macrodactylus subspinosus is a north american beetle of the family scarabaeidae .

it is one of at least two beetles in this family known as the " rose chafer " , the other being the european cetonia aurata .

m. subspinosus occurs from eastern canada to colorado and is considered a pest of many crops and flowers .

it is given its common name of rose chafer because it eats the leaves of roses , although it also feeds on many other plants .

macrodactylus subspinosus

development of an attractant for the scarab pest macrodactylus subspinosus ( coleoptera : scarabaeidae ) .
the rose chafer ( macrodactylus subspinosus ) is a common pest in vineyards in the northeastern united states .
development of an attractant for the scarab pest macrodactylus subspinosus ( coleoptera : scarabaeidae ) . - pubmed - ncbi
the rose chafer ( macrodactylus subspinosus ) is commonly found in many areas of minnesota , particularly areas with sandy soil .
rose chafer ( macrodactylus subspinosus fabricius ) adults emerge at bloom and immediately attack grape blossoms and skeletonize leaves , eating all the tissue around large veins . they can cause extensive damage to foliage and completely destroy blossom buds and / or developing berries , resulting in reduced grape yields . damage can be severe in vineyards with sandy soils , the preferred habitat for these beetles .
in identifying the rose chafer ( macrodactylus subspinosus syn . cetonia aurata ) , one will note that it is a tan , long - legged , slender beetle from 5 / 16 to 15 / 32 inches long ( 8 - 12 mm ) . as you can see , this beetle is smaller than the japanese beetle and does differ in appearance . they are , however , alike in the appetite and the damage they do .
in the past few years , populations of the rose chafer beetle , macrodactylus subspinosus , have been abundant in some regions of southwest michigan . these beetles can be found in grape growing regions across michigan and beyond ; however , outbreaks in the lawton , mich . , region in recent years have been causing some high levels of damage to clusters during the bloom period . with this recent history , it is a good time to review the biology and lifecycle of this pest and review the available options for control .
we analyzed the flight of the rose chafer macrodactylus subspinosus ( f . ) ( melolonthinae : scarabaeidae ) and the japanese beetle popillia japonica newman ( rutelinae : scarabaeidae ) in a wind tunnel with controlled humidity , temperature , light , and airflow . the data indicate that an optimum combination of light and temperature dramatically improves their response to lures . both species took off upwind , oriented to , and contacted the odor source well ( 40–60 % ) when light intensity was > 50 % of a clear day , temperature was 26–27°c , and relative humidity was 65–75 % .
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integrated pest management ( ipm ) is a sustainable approach to managing pests that combines multiple approaches including prevention , avoidance , pest monitoring and suppression in a manner that minimizes public health , economic , and environmental risk . ipm serves as a framework to provide an effective , comprehensive , low - risk approach to protect people and resources from pests .
ultimately , the goal with an ipm program is to help stakeholders deal with pests—insects , plant diseases , weeds , and more—with methods that reduce risks to public health and the environmental while saving money .
the north carolina extension ipm program serves as a focal point for team building , communication and stakeholder participation in integrated pest management ( ipm ) within the state . program goals include promoting effective and economical management of pests , reducing risks to human health from pests and pest management practices , and minimizing environmental effects through the adoption of ipm on a variety of crops and settings in north carolina . these goals are achieved by the timely delivery of ipm technology and research information to stakeholders in all regions of the state .
leadership of the extension ipm program in north carolina is provided by an ipm coordinator ( designated by the director of the north carolina cooperative extension service ) , an established advisory board , and working groups . dr . danesha seth carley , the current ipm coordinator , accepted the nc ipm coordinator position in 2013 .
the extension ipm coordinator involves faculty and staff from north carolina state university and north carolina a & t state university in ipm activities across the state , communicates program successes , and maintains stakeholder input via the advisory board and the ipm portal , as well as multiple training sessions and meetings .
the advisory board provides advice to the extension ipm coordinator regarding the direction of the ipm program in the state . it is composed of a wide variety of ipm stakeholders , including north carolina state university and north carolina a & t state university faculty , non - governmental agencies , environmentalists , north carolina department of agriculture & consumer services personnel , farmers and agricultural consultants .
the north carolina extension ipm program focuses on delivering ipm content to our stakeholders and community members through a variety of activities . the primary activities include information delivery , pest monitoring and data managment , evaluation and needs assessment tools development , and programs to magnify statewide ipm impacts .
the extension ipm program is a cooperative effort of the usda nifa , north carolina cooperative extension service , north carolina state university , and north carolina a & t state university . it also works in partnership with the southern ipm center , located at north carolina state university in raleigh , and acts to promote ipm in north carolina and the southern united states .
funding for the north carolina extension ipm program is provided by competitive grants from the u . s . department of agriculture’s national institute of food and agriculture ( nifa ) .
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adult rose chafers feed primarily on flower blossoms , especially roses and peonies , causing large , irregular holes .
rose chafers also feed on the foliage of many trees , shrubs and other plants , such as rose , grape , apple , cherry and birch .
they damage leaves by eating the leaf tissue between the large veins , a type of injury known as skeletonizing .
it has short antennae that have a series of flat plate - or page - like segments .
adult beetles are seen coming out of the ground in late may and early june .
they feed on plants for three or four weeks , generally until late june .
two to three weeks later , the eggs hatch into small , white grub‑like larvae which feed on the roots of grasses and weeds .
plants located on sandy sites are have more chances of being attacked as rose chafers prefer sandy soil to lay eggs .
adult beetles damage leaves and the larvae feed on the roots of grasses and non - crop plants .
rose chafers contain a toxin and can be deadly to birds ( including chickens and small animals ) when they eat these beetles .
protecting plants from rose chafers can be challenging , especially when large numbers are present .
check for rose chafers in your garden starting in late may , especially if you have a history of rose chafer infestations .
when small numbers are present , pick rose chafers from plants and drop into pails of soapy water to kill them .
you can also use a physical barrier , like a cheesecloth or floating row cover .
place the barriers around the plants just as rose chafers become active and take them down after the rose chafers are done feeding ( after june ) .
if large numbers of rose chafers are present , you can treat plants with a garden pesticide . you may need to treat plants more than once when rose chafers are numerous .
you can find the common name for a pesticide by looking under active ingredients . look closely as this is usually in small print .
examples of common names of active ingredients include : bifenthrin , esfenvalerate , cyfluthrin , imidacloprid , permethrin and carbaryl .
caution : mention of a pesticide or use of a pesticide label is for educational purposes only . always follow the pesticide label directions attached to the pesticide container you are using . remember , the label is the law .
be sure that the fruit / vegetable you wish to treat is listed on the label of the pesticide you intend to use . also be sure to observe the number of days between pesticide application and when you can harvest your crop .
these diagnostic tools will guide you step - by - step through diagnosing a plant problem or identifying a weed or insect .
