohio medical university ( neomed ) and are catalogued using the h - gsp acronym and accessioned into the collections of the geological survey of pakistan ( gsp ) . since the gsp is a branch of the pakistani government , all collected fossils are the property of the pakistani state . at present , fossils are for study in the u . s . a . , but they will be returned to pakistan and will be permanently housed at the natural history museum of pakistan , located in islamabad . gsp geologists are collaborators on all field work as part of their official assignment and all permitting is internal to the different pakistani government agencies involved . all necessary permits were obtained for the described study , which complied with all relevant regulations .
we also studied fossils from the locality kalakot in jammu & kashmir state , india , the location of which is described by russell and zhai [ 32 ] . this locality was discovered by a . ranga rao , a geologist of the oil and natural gas commission , and he quarried large quantities of fossiliferous rock matrix from this locality . after his death , we were given access to these by his widow , f . obergfell . fossils were prepared at neomed in the u . s . a . , and are the property of the rangarao - obergfell trust for geosciences , located in dehra dun , india , and are catalogued using the acronym rr . they will be housed on the trust campus on rajpur road in dehra dun .
the electronic edition of this article conforms to the requirements of the amended international code of zoological nomenclature , and hence the new names contained herein are available under that code from the electronic edition of this article . this published work and the nomenclatural acts it contains have been registered in zoobank , the online registration system for the iczn . the zoobank lsids ( life science identifiers ) can be resolved and the associated information viewed through any standard web browser by appending the lsid to the prefix “ urltoken ” . the lsid for this publication is : urn : lsid : zoobank . org : pub : cd33f507 - 0d06 - 4de1 - a3c8 - 8bf9e7697114 . the electronic edition of this work was published in a journal with an issn , and has been archived and is available from the following digital repositories : pubmed central , lockss .
whenever possible , characters were scored based on analysis of original fossil material and / or casts housed in either the department of vertebrate paleontology at the american museum of natural history , the department of anatomical sciences at stony brook university , or the department of anatomy and neurobiology at northeast ohio medical university . as noted above , some taxa and / or characters were scored from the literature . some dental characters were rescored for arsinoitherium , taking into account new evidence for cusp homologies provided by the more basal embrithopod namatherium [ 52 ] . furthermore , a new character state ( “large , inflated” ) was added for the character “m1 - 2 metaconules” ( number 18532 ) to take into account the condition seen in anthracobunids and some other ingroup taxa , and the character “shape of astragalar ectal facet” was excluded , because it could not be consistently scored across the new taxa that were sampled .
midshaft cross - sections of fossil and extant large - bodied ungulate long bones and ribs [ 71 ] were visualized via paleohistological sections or high resolution micro - ct scans . bone compactness ( total amount of bone per section ) was calculated via published methods [ 72 ] .
for the analysis of stable isotopes , three or more specimens of each species were analyzed ( when available ; see tables s8 and s9 ) to provide an estimate of the population mean ± s . d . for carbon and oxygen isotope values [ 73 ] urltoken - b33 . about 5 mg of enamel powder was collected from each specimen , either by drilling directly from the tooth or by grinding enamel chips in an agate mortar and pestle . before collection , contaminants were removed by abrading the outer surface of the specimen after cleaning the outer surface to a depth of ∼0 . 5 mm using a dremel drill with a 1 mm diameter diamond - coated bit .
preparation of powders for stable isotope analysis followed published methods [ 74 ] , [ 75 ] . powders were first transferred to 1 . 5 ml microcentrifuge vials and then soaked sequentially overnight in about 0 . 20 ml of a sodium hypochlorite solution ( 1–2 g dl - 1 ) and then in about 0 . 20 ml of calcium acetate buffered acetic acid ( ph about 5 ) following published procedures [ 31 ] . on addition of each reagent , samples were agitated for 1 min on a vortex genie vortex mixer . after each soak , the supernatant was removed by aspiration and the residual powder was rinsed five times with deionized water . samples were then freeze - dried overnight and about 1 . 5 mg of powder from each was weighed into individual test tubes for analysis on a thermo - finnigan gas bench autosampler attached to a thermo - finnigan deltaplusxp continuous - flow isotope - ratio mass spectrometer at the university of wyoming stable isotope facility .
all values for stable isotopes are reported in delta ( δ ) notation , using the equation δ ( ‰ ) = 1 , 000 × ( rsample / rstandard - 1 ) , where rsample is the observed isotope ratio of the sample ( 13 c / 12 c or 18 o / 16 o ) and rstandard is the accepted ratio for an appropriate international standard ( vienna pee dee belemnite for δ 13 c ; vienna standard mean ocean water for δ 18 o ) . analytical precision is typically better than 0 . 1‰ for δ 13 c values and 0 . 2‰ for δ 18 o values ( ±1σ ) .
enamel δ 18 o values are influenced by the oxygen isotope compositions of oxygen sources ( atmospheric o 2 , food and water ingested by the animal ) , certain physiological processes that affect intake or loss of oxygen by an animal ( sweating , panting , respiration , and elimination of urine and feces ) , and body temperature [ 76 ] – [ 78 ] . for aquatic and semi - aquatic mammals , the flux of environmental water by means of direct ingestion and transcutaneous exchange [ 79 ] overwhelms all other oxygen sources , which reduces inter - individual variation [ 76 ] , [ 80 ] and causes enamel δ 18 o values of freshwater taxa ( for example hippopotamus ) to be lower than those for terrestrial mammals ( ∼2–3‰ ) ; [ 81 ] – [ 83 ] . the magnitude of offset between freshwater and terrestrial mammals can vary in response to changes in humidity , such that faunas experiencing drier conditions show a greater offset ( ∼3 . 0‰ ) than those living under wetter conditions ( < 1 . 0‰ ) [ 84 ] .
in combination with enamel δ 18 o values , we also determined enamel δ 13 c values of fossil species , which were used to infer diet and past environmental conditions . for herbivorous ungulates , enamel carbon isotopic compositions are enriched in 13 c relative to diet by ∼14‰ [ 85 ] . variation in enamel δ 13 c values among ungulate species is most strongly affected by the carbon isotopic composition of the plants they ingest . in the eocene , terrestrial ecosystems were dominated by plants performing c3 photosynthesis , a process that discriminates against the heavier carbon isotope ( 13 c ) , causing plant tissue δ 13 c values to be considerably lower than those of plants exploiting other photosynthetic pathways ( c4 plants , cam plants ) [ 86 ] , [ 87 ] . the carbon isotopic compositions of c3 plants are more variable than other plant types and are strongly affected by temperature and aridity , causing c3 plants grown under hot dry conditions to have higher δ 13 c values than those grown in cooler and wetter environments [ 88 ] , [ 89 ] . since enamel δ 13 c values reflect the carbon isotopic composition of the plants in an animal ' s diet , consumer enamel δ 13 c values can be used to infer past environmental conditions ( wet vs . dry ) as well as dietary information for extinct species .
included genera . jozaria wells and gingerich , 1983 ; and obergfellia , new genus .
diagnosis . stem perissodactyls with premolars increasing in complexity from anterior to posterior , but never molariform ; molars brachyodont , but with high cusps and low valleys between cusps ; upper molars increasing in size from m1 to m3 ; lower molars with a distinct trigonid elevated above the talonid ; large caudally - projecting angular process of the mandible .
description . unlike tethytheres and other paenungulates , anthracobunids have small and simple upper and lower incisors and relatively large canines ( figs . 1 , 2 ) . cheek teeth are bunodont and brachyodont and each cusp has a high apex , but there are deep valleys between cusps . premolars are never totally molariform , and diastemata , if present at all , are short . upper p3 and p4 are shorter than the molars , and have a paracone , protocone , and metacone , but no hypocone . the m1 and m2 have four well - developed cusps , as well as strong para - and metaconules , without lophs . the distance between proto - and paracone is longer than that between the hypo - and metacone , giving the tooth a posteriorly pinched appearance . lower molars have a trigonid with two large cusps , and large , sometimes twinned , hypoconulid on the third lobe ( figs . 1g–i , s1 ) .
( a ) crushed skull of a . pinfoldi ( h - gsp 97106 ) in ventro - lateral view ( left maxilla detached ) and ( b ) dorsal view ; ( c ) p2 of a . wardi ( h - gsp 30229 ) in lingual view and ( d ) occlusal view ; ( e ) skull fragment of a . wardi ( rr 411 ) in occlusal view , and ( f ) lateral view ; ( g ) mandible of a . wardi ( h - gsp 96434 ) in occlusal view ; ( h ) mandible of a . wardi ( h - gsp 96258 ) in lateral view , and ( i ) occlusal view . ( j ) proximal phalanx of a . pinfoldi ( h - gsp 97106 . 105 ) in dorsal view ; ( k ) proximal phalanx of a . pinfoldi ( h - gsp 97106 . 101 ) in ventral view , and ( l ) lateral view ; ( m ) head of a metapodial of a . pinfoldi ( h - gsp 97106 . 250 ) in dorsal view ; ( n ) phalangeal fragment of a . pinfoldi ( h - gsp 97106 . 257 ) in dorsal - superior view ; ( o ) terminal phalanx of a . pinfoldi ( h - gsp 97106 . 302 ) in dorsal - superior view . scale bar is 1 cm in length .
