shenshou lui is an early mammal dating from the oxfordian stage of the late jurassic , approximately 160 million years ago .

fossils were recovered from the tiaojishan formation in the liaoning province of china .

shenshou

figure 6 . shenshou pes reconstrution . the short calcaneum also indicates an arboreal taxon .
figure 8 . shenshou nests basal to rattus and the former allotheria ( = multituberculata + haramiyida ) .
figure 5 . manus of shenshou in situ and reconstructed . compare this the original reconstruction shown in figure 7 .
figure 4 . scapula and vertebrae of shenshou . one centrum was misidentified as an unusually shaped coracoid by bi et al .
crunchyroll ' s translation names the relic
shenshoujin
. fansubs use
shenshou jing
or
shen shou jing
.
figure 3 . shenshou skull traced in colors . the lateral temporal bar is gracile . the vomers are visible . there is a nice ectotympanic hemicylinder there .
xiamen shenshou science & technology co . ltd does not currently advertise comprehensive company & product information with global sources . we cannot guarantee the accuracy of company and product information .
shenshou is thought to be arboreal because it had a light frame , a prehensile and elongated tail , and hands and feet which had evolved for clutching and enabled the animal to climb . these features , including the large incisors of shenshou , made the animal resemble a squirrel . however , shenshou are not the direct ancestors of squirrels , the resemblance being purely due to convergent evolution . individuals are believed to have weighed 300 grams ( 11 oz ) . the presence of a three - boned middle ear suggests these animals were mammals . the teeth , which have a number of cusps , suggest that shenshou was probably an omnivore , most likely with a diet of fruits , nuts and insects .
a fossil of shenshou lui , a newly discovered species of ancient mammal , shows the animal ' s gracile body with long fingers and tail adapted for life in the trees .
figure 1 . the jurassic mammal shenshou , which nests within allotheria ( haramiyida + mutituberculata ) within the mammalia , as i proposed based on the lrt without knowledge of this paper .
“where they go in the consensus ( e . g . “sister” to shenshou in your terminology - bi et al . , 2014 ) , based on more characters and taxa than you use . ”
figure 1 : the holotypes of three euharamiyidan species . a , holotype ( ldnhmf2001 ) of shenshou lui . b , holotype ( ivpp v16707a ) of xianshou linglong . c , holotype specimen ( bmnhc - pm003253 ) of xianshou songae . doi : urltoken
genus : shenshou bi , wang , guan , sheng , & meng , 2014 etymology : shen , from pinyin of the chinese word , meaing deity , divinity or cleaver , shou , from pinyin of the chinese word for creature , animal or beast .
the largest of the three species is named shenshou lui , with
shenshou
meaning
divine beast
in chinese and
lui
referring to the collector of the specimen , lu jianhua . the other two species are named xianshou songae and xianshou linglong , with
xianshou
meaning
celestial beast
in chinese ,
songae
referring to rufeng song , the collector of the specimen , and
linglong
meaning
exquisite
in chinese and also referring to the town of linglongta , where the specimen came from .
figure 7 shenshou original art by or traced from bi et al . 2014 , compared to rattus , the rat . note the extremely deep posterior dentary here is not a good reflection of the actual fossil ( fig . 3 ) . note the original interpretation of a coracoid . even that looks like a vertebral centrum . note , the large reconstruction is a chimaera .
all specimens described here are considered to be molariforms , by comparison with haramiyids for which premolariforms in situ are known ( haramiyavia jenkins et al . , 1997 , megaconus zhou et al . , 2013 , arboroharamiya zheng et al . , 2013 , shenshou bi et al . , 2014 , and xianshou bi et al . , 2014 ) . indeed , theroteinus material does not show neither the hypertrophied mesial cusp of lower premolariform of haramiyavia ( jenkins et al . , 1997 ; luo et al . , 2015 ) , megaconus ( zhou et al . , 2013 ) , arboroharamiya ( zheng et al . , 2013 ; meng et al . , 2014 ) , shenshou , and xianshou ( bi et al . , 2014 ) ; nor the circular arrangement of cusps of upper premolariform of megaconus ( zhou et al . , 2013 ) , arboroharamiya ( zheng et al . , 2013 ; meng et al . , 2014 ) , shenshou , and xianshou ( bi et al . , 2014 ) . moreover , theroteinus teeth show no character that could give information about their position in the dental series ( see below ) .
the life science identifiers ( lsids ) for this publication have been deposited at urltoken and include : urn : lsid : zoobank . org : pub : 766ebc08 - ef77 - 41e5 - ac59 - 9f69e9f59baa ( for this publication ) , urn : lsid : zoobank . org : act : 32b0742d - 2dd3 - 47f9 - a35b - e1fb284e9ea5 ( shenshou ) , urn : lsid : zoobank . org : act : ba8bf69d - 0f21 - 4de7 - 9ea5 - ba6082973940 ( shenshou lui ) , urn : lsid : zoobank . org : act : 4c10f9f1 - a920 - 450b - 8e5f - c88f3bd0920c ( xianshou ) , urn : lsid : zoobank . org : act : 3db1d738 - 70e9 - 4966 - 93fe - 3fe5269f9c91 ( xianshou linglong ) , urn : lsid : zoobank . org : act : 1f7eee49 - 5e88 - 4941 - ae8b - b2c7c31f9788 ( xianshou songae ) .
the orientation of theroteinus teeth follows the orientation described for haramiyavia ( jenkins et al . , 1997 ; luo et al . , 2015 ) , arboroharamiya ( zheng et al . , 2013 ; meng et al . , 2014 ) , shenshou ( bi et al . , 2014 ) , xianshou ( bi et al . , 2014 ) , and thomasia ( e . g . , butler , 2000 ) . the lower molariforms are distinguished from the upper molariforms by the presence of , respectively , two or three rows of cusps and by the form of row a / a . in lower molariforms , the lingual row a includes the largest cusps . cusp a1 is especially much larger than the other cusps , and it is located on the mesiolingual side of the crown . in upper molariforms , the labial row a shows three subequal cusps , when the central row b displays a cusp b2 larger than the other cusps of the row and located on the distal side of the crown .
here , a reconstruction of arboreal mammals in a jurassic forest . the three animals on the left side represent three newfound species of euharamiyidan mammals that lived some 160 million years ago .
extinct squirrel - like creatures from china suggest the earliest mammals originated more than 200 million years ago , much earlier than often previously thought , researchers say .
the fossils were discovered in the last three years by private collectors and amateur paleontologists in a liaoning province cornfield in northeastern china . liaoning has become famous for the trove of feathered dinosaurs and winged reptiles known as pterosaurs unearthed there over the last decade . the province is also known for a fossil of a baby dinosaur inside a mammal ' s gut , the first direct proof that mammals dined on dinosaurs .
the newfound fossils are about 160 million years old , dating back to the triassic period . back then , the area was a warm , wet forest populated by dinosaurs , mammals and pterosaurs , all living on thelost supercontinent laurasia , which once included what are today ' s northern continents . [ see images of squirrel - like creatures and other ancient mammals ]
the six well - preserved fossil specimens are from three different extinct species . the animals ranged in size from 1 to 10 ounces , or from
a house mouse to a small squirrel ,
said study co - author jin meng , a paleontologist at the american museum of natural history in new york . their teeth suggest
they were most likely omnivorous , eating insects , nuts and fruits ,
he said .
the holotype specimen of senshou lui , which represents a new species of euharamiyidan mammal , described in the sept . 11 , 2014 , issue of the journal nature .
judging from their slender builds , long tails , hands and feet adapted for grasping and climbing , and enlarged incisor teeth , these animals would have been tree - dwellers that looked similar to squirrels . however ,
don ' t confuse these new animals with any living species ,
meng said . any similarities between these creatures and squirrels are due to convergent evolution , just as fish and dolphins both have streamlined bodies to better swim in the water but are only distantly related .
the bones were of mysterious animals known as haramiyids . scientists learned of haramiyids back in darwin ' s time , but researchers knew about them only from their teeth and fragments of their jaws .
for over a century , paleontologists were puzzled as to whether these creatures were mammals or animals closely related to mammals ,
meng told live science .
these fossils reveal new details about the skulls , teeth and skeletons of three previously unknown species of haramiyid , revealing that haramiyids were probably mammals . for example , the fossils show evidence of a typical mammalian middle ear , the area just inside the eardrum that turns vibrations in the air into electrical signals that get transmitted to the brain . the middle ears of mammals are unique in that they have three bones .
the new findings suggest their closest known relatives were rodentlike creatures known as multituberculates . both of these groups have no living descendants , having long ago separated from the lineage leading to modern mammals .
the oldest known haramiyids date to the late triassic period about 220 million to 200 million years ago . this suggests mammals are at least that old ,
earlier than much previous research predicted ,
meng said .
many scientists had suggested that mammals originated in the middle jurassic , which ranged from 174 million to 164 million years ago .
haramiyids are one of the oldest group of mammals , if not the oldest ,
meng said .
the scientists detail their findings in tomorrow ' s ( sept . 11 ) issue of the journal nature .
follow us @ livescience , facebook & google + . original article on live science .
charles q . choi is a contributing writer for live science and space . com . he covers all things human origins and astronomy as well as physics , animals and general science topics . charles has a master of arts degree from the university of missouri - columbia , school of journalism and a bachelor of arts degree from the university of south florida . charles has visited every continent on earth , drinking rancid yak butter tea in lhasa , snorkeling with sea lions in the galapagos and even climbing an iceberg in antarctica .
elon musk ' s plan to rescue trapped thai boys ? a kiddie submarine that looks like a coffin .