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( fabricius ) and is found from eastern canada south to colorado . the second species is the western rose chafer ,
horn and it occurs in arizona , new mexico , texas and south into central mexico . both species are very similar in size , body shape , and habits . control measures for one species should also apply to the other .
( fabricius ) , is a tan , long - legged slender beetle from 8 - 12 mm long . the rose chafers damage plants by feeding on the flowers , newly set fruit and foliage . on roses it skeletonizes the leaves in the same way as other scarab beetles like the japanese beetle , chinese rose beetle , and the serica garden beetle . it feeds on on the foliage of many different plants and it is greatly attracted by flowers . on june 27 , 1997 i observed numerous beetles on milkweed flowers in an area near galesville , wi . i also swept this beetle from the blooms and foliage of several plants in the area . the rose chafer is native to the northeast from eastern canada south to colorado . i have seen samples from fenton , mi and galesville , wi .
adult beetles emerge from the soil in late may through early june and and they live for about one month . mating occurs soon after emergence and the females lays her 24 - 36 eggs continuoulsy for about two weeks in the soil in grassy sandy areas . upon hatching the larvae burrow in the soil and feed on the roots of grasses and weeds . the overwinter as a larvae and continues development in the spring . the full grown larva or grub is white and measures up to 18 mm in length . pupation occurs in the spring . the rose chafer has only one generation per year .
scouting or monitoring for the presence of rose chafers is very important for adequate control . rose chafers are most prevalent in areas with sandy soil . adults readily attracted to many types of flowers and feed on the foliage many plants . the larvae or white grubs feed on the roots of grasses and many cultivated and wild plants . from late may through july , the foliage should be inspected for skeletonized leaves and the presence of adult beetles on the leaves and flowers .
rose chafers can be handpicked and destroyed if the infestations are light . rose chafers can be very numerous especially in areas with sandy soils . in these cases insecticides may not give satisfactory control as rose chafers can move in from surrounding untreated areas or the insecticides do not seem to prevent feeding activity for very long . however , after about 2 - 3 weeks of heavy damage the beetle numbers appear to subside . insecticides used in the control of rose chafers include carbaryl ( sevin ) , acephate ( orthene ) , diazinon , and chlorpyrifos ( dursban ) , tempo , talstar , mavrik , rotenone , etc .
it was interesting to read that birds sometimes die from eating adult rose chafers . the beetles apparently have a chemical that affects the heart of small , warm - blooded animals .
horn occurs primarily in arizona and new mexico south into mexico . it resembles the rose chafer and its damage is similar . it is 10mm long and the body is rather slender , long - legged , and yellowish brown in color . its pubescence is longer and denser than that of the rose chafer . i have collected numerous beetles in southern arizona and central mexico feeding on many kinds of plants .
essig , e . o . 1926 . insects of western north america . the macmillan company , ny , p . 445 .
johnson , warren t . and howard h . lyon . 1988 . insects that feed on trees and shrubs , 2nd edition , cornell university press , ithaca , ny . , pp . 236 - 37 .
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these systems aim to suppress pest populations below a predetermined economic injury level ( eil ) .
ipm extends the concept of integrated control to all classes of pests including plant pathogens , insects , and weeds .
it also allows for safer pest control and is expanded to include all control tactics such as pesticides , host - plant resistance , and cultural manipulations .
the ohio grape ipm program will provide information and resources for managing diseases , insects , and weeds . it is a comprehensive program designed to encourage collaboration among ohio agricultural research and development center ( oardc ) scientists with different specializations to better address the pest management needs of the grape industry .
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upcoming events 2018 bugguide gathering in virginia july 27 - 29 : registration and discussion photos of insects and people from the 2015 gathering in wisconsin , july 10 - 12 photos of insects and people from the 2014 gathering in virginia , june 4 - 7 . photos of insects and people from the 2013 gathering in arizona , july 25 - 28 photos of insects and people from the 2012 gathering in alabama photos of insects and people from the 2011 gathering in iowa photos from the 2010 workshop in grinnell , iowa photos from the 2009 gathering in washington
very similar to and hard to distinguish from m . angustatus , with which it used to be considered synonymous
suspect ( art evans , pers . comm . to = v = 5 . iv . 2011 )
adults emerge in early summer and feed on flowers , some leaves . they live for up to 6 weeks . mating occurs on food sources . eggs are laid deep ( 13 - 15 cm ! ) in soil and hatch in one to three weeks . larvae feed on roots and overwinter deep in soil . pupation in early spring in the soil , just under the surface .
the beetles of northeastern north america , vol . 1 and 2 . downie , n . m . , and r . h . arnett . 1996 . the sandhill crane press , gainesville , fl .
scarab beetles ( coleoptera : scarabaeidae ) of south carolina phillip j . harpootlian . 2001 . clemson university public service .
disclaimer : dedicated naturalists volunteer their time and resources here to provide this service . we strive to provide accurate information , but we are mostly just amateurs attempting to make sense of a diverse natural world . if you need expert professional advice , contact your local extension office .