( a ) right p2 - m2 of a . wardi ( luvp - 15006 ) in occlusal view , and ( b ) labial view ; ( c ) left m1 - m3 of a . wardi ( rr - 411 ) in occlusal view , and ( d ) labial view ; ( e ) left p3 - m3 of a . pinfoldi ( h - gsp 82 - 31p ) in occlusal view ; ( f ) left m3 of a . pinfoldi ( h - gsp 82 - 31p ) in labial view ; ( g ) right c - m3 of the cambaythere kalitherium ( iitr - sb - vlm 931 ) in occlusal view ; ( h ) right m3 of kalitherium ( iitr - sb - vlm 931 ) in labial view ; ( i ) left p4 - m2 of nakusia [ 6 ] in occlusal view ; ( j ) left m2 of nakusia [ 6 ] in labial view ; ( k ) mx of cambaytherium ( iitr - sb - vlm - 521 ) in occlusal view ; ( l ) right m3 of the phenacodontid ‘condylarth’ tetraclaenodon ( ku - 8052 ) in occlusal view ; ( m ) left p1 - m3 of a . wardi ( wif / a 1101 ) [ 15 ] in occlusal view ; ( n ) left c - m3 of a . wardi ( h - gsp 96258 ) in occlusal view , and ( o ) labial view ; ( p ) left p1 - m3 of obergfellia occidentalis ( h - gsp 1981 ) in occlusal view , and ( q ) labial view ( m1 inverted from right side ) . scale bar is 1 cm in length . illustrations by jacqueline dillard .
partial skulls are known for two anthracobunids ( h - gsp 97106 and rr 411 ) , and both show the presence of a very thick bony shelf over the orbits ( fig . 1b , f ) . the palate is concave mediolaterally , with the areas near the midline much higher than the alveolar processes . the lateral sides of the basisphenoid and basioccipital are recessed ; this is particularly true for the basioccipital , which suggests that the petrosal , which is not preserved , was also deeply recessed . the mandibular fossa is concave and is located just lateral to the ear region . the postglenoid process is oval and does not extend along the entire width of the mandibular fossa .
h - gsp 97106 includes , among other elements , metapodials and phalanges ( fig . 1j - o ) . the heads of most metapodials are wider than deep , strongly convex , and the posterior side bears a strong crest . proximal phalanges show a broad , oval articular surface for the metapodial and are squat : nearly as wide mediolaterally as they are long proximodistally . their distal articular facet is rectangular in outline and deeply incised caudally , indicating the presence of a caudal crest on the middle phalanx . most prominent on the proximal phalanges is the deeply excavated posterior side which is concave both mediolaterally and proximodistally .
the genera of the family anthracobunidae . we restrict the family anthracobunidae to the genera anthracobune ( which includes most specimens previously referred to pilgrimella and lammidhania ) , jozaria , and the new genus obergfellia . there are several other families of medium - sized bunodont “ungulates” from the eocene of india and pakistan , such as quettacyonids [ 90 ] and cambaytheres [ 4 ] , [ 91 ] . while all of these groups are brachyodont , anthracobunids are unique among them in having tall cusps on their molars , while retaining valleys between the cusps that are low and close to the cingulum ( fig . 2 ) . the new genus obergfellia differs from other anthracobunids in exhibiting the following combination of features : ( i ) lower molars broad , ( ii ) lower m3 relatively short , and ( iii ) angular process of the mandible long but shorter than that of anthracobune ( figs . 1 , 2 , s1 . ) . the last of these features is not known in jozaria .
we exclude ishatherium subathuensis from anthracobunids . its holotype consists of the lingual side of an upper molar , referred originally to sirenia [ 15 ] , [ 92 ] . wells and gingerich [ 1 ] referred ishatherium to anthracobunidae . the holotype and only specimen shares with anthracobunids the strong development of conules and the deep transverse wear along the protocone - paraconule - paracone and along the hypocone - metaconule - metacone . the specimen is decidedly unlike anthracobunids in that the distance between protocone and paracone is similar to that between hypocone and metacone , and in that its cusps are not highly raised . we also exclude nakusia shahrigensis [ 6 ] from anthracobunids ( fig . 2 i , j ) . its holotype is a maxilla with p4 - m2 and the base of p3 . the base of p3 is elongate , the cusps on the relatively unworn m2 are low , and the molars are relatively wide . anthracobunids have a short p3 , high molar cusps , and squarish upper molars . nakusia is more likely a quettacyonid or cambaythere than an anthracobunid .
the genus indobune was included in anthracobunidae [ 4 ] , but it has low cusps on its teeth and is better classified as a cambaythere . ducrocq et al . [ 7 ] described hsanotherium parvum on the basis of two specimens with upper molars from myanmar . these specimens are bunodont , with small conules and lack a hypocone . the transverse wear pattern of anthracobunids is also absent . hsanotherium displays similarities to medium - sized bunodont artiodactyls , such as haqueina [ 22 ] , and we do not include it in anthracobunids .
referred species . anthracobune wardi ( dehm and oettingen - spielberg , 1958 ) .
diagnosis . anthracobunid with narrow lower molars and a very large angular process of the mandible ( fig . 1 , fig . s1 ) .
discussion . we refer to most of the specimens formerly included in lammidhania and pilgrimella in the past to anthracobune .
holotype . bmnh m . 15792 , left m2–3 and right m3 , from ‘ lammidhan ’ and ‘ planorbis freshwater beds . ’ pilgrim never visited the type region , and the fossils were collected by a geological surveyor ( t . g . b . davies ) . lammidhan appears on davies’ unpublished map and is located on a broad alluvial plain . however , it does not match a modern topographical landmark and its meaning is not known to local people . eocene deposits occur to the north and south of the plain , with marine limestones and muds forming ledges usually towering over the unconsolidated muds that form the bottoms of valleys . these muds are the “ planorbis freshwater beds , ” and it is our contention that the specimen may have been found on these beds , but that it rolled down from overlying marine beds . gingerich [ 23 ] improved the holotype by fitting bmnh m . 15794 ( the trigonid of the left m2 ) to it .
description . h - gsp 97106 is the most complete specimen known for any anthracobunid . upper incisors are similar in size ( based on their alveoli ) ; the crown of i3 shows that there is a single , large pointed cusp , with small basal thickening on the cingulum anterior and posterior to it . canine is long and compressed mediolaterally and p1 is two - rooted , with single cusp . p2 has a small protocone and two cuspules on the posterior crest of the paracone . p3 has a protocone , paracone , metacone and two conules , and a strong postprotocrista . upper molars with four main cusps as well as strong para - and metaconule . instead , protocone - paraconule - paracone have aligned wear surfaces , and similar wear surfaces on hypocone - metaconule - metacone . with wear , these surfaces resemble crests .
lower p1 with two roots , strong protoconid , high but narrow paraconid and low metaconid , lacking talonid . lower p2 is similar to p1 but larger . lower p3 and p4 are similar with a well - developed trigonid having three cusps , and two cusps on the talonid . lower molars with strong crests between protoconid and metaconid , and between hypoconid and entoconid ; weak hypoconulid on anterior molars . these crests are stronger than the cristid obliqua . paraconid usually absent on molars .
upper anterior teeth lined up in a parasagittal row , and rostrum ends in a narrow point . the caudal edge of the nasal opening of h - gsp 97106 is over the upper canine , and the infraorbital foramen is dorsal to p3 . the zygomatic arch projects lateral from the base of m1 and m2 , and extends caudally from there . the orbit is located over m2 and m3 . the skull roof dorsal to the orbits is a thick bony plate that is flat and stretches rostrally to the nasal aperture , and caudally to the temporal crests . left and right temporal crests form the caudal extent of this plate and join into a strong sagittal crest that delineates the large temporal fossae . this crest ends at the left and right nuchal crests , which extend ventral to the ear region .
the palate is narrow rostrally , and concave mediolaterally . a weak postpalatine torus is present , and the hard palate is indented caudally with the choana reaching rostrally medial to m3 . the pterygoid process is thick . the periotics are deeply recessed between basioccipital and mandibular fossa . basisphenoid and basioccipital slope strongly from medial to lateral . the postorbital process is strong and oval in cross - section , the mandibular fossa is cylindrical , its rostrocaudal extent is as large as its mediolateral extent . the external auditory meatus is immediately caudal to the postorbital process , there is no postglenoid foramen . the external auditory meatus is directed caudolaterally . the suture between mastoid and squamosal is on the nuchal crest , and there is a mastoid emissary foramen just below it . a large emissary foramen also occurs on the parietal in the temporal fossa . the foramen magnum faces more caudal than ventral and the supraoccipital immediately dorsal to it is flat . near the sagittal crest , the supraoccipital plane is concave .
discussion . specimens referred to a . pinfoldi are known from the upper part of the kuldana formation . these are coastal beds , as indicated by the sedimentology and invertebrate paleontology of their localities , although isotopic evidence implies that the individuals fed on land . the upper part of the kuldana formation is well known for its fossil whales : ambulocetus and attockicetus .
holotype . bmnh m . 15799 , isolated talonid , most likely of m1 based on its size . locality given as ‘ lammidhan … planorbis freshwater beds of chharat stage’ ( see discussion under holotype of anthracobune pinfoldi ) . unlike other specimens from pilgrim ' s ‘ lammidhan , ’ this specimen is probably indeed from the planorbis beds , an inference based on its preservation .
diagnosis . lower molars with high length / width ratio , the m3 short , paraconid and metaconid of lower premolars small , anterior premolars long .
description . new material referred to this species includes two complete lower jaws that include the entire post - incisor dentition ( h - gsp 96258 and 96434 ) , a skull fragment that includes orbit and supraorbital region ( rr 411 ) , and part of the deciduous dentition .