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we thank h . - j . li and z . - j . gao for using the specimens housed at the jizantang paleontological museum ; z . - y . sun and d . - y . sun for using the specimen housed at the museum of wuyishan mountain ; j . - c . lü for assistance in specimen collecting ; s . - h . xie for specimen preparation ; m . a . klingler and f . - x . wu for specimen drawing and photography ; w . - d . zhang for sem imaging ; z . - h . zhou , x . xu , f . - c . zhang and x . - l . wang for discussion on stratigraphy and faunal compositions ; and a . weil and g . w . rougier for instructive comments . the study was supported by the national basic research program of china ( 973 program , 2012cb821906 ) , the strategic priority research program of chinese academy of sciences ( xdb03020501 ) , the national science foundation of china ( 41128002 ) and the hundred talents programs of the chinese academy of sciences .
s . b . , j . m . and y . w . designed the study , performed the comparative and analytical work and wrote the paper . j . g . and x . s . collected data and contributed to the writing and discussion .
holotype and close - up views of craniodental features of xianshou linglong ( ivpp v16707b ) .
close - up views of craniodental features of xianshou songae ( bmnhc - pm003253 ) .
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jurassic skeletons show that early mammals didn ' t just hide in the undergrowth .
squirrel - size mammals scampered through the trees above dinosaurs ' heads , newfound chinese fossils show , revising our image of the first furry beasts . three newly described species suggest that mammals evolved earlier , and faster , than previously thought .
called haramiyids , the recently discovered mammals lived in jurassic china around 160 million years ago . slender and graceful , the animals appear to have been specialized for life in the trees , with hands and feet that could grasp branches and a long prehensile tail like today ' s monkeys .
the picture that mesozoic mammals were shrew - like insectivores that lived in the shadow of the dinosaurs needs to be repainted ,
says american museum of natural history paleontologist jin meng , a coauthor of the new study . discoveries during the past few decades , including the haramiyids , have shown that early mammals occupied a variety of habitats .
they walked on the ground ; they also swam , dug to burrow , and glided in the forests ,
meng says .
the strange mammals were known from teeth and a bit of jaw , but last year meng and colleagues reported on the first complete haramiyid skeleton . the haramiyid wasn ' t quite like any living mammal , but was closely related to an extinct group called multituberculates , named for the complex ridges , or cusps , on their teeth .
the mouse - size animal xianshou songae , shown in an artist ' s illustration , was a tree - dweller in jurassic forests .
in the new study , meng and collaborators report six more haramiyid skeletons from three new species in the journal nature on wednesday . the find adds to the shifting picture of mammal evolution during the mesozoic era , a time when dinosaurs dominated the land .
the emerging picture is that mammal evolution spun off different body types early on , says oklahoma state university paleontologist anne weil .
you can look at small rodents today and see that a mouse and a squirrel are different animals ,
weil says , and mammals like the haramiyids show that the same was true in the mesozoic .
and the new haramiyids do more than expand the image of how our ancient mammalian cousins lived . the relationships among these long - enigmatic creatures suggest that the very first mammals originated early .
the traditional view is that the first true mammals evolved sometime during the jurassic . haramiyids , while roughly mammal - like , were thought to fall outside the group .
with complete skeletons to work from , however , meng and colleagues found that haramiyids were true mammals after all . and given the age of the earliest known haramiyid , meng says that mammals originated
at least in the late triassic ,
between 220 million and 201 million years ago .
i expect this will be contentious ,
weil says , but the study is an important addition to investigations of where mammals came from .
i think it ' s going to be part of an argument that will be going on for some time ,
weil says ,
and i expect paleontology as a whole will learn a lot from questions gleaned from these animals about the antiquity of mammalia .
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( meng et al . 2017 ; jurassic , 160 mya ) was originally described as a pre - mammal haramiyid close to
. here it nests with between interatherium and vintana + groeberia . it is also not related to
the mandible is not known . the ventral jugal has a robust process . the clavicle is robust here and fused with the interclavicle . the coracoid is tiny , but present here as in paedotherium ( above ) . a patagium is not seen , but a halo of thick fur is preserved .
( bi et al . 2014 ; oxfordian , earliest late jurassic ) was described as a member of the early mammalian clade , allotheria . the
is an ancestral taxon . a purported coronoid was actually two vertebral centra . grasping hands and feet suggest
( luo et al . 2017 ; jurassic , 60 mya ) was originally considered a relative of
as a flying squirrel mimic . it has extended patagia for gliding . tiny middle ear bones fail to develop here . rather they remain as post dentary bones . this caused meng et al . 2017 to consider this a pre - mammal . they did not realize this trait was a reversal in a rodent - related mammal .
bi s , wang y - q , guan j , sheng x and meng j 2014 . three new jurassic euharamiidan species reinforce early divergence of mammals . nature 514 : 579 - 584 . online here .
luo z - x , meng q - j , grossnickle dm , neander ai , zhang y - g and ji q 2017 . new evidence for mammaliaform ear evolution and feeding adaptation in a jurassic ecosystem . doi : 101 . 1038 / nature 23483 \
meng q - j , grossnickle dm , liu d , zhang y - g , neander ai , ji q and luo z - x 2017 . new gliding mammaliaforms from the jurassic . nature ( advance online publication ) doi : 10 . 1038 / nature23476
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species : lui bi , wang , guan , sheng , & meng , 014 etymology : in honor of lu jianhua , the collector of the holotype .
locality : daxishan site , linglongta , jianchang county , liaoning province , china .
is an ancient type of round bronze mirror decorated with images of gods and animals from chinese mythology . the obverse side has a polished mirror and the reverse has relief representations of legendary chinese
during the relic excavation at mt . minakami in nagano that kanade ' s family died in , finé found and left a fragment of the shénshòujìng in the care of the us government . the mirror relic , shénshòujìng , was originally developed under the assumption it would be worn by a user . but with the absence of a linked user and the fact that it could be utilized by the fis ' s mechanical control process , it was led to being equipped to the air carrier using mechanical methods that were derived from the symphogear system , where the shénshòujìng enabled stealth capabilities in addition to acting as the source of power for the remover ray that can invalidate the power of relics .
although miku successfully activates the frontier with the shénshòujìng ' s beam with maria ' s assistance , the shénshòujìng was destroyed by its own beam along with hibiki ' s gungnir fragments in their battle .
like all mirrors , the shénshòujìng ' s special properties are based on its ability to reflect light . one of those properties ,
wizardry stealth
, can not only make the helicopter invisible , but also give off oscillations that cut off its signal from any enemy monitoring equipment .
in ancient times , the mirror was said to have the properties of driving away curses and purifying bad luck . as a result , the shénshòujìng has the ability to destroy relics with its beams .
as a mirror , its most potent and basic ability is projecting a brilliant radiance and reflecting back an
ideal form
. the mirror ' s most important role is releasing the sealed frontier that lies in the middle of the ocean . however , that is not an easy task , as mechanically amplifying its power is an imperfect implementation of the relic ; however , the shénshòujìng symphogear equipped by a human proves to be able to remove the seal the frontier .
the special function of the shénshòujìng symphogear is its
direct feedback system
. functioning similarly to a mirror , this system has the ability to project images and information directly into the brain of its user . if you take a previously - prepared program and install it , even a user with a very weak battle sense will mechanically have their potential upgraded and for a short time be enabled to fight rationally with their battle power significantly enhanced .
however , this ability to directly project images and information into the brain is controlled by a third - party . they can enter in whatever information they like , overriding the user ' s own personality and utilizing the symphogear as a brainwashing tool , thereby making it significantly dangerous to use normally .
compared to the ame no habakiri , ichaival , shul shagana , and igalima , the shénshòujìng was created from a sacred relic that is several levels inferior to the others , and thereby , inevitably , part of its characteristics are lower specs .
it was for this reason that it was integrated into the air carrier to function as the power source for wizardry stealth ; however , the free - thinking dr . ver once again led it on the path of a symphogear that encases the mind and body of its user .
the strong feelings of miku , who wanted to
prevent her precious friend hibiki from fighting , so that the fusion with gungnir will not spread any further
, were warped by the mirror ' s property of reflecting an ideal form by utilizing
exorcism of curses
and amplified its manifestation to the point where it became able to eradicate the base power of relics themselves . furthermore , the electromagnetic force of the light brought forth by the mirror ' s ability to emit an overwhelming radiance , combined with the biefield - brown effect , granted the symphogear the ability of flight implemented as an ionocraft .
even with its low specs , having abilities which deviate from its mirror base , including others that have been omitted at this point , makes it so when fighting against other relics , the shénshòujìng has an overwhelming killing potential and a flight ability that is unmatched . despite being the weakest , it is the symphogear utilized as a supreme evil , literally a
distorted mirror
. [ 3 ]
the shénshòujìng reappears in the xdu event kageri saku senkō , where miku gains an x - drive form for the symphogear .
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the generic name is derived from mandarin shen , meaning deity , and shou , meaning animal , while the specific name is in reference to lu jianhua , the scientist who collected the holotype specimen .
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the three animals on the left side represent the three new species of euharamiyidan mammals . the other two represent a gliding species and another euharamiyidan , respectively , that were reported earlier .