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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
n common north american beetle : larvae feed on roots and adults on leaves and flowers of e . g . rose bushes or apple trees or grape vines
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- common north american beetle : larvae feed on roots and adults on leaves and flowers of e . g . rose bushes or apple trees or grape vines
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detailing the physical features , habits , territorial reach and other identifying qualities of the rose chafer .
rose chafers love to congregate on their namesake and chip away at a rose bush ' s health , eating its blooms and leaves . .
antennae : beetles have a pair of antennae on the head used as sensors .
head : the head is home to the insect ' s eyes , antennae , and mandibles ( jaws ) .
thorax : holds the three pairs of legs as well as vital internal organs .
elytron : one of two wing cases on a beetle that protects its wings ( plural : elytra ) .
legs : beetles have three pairs of legs located at the thorax , numbering six legs in all .
an insect ' s reach is not limited by lines drawn on a map and therefore species may appear in areas , regions and / or states beyond those listed below as they are driven by environmental factors ( such as climate change ) , available food supplies and mating patterns . grayed - out selections below indicate that the subject in question has not been reported in that particular territory . u . s . states and canadian provinces / territories are clickable to their respective bug listings .
the map below showcases ( in red ) the states and territories of north america where the rose chafer may be found ( but is not limited to ) . this sort of data can be useful in seeing concentrations of a particular species over the continent as well as revealing possible migratory patterns over a species ' given lifespan . some species are naturally confined by environment , weather , mating habits , food resources and the like while others see widespread expansion across most , or all , of north america .
it is important to identify the type of grub in your lawn before treating . it is usually necessary to consult an expect for grub identification . adults are pictured to help with beetle identification . keep in mind , however , that adults do not always stay in the same area as the larvae .
early indications of grub infestation are irregular patches of dry grass , flocking birds , or areas of turf being torn up by raccoons , possums and skunks looking for a tasty treat .
if grubs have been eating the root system , patches of turf will come up easily from the soil surface , like pulling up a corner of carpeting , and the soil will be full of grubs .
it is important to identify the type of grub in your lawn . above , left to right : japanese beetle , european chafer and june beetle larvae .
japanese beetle larvae are typical white grubs that are c - shaped when disturbed . first instar larvae are about 1 / 16 inch long while the mature third instars are about 1 - 1 / 4 inch long .
adult japanese beetles are 7 / 16 - inch long metallic green beetles with copper - brown wing covers . a row of white tufts ( spots ) of hair project from under the wing covers on each side of the body .
effective biocontrol of japanese beetles includes winsome fly istocheta ( = hyperecteina ) aldrichi . adult beetles with eggs on the pronotum , like the one pictured , should not be destroyed .
european chafer larvae are typical c - shaped white grubs , reaching a maximum size of 1 / 4 inch wide and 1 inch long .
european chafer adults are 1 / 2 inch long . males and females are a uniform tan or light brown color .
grubs are whitish with brown heads and range from 1 / 2 to 1 inch in length . these are the largest grubs found in turf .
all species of phyllophaga are called may or june beetles . adults are about 1 inch long and a chestnut brown color and fly to lights in the early summer .
oriental beetle grubs cause considerable damage to turf grasses and nursery plants . they eat and destroy the roots of the grass and are found in nursery stock , strawberry beds and some outside potted plants .
oriental beetle adults are about 7 / 16 inch . they vary in color from light brown to black , often with darker mottling on the wing covers .
rose chafer larvae have a brown head and conspicuous legs . fully grown , a rose chafer larva is about 3 / 4 - inch long . the larvae feed on the roots of grasses and non - crop plants . they do not cause damage to home lawns or landscape plants .
rose chafers are scarab beetles approximately 3 / 8 inch long , slender , and light tan in color . they contain a toxin that can be deadly to birds , including chickens , and small animals .
rose chafers are generally found in areas with sandy soil . adults feed on rose flowers and foliage , skeletonizing leaves .
asiatic garden beetle larvae has a brown head capsule and six legs . it has three instars .
asiatic garden beetles are less than one - half inch long . they are cinnamon in color and have an iridescent sheen in the sunlight . they are attracted to porch lights on summer nights and feed at night , chewing irregular holes in many different plants . during the day , they rest in the soil .
( japanese beetle larvae ) jim baker , north carolina state university , bugwood . org ;
clemson university - usda cooperative extension slide series , bugwood . org ; ( rose chafer adults on leaf ) university of wisconsin entomology ; ( asiatic garden beetle larvae ) msu ipm resources ; ( asiatic garden beetle adult )
it is the policy of the state of maine to minimize reliance on pesticides . the maine department of agriculture and the maine ipm council encourage everyone to practice integrated pest management and to use pesticides only as a last resort . the mention of pesticides in the fact sheets linked to these pages does not imply an endorsement of any product . be sure that any product used is currently registered and follow all label directions .
adult rose chafers feed primarily on flower blossoms , especially roses and peonies , causing large , irregular holes . they damage fruits , particularly grape , raspberry , and strawberry . they also feed on the foliage of many trees , shrubs and other plants , such as rose , grape , apple , cherry , and birch , typically damaging leaves by eating the leaf tissue between the large veins , a type of injury known as skeletonizing .
they contain a toxin that can be deadly to birds , including chickens , and small animals . the larvae feed on the roots of grasses and non - crop plants ; they do not cause damage to home lawns or landscape plants .
rose chafer larvae have a brown head and conspicuous legs . fully grown , a rose chafer larvae is about 3 / 4 - inch long .
rose chafers are scarab beetles approximately 3 / 8 inch long , slender , and light tan in color .
clemson university - usda cooperative extension slide series , bugwood . org ; ( rose chafer adults on leaf ) university of wisconsin entomology
h . e . hummel and t . a . miller ( eds . ) . techniques in pheromone research . springer - verlag , new york .
, jr . 1984 . wind tunnels in pheromone research , pp . 75 - 110 ,
smith and lawrence to stereoisomers of 8 - methyl - 2 - decyl propanoate under laboratory conditions .
( l . ) ( coleoptera : scarabaeidae ) to female volatiles in a flight tunnel .
1994 . sex pheromone responses of the oriental beetle ( coleoptera : scarabaeidae ) .
1990 . effect of light exposure and carbohydrate content of snap bean leaves on chinese rose beetle ( coleoptera : scarabaeidae ) feeding .
( coleoptera : scarabaeidae ) : response to synthetic sex attractant plus phenethy proprionate : eugenol .
1994 . the influence of climatic factors on the flight activity of the japanese beetle ( coleoptera : scarabaeidae ) : implications for use of a microbial control agent .