lower incisors are similar in size to each other , based on their alveoli . canines with short crown , larger than the incisors , small diastemata on either side of p1 , but no other diastemata present . lower p1 and p2 with one main cusp and two roots , and small cusp on pre - and postprotocristid . lower p3 and p4 with well - developed trigonid and talonid . protoconid and metaconid on these teeth similar in height and paraconid lower , much lower on p4 . talonid of posterior premolars with high hypoconid , and entoconid high or barely distinguishable . lower molars with distinct trigonid and talonid . protoconid and metaconid similar in height , paraconid absent , but paracristid well developed . hypoconid and entoconid similar in height , cristid obliqua strong , hypoconulid weak on m1 and m2 , but strong and placed on third lobe in m3 , and sometimes twinned . coronoid process of the mandible small and with steep anterior slope . length of mandible anterior to anterior edge of coronoid process ( to rostral end of the fused symphysis ) is approximately two times as long as that posterior from this edge ( to the tip of the angular process ) . this indicates that the angular process is enormous . lower p3 with three cusps on trigonid , and two on talonid ; trigonid long . lower p4 with shorted trigonid , resembling lower m1 ( h - gsp 96052 ) . mandibular angle of juveniles small ( h - gsp 30349 , fig . s1 ) .
upper dentitions of a . wardi are mostly known from india ( luvp 15006 , rr 411 , wif / a 616 ) . the p2 is longer than wide , unlike the p3 and p4 . p2 has connate paracone and metacone , and a lingual bulge with a small protocone ( luvp 15006 , h - gsp 30229 ) . p3 and p4 are similar in shape , with connate para - and metacone , and large protocone . paraconule and metaconule are strong in p3 ( luvp 15006 ) , whereas in p4 transverse crests are well developed and conules less so . upper molars have four strongly developed cusps , with smaller but distinct paraconule and metaconule . transverse crests are absent , but cusps are lined up in such a way that they form an anteriorly convex arch .
tooth wear in anthracobunids tends to be distinct on the posterior side of the protoconid - metaconid , and the anterior side of the protocone - protoconule - paracone , and is accentuated by the deep valleys behind these rows . this transverse wear pattern is consistent with the large and caudally - projecting angular process of the mandible , the area of insertion for the medial pterygoid and masseter muscles .
specimen rr 411 reveals some details about the orbit and zygomatic arch . the zygomatic arch has a weak postorbital process and the dorsal part of the zygomatic arch is made up of the maxilla in the orbital rim , but not posterior to the orbit . the ventral edge of the zygomatic arch is made up of the jugal . the morphology of the root of the zygomatic arch , the postorbital process of the frontal , and the mandibular fossa in this specimen resembles that of h - gsp 97106 .
the anterior edge of the ascending ramus of the mandible of h - gsp 96259 is vertical , whereas it slopes slightly caudal in h - gsp 96434 : the superior part overhanging the inferior part . h - gsp 30349 is a juvenile in which the anterior edge of the ramus slopes slightly rostral . in h - gsp 96149 this ramus is vertical . the mandibular symphysis of all jaws , except h - gsp 30349 , is strongly fused , and slopes caudal ending ventral to the premolars .
discussion . most specimens of anthracobune wardi are known from the lower redbeds of the kuldana formation of pakistan , where pakicetid cetaceans are common . these are freshwater deposits and the specimens of anthracobune found here pertain to a . wardi . additional specimens of a . wardi are from the freshwater deposits of the subathu formation of india .
obergfellia , new genus . urn : lsid : zoobank . org : act : 952704f2 - f029 - 4643 - b791 - b30e7c71aa0a .
anthracobune ( in part ) , west , 1980 , 1981 , 1983 , 1984 .
pilgrimella ( in part ) , wells and gingerich , 1983 ; kumar , 1991 .
etymology . in honor of the late vertebrate paleontologists dr . friedlinde obergfell and her husband a . ranga rao . both were instrumental in the initial discovery and description of the kalakot fauna . this fauna became the best - known eocene land mammal fauna from india , mostly due to the efforts of a . sahni and k . kumar .
obergfellia occidentalis , new species , urn : lsid : zoobank . org : act : b5a9a6da - db8f - 4498 - 8bf2 - 3812b98458d9 .
anthracobune pinfoldi ( in part ) , west , 1980 , p . 518 , pl . 2 . 5 .
anthracobune pilgrimi , west , 1981 ; west , 1983 ( in part ) , p . 367 , fig . 5 , fig . 6 ; west , 1984 ( in part ) , p . 187 , fig . 4 , fig . 5 .
adams consensus of all trees recovered from parsimony analyses that included some ordered multistate characters , with continental geography ( see states in upper left - hand corner ) optimized onto the tree using parsimony ( relationships among extant taxa were constrained by a “molecular scaffold” ) .
bootstrap support for clades derived from each analysis contributing to the consensus is depicted by colored circles . a1 = atlantogenata constraint , transitions between polymorphic and “fixed” states in ordered morphoclines weighted as 0 . 5 steps ; a2 = atlantogenata constraint , transitions between polymorphic and “fixed” states in ordered morphoclines weighted as one step ; e1 = exafroplacentalia constraint , transitions between polymorphic and “fixed” states in ordered morphoclines weighted as 0 . 5 steps ; e2 = exafroplacentalia constraint , transitions between polymorphic and “fixed” states in ordered morphoclines weighted as one step . across all trees , anthracobunids and desmostylians were placed as perissodactyls , along with two enigmatic asian taxa , the late paleocene “condylarth” radinskya and early eocene cambaytherium .
( d ) bar diagram of bone compactness , a quantification of the amount of bone per midshaft cross - section , compared here between fossil and extant ungulates .
c values for enamel samples of early and middle eocene mammals from india and pakistan .
results shown as mean ±s . d . for the sample populations . data from ( a ) early eocene and ( b , c ) middle eocene taxa from india and pakistan . circles , perissodactyls ; red circles , anthracobunids ; squares , artiodactyls ; triangles , cetaceans ; inverse triangles , creodonts ; diamonds , condylarths . see suppl . info . for details .
pilgrimella pilgrimi , wells and gingerich , 1983 ( in part ) , p . 122 , fig . 2c–d ; kumar , 1991 ( in part ) , p . 230 .
etymology . specific epithet is from occidentalis , latin for west , in honor of robert m . west , who discovered and described the holotype .
holotype . h - gsp 1981 , mandibles with left p1 - m3 and right p4 - m3 ( fig . 2p–q ) .
type locality . h - gsp locality 62 , ganda kas area , punjab province , pakistan .
formation and age . kuldana formation , kala chitta hills of northeastern pakistan , early lutetian in age ( ≈48 ma ) .
diagnosis . differs from other anthracobunids in exhibiting the following combination of features : ( i ) lower molars broad , ( ii ) lower m3 short , and ( iii ) angular process of the mandible long but shorter than that of anthracobune .
description . the most complete specimen for this species is h - gsp 1981 , a left and right mandible not larger than a . wardi ( h - gsp 96258 and 96434 ) , but with lower molars much broader than a . wardi . h - gsp 1981 lacks the angular process , due to post - mortem damage but it is preserved in h - gsp 96214 , a specimen with an erupting m3 . in this specimen , the angular process is 80 % as long , rostro - caudally , as in a . wardi . the angular process is not preserved in h - gsp 96149 , but its root is preserved and the slope thereof indicates that the process is small h - gsp 96149 has extremely worn teeth , but its molars are similar in size to h - gsp 1981 ( fig . s1 ) , premolars and canine are larger and the depth and robusticity of the jaw is much greater than in h - gsp 1981 . the wear stage is most consistent with advanced age , and makes it unlikely that the smaller angular process is the result of young age . eventually this specimen may be shown to pertain to a new species of obergfellia .
few upper molars have been discovered for o . occidentalis . h - gsp 538 is a large tooth , and an interstitial facet on its posterior side indicates that it is not an m3 . m1 and m2 of p . pilgrimi do not reach this size , and we therefore refer this tooth to p . obergfelli .
discussion . west [ 10 ] , [ 93 ] recognized that his collection of specimens from the ganda kas region of pakistan contained two species of anthracobunids from the freshwater part of the formation and used the names anthracobune pilgrimi and lammidhania wardi for them . unfortunately , the type specimen for l . wardi is an isolated trigonid , and pertains to the same species as several large complete lower jaws collected by us ( h - gsp 96258 and 96434 ) , and these match upper dentition material referred to pilgrimella pilgrimi ( the type of which is an upper molar ) . hence a new genus and species name is needed . a specimen described by west , h - gsp 1981 , is the most appropriate holotype for this species .
discussion . wells and gingerich [ 1 ] described jozaria palustris on the basis of a single specimen which shows the diagnostic features of anthracobunids . the lower premolars of this specimen are large compared to the molars , confirming it as distinct from other anthracobunid species and genera .
parsimony analyses were first run to determine the most parsimonious placement of xenarthra ( represented in this analysis by dasypus ) given different placements in placental phylogeny – i . e . , as either the sister group either of afrotheria ( the atlantogenata hypothesis ) or of laurasiatheria + euarchontoglires ( the exafroplacentalia hypothesis ) , as this is one of the nodes that was not convincingly resolved by meredith et al . ' s [ 27 ] analysis , and continues to be debated in the literature on placental phylogenetics [ 94 ] . a placement with afrotheria was more parsimonious [ ( 3704 . 5 versus 3716 steps ( i . e . , constraint a1 versus e1 , with 0 . 5 weighting of ordered multistate characters with intermediate polymorphic states ; fig . s2a versus fig . s2c ) and 5758 versus 5773 steps ( a2 versus e2 , no weighting of ordered multistate characters with intermediate polymorphic states , fig . s2b versus fig . s2d ) ] , and all subsequent alternative constraints accordingly employed an atlantogenata , rather than an exafroplacentalia , constraint .