, mouse - sized animal was a tree dweller in the jurassic forests and belonged to an extinct group of mesozoic mammals called euharamiyida .
etymology . shen , from pinyin of the chinese word , meaning deity , divinity or cleaver ; shou , from pinyin of the chinese word for creature , animal or beast ; specific name after lu jianhua , the collector of the holotype .
etymology . xian , from pinyin of the chinese word meaning celestial being or immortal .
etymology . linglong , from pinyin of the chinese word , meaning ‘exquisite’ , and also after the town name linglongta where the specimen came from .
etymology . the specific name is after rufeng song , the collector of the holotype specimen .
bi , shundong ; wang , yuanqing ; sheng , xia and meng , jin 2014 . three new jurassic euharamiyidan species reinforce early divergence of mammals . nature . doi : urltoken .
wuodendron b . xue , y . h . tan & chaowasku wuodendron praecox ( hook . f . & thomson ) b . xue , y . h . tan & x . l . hou in xue , tan . . .
[ botany • 2017 ] begonia fulgurata | ดาดดารารัศมี • a new species ( sect . diploclinium , begoniaceae ) from chiang mai , northern thailand
begonia fulgurata c . - i peng , c . w . lin & phutthai ดาดดารารัศมี | | doi : 10 . 3767 / blumea . 2017 . 62 . 03 . 01 urltoken be . . .
chamaelirium viridiflorum l . wang , z . c . liu & w . b . liao in liu , feng , wang & liao , 2018 . doi : 10 . 11646 / phytotaxa . 357 . . . .
great - billed seed - finch sporophila maximiliani ( cabanis , 1851 ) in ubaid , silveira , medolago , et . al . , 2018 . doi : 10 . 11646 / . . .
endocerids with their filtering apparatus in mironenko , 2018 . doi : 10 . 1080 / 08912963 . 2018 . 1491565 reconstruction by andre . . .
aristolochia tongbiguanensis j . y . shen , q . b . gong & s . landrein in gong , landrein , xi , et al . , 2018 . doi : 10 . 6165 / tai . . . .
bagualosaurus agudoensis pretto , langer & schultz , 2018 doi : 10 . 1093 / zoolinnean / zly028 illustration : jorge blanco c . . .
the hypothetical phylogenetic relationships of ceratosaurs based on current topologies . the main source is from wang et al . ( 2016 . . .
nipponosaurus sachalinensis nagao , 1936 in takasaki , chiba , kobayashi , et al . , 2018 ニッポノサウルス | | doi : 10 . 1080 / 08912963 . 2017 . . . .
begonia medogensis jianw . li , y . h . tan & x . h . jin in li , tan , wang , et al . , 2018 . doi : 10 . 3897 / phytokeys . 103 . 25392 . . .
[ herpetology • 2014 ] a checklist and key to the ho . . .
[ mammalogy / evolution • 2013 ] mad , bad and danger . . .
on this day ( july 10th ) . . . . . . . . . . . . . . .
anomaloglossus meansi : a new species of cryptic forest frog from the wokomung and ayanganna sky islands of southern guyana .
the benefits and costs of academic travel . or
there and back again ; again and again
canon renueva su gama de 70 - 200 mm f : 2 . 8 y f : 4
plants go extinct , but sometimes species are rediscovered . this one after 151 years .
i ' m killing antediluvian salad but even in death there is rebirth . . .
necps carnivorous plant show : sept . 9 - 10 at tower hill botanical garden
this is a particularly beautiful species of centrolenid - the granular glass frog , cochranella granulosa .
x . linglong wang et al . 2014 x . songae meng , et al . 2014
by the sharper cusps and ridges of its upper molars , and by larger size .
bi , shundong ; wang , yuanqing ; sheng , xia ; meng , jin ( 10 september 2014 ) .
this article is issued from wikipedia - version of the 11 / 26 / 2016 . the text is available under the creative commons attribution / share alike but additional terms may apply for the media files .
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this is an open access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , reproduction and adaptation in any medium and for any purpose provided that it is properly attributed . for attribution , the original author ( s ) , title , publication source ( peerj ) and either doi or url of the article must be cited .
hahn , sigogneau - russell & wouters ( 1989 ) included theroteinus , other haramiyids , and multituberculata cope ( 1884 ) within allotheria marsh ( 1880 ) . they erected the order theroteinida hahn , sigogneau - russell & wouters ( 1989 ) and raised the suborder haramiyoidea hahn ( 1973 ) to ordinal rank as haramiyida hahn , sigogneau - russell & wouters ( 1989 ) . their classification was illustrated by a phylogenetic tree in which theroteinus is the sister - group of all other allotherians ( hahn , sigogneau - russell & wouters , 1989 : text - fig . 12 ) . butler ( 2000 ) modified the classification of allotheria . within the order haramiyida of hahn , sigogneau - russell & wouters ( 1989 ) , he changed the rank of theroteinida hahn , sigogneau - russell & wouters ( 1989 ) to suborder and put back the suborder haramiyoidea hahn ( 1973 ) ( butler , 2000 ) . hahn & hahn ( 2006 ) published the most recent classification of haramiyida including theroteinus . they modified the names of the sub - orders of butler ( 2000 ) as theroteinina and haramiyina , respectively , and included millsodon butler & hooker , 2005 ( middle jurassic , england ) into the family theroteinidae ( hahn & hahn , 2006 ) .
in these classifications , theroteinus is always considered as more basal than other haramiyids upon one main feature : in centric occlusion , one tooth of theroteinus is in contact with two opposite teeth ( ‘one - to - two’ occlusion ) . this feature is shared by other mammaliaforms such as morganucodonts and kuehneotheriids but not by other haramiyids , which are characterized by an occlusal mode where one tooth is in contact with only one opposite tooth in centric occlusion ( ‘one - to - one’ occlusion ) .
this study describes 20 isolated teeth of haramiyids from saint - nicolas - de - port . denise sigogneau - russell and her co - workers have excavated only one stratigraphic level in the sand pit . specimens collected during this fieldwork are kept both in the mnhn , with the acronym ‘snp , ’ and in the rbins , with the acronym ‘ras . ’ several amateur palaeontologists gathered their own collections alongside sigogneau - russell’s team and donated them to mnhn and rbins . the collection of georges wouters is identified by the suffix ‘w’ or ‘fw , ’ and the collection of m . marignac is identified by the suffix ‘ma . ’ however , there are no data on the exact , original stratigraphic level within the quarry of these collections . all the specimens described by sigogneau - russell , frank & hemmerlé ( 1986 ) and hahn , sigogneau - russell & wouters ( 1989 ) are considered here , alongside with eight new specimens ( mnhn . f . snp 14 fw , mnhn . f . snp 787 , rbins ras 3 fw , rbins ras 11 fw , rbins ras 62 fw , rbins ras 74 fw , rbins ras 77 fw , rbins ras 103 fw ) .
the nomenclature used here to describe the haramiyid teeth is derived from parrington ( 1947 : fig . 3 ) , hahn ( 1973 : p . 5 ) , butler & macintyre ( 1994 : p . 435 ) and butler ( 2000 : p . 319 ) . the row of cusps named a / a is characterized by less numerous , and well - individualized cusps . this row is lingual on lower teeth , but labial on upper teeth . the second row of cusps is named b / b . this second row is labial on lower teeth , but lingual on upper teeth . both rows define a central basin . additional cusps are named aa / aa when they are on the lateral flank of row a / a , and bb / bb when they are on the lateral flank of row b / b . in each row , cusps are numbered starting from number 1 . on lower teeth , the numbering starts from the mesial extremity , while on upper teeth , it starts from the distal extremity . the term ‘u - ridge’ refers to the junction of crests which close the basin at its distal extremity on lower teeth and its mesial extremity on upper teeth . the term ‘saddle’ refers to the junction of two crests which delimits the basin at its open extremity , respectively mesial on lower teeth and distal on upper teeth . this nomenclature is used only in a descriptive purpose . the homonymy does not necessarily imply homology . capital letters are used for upper teeth and lower case letters for lower teeth .
the descriptions of the wear facets are based on the nomenclature of von koenigswald et al . ( 2013 : p . 146 ) for jaw movements . this nomenclature is used to define the direction and the angle of the slope of the wear facets . the process and the pattern of the occlusion are beyond the scope of this article and will be dealt with in detail later on .
the haramiyid teeth are distinguished from other contemporary mammaliaforms by the presence of longitudinal rows of cusps separated by basins . for now , only two haramiyids are known in the saint - nicolas - de - port material , theroteinus and thomasia poche , 1908 . the material referred to theroteinus is distinguished from material referred to thomasia by two main characteristics : low and obtuse cusps and a basin smaller in length and width .
type - family : theroteinidae sigogneau - russell , frank & hemmerlé , 1986 , by monotypy .
type - genus : theroteinus sigogneau - russell , frank & hemmerlé , 1986 .
type - species : theroteinus nikolai sigogneau - russell , frank & hemmerlé , 1986 .
emended diagnosis : haramiyids with lower and upper molariforms showing low cusps with more extended base and more massive aspect , short and narrow basins in relation to the size of the crown , presence of only two cusps in row a on lower molariforms ( shared with some specimens of thomasia ) , presence of a row bb on upper molariforms ( potentially shared with eleutherodon , megaconus and millsodon —see below ) , and an essentially vertical , masticatory movement .
distribution : upper triassic ( rhaetian ) : france , lorraine , saint - nicolas - de - port ( “grès infraliasiques” formation ) .