: a superior attractant , phenylethyl propionate + eugenol + geraniol , 3 : 7 : 3 .
1978 . sustained - flight tunnel for measuring insect responses to wind - borne sex pheromones .
1995 . bioassay approaches to assessing behavioral responses of plum curculio adults ( coleoptera : curculionidae ) to host fruit odor .
( coleoptera : chrysomelidae ) to corn silks and analysis of the host - finding response .
1977 . identification of the female japanese beetle sex pheromone : inhibition of male response by an enantiomer .
1999 . turfgrass insects of the united states and canada . comstock pub . associates , ithaca , new york .
1990 . tansley review no . 27 . the control of carbon partitioning in plants .
1989 . systat : the system for statistics . systat , evanston , illinois .
( f . ) , containing alpha - ionone . united states department of agriculture , patent 5 , 202 , 124 ; april 13 .
heath , j . j . , williams , r . n . & phelan , p . l . j chem ecol ( 2001 ) 27 : 419 . urltoken
michigan state university extension helps people improve their lives by bringing the vast knowledge resources of msu directly to individuals , communities and businesses .
rose chafers can quickly damage vines . to better manage this pest , know the biology and lifecycle , and available options for control .
the rose chafer is a light - tan beetle with a darker - brown head and long , spiny legs . it is about 0 . 5 inches long . the adult beetles have only one generation per season with emergence from the soil starting in late may and june , and with the beetles living for up to a month . while grapes are a preferred host , there are also many other plants it will feed on including roses , strawberries , peaches , cherries , apples , raspberries , blackberries , clovers , hollyhocks , corn , beans , beet , peppers , cabbage , peonies and many more plants , trees and shrubs .
this vineyard pest is distributed throughout the eastern united states with greatest abundance in areas with sandy soils and grass . this is because the female beetle , once mated , selects grassy and sandy areas for laying her eggs . those eggs hatch into larvae that feed on grass roots through the summer , moving down away from the frost line during the winter . in spring they feed again , pupate , and then emerge in late may and june . emergence of adult beetles typically coincides with bloom of grapevines . the beetle’s ability to skeletonize leaves until only the midribs are left , and consume the young tender clusters , makes it an economically - significant pest of grape production
as mentioned above , rose chafer beetles are attracted to sandy and grassy areas during their egglaying period . it may not be feasible , but if areas that are obvious sources of the beetle can be changed into a non - grass cover crop or can be fallow for the period of late may until early june , this might force the beetles to seek egglaying sites elsewhere . small numbers of beetles can also be hand - removed and placed into soapy water if you have a small enough vineyard that this manual control is possible .
for chemical control , there are a number of options to consider . assail , sevin and danitol are all ranked as providing excellent control of this pest . these have some different properties with the neonicotinoid insecticide assail providing protection due to it knocking down the beetles and also because it is a systemic insecticide that is taken into the vine making it resistant to wash - off and providing good residual activity .
in a 2003 msu research trial in leelanau county , we found that assail provided longer control than sevin . the carbamate insecticide sevin and the pyrethroid insecticide danitol both have contact activity against rose chafers , providing knockdown of the beetles on contact and with sevin also having some activity as a stomach poison after being eaten by the beetles .
additional broad spectrum insecticides such as the pyrethroids baythroid and mustang max are expected to also have good activity , as is the organophosphate imidan . under the high spring temperatures we have been having recently , the residual control of pyrethroids is expected to be shorter than under cool conditions . however , we also expect there to be a shorter period of rose chafer activity during hot conditions , allowing for a shorter period needed for vine protection against this pest .
the timing of rose chafer activity also can overlap with early season grape berry moth activity and early potato leafhopper infestation . the insecticides mentioned above will also provide some control of these other insect pests at the same time .
this article was published by michigan state university extension . for more information , visit urltoken . to have a digest of information delivered straight to your email inbox , visit urltoken / newsletters . to contact an expert in your area , visit urltoken , or call 888 - msue4mi ( 888 - 678 - 3464 ) .
june 7 , 2018 | john wise | precipitation can impact the performance of insecticides on fruit crops , but some compounds resist wash - off .
may 8 , 2018 | heidi lindberg | the new respirator requirements for the updated worker protection standard requires growers complete a respirator fit test annually .
may 8 , 2018 | heidi lindberg | as of january 2018 , three requirements were added to the worker protection standard .
may 1 , 2018 | rufus isaacs | with fruit crop bloom season kicking in , it’s a good time to review these recommendations .
april 30 , 2018 | john wise | there are many options available for mite control in fruit crops in this 2018 update .
physical removal and destruction of this insect pest can manage the population when only a few beetles are present ; however , when greater than two beetles per vine are seen during vineyard monitoring , a chemical spray is recommended . intensive baited trapping of the beetles over multiple years has shown to reduce populations to below threshold levels . an insecticide application in addition to trapping may be needed when populations are high . chemical applications should continue through the first or second post - bloom spray when pressure is severe . refer to a grape pest management guide ( see recommended links ) for insecticide specifics and efficacies . cultivation between rows to destroy the larvae may reduce the population but will not provide sufficient control of the pest . rose chafers contain a toxin that can be deadly to birds ( including chickens ) and small animals if the beetle is ingested .
the rose chafer is a light tan beetle with a dark brown head , about 12 mm ( ½ inch ) long and has one generation a year . larvae ( c - shaped grubs ) overwinter underground in the soil . they move closer to the surface and begin to feed on grass roots until they pupate in the spring . adults emerge during late may or june in most areas , around grape bloom , and congregate on plants to mate and feed . the females lay eggs just below the ground surface . they prefer grassy areas of sandy , well - drained soil . the adults live for three to four weeks after emergence . the eggs hatch into small , white grub - like larvae , which feed on roots of grass , weeds , grains , and other plants throughout the summer . they move down in the soil as the temperatures decline , where they will overwinter until the next growing season .
eviticulture and the grape community of practice are funded by the nifa - usda specialty crops research initiative ( scri ) and extension .
this work is supported by the usda national institute of food and agriculture , new technologies for ag extension project .