the placement of cambaytherium within perissodactyla is consistent with bajpai et al . ' s [ 38 ] , [ 91 ] assessment of the genus . confusion surrounding the dental similarities that cambaytherium shares with perissodactyls and anthracobunids [ 102 ] is resolved by the placement of cambaytheriids and anthracobunids as closely related sister taxa of crown perissodactyla , thereby indicating that similarities to tethytheres are entirely due to convergence . bajpai et al . [ 38 ] argued that the middle eocene european genus hallensia , which was originally identified as a “condylarth” [ 50 ] and later placed within perissodactyla as either a possible equoid [ 51 ] , [ 103 ] , might be a cambaytheriid . in our analyses hallensia was usually placed as a stem perissodactyl ( a1 , e1 , e2 ) or in an unresolved position relative to these taxa ( a2 ) . the monophyly of a clade containing anthracobunidae , cambaytherium , hallensia , radinskya , and “advanced” perissodactyls is supported by 11 unambiguous synapomorphies ( table s2 ) . the clade containing anthracobunids , cambaytheriids , radinskya , crown perissodactyla , and hallensia was reconstructed as having been equivocally of either asian or european origin ( table s3 ) , while the anthracobunid - desmostylian - hippomorph - tapiromorph clade was optimized as having been of asian origin .
we quantified the midshaft cross - sectional geometries of limb and rib bones of anthracobunids and other ungulates . long bone and rib cross - sectional phenotypes are a reliable indicator of vertebrate habitat , with most taxa occupying a shallow - water habitat displaying the greatest amount of bone per cross - section via thickened cortices and / or bone - filled medullary cavities [ 71 ] , [ 104 ] , [ 105 ] . in contrast , bones of terrestrial taxa typically display thin cortices and vacant medullary cavities . bone compactness values , a combined measure of cortical and medullary bone area [ 72 ] , indicate that anthracobune limb bones display values between 0 . 85–0 . 96 . this range is greater than values observed in a sample of fossil and extant artiodactyls and perissodactyls ( 0 . 42–0 . 83 ) , with the exception of modern hippopotamus ( 0 . 94 , 0 . 96 ) and rhinoceros ( 0 . 64–0 . 85 ) , and semi - aquatic pakicetid and remingtonocetid eocene cetaceans ( 0 . 93–0 . 96 ) ( fig . 4 ) . similar results were documented in rib elements ( table s7 ) .
we also studied the oxygen and carbon isotope composition of structural carbonate in tooth enamel [ 75 ] , [ 106 ] for anthracobunids and coeval taxa from the early and middle eocene of india and pakistan ( fig . 5 , table s8 , table s9 ) . mammal fossils sampled from early eocene localities ( fig . 5c ) , including cambaytherium , spanned a narrow range of δ 18 o values , and are consistent with occupation of a wet and forested habitat . enamel δ 13 c values showed more than 3‰ range in individual values ( −12 . 4‰ to −9 . 1‰ ) , and , when corrected to present - day atmospheric δ 13 c values [ 107 ] , suggest foraging in a relatively wet environment . conversely , taxa sampled from middle eocene deposits of northern pakistan displayed a much wider range of δ 18 o values ( 21 to 27‰ ) and much higher enamel δ 13 c values ( fig . 5b , c ) .
two specimens of a . wardi collected from middle eocene deposits of northern india had low enamel δ 18 o values ( 21 . 3‰ and 21 . 8‰ ) that approached values for aquatic taxa , but also overlapped with values for terrestrial rhinocerotoids ( 20 . 6‰ to 24 . 7‰ ) from the same formation . these low enamel δ 18 o values are more consistent with a freshwater habitat for these individuals , but , again , low sample size complicates differentiating δ 18 o values for these specimens from δ 18 o values for taxa assumed to be fully terrestrial .
anthracobunids have not previously been incorporated into taxonomically broad phylogenetic analyses of morphological data that incorporate the molecular evidence for placental supraordinal relationships . we tested for the possibility that anthracobunidae and various laurasian “condylarths” ( phenacodontids , louisinids ) are not members of afrotheria by adding a number of living and extinct laurasiatherians to a morphological character matrix that has previously been employed to reconstruct afrotherian phylogeny [ 19 ] , [ 35 ] , [ 108 ] . we constrained relationships of extant taxa , based largely on the results of meredith et al . [ 27 ] , using the “molecular scaffold” technique proposed by springer et al . [ 109 ] . we also employed several alternative constraints to obtain tree lengths for competing topologies .
when continental geography is mapped onto the adams consensus in fig . 3 , crown and advanced stem perissodactyls ( including cambaytherium and radinskya ) are optimized as having originated in asia ( fig . 3 ) , while the perissodactyl stem lineage is of ambiguous ( but certainly laurasian ) continental origin . later colonization of north america by crown perissodactyls was presumably facilitated by global warming at the paleocene - eocene boundary [ 111 ] , [ 112 ] . afrotheria is unambiguously of afro - arabian origin on the adams topology , and the recently described paleocene african “condylarth” ocepeia [ 58 ] was placed as either a stem paenungulate or as a stem atlantogenatan , depending on whether atlantogenata or exafroplacentalia was constrained ( respectively ) . the consistent recovery of the european louisinids paschatherium and teilhardimys as stem perissodactyls challenges the view that they are afrotherian macroscelideans [ 62 ] .
some early tethytheres are thought to have had a semi - aquatic lifestyle ( proboscideans [ 81 ] , [ 116 ] , desmostylians [ 117 ] , sirenians [ 104 ] , [ 118 ] – [ 120 ] ) , but no evidence suggests that stem perissodactyls occupied an aquatic habitat . to reconstruct the ancient habitat preferences of anthracobunids , we analyzed bone geometry of the postcranial skeleton as well as stable isotope values within tooth enamel .
we quantified the midshaft cross - sectional geometries of limb and rib bones of anthracobunids and other ungulates . anthracobune limb bones displayed bones that were more hyperostotic and therefore more compacted compared to most extant ungulates . exceptions included extant taxa that are known to frequently swim or wallow in shallow water habitats ( i . e . , hippopotamus and rhinoceros ) [ 121 ] , and fossil eocene cetaceans that are thought to be semi - aquatic and also occupy shallow water habitats ( i . e . , pakicetid and remingtonocetids ) ( fig . 4 ) . thus , anthracobunids display a variety of skeletal modifications consistent with taxa that exploit shallow - water habitats , including thickened postcranial elements and widened elements of the autopodium , in addition to large body size .
we also studied the oxygen and carbon isotope composition of structural carbonate in tooth enamel [ 75 ] , [ 106 ] for anthracobunids and coeval taxa ( fig . 5 ) . the anthracobunids recovered from the kuldana formation ( anthracobune and obergfellia ) displayed enamel δ 18 o values suggesting some evidence of aquatic habits , and enamel δ 13 c values showed they fed on land . anthracobunids recovered from northern india displayed enamel δ 18 o values consistent with occupation of freshwater habitats . although all sampled anthracobunids may have had some aquatic preferences , stable isotopic evidence alone is inconclusive as there is overlap in range of isotopic values between terrestrial and aquatic taxa .
taken together , skeletal and isotopic evidence are most consistent with an interpretation of anthracobunids ( especially anthracobune ) as having had ecological preferences similar to those of modern rhinos and tapirs , but not exhibiting the same degree of commitment to aquatic habits as hippopotamus [ 84 ] , [ 122 ] . most rhinos and tapirs have a restricted range near permanent bodies of water in which they frequently wallow and wade [ 121 ] , and obtain most of their drinking water [ 84 ] , [ 122 ] . modern rhinos may be a good model for the anthracobunid lifestyle as both display thickened limbs and intermediate oxygen isotope values relative to hippopotamus and terrestrial ungulates .
stable isotopic evidence also showed mammal fossils sampled from early eocene localities ( fig . 5c ) occupied and foraged within a wet and forested habitat . conversely , taxa sampled from middle eocene deposits of northern pakistan ( fig . 5b , c ) indicated shifts in the climate toward increased aridity . this climate change was most likely caused by the northward movement of the indian plate , which pushed these areas out of an equatorial zone of high precipitation in the middle eocene [ 123 ] , [ 124 ] . vegetation would have opened up , and shifted to a drier , tropical forest or savanna habitat .
based on new dental , cranial , and postcranial material , our phylogenetic analysis is the first to suggest that anthracobunids are stem members of the laurasiatherian order perissodactyla , rather than members of the afrotherian clades tethytheria , proboscidea , or sirenia . the strictly asian distribution of anthracobunids is consistent with the isolation of afrotherian and laurasiatherian clades through the early paleogene [ 19 ] , [ 113 ] , while the phylogenetic result is consistent with the recent recovery of very primitive basal stem proboscideans such as phosphatherium [ 125 ] and eritherium [ 18 ] , from the early eocene and paleocene of africa . analyses of postcranial bones and stable isotopes also indicate that anthracobunids were large and lumbering , and suggest at least some taxa spent time in the water similar to modern rhinos , and that all known anthracobunids fed on land . our results therefore identify an old world radiation of large , non - cursorial , partly aquatic perissodactyls that convergently came to occupy a basal tethythere - like niche on the northern coast of the tethys sea .


the german cockroach ( blattella germanica ) is a small species of cockroach , typically about 1.1 to 1.6 cm ( 0.43 to 0.63 in ) long .

in colour it varies from tan to almost black , and it has two dark , roughly parallel , streaks on the pronotum running anteroposteriorly from behind the head to the base of the wings .

although blattella germanica has wings , it can barely fly , although it may glide when disturbed .

of the few species of cockroach that are domestic pests , it probably is the most widely troublesome example .

it is very closely related to the asian cockroach , and to the casual observer the two appear nearly identical and may be mistaken for each other .

however , the asian cockroach is attracted to light and can fly rather like a moth , while the german cockroach can not .