( a ) mnhn . f . snp 78 w , right upper , holotype ; ( b ) mnhn . f . snp 722 , right upper ; ( c ) mnhn . f . snp 226 w , left lower . 1 , occlusal view ; 2 , distal view ; 3 , labial view ; 4 , mesial view ; 5 , lingual view . ‘me’ indicates mesial extremity and ‘li’ indicates lingual side .
( a ) mnhn . f . snp 78 w , right upper , holotype ; ( b ) mnhn . f . snp 61 w , right lower ; ( c ) mnhn . f . snp 366 w , right lower ; ( d ) mnhn . f . snp 497 w , right lower ; ( e ) mnhn . f . snp 787 , right lower . right - angled arrow indicates mesial extremity and lingual side . letters in italics correspond to cusp nomenclature .
synonymy : theroteinus sp . hahn , sigogneau - russell & wouters , 1989 : p . 210 .
( a ) mnhn . f . snp 61 w , right ; ( b ) mnhn . f . snp 366 w , right ; ( c ) mnhn . f . snp 497 w , right . 1 , occlusal view ; 2 , distal view ; 3 , labial view ; 4 , mesial view ; 5 , lingual view . ‘me’ indicates mesial extremity and ‘li’ indicates lingual side .
dental measurements ( in mm ) of theroteinus molarifoms from saint - nicolas - de - port ( upper triassic , france ) .
l , mesiodistal length ; w , labiolingual width ; r , length / width ratio .
normality of the data has been tested by shapiro - wilk test ( table s2 ) , p - values have been corrected by holm - bonferroni method , the alternative hypothesis is “true difference in means is not equal to 0 . ”
) , right upper molariform , from saint - nicolas - de - port ( upper triassic , france ) .
the crown is dominated by two longitudinal rows of cusps which delimit a central basin , the lingual row a and the labial row b . the central basin is confined mesially by the saddle which joins cusps a1 and b2 , and distally by the u - ridge which joins rows a and b . the central basin gets deeper and narrower from the mesial extremity to the distal extremity .
cusp row a includes two cusps . cusp a1 is the largest one of the teeth and rises vertically in lateral view . this cusp extends over the mesial half of the tooth . cusp a1 shows a weak mesial carina which splits into two segments . one segment runs mesially and the other bends labially to join cusp b1 . at the level of the base of cusp b1 , the mesial segment turns into a short , horizontal cingulum to join cusp b1 . a distal crest starts from the distolabial side of the apex of cusp a1 to join cusp a2 . this crest is straight in lateral view , but it is curved labially in occlusal view . a sulcus underlines the lingual side of this crest and descends to the base of cusp a1 . a second crest , straight in occlusal and lateral views , starts from the labial side of the apex of cusp a1 to the base , where it takes part in the saddle . the distal and labial crests delimit a concave , narrow surface on the distolabial flank of cusp a1 , which extends from the apex to the central basin . cusp a2 is twice lower and labiolingually narrower , and much mesiodistally shorter than cusp a1 . cusp a2 is more lingual than cusp a1 . the lingual flanks of cusps a1 and a2 are aligned and parallel to the mesiodistal axis of the tooth on mnhn . f . snp 61 w , but deviate distolabially on mnhn . f . snp 366 w , and mnhn . f . snp 787 . the occlusal outline of cusp a2 is semicircular with a convex , lingual side and nearly flat , labial side . the labial side shows a vertical , weak ridge in the middle . in distal view , the slope of the labial flank is more vertical than the slope of the lingual flank . the latter is slightly convex . in labial view , the mesial base of cusp a2 is higher than the distal base of the cusp . in lingual view , the bases of cusps a1 and a2 are at the same level . cusp a2 shows two crests , respectively mesial and distal , straight in lateral and occlusal views , and aligned mesiodistally . the first crest starts from the mesiolabial side of the apex to join the distolingual crest of cusp a1 . the second crest starts from the distolabial side of the apex to the extremity of row a . the distal crest is much longer than the mesial crest . the slope of the mesial crest of cusp a2 is weaker than the slope of the distal crest of cusp a1 and the slope of the distal crest of cusp a2 . the slope of the latter is more vertical than the slope of distal crest of cusp a1 .
cusp row b includes four cusps , less distinguished from each other than the cusps of row a . cusp b1 is the most mesial of the tooth . this cusp is subequal in size with cusp b4 , or larger in mnhn . f . snp 61 w . cusp b1 is located in front of the saddle , but tends to rise lingually to join the mesiolabial carina of cusp a1 . cusp b2 is the largest cusps of row b . this cusp is slightly smaller than cusp a1 , except in mnhn . f . snp 366 w where cusp b2 is slightly larger and higher than cusp a1 . cusp b2 is labial to cusp a1 , its base extends as mesially but much less distally , and its apex is slightly more distal , or much more distal in mnhn . f . snp 61 w . cusp b2 shows two crests , straight in occlusal and lateral views . the first crest runs labially , but mesiolabially in mnhn . f . snp 61 w , to take part in the saddle . the second crest runs distally and joins cusp b3 . both crests define on the one side a slightly convex , distolingual occlusal outline , and on the other side a large arc of a circle . cusp b3 is much smaller than cusps a1 , a2 , and b2 and slightly smaller than cusps b1 and b4 . cusp b3 is directly distal to cusp b2 , except in mnhn . f . snp 787 where it is slightly more labial . this cusp is more mesial than a1 – a2 notch in mnhn . f . snp 787 , aligned with a1 – a2 notch in mnhn . f . snp 366 w , and more distal than a1 – a2 notch in mnhn . f . snp 61 w . the apex of cusp b3 is slightly higher than the apex of cusp a2 , or at the same level in mnhn . f . snp 61 w . the long axis of cusp b3 slightly deviates distolingually from the mesiodistal axis of the tooth , except in mnhn . f . snp 787 where both axes are parallel . cusp b4 is distal to cusp b3 and slightly more lingual . consequently , the long axes of both cusps are aligned , except in mnhn . f . snp 787 . the apex of cusp b4 takes place slightly lower than the apex of cusp a2 and faces the distal crest of cusp a2 . cusp b4 shows a lingual carina , which is well developed in mnhn . f . snp 787 . the u - ridge is a low crest which extends row b and bends lingually to join the extremity of row a .
comments on mnhn . f . snp 266 w and mnhn . f . snp 497 w
mnhn . f . snp 266 w and mnhn . f . snp 497 w are difficult to describe because they are extensively damaged . the surface of mnhn . f . snp 266 w is not well preserved (
) and mnhn . f . snp 497 w is heavily worn . for these reasons , they have not been incorporated in the description above . about mnhn . f . snp 497 w , it may be noticed that row
the crown is dominated by three longitudinal rows of cusps : labial row a , central row b and lingual row bb . rows a and b define a labial basin delimited distally by the saddle , constituted only by the lingual crest of cusp a2 , and mesially by the u - ridge which joins rows a and b . rows b and bb define a lingual basin , smaller than the labial basin , delimited distally by the meeting of cusps b2 and bb1 , and mesially by the crest which joins rows b and bb . both basins get deeper and larger mesially .
row a includes three mesiodistally aligned cusps . the three cusps take place at the same level on the crown . cusps a1 and a3 are subequal in length and width , cusp a1 is slightly higher than cusp a3 . cusp a2 is twice mesiodistally longer and higher , and labiolingually much wider than cusps a1 and a3 . the occlusal outlines of cusps a1 and a3 show a semicircular , labial flank and a relatively flat , lingual flank , sometimes concave because of wear . cusp a1 shows two crests , straight in occlusal and lateral views . the longest crest runs distolingually from the apex to cusp b1 . the other crest runs mesially to cusp a2 . the slopes of these crests are subequal . cusp a3 shows two crests , straight in occlusal and lateral views . the longest crest runs mesiolingually from the apex to take part in the u - ridge . the other crest runs distally to cusp a2 . the slopes of these crests are subequal . cusp a2 shows three crests , straight in occlusal and lateral views . the first crest runs distally to cusp a1 . the second crest runs mesially to cusp a3 . the third crest runs lingually but does not join another structure . the distal crest is the shortest and shows the strongest slope . the lingual crest is much wider than both other crests . the lingual and mesial crests define a flat surface on the mesiolingual flank of cusp a2 . a1 – a2 and a2 – a3 notches are equal in depth , but a1 – a2 notch takes place higher than a2 – a3 notch .
a small , supplementary cusp takes place under the labial flank of cusp a3 .
row b includes five cusps . cusp b1 looks like a distally curved semicircle . this cusp is slightly smaller in length and width than cusp a1 , but much smaller in height . cusp b1 is more distal and more lingual than cusp a1 and is mesiodistally aligned with the saddle . cusp b2 is slightly smaller than cusp a2 . cusp b2 is much more lingual than cusp b1 and is labiolingually aligned with the a1 – a2 notch . this cusp is cone - shaped and does not show any crest . three small cuspules take place at the base of the mesiolingual flank of cusp b2 . cusp b3 is directly mesial to cusp b2 . the apex of cusp b3 is slightly more labial the apex of cusp b2 and slightly more mesial than apex of cusp a2 . cusp b3 is subequal in size with cusp b1 and take place slightly lower than cusp b2 . cusp b4 is directly mesial to cusp b3 . this cusp is smaller in all dimensions and takes place lower than cusp b3 . cusp b4 is labiolingually aligned with a2 – a3 notch . cusp b5 is the most mesial cusp of the tooth . this cusp is smaller in all dimensions , takes place lower , and is slightly more labial than cusp b4 . the mesial extremity of cusp b5 shows two crests . one crest runs labially to take part in the u - ridge of the labial basin . the other crest runs lingually to mesially close the lingual basin .