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rose chafers are commonly found in many areas , particularly those with sandy soil . they are pests on many different types of flowers , fruits , trees and shrubs but they are especially damaging to young , developing rosebuds , grapes and cherries .
an adult rose chafer is a moderate - sized beetle , 1 / 3 to 3 / 4 - inch ( 8 to 19 mm ) long , with wing covers having small yellowish hairs . it ' s slender and can be pale green to tan in color with reddish - brown or orangish spiny legs , and the body is black on the underside . larvae are small white grubs with a brown head that are found in the soil .
rose chafer larvae overwinter in the soil and pupate in the spring . adults emerge in late may to early june . because the rose chafer prefers sandy soil to lay eggs , plants located on sandy sites are most likely to be attacked . adult beetles feed on plants for three or four weeks , generally until late june . females lay eggs in groups of 6 to 25 in the soil among the weeds and grasses until early july , and then die shortly afterwards . eggs hatch in about two weeks , and larvae feed on the roots until fall , then burrow deeper into the soil to spend the winter in the soil below the frost line . they pupate the following spring and then emerge as adults . there is one generation a year .
rose chafer feed on the foliage of many trees , shrubs , also blackberry , grape , raspberry , strawberry , and tree fruit such as apple , and cherry . also garden vegetables , and ornamentals : birch , dahlia , hollyhock , iris , peony , poppy and rose .
adult rose chafers chew on flowers , especially roses and peonies , causing large , irregular holes by eating the leaf tissue between the large veins , a type of injury known as skeletonizing . they also chew on leaves , and fruits , but are usually damaging only when exceptionally numerous . damage is more severe in areas with sandy soil . larvae feed on roots of grass and weeds doing little noticeable damage .
prevention : gardeners should regularly monitor their gardens starting in late may , especially if you have a history of rose chafer infestations . cover smaller plants with floating row covers until july control : 1 . cultivation destroys eggs and pupae in the soil . it is especially effective against pupae if cultivating is continued into early june 2 . if large numbers of chafers are around , drench the soil with insect parasitic nematodes to kill the larvae . 2 . when small numbers are present , hand pick to remove rose chafers . put them into pails of soapy water to kill them . because rose chafers are good fliers , more can fly into your garden and you will need to check your plants routinely for any additional rose chafers .
if you love fruit tress like apples , peaches , pears and plums , but don ' t have the room , plant a dwarf variety . most grow from 3 feet to 8 feet . they product tons of fruit and are easier to harvest because they are low to the ground .
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and the life cycle is similar . rose chafer was first reported as a grape pest as early as 1810 , later extending its host range to include a wide assortment of host plants . grape remains among the most severely injured crops . larvae overwinter in soil , resuming development in the spring . adults emerge in late may or early june , near the time of grape bloom . these are tan , long - legged beetles about 12 mm long . mating and egg laying occur continuously for about two weeks with each female depositing 24 to 36 eggs . the average life span of the adult is about three weeks , when they feed on blossoms , newly set fruit and leaves . in about two weeks , eggs hatch and larvae begin feeding on grass roots . this pest is more common in areas with light sandy soils . there is one generation per year .
control is seldom needed for this pest , but vigilance should be maintained early in the season in case of high numbers . in severe cases , blossom buds are often completely destroyed , resulting in little or no grape production . population levels vary from year to year . petal - fall sprays for grape berry moth will also control rose chafers .
this is taken primarily from an extension bulletin by d . g . pfeiffer & p . b . schultz , entitled
major insect and mite pests of grape in virginia
( va . coop . ext . serv . 444 - 567 ( 1986 ) )
rose chafer adults attack grapes at bloom as they emerge from the soil . not only do they destroy the fruit at blossom , in addition , they frequently skeletonize the leaves , leaving only the large veins intact ( see figure 22 . 21 ) .
females deposit eggs below the soil surface and soon after hatch into tiny , white , grubs . the larvae feed on the roots of grasses and weeds throughout the summer becoming fully developed by fall .
scouting for rose chafer within the vineyard should begin when newly emerged adults may be found feeding primarily on newly formed blossom buds .
cultivating between rows may be effective in destroying a good number of chafers since the pupal stage is extremely vulnerable to disturbance .
assail ( acetamiprid ) , sevin ( carbaryl ) , imidan ( phosmet ) , and danitol ( fenpropathrin ) are all ranked as providing excellent control of this pest .
treatment with an insecticide should be when the first newly emerged beetles are detected in adequate numbers .
click on the following topics for more information on managing vineyard insect and mite pests .
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thrips ( several species including the flower thrips , western flower thrips ) ( univ . of california site )
root weevils ( otiorhynchus spp . , polydrusus spp . ) ( univ . of california site )
kovach , j . , w . wilcox , a . agnello an m . pritts . 1993 . strawberry ipm scouting procedures . cornell univ . n . y . state ipm program . publ . no . 203b . 33p .
a . refer to refer to handout on
insects and mites associated with small fruits in the u . s .
anon . 1972 . strawberry insects : how to control them . usda farm . bull . 2184 . 17 p .
butcher , m . r . , d . r . penman & r . r . scott . 1987a . a binomial sequential decision plan for control of two - spotted spider mite on strawberries in canterbury . n . z . j . exp . agric . 15 : 371 - 374 .
butcher , m . r . , d . r . penman & r . r . scott . 1987b . population dynamics of two - spotted spider mites in multiple year strawberry crops in canterbury . n . z . j . zool . 14 : 509 - 517 .
butcher , m . r . , d . r . penman & r . r . scott . 1987c . the relationship between two - spotted spider mite and strawberry yield in canterbury . n . z . j . exp . agric . 15 : 367 - 370 .
chagnon , m . , j . gingras & d . deoliveira . 1991 . honey bee ( hymenoptera : apidae ) foraging behavior and raspberry pollination . j . econ . entomol . 84 : 457 - 460 .
cooley , d . r . & s . g . schloemann . 1990 . development and implementation of a strawberry ipm program . 1990 reg . ext . fruit school . middleway wv . oct . 30 - nov . 1 .
davidson , r . h . & w . f . lyon . 1979 . insect pests of farm , garden , and orchard . wiley , n . y . 596 p .