german cockroach

german cockroach diseases german cockroach life cycle german cockroach nests german cockroach nymphs german cockroach vs . american cockroach
insect management guide for cockroaches . least toxic methods of cockroach control . german cockroach management in low income housing
the german cockroach is the most common species of the cockroach . german cockroaches can breed at a rate of up to six generations per year . the german cockroach can fit through an opening as small as 3 / 8 inch in width .
german cockroaches can be found all over the world . they are the most common cockroach in the united states . each german cockroach can live about 100 - 200 days .
the german cockroach eats almost anything but prefers starchy foods like potatoes , rice and cereal .
i saw one small german cockroach in my home and now i am worried . how long before they overrun my home ? keep reading to learn how fast german cockroach reproduction occurs .
the german cockroach , which originally came from africa , is the smallest of the pest species .
differential development and reproduction of the german cockroach ( dictyoptera : blattellidae ) on three laboratory diets .
the american cockroach is the largest cockroach found in houses . despite its name , the american cockroach is not native to north america , but was likely introduced via ships from africa in the 1600s .
prof essional roach baits , pesticide dusts , aerosols or residual spray concentrates can be used to eliminate german cockroach infestations .
differential development and reproduction of the german cockroach ( dictyoptera : blattellidae ) on three laboratory diets . - pubmed - ncbi
wondering how to get rid of german cockroaches ? the best advice for german cockroach control is to practice good sanitation . to prevent german cockroaches from infesting the space , vacuum often , keep a spotless kitchen , seal all entrances around utility pipes and ventilate crawl spaces to prevent moisture buildup . if there is evidence of a cockroach infestation , contact a licensed pest professional to inspect and treat the german cockroach problem .
robinson , w . 1999 . worries over german cockroach resistance fades . pest control 67 ( 7 ) : 40 - 42
the smallest cockroach known is attaphila fungicola , measuring 1mm wide and 3mm long . this cockroach lives in the nest of leaf cutter ants .
the german cockroach is the cockroach of concern , the species that gives all other cockroaches a bad name . it occurs in structures throughout florida , and is the species that typically plagues multifamily dwellings . in florida , the german cockroach may be confused with the asian cockroach , blattella asahinai mizukubo . while these cockroaches are very similar , there are some differences that a practiced eye can discern .
figure 4 . third instar nymph of german cockroach blattella germanica ( linnaeus ) . photograph by james castner , university of florida .
rust mk , owens jm , reierson da . 1995 . understanding and controlling the german cockroach . oxford university press , oxford .
nalyanya g . , gore j . c . , linker h . m . , schal c . german cockroach allergen levels in north carolina schools : comparison of integrated pest management and conventional cockroach control .
robinson , w . 1996 . resistance remediation in the german cockroach . resistant pest management 8 ( 1 ) : 34 - 35 .
wang c . , bennett g . w . comparative study of integrated pest management and baiting for german cockroach management in public housing .
figure 1 . adult female german cockroach , blattella germanica ( linnaeus ) , with ootheca . photograph by james castner , university of florida .
figure 6 . adult male german cockroach , blattella germanica ( linnaeus ) . photograph by p . g . koehler , university of florida .
figure 7 . adult female german cockroach , blattella germanica ( linnaeus ) . photograph by p . g . koehler , university of florida .
tsai , y . , k . cahill . 1970 . parasites of the german cockroach ( blattella germanica l . ) in new york city .
figure 2 . oothecae ( egg cases ) of the german cockroach , blattella germanica ( linnaeus ) . photograph by james castner , university of florida .
the german cockroach is also the most sociable of the pest species and is often found in large groups , especially near warm areas like water heaters .
check out the tips in this video to help you prevent and treat cockroach infestations .
find more facts about and information on cockroach pest control at the official npma website .
ithe german cockroach is the hardest pest cockroach to get rid of and is the most troublesome in sydney . it can be found anywhere in the home but is probably most common in the kitchen , frequently inside cupboards , drawers and electrical appliances .
robinson , w . 1996 . how the german cockroach has adapted to the household environment . pest control 64 ( 7 ) : 43 , 44 , 46 .
2004 .
german cockroach - blattella germanica
( on - line ) . north carolina museum of natural sciences . accessed october 18 , 2008 at urltoken .
the world ' s smallest cockroach is only 0 . 3mm long and lives in ant nests
cornwell pb . 1968 . the cockroach , volume i . hutchinson of london , london .
survey - to control german cockroaches , it is important to do a thorough inspection . a cockroach survey ( trapping ) is sometimes necessary to determine the extent of an infestation , as even a thorough inspection will not reveal all cockroach harborages or foraging areas .
the german cockroach is one of the most common roaches found in apartment houses , restaurants , and hotels . german cockroaches ( eggs included ) , are brought inside or hitchhike on man ' s belongings , luggage , boxes or packages . a large german roach population will produce a foul odor . these roaches prefer moisture , food , and warmth . sanitation measures are critical in controlling german roaches . german roaches can be a serious health hazard in restaurants .
fasulo tr . ( 2002 ) . stored product pests and german cockroach . bug tutorials . university of florida / ifas . cd - rom . sw - 165 .
chai r . y . , lee c . y . insecticide resistance profiles and synergism in field populations of the german cockroach ( dictyoptera : blattellidae ) from singapore .
if you have ever seen a particularly long cockroach , then you may have seen a female carrying its egg case on its abdomen . the egg case , an ootheca , of a german cockroach typically carries 30 to 40 eggs . the german cockroach will carry this egg case up to nearly the time when the eggs are ready to hatch , and once the new baby roaches hatch , they are initially bright , white nymphs .
the german cockroach is found throughout the world in association with humans . they are unable to survive in locations away from humans or human activity . the major factor limiting german cockroach survival appears to be cold temperatures . studies have shown that german cockroaches were unable to colonize inactive ships during cool temperatures and could not survive in homes without central heating in northern climates . the availability of water , food , and harborage also govern the ability of german cockroaches to establish populations , and limit growth .
droppings german cockroach droppings may appear as small , dark , “pepper - like” material left on countertops or in drawers . fecal staining may appear as dark spots or smears :
egg capsules since german cockroach females carries their egg case until 1 to 2 days before depositing it , empty egg cases may be found in areas that the females frequent .
snell , e . and w . h robinson . 1991 . bait stations and german cockroach management programs . pest control technology 19 ( 8 ) : 55 - 57 .
robinson , w . 1996 . more good news from the field ( german cockroach resistance ) . pest control technology 24 ( 6 ) : 56 , 61 , 62 .
watch this video to learn more about the most common cockroach species that can infest your home .
koehler pg , strong ca , patterson rs . 1994 . rearing improvements for the german cockroach ( dictyoptera : blattellidae ) . journal of medical entomology 31 : 704 - 710 .
zhai , j . and w . h robinson . 1992 . chemical control of the german cockroach . pest control technology 20 ( 3 ) : 40 , 41 , 44 .
since the german cockroach is considered an aesthetic pest , the action threshold for this insect depends upon the tolerance of the people living in the infested dwelling . however , most people associate cockroach infestations with poor sanitary conditions and typically go to excessive lengths to eradicate them from their houses .
the german cockroach is a widely distributed urban pest . it is also the most common cockroach species in houses , apartments , restaurants , hotels , and other institutions . this is true not only in pennsylvania but also throughout the united states and in most parts of the civilized world .
figure 3 . first instar nymphs emerging from the oothecae ( egg case ) of a german cockroach , blattella germanica ( linnaeus ) . photograph by james castner , university of florida .
the german cockroach ranges in size from about one - eighth of an inch in length when it hatches , to a little over one - half of an inch as an adult .
zhai , j . and w . robinson . 1996 . antennal and leg grooming in two strains of the german cockroach ( orthoptera : blattellidae ) . med . entomol . zool .
mccandless , l . 2005 .
cu scientists unravel mating clues of the german cockroach
( on - line ) . cornell chronicle . accessed november 14 , 2008 at urltoken .
lifespan as adults , german cockroaches can survive anywhere from 100 to 200 days .
figure 5 . newly molted adult german cockroach , blattella germanica ( linnaeus ) . within a few hours the cuticle will harden and darken . photograph by james castner , university of florida .
robinson , w . h and j . zhai . 1992 . insecticide rotation and german cockroach control . pest control technology 20 ( 12 ) : 24 , 28 , 30 . 31 .
robinson , wm . h and j . zhai . 1992 . insecticide rotation and german cockroach control . pest control technology 20 ( 12 ) : 24 , 28 , 30 , 31 .
german cockroaches produce odorous secretions that can affect the flavor of various foods . when cockroach populations are high , these secretions may result in a characteristic odor in the general region of the infestation . disease - producing organisms such as bacteria , protozoans , and viruses have been found on cockroach bodies .
when beginning your german roach control program , sanitation is one of the first things to consider . german roaches only need a small amount of food to sustain them .
nature has provided german roaches the means to ensure their survival in a splendid way . german roaches protect their eggs to help ensure hatching and reproduce at astounding rates .
german cockroaches are one of the most common cockroach species found in households . german cockroaches undergo three distinct life phases : egg , nymph and adult . their entire life cycle spans approximately 100 days , although this is dependent on factors such as temperature , diet and injuries .
asian cockroaches , a pest of the southeastern united states , are often mistaken for the german cockroach . similar in appearance , the main differences between the two cockroaches are evident in their behavior .
durier v , rivault c . 2003 . improvement of german cockroach ( dictyoptera : blatellidae ) population control by fragmented distribution of gel baits . journal of economic entomology 96 : 1254 - 1258 .
phantom aerosol may be used with roach baits and will enhance your german roach control .
breeding season german cockroaches breed continuously year round , though breeding slows during colder months .
advion arena roach bait advion cockroach bait arena can be used to control cockroach populations indoors and outdoors with the active ingredient indoxacarb , an insecticide that acts through ingestion by cockroaches . advion roach bait arena ( 60 ctn )
the german cockroach was once the no . 1 pest in residential and commercial buildings . that changed with the introduction of cockroach baits , which resulted in one of the great success stories in insecticide development . cockroaches became almost a second tier pest , and remained so for more than a decade . but have you noticed ? they’re back ! it’s time for pest management service technicians to review some of the critical aspects of german cockroach biology and habits that can make it one of our more challenging pests .
zhai , j . and w . h robinson . 1992 . measuring cypermethrin resistance in the german cockroach ( orthoptera : blattellidae ) . j . econ . entomol . 85 : 348 - 351 .