row bb includes two cusps . cusp bb1 is sub - equal in size to cusp b4 and takes place at the same level . cusp bb1 is placed right next to cusps b2 and b3 , directly lingual to the b2 – b3 notch . cusp bb2 is mesial to cusp bb1 but slightly more lingual . this cusp is subequal in length and width with cusp b5 , but slightly higher . a cusp bb3 was possibly present , but this part of the crown is broken . row bb is extended by a crest which runs mesially and then bends labially to close the lingual basin .
only the distal part of mnhn . f . snp 722 is preserved , with cusps
is fragmentary . mnhn . f . snp 722 differs from mnhn . f . snp 78 w by a smaller cusp
in mnhn . f . snp 787 , only the apices of the cusps are abraded by wear . in mnhn . f . snp 366 w , the apices of cusps a1 and a2 show a shallow , distal facet . the apex of cusp b1 shows a steep , mesiolabial facet . the apex of cusp b2 shows a shallow , distal facet . cusp b3 shows a steep , labial facet , which slightly extends on the mesial part of cusp b4 . cusp b4 shows a shallow , distal facet . the sides of the basin show traces of wear but do not develop clear facets . in mnhn . f . snp 61 w , the facets of apices of the cusps a1 and b2 are more extended labially and the carina of cusp a1 is flattened . mnhn . f . snp 61 w differs from mnhn . f . snp 366 w by a horizontal facet on cusp b3 . in mnhn . f . snp 497 w , the distal parts of cusps a1 and a2 each show a shallow , distal facet . the distal part of cusps b2 , b3 , and b4 shows one shallow , distal facet . the remains of cusps a1 and b2 , and cusp b1 are abraded by wear .


tarantula hawks are a large , conspicuous family of long legged wasps which prey on tarantulas .

they use their long legs to grapple with their tarantula prey before paralyzing them with a powerful sting .

their stings are ranked second most painful in the insect world .

the tarantula hawk is also the state insect of new mexico .

they are a solitary wasp species , displaying lekking territorial behavior in their mating rituals .

hemipepsis ustulata

hemipepsis ustulata is a species of tarantula hawk native to the southwestern united states .
bradykinin in hemipepsis ustulata : a novel method for safely milking wasps . - pubmed - ncbi
success in territorial defence by male tarantula hawk wasps , hemipepsis ustulata : the role of residency .
hemipepsis ustulata dahlbom , 1843 . hym . europaea , v . 1 , p . 123 . ♀ .
total ion chromatogram for hemipepsis ustulata venom after trypsinolysis . wasps were restrained in a supine position and agitated to sting parafilm .
an experimental study of the attractiveness of artificial perch territories to male tarantula - hawk wasps , hemipepsis ustulata ( hymenoptera : pompilidae ) .
john alcock , d kemp . the behavioral significance of male body size in the tarantula hawk wasp hemipepsis ustulata ( hymenoptera : pompilida ) . ethology ( 2006 ) .
john alcock . possible causes of variation in territory tenure in a lekking pompilid wasp ( hemipepsis ustulata ) ( hymenoptera ) . journal of insect behavior ( 2000 ) .
type - species : hemipepsis capensis dahlbom . desig . by ashmead , 1900 .
type - species : hemipepsis ( xenopepsis ) commixta arnold . orig . desig .
size of hemipepsis ustulata , but color of body different . seems more like a small pepsis , but i can ' t match to any known to occur in co . help please !
alcock , j . & kemp , j . 2006 . the behavioral significance of male body size in the tarantula hawk wasp hemipepsis ustulata ( hymenoptera : pompilidae ) . ethology 112 : 691 - 698 .
the tarantula hawks or hemipepsis ustulata is the only potential predator of tarantulas . these wasps are found in americas and they are able to kill large tarantulas . giant centipedes are also believed to be the predators of tarantulas .
hemipepsis ochroptera stal , 1857 . ofvers . svenska vetensk . akad . , forh . 14 : 64 .
tarantula hawks are diverse , too . i have personally collected seven species in tucson alone : hemipepsis ustulata , pepsis grossa , p . chrysothemis , p . mexicana , p . thisbe , p . cerberus , and p . mildei .
hemipepsis ustulata ranges from southern california east through nevada to texas , oklahoma , and kansas . it would be well worth it to accumulate more records of this species . more county records are certainly possible , and perhaps state records , too .
alcock , j . and m . carey . 1988 . hilltopping behavior and mating success of the tarantula hawk wasp , hemipepsis ustulata ( hymenoptera : pompilidae ) , at a high elevation peak . journal of natural history 22 : 1173 - 1178 .
wing ventaion is very important for identification of pepsis especially distinction between pepsis and hemipepsis . those characters are as follows :
genera pepsis and hemipepsis have similar biology and can be very difficult to distinguish in the field ( but see
see also
below ) . they are discussed here together under pepsis , though there is a brief account for hemipepsis .
the term “tarantula hawk” applies to more than one species of large wasp . it even applies to more than one genus . here in the united states , the genera pepsis and hemipepsis are the only tarantula hawks , and the largest of our spider wasps in the family pompilidae . hemipepsis ustulata is a common species across much of the western u . s . , one i have encountered in both arizona and colorado .
the method was tested on three individuals of hemipepsis ustulata - large wasps ( up to 5 cm ) with rust colored wings and iridescent blue bodies . hemipepsis ustulata are ectoparasites that use paralyzed spiders as a food source for their young . these wasps are not aggressive , in general , but their sting has a fierce reputation ( schmidt , 2004 ) causing extreme , but short - lived pain . although they are distributed worldwide , they are particularly well known in the southwest u . s . , where they enjoy the status of being the state insect of new mexico .
hemipepsis subg . moropepsis banks , 1934 . amer . acad . arts and sci . , proc . 69 : 6 , 8 .
wahis , r . 1960a . recherches systematiques sur les hemipepsis indo - orientales et australiennes hym . pompilidae , pepsinae . 1 . sur hemipepsis ichneumonea guerin , 1831 , et sus varietes . societe royal d ' entomologie belge bull . ann . 96 : 52 - 58 .
it is not easy to separate hemipepsis from pepsis , but i find that hemipepsis are matte black in color , not the iridescent blue - black that their relatives are . hemipepsis also tends to fly earlier in the season than pepsis , especially in the southwest , but both genera can be found together in the fall , hemipepsis being in the second generation in autumn . wing venation is apparently the only sure way to tell the two apart , but most folks understandably don’t want to get that close to such a large , stinging insect to find out which is which .
wahis , r . 1960b . recherches systematiques sur les hemipepsis indo - orientales et australiennes . 2 . - sur hemipepsis fulvipennis fabricius 1793 et 5 especes voisines , nouvelles ( hymenoptera pompilidae , pepsinae ) . institut royal des sciences naturelles de belgique , bull . 36 : 1 - 16 .
ustulata dahlbom . tex . , okla . , kans . , colo . , n . mex . , utah , ariz , in l . austr . and u .
wahis , r . 1960c . recherches systematiques sur les hemipepsis indo - orientales et australiennes . 3 . - sur hemipepsis robertiana et misera cameron et quelques especes voisines , nouvelles ou peu connues ( hymenoptera pompilidae , pepsinae ) . institut royal des sciences naturelles de belgique , bull . 36 : 1 - 14 .
. . . 1 . some male tarantula hawk wasps hemipepsis ustulata defend landmark shrubs growing on mountain tops , while others only visit a succession of territory sites , generally fleeing from territorial males . to determine whether residency asymmetries contribute to the short duration of most interactions between territorial males and visiting intruders , established residents were held captive for varying . . .
wahis , r . 1961b . recherches systematiques sur les hemipepsis indo - orientales et australiennes . 5 . - sur plusiers especes we sous - especes voisines de hemipepsis speculifer lepeletier de st fargeau . dont deux nouvelles ( hymenoptera pompilidae , pepsinae ) . institut royal des sciences naturelles de belgique , bull . 37 : 1 - 10 .
wahis , r . 1961a . recherches systématiques sur les hemipepsis indo - orientales et australiennes ( hym . pompilidae : pepsinae ) . 6 . a propos de hemipepsis flava ( dahlbom ) , sensu wahis , 1960 ( hym . pompilidae : pepsinae ) . bull . inst . roy . sc . nat . belgique 32 : 1 - 3 .
wasp venom characterization is of interest across multiple disciplines such as medicinal chemistry and evolutionary biology . a simple method is described herein to milk wasp venom without undue risks to the researcher . the wasps were immobilized by cooling for safe handling , restrained , and their venom was collected on parafilm . bradykinin from hemipepsis ustulata was identified by lc - ms / ms during method verification .
van der vecht , j . a . j . w . 1953 . the hemipepsis species of java ( hym . , pompilidae ) . treubia 21 : 685 - 724 .
wahis , r . 1960d . recherches systematiques sur les hemipepsis indo - orientales et australiennes . 4 . sur l ' identite de hemipepsis flava dahlbom , 1843 et description d ' une espece indienne nouvelle h . eximia nov . sp . ( hymenoptera : pompilidae , pepsinae ) . institut royal des sciences naturelles de belgique , bull . 36 : 1 - 10 .
williams , f . 1956 . life history studies of pepsis and hemipepsis wasps in california ( hymenoptera , pompilidae ) . entomological society of america , annals . 49 : 447 - 466 .