english - loeb , g . , m . pritts , j . kovach , r . rieckenberg and m . j . kelly . 1999 . compensatory ability of strawberries to bud and flower removal : implications for managing the strawberry bud weevil . j . econ . entomol . 92 : 915 - 921 .
hansen , r . w . & e . a . osgood . 1983 . insects visiting flowers of wild red raspberry in spruce - fir forested areas of eastern maine . entomol . news 94 : 147 - 151 .
johnson , d . t . & r . l mayes . 1986 . biology and control of the rednecked cane borer . arkansas farm res . 35 ( 3 ) : 6 .
mailloux , g . & n . j . bostanian . 1988 . economic injury level model for tarnished plant bug , lygus lineolaris ( palisot de beauvois ) ( hemiptera : mirideae ) , in strawberry fields . environ . entomol . 17 : 581 - 586 .
mcgregor , s . e . 1976 . insect pollination of cultivated crop plants . usda - ars agric . handbk . 496 . 411 p . miller , j . c . , p . e . hanson & r . v . dowell . 1987 . the potential of gypsy moth as a pest of fruit and nut crops . calif . agric . nov . - dec . 87 : 10 - 12 .
pennsylvania . 1991 . small fruit and pest management guide 1991 - 1992 . penn . state coll . agric .
price , j . f . & j . b . kring . 1991 . response of twospotted spider mite , tetranychus urticae koch , and fruit yield to new miticides and their use patterns in strawberries . j . agric . entomol . 8 : 83 - 91 .
raine , j . 1962 . life history and behaviour of the raspberry crown borer bembecia marginata ( harr . ) ( lepidoptera : aegeriidae ) . can . entomol . 94 : 1212 - 1222 .
schaefers , g . a . 1964 . control of the strawberry leaf roller , ancylis comptana fragariae ( lepidoptera : tortricidae ) . j . econ . entomol . 57 : 983 - 986 .
schaefers , g . a . 1981 . pest management systems for strawberry insects . p . 377 - 393 . in : d . pimentel ( ed . ) . crc handbook of pest management in agriculutre . vol . iii . crc press , boca raton fl . 656 p .
schloemann , s . g . & d . r . cooley . 1988 . the initiation of a new ipm program in strawberries in massachusetts . p . 67 - 81 . in : proc . new engladnd small fruit school . jan . 5 , 1988 .
stuart , l . c . , b . a . butt & f . takeda . 1986 . leaf curl of eastern thornless blackberry caused by blackberry psyllid , trioza tripunctata fitch . proc . 62nd cumberland - shenandoah fruit workers conf . , harpers ferry wv . nov . 20 - 21 .
waite , g . k . 1988 . integrated control of tetranychus urticae in strawberries in south - east queensland . exp . appl . acarol . 5 : 23 - 32 .
williams , r . n . 1977 . sap beetles on raspberries and strawberries . pest management tips . fruit insect pests no . 2 . 2 p .
williams , r . n . , m . j . weiss , k . v . miller & j . j . werner . 1984 . a summary of experiments for control of sap beetles which attack fruit crops . oardc res . circ . 283 : 66 - 68 .
zajac , m . a . & m . c . wilson . 1984 . the effects of nymphal feeding by the meadow spittlebug , philaenus spumarius ( l . ) on strawberry yield and quality . crop protect . 3 : 167 - 175 .
by : stan v . griep , american rose society consulting master rosarian – rocky mountain district
the rose chafer and the japanese beetle are both true villains of the rose bed . both appear to have the same habits and life cycles , going from eggs laid in the ground by the mature female beetles , hatching out to larvae / grubs in the ground and maturing to beetles that attack plants and blooms without mercy . read on for more rose chafer facts and control info .
the adult rose chafer’s main diet is flower blossoms , especially those of peonies and roses . the damage they do to the blooms can be devastating . rose chafer damage can be recognized by the large irregularly shaped holes all over the flowers , ruining the beauty of the blooms completely .
these bad guy beetles also include some fruits in their diet , seeming to prefer the raspberry , grape and strawberry . they will also feed upon the foliage of many trees and shrubs , such as apple trees , cherry trees and birch trees . this rose chafer damage is created by eating the leaf tissue between the large veins and results in what is known as “skeletonizing” of the leaves .
treating rose chafers is important , not only for the health of your rose and other susceptible ornamentals , but for wildlife too . the rose chafer contains a toxin within its body chemistry that can be deadly to birds , including chickens . the same toxin can be deadly to other small animals when they eat these beetles .
as a part of keeping an eye on things in our gardens and rose beds , we need to keep an eye out for the rose chafers starting around late may ( early spring ) , especially if there is a history of having rose chafer problems in the area or in our own gardens and rose beds . many gardeners feel there is little difference between the rose chafer and the japanese beetle , as attempting to protect our plants and rose bushes from them is a daunting task , especially when there are large numbers of them !


this group generally corresponds with the order blechnales described by j. l. reveal in 1993 .

however , the blechnales families were found to be embedded within the larger definition of the order polypodiales describe by smith et al. , 2006 , so according to their system , the group is treated as an informal clade .

this clade includes some important ferns , including the sensitive fern , onoclea sensibilis , which grows as a virtual weed throughout much of its temperate north american range , and ferns of the genus thelypteris , a genus that has shown remarkable speciation .

it also includes one of the more common horticultural ferns , the ostrich fern , matteuccia struthiopteris .