] . the german cockroach has been reported to have developed resistance to 42 unique insecticide active ingredients in 219 documented cases worldwide and is ranked as the world’s no . 2 insecticide resistant urban pest [
day , e . august 1996 .
german cockroach .
( on - line ) . virginia cooperative extension , virginia polytechnic institute and state university . accessed october 14 , 2008 at urltoken .
valles , s . 2008 .
german cockroach
( on - line ) . university of florida institute of food and agricultural sciences – featured creatures . accessed october 10 , 2008 at urltoken .
highly magnified footage of these experiments clearly shows a glucose - averse cockroach reacting to a dose of the sugar .
in addition to being a nuisance , the german cockroach has been implicated in outbreaks of illness and allergic reactions in many people . cockroaches have been reported to spread at least 33 kinds of bacteria , six kinds of parasitic worms and at least seven other kinds of human pathogens . they can pick up germs on the spines of their legs and bodies as they crawl through decaying matter or sewage and then carry these into food or onto food surfaces . medical studies have shown that german cockroach allergens cause allergic reactions and can exacerbate asthma attacks , especially in children . this makes german cockroach control incredibly vital .
reproduction & life cycle the female german cockroach produces an egg capsule every 3 to 4 weeks , with each capsule containing 25 - 45 eggs . rather than depositing the sack , the female carries the capsule with her and protects it until it hatches . this is another reason the german cockroach is so difficult to get rid of - its nymphs are well protected . these young nymphs will begin breeding again in as little as 36 days . adult german cockroaches can live up to 365 days .
female german cockroaches carry the oothecae attached to their abdomens until about two days prior to hatching and then deposit them in a protected location . oothecae may be seen protruding from the abdomens of german cockroach females . nymphs emerge from the oothecae as tiny insects . they gradually darken into dark brown or black cockroaches with parallel lines visible upon the pronotum . german cockroach nymphs are wingless and incapable of reproduction . nymphs molt six to seven times and can develop completely within 100 days under optimal conditions .
not applying insecticides effectively . whether baits , liquids , dusts , or other formulations , cockroach insecticides need to be applied in cracks , crevices , voids , and other sites where german cockroaches are aggregating .
zungoli , p . a . and w . h robinson . 1984 . feasability of establishing an aesthetic injury level for german cockroach pest management programs . environ . entomol . 13 : 1453 - 58 .
a flashlight would be a handy tool to inspect evidences of german roaches the following places .
wings : while adult german cockroaches have wings , they rarely fly , preferring to run .
robinson , w . 1996 . antennal grooming and movement behavior in the german cockroach , blattella germanica ( l . ) . proc . internat . conf . insect pests urban environment 1996 : 361 - 369 .
not rescheduling problem apartments . when cockroach populations become large , service needs to be repeated every few weeks to succeed .
the largest winged cockroach in the world is megaloblatta blaberoides from costa rica ( wingspan of 7 . 3 inches ) .
maxforce® forte contains fipronil 0 . 03 % along with gel that delivers quick results for treatment of cockroach . . .
the german cockroach is a highly pestiferous insect that produces asthma - inducing allergens , and is closely associated with areas of human habitation . as such , the german cockroach is transported through human activities , such as food and equipment dispersal . in connected apartments and buildings , german cockroaches can also move between units , making control difficult in heavy infestations . it has previously been shown that the bacterial flora transmitted by german cockroaches presents a serious epidemiological problem for human health and well - being , especially to those living in low - income housing . because german cockroaches are able to vector antibiotic - resistant bacteria , its presence must be controlled to reduce the threat of bacterial contamination in hospitals and swine farms .
for effective control , cockroaches must prefer to eat the bait product over surrounding , non - toxic foods . that means cleaning the dishes in the sink , removing trash in a timely fashion , properly storing food , cleaning spills quickly , and identifying potential water sources ( such as leaky pipes ) . cleaning and sanitation has been shown to be effective at reducting german cockroach population numbers , and reducing cockroach allergen levels . reduce clutter to reduce possible cockroach harborages .
the german cockroach , blattella germanica ( l . ) , is a common indoor cockroach species . infestations of this pest are associated with poor sanitation , particularly in and around food - handling facilities , and also tend to be associated with lower socioeconomic status . cockroach infestations lead to food damage and contamination because cockroaches can vector human and domesticated animal pathogens . cockroach feces , saliva and cast skins contain allergens that may trigger allergic reactions and psychological distress in sensitive individuals [ 46 ] . cockroaches are among the most problematic urban pests in initiating asthmatic and allergic reactions in children [ 47 ] .
farmer , b . r . and w . h robinson . 1984 . harborage limitation as a component of a german cockroach pest management program . proc . entomol . soc . wash . 86 : 269 - 73 .
teachers can find additional information on german cockroaches to share with students at the official npma website .
the adult stage begins with the last successful molting . at this point , german cockroaches are approximately 15 mm in length and are winged . adult german cockroaches are nocturnal insects that hide during the day and scavenge at night . despite their fully developed wings , german cockroaches very rarely fly .
3 . german cockroaches spend most of their lives hidden and protected . for every cockroach in view in daytime , hundreds can be hidden inside wall and ceiling voids , behind cabinets , inside appliances , and the like . a german cockroach’s favorite location is inside a narrow crevice with its antennae extended out . typical “prime” locations are the cracks between cabinets and walls , the flange under the kitchen sink , and the space between door corners and jambs .
zhai , j . and w . h robinson . 1996 . instability of cypermethrin resistance in a field population of the german cockroach ( orthoptera : blattidae ) . j . econ . entomol . 80 : 332 - 336 .
gondhalekar a . d . , scherer c . w . , saran r . k . , scharf m . e . implementation of an indoxacarb susceptibility monitoring program using field - collected german cockroach isolates from the united states .
zhai , j . and w . h robinson . 1991 . pyrethroid resistance in a field population of german cockroach , blattella germanica ( l . ) . jpn . j . sanit . zool . 42 : 241 - 244 .
ajjan , i . and w . h robinson . 1996 . measuring hydramethylnon resistance in the german cockroach , blattella germanica ( l . ) . proc . internat . conf . insect pests urban environment 1996 : 135 - 144 .
robinson , w . 1999 . the changing pest status of the german cockroach in the urban environment . proceedings of the fifty second new zealand plant protection conference , new zealand plant protection society inc . , 1999 : 16 - 21
german cockroaches breed continually . in a lifetime , a female cockroach is capable of producing almost 400 eggs . populations grow quickly in optimal conditions . a typical thriving population is comprised of 75 percent nymphs and 25 percent adult roaches .
german cockroaches do not have a specific home range . they usually dwell in homes and in garbage .
if insecticides have been used in a building , there is a possibility that the german cockroach simply scattered to another area . these harborages need to be located and treated . it may not be possible to eliminate all the german cockroaches in a building completely if a consistent supply of cockroaches is carried into the premises via packages or food shipments .
what are the most common areas in the home where cockroaches are found ? dr . jim fredericks discusses the most popular cockroach hangouts .
robinson , w . h and j . zhai . 1993 . insecticide program to manage a cypermethrin - resistant population of the german cockroach , blattella germanica ( l . ) . resistance pest managt . 5 ( 1 ) : 21 - 23
miller , d . m . , koehler , p . g . 2000 . novel extraction of german cockroach ( dictyoptera : blattellidae ) fecal pellets enhances efficacy of spray formulation insecticides . j . econ . entomol . 93 : 107 - 111
orkin can provide the right solution to keep german cockroaches in their place…out of your home , or business .
if you think you might have german roaches , then don’t wait ! call bug zapper pest control for a free inspection . let our trained technicians accurately identify possible german roaches and help you find the best solution .
temprid is a combination product suitable as residual spray for the control of american cockroaches and german . . .
miller , d . m . , koehler , p . g . 2000 . trail following in the german cockroach , blattella germanica ( l . ) ( dictyoptera : blatellidae ) . . j . econ . entomol . 93 : 1241 - 1246
thoms , e . m . and w . h robinson . 1987 . potential of the cockroach oothecal parasite prosevania punctata ( hymenoptera : evaniidae ) as a biological control agent for the oriental cockroach ( orthoptera : blattidae ) . environ . entomol . 16 : 938 - 944 .
german cockroaches will feed on almost anything , including soap , glue and toothpaste . german cockroaches are good hitchhikers and often find their way into new structures via grocery bags , cardboard boxes , drink cartons and secondhand appliances .
the german cockroach is the most successful of the species infesting buildings in pennsylvania . there are several reasons for this cockroach’s persistence and the difficulty of controlling it . german cockroaches produce a larger number of eggs per capsule and they undergo the shortest time from hatching until sexual maturity , resulting in a rapid population growth . a greater number of nymphs hatch successfully because the female carries the egg capsule during the entire time the embryos are developing within the eggs . also , and most importantly , german cockroaches are smaller than most other cockroaches and can conceal themselves in many places inaccessible to individuals of the larger species .