van der vecht , j . a . w . , j . 1953 . the hemipepsis species of java ( hym . pompilidae ) . treubia reinwardtia annales bogoriensis 21 : 685 - 724 .
following my last post , here ' s a range of spider hunting wasps - from asia , south america , north america , philippines and africa . the largest here is an 11cm wingspan female hemipepsis . # giant # wasp # entomology # specimen # pompilidae # spiderwasp # hawkwasp # tarantulahawk # tarantulahawkwasp # sting # venom # nature # hymenoptera # pepsis # hemipepsis # parasite # predator
this form of territoriality is called “landmark defense , ” and it is the strategy of choice in situations where defense of a resource used by females is impractical or impossible . while blooming palo verde trees might seem to be a nectar source for female tarantula hawks ( and they are , for pepsis chrysothemis ) , females of hemipepsis ustulata seem to prefer the flowers of creosote bush , which has a wide and scattered distribution in the landscape .
wahis , r . 1959 . nouvelle contribution a l ' etude des hemipepsis de java ( hymenoptera pompilidae , pepsinae ) . institute royal des sciences naturelles de belgique bull . 35 : 1 - 14 .
there are many similarities in the behavior of the hairstreak and the lek - territorial wasp hemipepsis ustulata . males of both insects employ the same kind of chase and spiral flights in defense of many of the same palo verdes . each year they rank the available territorial sites in much the same order as measured by frequency of occupation records . they share many of the same ecological pressures that make it difficult for individual males to monopolize groups of females directly or indirectly .
some other flying insects , such as moths ( sarto i monteys et al . 2016 ) , odonates ( corbet 1999 ) , wasps and flies ( e . g . , kemp and alcock 2003 ) , also exhibit aerial territorial contests . in the territorial wasp hemipepsis ustulata , physical injury during aerial contests was never observed during 20 years of study ( kemp et al . 2006b ) . the erroneous courtship hypothesis may provide new insights into the contest behavior of these animals .
the related spider wasp genera , hemipepsis and entypus , are also quite large and can look very similar to pepsis . these three genera are best distinguished by details of wing venation . for details , see the posts thumbnailed below :
tarantula hawks in general have interesting behaviors , especially the males . dr . john alcock has studied in - depth the territorial behavior of male hemipepsis ustulata in the superstition mountains near phoenix , arizona . they are among the first insects seen in spring , but as the season wears on they become early risers , taking advantage of the brief window of time when sonoran desert temperatures are not too cool , but not incinerating , either . they are looking for prominent outposts , often palo verde trees atop hills .
alcock , j . and k . o ' neill . 1987 . territory preferences and intensity of competition in the grey hairstreak strymon melinus ( lepidoptera , lycaenidae ) and the tarantula hawk wasp hemiepsis ustulata ( hymenoptera , pompilidae ) . the american midland naturalist 118 : 128 - 138 .
wahis , r . 1968 . recherches systematiques sur les hemipepsis indo - oientales et australiennes . 9 . sur quelques especes nouvelles ou peu connues de sumatra ( hymenoptera : pompilidae , pepsinae ) . ann . naturhistor . mus . wien 72 : 391 - 403 .
the breeding season of h . ustulata extends over a 2 1 / 2 month period in the sonoran desert near phoenix , az . individual males are active on mountain ridges for periods averaging about 3 weeks for those wasps captured in the first half of the flight season . many males live more than 1 month .
wahis , r . 1966c . recherches systematiques sur les hemipepsis indo - orientales et australiennes . 8 . sur les especes du sous - genre moropepsis banks , 1934 ( hymenoptera pompilidae , pepsinae ) . institute royale des sciences naturelles de belgique , bull . 42 : 11 - 17 .
large size , to 40 mm + . with blue , blue - green , or blue - violet iridescence . only rarely does hemipepsis have iridescence and most in that genus lack it completely , or if it is present it is very vague . only rarely does entypus reach the size listed above .
wahis , r . 1961c . recherches systematiques sur les hemipepsis indo - orientales et australiennes . 7 . sur quelques especes indiennes de la collection henri de saussure et description d ' especes malaises nouvelles ( hymenoptera pompilidae , pepsinae ) . institut royal des sciences naturelles de belgique bull . 37 : 1 - 19 .
first recurrent vein meeting the second submarginal cell at its basal third , thus making the first discoidal cell rather short and rounded . in hemipepsis the first recurrent vein meets the second submarginal cell at its apical one - third or is occasionally interstitial with the second transervse cubital vein , thus making the first discoidal cell rather long and pointed
here in arizona , spider wasps reach the gargantuan proportions of the so - called “tarantula hawks” in the genera pepsis and hemipepsis . the largest females exceed two inches in body length . the standard appearance of these colorful insects is an iridescent blue - black body with bright orange wings , though some species have black wings with bluish reflections .
large
metallic - sheened
blue - black to greenish wasps , most with orange wings . some species have orange antennae . females have curled antennae , males straight . among the largest and most easily recognized genera of spider wasps , only hemipepsis is easily confused , but orange - winged individuals of entypus are superficially similar . this genus can be recognized by the characters given below :
pulvillar pad and comb in pepsis is rather weak , the pulvillar pad about half as wide as the apical tarsal segment . the pulvillar comb consists of eight to ten weak setulae . in hemipepsis this character is well developed with a large pulvillar pad that is approximately three - quarters the width of the apical tarsal segment and the puvillar comb that consists of 14 - 40 strong setulae .
hemipepsis are often identifiable from photographs to species , but pepsis are more difficult . although it is often possible to narrow down the choices to groups of similar species , it is often impossible to assign a species name with certainty . an extensive knowledge of local fauna helps , as well as extensive work with museum specimens . despite the large size of members of this genus many photographs will likely remain unidentified , as some of the species are very similar .
the droplets secreted by hemipepsis were around 1 μl with about 4 - 8 droplets able to be collected in a single session . venom samples were purified and preconcentrated with c - 18 spin columns , lyophilized , and stored frozen until the time of analysis . thawed venom was redissolved in 20 μl of 0 . 1 % formic acid solution and a trypsin digestion was performed to reduce coelution issues with proteins . a sample injection of 2 μl was then analyzed by liquid chromatography - tandem mass spectrometry ( lc - ms / ms ) on a q exactive mass spectrometer with an easy nanolc - 1000 system ( thermo fisher scientific , u . s . a ) . separation was conducted using a c18 column ( 75 μm x 150 mm ) . reverse phase chromatography was performed with ( a ) 0 . 1 % formic acid / water and ( b ) 0 . 1 % formic acid / acetonitrile . a four - step , linear gradient was used for the lc separation ( column pre - equilibration with 2 % b for 10 min ; 2 % to 30 % b in the first 47 min . ; followed by 80 % b for 13 min . ) . bradykinin was targeted in the data sets since bradykinin and related compounds have been found in other solitary wasp species ( konno et al . , 2002 ; picolo et al . , 2010 ) .
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once a male wasp claims a lookout tree , it vigorously defends it from competing males . most interactions consist of the resident male flying off his perch in a brief foray to chase away the intruder . there is usually only feeble resistance , if any , from the interloper . males that have been displaced from their own territories , however , are more likely to try and displace another male . this takes the form of long , ascending flights in which the wasps spiral around each other as they climb to heights of thirty feet or more . sometimes they escalate to altitudes where they can no longer be observed without binoculars .
these aerial duels are contests of both endurance and speed . the faster and fitter male is thus the one more likely to win .
female wasps cruising past an occupied tree elicit an entirely different response . the resident male gives chase and , if he successfully reaches her , attempts to couple with her in the air and descend to the ground . there , they mate briefly if she is receptive . the male then returns to his perch , which he hopes has not been claimed by another male while he was um , otherwise engaged .
once mated , a female tarantula hawk goes about the business of finding occupied tarantula burrows and baiting the spider into battle aboveground . these burrows are also widely scattered . so , male wasps can’t stake out a tarantula “colony” because there is no such thing , at least not for spiders the size needed by the female wasp .
some of the hunting activities of these wasps may take place at night , or at least at dusk , for i have found females in southeast arizona coming to blacklights at night . i have also found females feeding on oozing sap of desert broom ( baccharis sarothroides ) . here in colorado springs i have seen both genders at a blooming saltcedar tree ( tamarix sp . ) , though the female simply circled the tree and flew off again .
sources : alcock , john . 1984 . “ridgetop rendezvous , ” natural history 93 : 42 - 47 . evans , howard e . 1997 . “spider wasps of colorado ( hymenoptera , pompilidae ) : an annotated checklist , ” great basin naturalist 57 ( 3 ) : 189 - 197 .
hi , aaron : the wasp in your image is clearly in the family sphecidae , not pompilidae . if it was that large , then it can only be sphex tepanecus , which is not uncommon in arizona . . . . but idaho ?