eupolypods ii

the following diagram shows a likely phylogenic relationship between the families of eupolypods ii , based on lehtonen , 2011 , and rothfels & al . , 2012 .
plastid phylogenomics resolve deep relationships among eupolypod ii ferns with rapid radiation and rate heterogeneity .
challenges inherent in resolving the eupolypod ii phylogeny . eupolypods ii phylogram modified from schuettpelz and pryer ( 2007 ) , in ( a ) unrooted and ( b ) rooted form . ( i ) outgroup taxa are on long branches . ( ii ) backbone internodes are very short , suggesting an “ancient rapid radiation . ” ( iii ) the ingroup is marked by significant heterogeneity in rates of evolution , with the members of aspleniaceae on much longer branches than other eupolypod ii taxa .
wherever possible , we included generic and familial types , to facilitate future taxonomic revisions . based on data from schuettpelz and pryer ( 2007 ) and liu et al . ( 2007 ) , our broad outgroup sample included 10 representatives from the sister group to the eupolypods ii ( eupolypods i , see fig . 2 ) . to better evaluate the effect of uncertainty in outgroup placement on the ingroup topology and to better understand the divergence between eupolypods i and ii , we also included two representatives from each of the two potentially successive sister groups to the eupolypods ( notholaena and cryptogramma from pteridaceae ; dennstaedtia and pteridium from dennstaedtiaceae ; see fig . 2 and schuettpelz and pryer 2007 ) . our total sample has 81 terminal taxa ( appendix 1 ) .
plastid phylogenomics resolve deep relationships among eupolypod ii ferns with rapid radiation and rate heterogeneity . - pubmed - ncbi
eupolypods were separated into two clades , corresponding with eupolypods i and ii [ 23 ] . diplaziopsidaceae was resolved as sister to eupolypods ii . christenhusz et al . [ 8 ] included diplaziopsis , homalosorus and hemidictyum within the diplaziopsidaceae , but here hemidictyum was supported as sister to aspleniaceae as in schuettpelz & pryer [ 18 ] and in kuo et al . [ 20 ] . hemidictyum is therefore considered here as a member of aspleniaceae and only diplaziopsis and homalosorus are within diplaziopsidaceae , as in a recent analysis of the matk gene [ 20 ] . family - level relationships mostly remained poorly supported or unresolved within both of the two large eupolypod clades .
eupolypods i is a clade of ferns in the order polypodiales , class polypodiopsida . [ 2 ] in the classification of christenhusz & chase ( 2014 ) this is equivalent to the family polypodiaceae . [ 1 ] it probably diverged from eupolypods ii ( = aspleniaceae of christenhusz & chase 2014 [ 1 ] ) during the mid - cretaceous . the divergence is supported by both molecular data and an often overlooked morphological characteristic which lies in the vasculature of the petiole . most species that make up the eupolypods i clade have three vascular bundles . the only exceptions are the grammitid ferns which have one , and the genus hypodematium which has two . this differs from eupolypods ii which mostly have two vascular bundles ( except the well - nested blechnoid ferns which generally have at least three ) . [ 3 ]
the following phylogram for the eupolypods i is based on lehtonen , 2011 [ 4 ] and christenhusz et al . , 2011 . [ 5 ]
the eupolypods ii , together with its sister group , eupolypods i , comprise the large eupolypod clade , which encompasses two - thirds of living fern species ( fig . 2 ; pryer et al . 2004 ; schneider et al . 2004b ; smith et al . 2006 ) . the ancestors of eupolypods i and ii diverged from each other in the early cretaceous ( pryer et al . 2004 ; schneider et al . 2004b ; schuettpelz and pryer 2009 ) . the eupolypod ii clade started to diversify shortly thereafter ( its crown group is approximately 100 million years old ; schuettpelz and pryer 2009 ) and has subsequently grown into a lineage - rich clade comprising nearly 30 % of extant fern diversity . eupolypods ii includes some of the most familiar groups of ferns ( the lady ferns , ostrich fern , sensitive fern , marsh ferns , and spleenworts ) , as well as some of the most species - rich genera : thelypteris s . lat . ( ∼950 species ) ; asplenium ( ∼700 species ) ; diplazium ( ∼350 species ) ; athyrium ( ∼220 species ) ; and blechnum s . lat . ( ∼150 species ) .
not surprisingly , given the clade ' s size and age , eupolypod ii taxa are morphologically disparate and seemingly incohesive . however , early workers did tend to recognize the close affinities among many of the taxa in this clade , although frequently with members of eupolypods i interdigitated among them ( holttum 1947 ; sledge 1973 ; mickel 1974 ; tryon and tryon 1982 ) . the cohesiveness of the eupolypods ii started to become apparent with the earliest applications of molecular phylogenetic techniques to ferns ( wolf et al . 1994 ; hasebe et al . 1995 ; sano et al . 2000a ) and has been strongly supported in recent broad studies ( schneider et al . 2004b ; schuettpelz and pryer 2007 ; kuo et al . 2011 ) . none of these studies , however , found support for the backbone relationships within eupolypods ii , and only kuo et al . ( 2011 ) attempted to sample its major lineages . it remains one of the few areas of the fern tree - of - life where the backbone relationships remain elusive ( smith et al . 2006 ; schuettpelz and pryer 2007 ) .
relative plastid substitution rates among clades / branches of the eupolypod ii ferns , based on the “83 - gene matrix , ” resulting from the random local clock analyses in beast v1 . 8 . 1 . numbers above the branches indicate relative , median rates with no measured units , scaled by dividing all rates by that of the crown node of eupolypods ii . colors of branches represents to relative rate . light red indicates a relative rate of 0 , whereas a trend of blue color marks a relative rate approaching 3 . 0 .
eupolypods ii is a clade of ferns in the order polypodiales , under the class polypodiopsida . this group generally corresponds with the order blechnales described by j . l . reveal in 1993 . however , the blechnales families were found to be embedded within the larger definition of the order polypodiales describe by smith et al . , 2006 , so according to their system , the group is treated as an informal clade .
hypotheses on the backbone relationship of eupolypod ii ferns with special reference to the positions of rhachidosoraceae and athyriaceae . ( a ) topology resolving rhachidosoraceae as sister to the thelypteridaceae - blechnaceae clade in ; ( b ) topology resolving rhachidosoraceae as sister to the aspleniaceae - diplaziopsidaceae clade in ) ; ( c ) topology showing athyriaceae as nonmonophyletic in , ) and ; and ( d ) topology showing unresolved relationships within eupolypod ii ferns in based on 25 low - copy nuclear genes .
shaw j , lickey eb , beck jt , farmer sb , liu w , et al . ( 2005 ) the tortoise and the hare ii : relative utility of 21 noncoding chloroplast dna sequences for phylogenetic analysis . amer j bot 92 : 142–166 .
comparison of variability in nonoverlapping 1 , 000 - bp sliding windows for eupolypod ii ferns . genes with interests are indicated , that is , genes with pi > 0 . 1 are marked in black and genes with pi < 0 . 1 are marked in red .