5 . german cockroaches protect their egg cases . the female german cockroach carries her egg case for as long as a month , dropping it just before it hatches . while she is carrying the egg case , the female german cockroach is less active and tends to stay hidden away in cracks and crevices and protected voids . for this reason , when applying residual insecticides , it is especially important to ( 1 ) treat deep inside cracks , crevices , and voids , and ( 2 ) schedule follow - up treatments from a few weeks to a month later in problem locations to control newly - emerged nymphs .
tekko pro stops the reproduction and reduces the survival of the german roach with two different modes of action . it effects the molting process , making the german roach vulnerable to survival and it stops it at the larvae state .
besides physiological resistance to bait insecticides , german cockroaches have developed behavioral resistance to various phagostimulants of bait formulations , typically d - glucose and d - fructose [ 65 , 66 , 67 ] . the glucose aversion in field german cockroach populations resulted in the failure of attracting cockroaches to toxic baits and protected them from receiving lethal doses of insecticides [ 65 , 66 ] .
the german cockroach has three life stages typical of insects with incomplete metamorphosis : the egg , nymph , and adult . the entire life cycle is completed in about 100 days . however , factors such as temperature , nutritional status , and strain differences may influence the time required to complete a life cycle . german cockroaches breed continuously with many overlapping generations present at any one time . under ideal conditions , population growth has been shown to be exponential . actively growing field populations are comprised of 80 percent nymphs and 20 percent adults . the german cockroach is omnivorous , eating table scraps , pet food , and even book bindings .
for german roach control in restaurants we recommend the use of insecticides , aerosols , dusts , and insect growth regulators instead of baits . there is too much competing food sources for effective control of german roaches in restaurants with bait .
many times , german roaches in restaurants require insecticides as opposed to baiting . using baits in german roach control requires that their is no competing food sources . in restaurants , it is almost impossible to eliminate all competing food sources .
robinson , w . h 1988 . bait stations in cockroach pest management programs . pest control 56 ( 8 ) : 56 , 57 , 60 .
general the german cockroach is the most common roach found in apartments , houses , hotels , and restaurants . german cockroaches and their eggs are usually transported into buildings on a person ' s belongings , luggage , or boxes . it only takes one small egg capsule and you have an infestation six months later . this species is also one of the most challenging to get rid of .
dust should only supplement any insecticide of bait control program for german roach control , not as a stand alone solution .
adult german roaches can live up to one year . they adapt quickly to changing environments , and evolve to survive .
as they are quite abundant , german cockroaches are not considered a species of concern in any part of their range .
german cockroaches live in warm and damp places , like kitchens , bathrooms , and places where people eat and drink .
4 - eliminating harborages german roaches prefer a tight crack or crevice , or a dark wall void in which to hide out of sight during daylight areas . eliminate as many harborages as you can and german roaches will hit the road .
german cockroaches adulterate food or food products with their feces and defensive secretions , physically transport and often harbor pathogenic organisms , may cause severe allergic responses , and in extremely heavy infestations have been reported to bite humans and feed on food residues on the faces of sleeping humans . in addition , some scientists suggest that german cockroach infestations may cause human psychological stress and that the stigma associated with infestations alters human behavior . for example , people with infested houses do less entertaining , and avoid the kitchen at night for fear of encountering a cockroach .
eliminating nesting sites by improving sanitation and reducing clutter and the elimination of food and water sources is imperative to eliminate german cockroach infestations and will help protect you from future infestations . you can also help prevent infestations by sealing any areas where roaches could enter from other units in multi - unit buildings and by checking items such as shipping boxes , school bags and grocery bags for cockroach hitchhikers before bringing them into your home .
the german cockroach is a small , noctural , dime / penny sized , fast moving cockroach that can quickly escape into cracks and crevices in the wall . adults are elongated , light carmel - colored brown , with two longitudinal black stripes running along the pronotum . adult females may be seen carrying an egg case ( shown to the right ) . nymphs are darker brown / black colored , smaller , and oval shaped .
miller , d . m . , meek , f . 2004 . cost and efficacy comparison of ipm strategies with monthly spray insecticide applications for german cockroach ( dictyoptera : blattellidae ) control in public housing . . j . econ . entomol . 97 : 559 - 569 .
miller , d . m . , koehler , p . g . , nation , j . l . 2000 . fecal extract trails enhance trap catch in german cockroach ( dictyoptera : blattellidae ) monitoring stations . j . econ . entomol . 93 : 865 - 870 .
the process of natural selection would strongly favour any chance genetic change that caused a cockroach to avoid the bait and therefore death . since individuals with the trait would have a greater chance of surviving and reproducing , their descendants with the trait would in time replace those that lacked the trait in the cockroach population .
tee h . , lee c . sustainable cockroach management using insecticidal baits : formulations , behavioral responses and issues . in : dhang p . , editor .
in addition to robotic intervention , there are several steps that people can take to reduce or eliminate cockroach populations . we ' ll look at these next .
the german cockroach can move well within a building . they also can travel from a neighboring apartment or location to another and can pass through small openings like light switch plates . many times they are brought in with grocery items , grocery bags , cartons , handbags , and luggage .
there are about 4 , 600 species of cockroach and fewer than 30 of these are considered pests . ( there are about 5 , 400 species of mammals )
avert dry flowable cockroach bait remains effective for more than one year after initial treatment . the dry formula reaches void areas that gels and bait stations cannot reach .
zhai , j . and w . h robinson . 1991 . cockroach distribution in urban buildings . pest control technology 19 ( 2 ) : 36 - 39 .
cypermethrin insecticides are the most popular for killing german cockroaches . mop up can also be used when mopping kitchen floors and is used primarily in restaurant roach control . the best professional baits for german roaches are invict gold cockroach gel and maxforce magnum . demon insecticide is the best residual spray and can best be used in conjunction with baygon aerosols . if you prefer to use professional dusts , delta dust and drione dust are the best dusts for roach control . for more help in ridding yourself of this pest , refer to german roach elimination .
ajjan , i . and w . robinson . 1996 . measuring hydramethylon resistance in the german cockroach , blattella germanica ( l . ) . in k . wildey and w . robinson , eds . proceedings international conf . on urban pests , prague , edinburgh , pp . 135 - 144
adult german cockroaches are light - brown and have two stripes on their pronotum , which is the “shield” just behind their head .
7 . german cockroaches regularly disperse to new areas . older , larger nymphs are most likely to disperse , followed by adult males and then adult females without egg cases . small nymphs are least mobile . during an outbreak of german cockroaches , technicians need to expand their search for cockroaches into adjacent rooms above and below , even into sites that are not typically infested with german cockroaches .
egg carrying patterns and rapid reproduction rates allow german roaches to quickly infest a home . don’t take chances with your family’s health .
empty cabinets and clean them out . another common food source for german roaches are the crumbs and food spills inside kitchen cabinets .
store food in sealed containers . german roaches are small enough to slip into the cardboard packaging that many foods are stored in .
females carry an egg capsule containing 30 - 48 eggs at the end of their abdomen . when the eggs are close to hatching , the females attach the capsule in a dark corner where the young can emerge safely . one female german cockroach can produce up to 20 , 000 young annually .
an allergy to cockroaches could set off adult or pediatric asthma . cockroach allergens can be environmental triggers for asthma . learn more about the health implications of cockroaches here .
the information for roaches should help not only identify pests but also help to differentiate types of roaches that are given common or incorrect names . to many people , any large roach , roaches that live primarily outdoors or any cockroach that is not a german cockroach is considered a woods roach . the wood roach is a distinct species of roach , not a group of bugs . however , many pest control experts do consider roaches that are not german roaches to belong to a generic
large roach
or
outdoor roach
because their primary source is the outdoors .
water bugs
is a term many people use to describe german cockroaches . there are no roach pests that are technically water bugs . when our customers say that water bugs are a problem in their home , we can usually assume that they have an infestation of german roaches .
in three to four months , those baby roaches will develop into fully grown adults . the lifespan of a cockroach is usually one year , and in any female roach’s lifetime , she can produce anywhere between 200 to 300 offspring or 6 generations a year . the number of eggs a single cockroach can produce will vary from species to species . potentially , and with optimal conditions , one female german cockroach and her offspring could produce 300 , 000 roaches in a year . this means that if you are not careful , a relatively small roach infestation can quickly develop into a much larger problem that needs to be dealt with quickly .
what do they eat ? german cockroaches are scavengers , capable of feeding on most any food source available . the pests will eat :
6 . german cockroach egg capsules are not susceptible to insecticides . most insecticides do not have the ability to penetrate the egg case , even if directly applied to the case . the unaffected egg cases will hatch days after treatment . this is another reason to schedule follow - up treatments in heavy infestations .
miller , d . m . , koehler , p . g . , patterson , r . s . 1997 . use of german cockroach ( blattella germanica ) fecal extract to enhance toxic bait performance in the presence of alternative food sources . j . econ . entomol . 90 : 483 - 487 .
feeding habits the german cockroach will eat practically anything and will live very close to food and water sources . they eat sweets , starches , grease , and meat products ; but will also feed on beer , leather , book bindings , hair , glue , dried skin , dead animals and plant material .
pest control effort ! failure to practice good housekeeping is the primary reason for outbreaks of german roaches . like most pests , german roaches require 3 things to thrive - food , water , and harborage . if you can eliminate even one of these things through proper sanitation ,
to inspect for the presence of german cockroaches , one can use pheromone sticky traps ( commonly found in hardware / supermarket stores ) to attract and trap german cockroaches . however , the use of these traps requires vigilant inspection over time ( traps are only as useful as the person that monitors them ) . for an easy - to - make homemade trap for a possible german cockroach infestation , take a baby food jar and coat the inside - upper walls with vasoline . put a small amount of beer and bread inside the baby food jar , and place jar along the wall underneath the kitchen sink . the smell of fermentation and yeast from this attractive mixture will lure nearby cockroaches , and when the cockroach falls into the jar it won’t be able to run up the vasoline !