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m . a . n . mourão 1 , 2 , a and p . e . c . peixoto 3 , b , *
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in many animal species , males fight for ownership of territories for mating purposes ( andersson 1994 ) . the fighting behaviors used by males differ greatly among species ( arnott and el wood 2009 ) , varying from disputes without any contact between the rivals ( e . g . , alcock and bailey 1997 ) to interactions with physical contact , injuries , and risk of death ( e . g . , bean and cook 2001 ) . such variation is common even in species that fight for the possession of mating sites during flight , such as butterflies , dragonflies , and damselflies ( marden and waage 1990 , peixoto and benson 2011a , junior and peixoto 2013 ) .
many theoretical models aim to explain the rules used by males to decide the winner of a territorial contest ( enquist and leimar 1983 , mesterton - gibbons et al . 1996 , payne 1998 ) . despite differences among these models about the ways by which disputes are decided , all assume that the chance of winning depends on two parameters : ( 1 ) the individual’s fighting ability ( rhp , resource holding potential ; parker 1974 ) , which can be determined by physical ( e . g . , size and weight ) or physiological traits ( e . g . , fat , muscle , and immunocompetence ) ( see suhonen et al . 2008 ) ; and ( 2 ) the individual’s motivation to invest in the contest ( rv , resource value ; parker 1974 ) , which is often determined by the previous experience of the owner of the contested resource and by the local quality of the defended site ( e . g . , briffa and elwood 2001 , bergman et al . 2010 ) .
because all the models previously mentioned postulate that rhp differences between rivals determine the chance of winning , to be able to evaluate such models empirically it is necessary to identify which morphological or physiological traits determine the males’ most often used for this purpose is to establish correlations between specific male traits and the chances of winning a territorial contest . however , this approach is unsuitable in establishing the mechanism that connects the trait to how costs are accrued during the contest , and it can lead to the establishment of spurious relationships ( lailvaux and irschick 2006 ) .
to avoid these results , it is necessary to establish the functional relationship between male traits and fighting behavior ( irschick et al . 2007 ) . one way to establish this functional relationship is to classify the fighting behaviors according to the occurrence of physical contact between rivals ( e . g . , peixoto and benson 2011b ) . if the fights occur with physical contact , injuries should be the main form of cost accrual during the fight . in this sense , features such as body mass should increase the individual’s resistance to injuries from rivals or the ability to inflict damage . therefore , traits related to physical strength must be functionally important to determine male rhp ( e . g . , forslund 2000 ) . if fights occur without physical contact , however , the costs should be determined by the energy depletion during the fight . the winner must be the individual that is able to persist for a longer period in the contest . consequently , physiological features such as energetic reserves may be more important in determining the individual’s fighting ability ( e . g . , marden and waage 1990 ) .
in many dragonfly species , males establish mating territories and fight for the ownership of these sites ( corbet 1980 , córdoba - aguilar 2008 ) . the fighting behaviors usually comprise a combination of circular chase flights , upward spiral flights , and straight back - and - forth persecutions ( córdoba - aguilar 2008 ) . in some species , males may exhibit physical contact during disputes ( corbet 1999 , switzer and schultz 2000 , junior and peixoto 2013 ) , but in others , the males exhibit displays without contact ( waage 1988 , switzer 2004 ) . features such as energy reserves ( marden and rollins 1994 ) and parasitism ( contreras - garduño et al . 2006 ) seem to predominate as rhp determinants in this group .
the study was conducted in a 130 - m stretch of urubú stream ( 15°42 ' s , 47°51 ' w ) , located about 9 . 5 km from the center of brasilia , df , brazil . the average depth of the stream at that location was 60 cm , and the average width was 3 . 5 m . the weather is tropical - high altitude , with two distinct seasons : a wet and rainy summer , with an average temperature of 29 . 7°c ( sd = 0 . 72 ) and average rainfall of 203 . 9 mm ( sd = 27 . 2 ) , and a dry and cold winter with an average temperature of 12 . 5°c ( sd = 2 . 5 ) and average rainfall of 24 . 9 mm ( sd = 76 . 7 ) ( inmet 2011 ) .
to evaluate whether disputes occur with or without physical contact , samples were collected in campaigns composed of two consecutive days between november 2011 and march 2012 . on the first day of each campaign , several males were marked with a number written on one wing with a black permanent marker that allowed visual identification of males without recapture . on the second day , behavioral observations of males and the removal experiment were done to evaluate the traits associated with the chance of winning a contest ( see “removal experiments” below for details ) . behavioral observations were made on 31 resident males ( males that were defending the territories at the beginning of our observations ) and 31 substitute males ( males that occupied the territories after the resident males were experimentally removed ) only on sunny days from 09 : 00 to 14 : 00 , when individuals are more active . during the observations , the behaviors of individuals were recorded for 10 min using a mini - portable recorder , with particular reference to the occurrence of physical contact during territorial disputes exhibited by the observed males (
) . afterwards , the recordings were used to time how long each individual spent in each behavior .
after the behavioral observation , we performed a removal experiment to evaluate male traits that may determine the winning chances . for this experiment , whenever a male exhibited signals of territorial defense ( i . e . , fights with conspecific males or flight patrols ) , it was removed from the defended site ( resident male ) . after the male was removed , the territory remained under observation until a new male arrived ( substitute male ) . upon arriving on the observed site , the substitute male was captured to receive a mark , consisting of a point on one wing . this approach was done with only one male at a time , and it never took more than 1 min to avoid stress on the manipulated animal ( all marked males resumed the territorial defense after being released ) . after releasing the marked substitute male , it was observed for 10 min and then captured . after this , both males ( resident and substitute ) were measured for weight , wing area , fat content , and residual muscle mass ( for a detailed description , see “morphological and physiological measures” below ) .
removal experiments have been used successfully to evaluate male traits that are important for contest resolution ( peixoto and benson 2008 , 2011a ) . in these experiments , it is expected that resident males are individuals that have disputed the ownership of territories and won , whereas substitute males represent weaker individuals that were unable to obtain territories . thus , if fights occur with physical contact , it is expected that resident males have a greater wing area or greater body mass than the rival males . alternatively , if fights occur without physical contact , it is expected that resident males have a greater fat content or greater investment in thoracic muscles for flight than substitute males .
a caliper was used to measure the length and width of each wing . the wing length was measured as the distance between the wing insertion on the thorax and its apex . the width was recorded as the longest distance between the wing edges on the perpendicular direction to the length axis . wing area was estimated by calculating the ellipse area , considering the wing length as the largest diameter and width as the smallest diameter . then , the wing area of the four wings was added to estimate the total wing area for each male .
to estimate fat content , males were kept in an oven at ∽60°c for 48 hr . later , the thorax ( without wings and legs ) and abdomen were separately weighed on a balance ( accuracy 0 . 0001 g ) . the summed mass of thorax and abdomen after dehydration was used as a measure of male size ( dry mass ) . after weighing , both parts of each animal were submerged in a closed bottle containing 10 ml of chloroform for 48 hr to extract lipids . after removing them from the chloroform , the body parts were kept in an oven at 60°c for an additional 48 hr before being weighed again . to estimate the amount of thoracic muscle mass available for flight ( flight muscle ratio ) , the same procedure was used . however , only the thorax weight ( after lipid extraction ) was measured before and after being immersed in 0 . 2 m potassium hydroxide ( plaistow and siva - jothy 1999 ) . the mass difference before and after immersion in chloroform was used to estimate fat content , and the mass difference before and after the immersion in potassium hydroxide was used to estimate muscle mass . because larger individuals tend to have more fat and muscle mass than smaller ones , we calculated the residuals of a linear regression between the fat content and the dry body mass of individuals before the lipid extraction ( residual fat ) and between the muscle mass and dry body mass ( residual muscle mass ) to obtain values of fat content and muscle mass independent of size ( marden 1989 , marden and chai 1991 , marden and rollins 1994 ) .
a logistic regression was used to assess the effect of each trait ( wing area , dry weight , residual fat content , and residual muscle mass ) on the probability of a focal male to be territorial ( hosmer and lemeshow 1989 ) . because data were collected from pairs of males ( resident and substitute ) , we adopted a modification to maintain the pairing ( kemp 2000 ) . for this , 16 pairs were randomly chosen . for each pair , the resident individual was designated as the focal male ( focal state 1 ) . for the remaining 15 pairs , the substitute male was designated as focal individual ( focal state 0 ) . when the focal state was 1 , the values of resident male traits were subtracted from the values of its substitute pair . inversely , when the focal state was 0 , the values of the substitute male were subtracted from the values of its resident pair . thus , if resident males possess a higher fighting ability than substitutes , it was expected to find positive values for the differences between male traits associated with focal pairs whose state was 1 and negative values associated with focal pairs whose state was 0 ( e . g . , peixoto and benson 2008 ) . to identify the most parsimonious model describing the relationship between the focal status and differences in the male traits , we used a corrected version for small samples of the akaike information criterion ( burnham and anderson 2002 ) .