another study suggests that cockroaches have a collective intelligence made up of the decisions of individual roaches . european scientists developed a robot called insbot that was capable of mimicking cockroach behavior . the researchers applied cockroach pheromones to the robot so real roaches would accept it . by taking advantage of roaches ' tendencies to follow each other , insbot was able to influence the behavior of entire groups , including convincing roaches to leave the shade and move into lighted areas . scientists theorize that similar robots could be used to herd animals or to control cockroach populations .
in the cockroach case , sugar actually tastes bitter - an effective way for natural selection to quickly produce cockroaches that won ' t accept the sugar baits that hide poison .
farmer , b . r . and w . h robinson . 1984 . is caulking beneficial for cockroach control . pest control 52 ( 6 ) : 28 , 30 , 32 .
this kit contains invict roach bait with gentrol point source igr for the best combination for german roach control , it kills the adult population and prevents new births . invict gold cockroach gel rapidly kills roaches with its bait matrix of 11 food attractants as a lure . it is known to maximize roach control and fight against bait aversion .
whether mother roaches care for their young also varies from one species to another . some mothers hide or bury their ootheca and never see their offspring . others care for their offspring after birth , and scientists believe that some offspring have the ability to recognize their mothers . the number of young that one roach can bear also varies considerably . a german cockroach and her young can produce 300 , 000 more roaches in one year . an american cockroach and her young can produce a comparatively small 800 new roaches per year .
robinson , w . h . 1985 . german cockroaches , facts or fiction . pest control 53 ( 10 ) : 70 , 75 - 76 .
the mating behavior of german cockroaches is driven by pheromones given off by females , which are detected by the antennae of males . german cockroaches breed continuously with many overlapping generations present at any one time . as a result of continuous breeding and promiscuity , population growth has been shown to be exponential .
male german cockroaches , on average , live 100 to 150 days . females live much longer , with an average lifespan of 190 to 200 days .
thoms , e . m . and w . h robinson . 1987 . distribution and movement of the oriental cockroach around urban apartment buildings . environ . entomol . 16 : 731 - 737 .
) . conventional cockroach control programs have used spray formulations containing carbamates , ops , organochlorines and pyrethroids . consequently , high levels of resistance to these insecticides have been documented in many field populations [
german cockroaches , believed to have originated in southeast asia , are the most widely distributed urban pests . they have been introduced to all parts of the globe including north america , australia , africa , and the oceanic islands . this ubiquity makes german cockroaches cosmopolitan , with the only deterrent being cold temperatures .
male adult german cockroaches are slender and a little smaller than the females , who are a little larger and wider in the abdomen . when female adult german roaches are ‟gravid , ” or pregnant , they have an egg capsule , or ootheca , that can be seen protruding about one quarter - inch from the end of the abdomen . the egg capsule contains about 40 german cockroach eggs , more or less , that will be carried with her until about 24 hours before the new roaches are ready to hatch . she then drops the egg capsule in a concealed location . when the new roaches open it and crawl out , they are on their own .
german cockroach females , unlike most other roaches , carry the egg capsule protruding from their abdomen until their capsules ready to split open . during the last three or four days prior to dropping her egg case , the female german cockroach does not forage for food or water . the case is then placed in a secluded location , with the nymphs emerging sometimes within the hour or as long as a week . a female may produce four to six cases during her lifetime , each containing 30 to 40 eggs . eggs hatch in 28 to 30 days , and nymphs develop in 40 to 125 days . female roaches live about 200 days , with males living not quite as long . the german roach produces more eggs and has more generations per year ( three to four ) than other roaches , and only a few individuals are needed to develop into troublesome infestations .
robinson w . h and j . zhai . 1990 . pyrethroid resistance in german cockroaches . pest control technology 18 ( 10 ) : 26 - 28 .
although german cockroaches are capable of living outdoors , they are often found in homes where food is widely available . learn more in the orkin pest library urltoken
it shakes its head and refuses to imbibe that liquid , at the end , you can see the [ glucose ] on the side of the head of the cockroach that has refused it .
the saliva , droppings and decomposing bodies of cockroaches contain allergen proteins which are known to trigger allergies and increase the severity of asthma symptoms . for more on cockroaches , check out the cockroach pest guide .
thoms , e . m . and w . h robinson . 1986 . the oriental cockroach : new insights on an old foe . pest control 54 ( 7 ) : 30 - 32 , 36 .
at rentokil , our british pest control association ( bpca ) certified technicians are able to provide customised , targeted treatments to guarantee complete control of your cockroach problem and help to prevent a re - infestation .
german cockroaches have three developmental stages : egg , nymph , and adult . females develop 4 to 8 capsules containing 30 to 48 eggs each in their lifetime . capsules hatch about 28 days after they begins to form . a few weeks thereafter , a new egg capsule begins to form . the egg stage varies in duration from 14 to 35 days . german cockroaches have 6 to 7 nymphal stages occurring over a period of 6 to 31 weeks . they express incomplete metamorphosis : zygotes develop within eggs and hatch directly into nymphs , which then grow into adult cockroaches . the complete life cycle of the cockroach is roughly 100 to 200 days for females , during which 10 , 000 descendants of a single cockroach can be produced .
the most common and effective igr used for german roach control is gentrol igr , which comes in an aerosol , concentrated liquid , and point - source tablet .
due to resistance to most insecticides on the market , professional roach baits are by far the most effective products to use when german cockroach extermination is needed . there is an insecticide concentrate that can be used against these resistant roaches but it is labeled for use only in commercial kitchens - and it has a very high odor . commercial kitchens should either bait or use orthene
the best way to get rid of german roaches is place out a high quality roach bait like invict gold to kill the adult german roaches and an insect growth regulator ( igr ) like gentrol point source igr to stop the reproduction of roaches . we have both products in a kit , invict gold roach kit for savings .
adult german cockroaches are 1 / 2 to 5 / 8 inch long and tan to light brown ( fig . 1 ) . although they have fully developed wings , they do not fly . nymphs are similar in appearance to adults except that they are smaller and lack wings . the german cockroach is best identified by its small size and by two dark parallel lines running from the back of the head to the wings . it is usually found in kitchens ( near dishwashers , stoves , and sinks ) and in bathrooms of homes .
german roach infestations are usually are generally found in kitchens and bathrooms . if the population is large enough , they will spill over to other rooms . these roaches are mostly active at night . if german roaches are seen active during the day , it most likely is due to overcrowding in their hiding places or a shortage of food and water supply . cracks and crevices are their harborage areas ; they spend about 75 % of their time in such harborages . first instar nymphs require a crack of about 1 / 32
whereas , adults require an inch width . german cockroach nymphs are smaller than most other cockroaches ; thus , they are able to conceal themselves in many places and remain protected .
non toxic and low toxic alternatives for german cockroach control are available . sticky traps can be used to monitor or reduce population size . improving sanitation by eliminating food and water sources and clutter can have a significant impact on reducing the chances of infestation population size . finally , exclusion practices such as sealing cracks and crevices will reduce harborage space and also negatively impact population size .
german cockroaches prefer to live in warm , humid places close to food and moisture sources . they are frequently found in residential and commercial kitchen environments , and bathrooms .
zhai , j . and w . h robinson . 1991 . the walking behavior of german cockroaches . pest control technology 18 ( 7 ) : 44 - 46 .
jacobs , s . 2007 .
german cockroaches
( on - line ) . entomology notes , pennsylvania state university . accessed october 15 , 2008 at urltoken .
bao , n . and w . h robinson . 1990 . morphogenetic effects of hydroprene on genitalia of the oriental cockroach ( dictyoptera : blattidae ) . j . econ . entomol . 83 : 1415 - 1421 .
if a german roach infestation is found , treatment will be applied in areas where roaches have been or may be hiding . knowledge about the german cockroach is vital for effective treatment . for example , young roaches often feed on the fecal droppings of the adult cockroaches in “aggregation” areas , while the adults are out looking for food . as grotesque as this may sound , it is a key to pest control that helps achieve timely and effective cockroach elimination . professional baits are slow - acting by design . when the adults start feeding on baits , they are preparing a final meal for the young roaches . they live just long enough to digest the bait , get back to their aggregation hangout , and leave their final legacy for the up and coming young roaches .
habitat german roaches require moisture to thrive and so tend to dwell in moist places , such as kitchens and bathrooms . however , they may inhabit any room of a structure or home , particularly if the infestation is severe . german cockroaches spend about 75 % of their time hanging out in cracks and crevices near food and water sources . adult german cockroaches can hide inside cracks as small as 3 / 16
wide , while the nymphs can fit into cracks as small as 1 / 32
wide .
if you have a german roach problem , it could be that you are at your wits ' end wondering how to get rid of one of the most stubborn household pests in existence . german roaches get into everything , multiply rapidly , and can survive for several months without food and up to two weeks without water . these little guys definitely pose a challenge , but with the proper tools and products , you can win the battle over german roaches by following this strict integrated pest management ( ipm ) program .
eliminating food sources is a constant battle in german roach control . while this effort involves a great deal of work and vigilance , the results are well worth the effort .
nymphs molt several times as they become adults . the period between each molt is known as an instar . each instar is progressively more like an adult cockroach . in some species , this process takes only a few weeks . in others , like the oriental cockroach , it takes between one and two years . the overall life span of cockroaches differs as well - - some live only a few months while others live for more than two years .