) . resident and substitute males stayed perched for the greatest proportion of the time ( resident males : mean = 519 . 2 s , sd = 41 . 1 ; substitute males : mean = 524 . 8 s , sd = 39 . 2 ) . when males were not perched , they spent most of their time in flight and flight patrol , respectively . although the 62 males ( resident and substitute combined ) had been involved in 180 disputes , the proportion of time spent in fights accounted for a smaller proportion of the time budget of males . on average , the fights lasted 10 . 1 sec ( sd = 5 . 3 ) for resident males and 8 . 09 sec ( sd = 3 . 88 ) for substitute males .
average proportion of time that the resident and substitute males of the dragonfly macrothemis imitans spent in each behavior during 10 min of behavioral observation . the proportion of time in which males remained resting is not shown . bars represent the standard error . high quality figures are available online .
fights between males always initiated when a male entered the site defended by a resident or by a substitute male . in both situations , the territory owners flew toward the intruders , colliding against them . after the collision , the territory owner ( for both resident and substitute males ) performed straight flights and / or circular chase or collision against the intruder . after expelling the invader , the winner returned to the perch of origin . the resident males were involved in 99 disputes and won 100 % of the contests , and the substitute males were involved in 81 disputes and also won 100 % of the contests . we observed two matings performed by resident males and eight matings by substitute males ( corresponding to the smaller proportion of the time budget of males ) . no individual without a territory was observed mating .
summary of the logistic model describing the probability of the focal male of macrothemis imitans ( n = 31 ) being the resident in relationship to the value difference between their traits and its non - focal pair .
the models are ordered according to their aicc values ( aicc represents the value of the akaike information criterion corrected for small samples ; äi is the value difference between the most parsimonious model and model i , wi is the weight of akaike model i ) .
probability of the focal male macrothemis imitans to be the resident in relation the residual muscle mass difference from its non - focal pair . overlapping points represent different samples with the same focal result and residual muscle mass difference . high quality figures are available online .
because resident and substitute males in ownership of the territories won all the fights , it is possible that the asymmetry in territorial status determines the winner ( davies 1978 ) . although there is a lack of clear empirical demonstrations , theoretical models suggest that males may determine the contest winner through uncorrelated asymmetries , such as the simple distinction between territory owners and intruders ( maynard smith and parker 1976 , kokko et al . 2006 ) . it is also possible that disputes are settled on the basis of differences in territory value for each rival ( parker and rubenstein 1981 , enquist and leimar 1987 ) . in many species , individuals in ownership of territories present greater investment in fights than intruder rivals ( switzer 2004 , takeuchi and honda 2009 , bergman et al . 2010 , peixoto and benson 2011b ) . consequently , it is possible that even individuals with lower fighting ability remain as residents because asymmetries in motivation associated with the value that each rival attributes to the territory .
the lack of correspondence between male traits that are important in determining residency and the occurrence of physical contact between rivals during the fights is intriguing . it is possible that injuries are , in fact , an important fighting cost ( e . g . , junior and peixoto 2013 ) . in this situation , there may be alternative individual traits that affect contest settlement . perhaps the fighting ability of males is not determined solely by a particular trait , but by a combination of different characteristics that determine the overall male condition ( kemp et al . 2006 ) . on the other hand , vieira and peixoto ( 2013 ) have demonstrated that the functional relationship between fighting behaviors and how individual traits affect cost accrual during the fight is not universal . in this sense , it is possible that collisions do not generate any fighting cost . if this is true , disputes should be mainly determined by asymmetries in male motivation associated with differences in resource value , as occur in some butterfly species ( takeuchi and honda 2009 , bergman et al . 2010 ) .
we thank the researchers from the department of ecology at the university of brasilia ( unb ) , rosana tidon , and regina macedo for the laboratory access for the physiological extractions of males of m . imitans and for the theoretical content related to sexual selection , respectively . we also thank lorenzo zanette ( federal university of ceará ) and márcio cardoso ( federal university of rio grande do norte ) for comments on the manuscript . m . a . mourão received a scholarship provided by funcap in the early months of work .
contreras - garduño j . , canales - lazcano j . , córdoba - aguilar a . . 2006 . .
irschick d . j . , herrel a , vanhooydonck b . , van damme r . . 2007 . .
kokko h , lópez - sepulcre a . , morrell l . j . . 2006 . .
mesterton - gibbons m . , marden j . h . , dugatkin l . a . . 1996 . .
peixoto p . e . c , benson w . w . . 2008 . .
peixoto p . e . c , benson w . w . . 2009 . .
suhonen j , rantala m . j . , honkavaara j . . 2008 . .
dragonflies and damselflies : model organisms for ecological and evolutionary research . pp . 203–217 . oxford university press
vieira m . c , peixoto p . e . c . . 2013 . .
( 2002 - 5 ) . listed below are the former names . . . as they appeared in main earlier references by hurd
p . angustimarginata
= p . basifusca , see part 3 : p . 141
p . arizonica
= p . cassiope , see part 2 : p . 92
p . azteca
= p . sommeri , see part 3 : p . 96
p . cerberus
= p . menechma , see part 3 : p . 131
p . elegans
= p . menechma , see part 3 : p . 131
p . formosa
= p . grossa , see part 2 : p . 58
p . saphirus
= p . ruficornis , see part 3 : p . 232
p . venusta
= p . terminata , see part 2 : p . 68
mostly southwest us , east to kansas . only three species occur in the east , the most common and wide - ranging one being p . menechma . . . the other two , p . marginata and p . ruficornis ( = saphirus ) , have been recorded from south florida ( possibly as accidentals from the west indies ? ) . the genus also extends far south into neotropics , where it is much more speciose .
larvae feed on tarantula and some other large spiders . adults take nectar , and are particularly fond of milkweed ( genus asclepias ) .
mating takes place on flowers , or sometimes near a
hilltop
, such as a large cactus , defended by the male . females seek out tarantula burrows , paralyze them , and bury it in a burrow , laying a single egg . larvae feed on the paralyzed spider .
these wasps are reputed to have a very powerful sting , though they are not aggressive .
the new world tarantula - hawk wasp genus pepsis fabricius ( hymenoptera : pompilidae ) . c . r . vardy . 2005 . zoologische verhandelingen / zoologische mededelingen .
revision of the nearctic species of the pompilid genus pepsis ( hymenoptera , pompilidae ) paul d . hurd . 1952 . american museum of natural history , new york .
nearctic wasps of the subfamilies pepsinae and ceropalinae henry k . townes . 1957 . smithsonian institute press ( bulletin 209 ) .
catalog of hymenoptera in america north of mexico karl v . krombein , paul d . hurd , jr . , david r . smith , and b . d . burks . 1979 . smithsonian institution press .
american insects : a handbook of the insects of america north of mexico ross h . arnett . 2000 . crc press .
the common insects of north america lester a . swan , charles s . papp . 1972 . harper & row .
pompilidae ( spider wasps ) spider wasps are usually black , sometimes with patches of yellow or red . they have stingers , which they use to paralyze spiders . the spider is then dragged back to its den or to a specially constructed chamber in the ground , whereupon a single egg is laid on the spider , and the chamber is sealed . the wasp larvae are parasitoid on spiders . spider wasps are fairly common visitors to flowers .
hilty , j . editor . 2015 . insect visitors of illinois wildflowers . world wide web electronic publication . illinoiswildflowers . info , version ( 09 / 2015 ) see : abbreviations for insect activities , abbreviations for scientific observers , references for behavioral observations
, though these names can be generally applied to any very large stinging wasps . furthermore , in some parts of venezuela and colombia , it is called
pompilids typically have long , spiny legs ; the hind femur is often long enough to reach past the tip of the abdomen . the tibiae of the rear legs usually have a conspicuous spine at their distal end . the first two segments of the abdomen are narrow , giving the body a slender look . the pompilid body is typically dark ( black or blue , sometimes with metallic reflections ) , but many brightly colored species exist . from a lateral view , its pronotum looks rectangular and it extends back to the tegulae , near the base of the wings . most species are macropterous ( having long wings ) , but a few brachypterous ( short - winged ) and apterous ( no wings ) species are known .
( the mesepisternal sclerite , a region on the side of middle segment of the thorax above the point where the legs join ) . like other vespoidea , they have antennae with 10 flagellomeres in the female and 11 in the male . most pompilidae have straight inner eye margins . the hind wings do not have a distinct claval lobe , but they have a distinctive jugal lobe . the hind leg has a tibial spur with a tuft or row of fine hairs . the legs are long and slender with the tips of the tibia ( metatibia ) long enough to extend beyond the tip of the abdomen ( metasoma ) . sexual dimorphism is not pronounced , although females are often larger than the males . coloring and wing appearance varying greatly among the many species , though black is the most common color , with contrasting
markings of orange , red , yellow , or white also being fairly common .
as a host for feeding their larvae . they paralyze the spider with a venomous
. once paralyzed , the spider is dragged to where a nest will be built – some wasps having already made a nest .
a single egg is laid on the abdomen of the spider , and the nest or burrow is closed . spider wasps protect their nests by putting dead ants into the outermost chamber , the chemicals within deterring predators .
the size of the host can influence whether the wasp ' s egg will develop as a male or a female ; larger prey yield the ( larger ) females .
a complex set of adult behaviors can then occur , such as spreading soil or inspecting the area , leaving the nest site inconspicuous . when the wasp larva hatches , it begins to feed on the still - living spider . after consuming the edible parts of the spider , the larva spins a silk
, usually emerging as an adult the next summer . some ceropalines lay the egg on a still - active spider , where it feeds externally on
. in time , that spider will die , and the mature wasp larva will then pupate .
currently , the pompilidae are split unequally into six subfamilies throughout most of the world . ceropalinae and notocyphinae contain two genera each and occur in central and south america and the oriental region . the epipompilinae contain a single genus and occurs in the neotropics , the australasian region , and the afrotropical region . the ctenocerinae contain two genera in the neotropics , four in australia and 11 in africa . pepsinae and pompilinae are the most diverse , and the remaining genera are split between them .
as mentioned above , the pompilidae get their common name , “spider wasps” , from their notable behavior of hunting and killing spiders , often larger than themselves , as food for their larvae . they provide each of their larvae with a single prey / host , which must be large enough to serve as its food source throughout its development .
adult is determined by the size of the host provided for it by its mother , the seasonal frequency of host sizes implicitly will determine the size variation in adult wasps .
because of the large body size of their prey , pompilids usually will either construct burrows near the site of attack or use the host’s own burrow or tunnel . the paralyzed spider is concealed in a burrow so larva can develop without disruption by other parasites or scavengers . some pompilids only temporarily paralyze their hosts , which regain activity before being killed by the maturing wasp larva .