the ml phylogram of 40 species based on 83 - gene matrix and codon partitioned strategy and bayesian divergence time estimation based on 83 - gene matrix with unpartitioned strategy . ( a ) maximum likelihood bootstrap values ( bss ) are 100 % and bayesian posterior probabilities ( pps ) are 1 . 0 , unless otherwise indicated . numbers above the branches indicate bss and pps based on whole plastome matrix , and bss ( unpartitioned , parititionfinder , gene partitioned ) and pps based on 88 - gene matrix , while numbers below the branches as bss ( unpartitioned , codon partitioned , parititionfinder , gene partitioned ) and pps based on 83 - gene matrix ; ( b ) chronogram with secondary calibration nodes indicated by numbers ; ( c ) bar chart indicating stem clade ages and hpd intervals of each family of eupolypods ii . blue bars indicate 95 % highest posterior density ( hpd ) intervals of the age estimates ; grey bars indicate the time - scale of eupolypod ii radiation .
carl j . rothfels , anders larsson , li - yaung kuo , petra korall , wen - liang chiou , kathleen m . pryer ( 2012 ) .
overcoming deep roots , fast rates , and short internodes to resolve the ancient rapid radiation of eupolypod ii ferns
. systematic biology 61 ( 1 ) : 70 .
in pteridaceae , all subfamilies accepted by christenhusz et al . [ 8 ] were found to be monophyletic , although the monophyly of cheilanthoidea had poor support . by contrast , numerous pteridoid genera , including adiantum , were not monophyletic . the need for a generic redefinition within pteridoids has already been well recognized by earlier studies [ 8 ] , [ 18 ] , [ 23 ] – [ 27 ] . the relationship between pteridoids and dennstaedtioids was still ambiguous to date [ 23 ] , and the present study also did not provide conclusive results . pteridaceae was positioned as a sister group to eupolypods by parsimony , whereas ml supported dennstaedtiaceae as sister to eupolypods . both hypotheses received less than 50 % support . the genus dennstaedtia was paraphyletic as in previous studies [ 18 ] , [ 28 ] , due to the inclusion of the monophyletic microlepia .
carl j . rothfels , anders larsson , li - yaung kuo , petra korall , wen - liang chiou , kathleen m . pryer ; overcoming deep roots , fast rates , and short internodes to resolve the ancient rapid radiation of eupolypod ii ferns , systematic biology , volume 61 , issue 3 , 1 may 2012 , pages 490 , urltoken
the trees obtained here were generally consistent with the prevailing view of the molecular phylogeny of ferns [ 1 ] , [ 18 ] – [ 20 ] . the taxonomic sampling employed here was almost seven - times broader than in the previous best - sampled fern phylogenetic analysis , hence providing a broader picture of fern phylogenetics , and enabling the investigation of the monophyly of currently accepted genera and families . however , despite the broad sampling , numerous fern groups remained poorly sampled and some phylogenetic relationships could not be completely resolved . for example , the families belonging to the eupolypods ii group are well supported as monophyletic entities , but the relationships between them remained poorly established . the relationships among some of the early diverging polypods ( saccolomataceae , cystodiaceae ) were not unambiguously resolved and questions about a pteridoid - dennstaedtioid relationship still remained unanswered . similarly to previous studies [ 1 ] , [ 20 ] , [ 27 ] – [ 30 ] , the phylogenetic position of horsetails ( equisetaceae ) remained controversial .
arthropteris and psammiosorus were mixed , but together formed a well - supported sister lineage to all other tectariaceae . the proposed subfamilies of polypodiaceae [ 8 ] were monophyletic in the ml analysis , except that synammia was resolved as sister to drynarioideae rather than being a member of polypodioideae . the parsimony analysis did not support monophyletic polypodiaceae , resulting in a largely unresolved topology within the eupolypods i . the current generic classification failed to delimit natural groups within drynarioideae , microsoroideae and polypodioideae . the subfamily loxogrammoideae was resolved as sister group to the remaining polypodiaceae in the ml analysis .
as stated above , earlier studies ( e . g . , schuettpelz and pryer 2007 ) indicate that the eupolypod ii phylogeny is likely to include several key challenges for phylogenetic inference , specifically a series of long branches among very short backbone internodes ( an ancient rapid radiation ) , lineage - specific rate heterogeneity , and a distantly related outgroup . given these concerns , we sought to explicitly evaluate our topology and support values against these potential artifacts , with particular emphasis on the support values along the backbone of the tree .
the eupolypod ii clade is cosmopolitan in distribution , with the subgroups primarily temperate to tropical , and the larger subclades each well represented in both areas . however , many of the phylogenetically enigmatic taxa in this clade are limited to the himalayas or southeast asia , and critical members of several genera are rare and / or infrequently collected . this pattern of rarity and endemism , in conjunction with the richness and geographical breadth of the clade as a whole , is undoubtedly a contributing factor to the incomplete sampling of these ferns in previous phylogenetic studies .
this trend of shared morphological syndromes across very deep splits in the tree by some members of the “woodsiaceae” is in contrast to the interdigitation , among those same taxa , of a series of distinct morphologically unique groups , including the aspleniaceae , blechnaceae , onocleaceae , and thelypteridaceae . the coarse picture of eupolypod ii morphological evolution , then , is marked by two seemingly opposing patterns . on the one hand are the autapomorphy - rich clades , whose individual phylogenetic coherence is strong , but whose deep relationships were obscure based on morphological data . and , on the other , the morphologically consistent yet phylogenetically incoherent members of the “woodsiaceae” .
dna was extracted from silica - dried or herbarium material , using either ( i ) a modified carlson - yoon protocol ( < 0 . 01 g dried plant material , silica beads , 750 μ l carlson buffer , and 20 μ l mercaptoethanol added to a 2 - ml tube and ground for 45 s using a mini - beadbeater ( biospec products ) , followed by incubation at 65 о c for 45 min ; yoon et al . 1991 ) or ( ii ) the protocol of pryer et al . ( 2004 ) or ( iii ) the protocol of kuo et al . ( 2011 ) . for material extracted under the carlson - yoon protocol , the extracted dna was purified by illustra gfx pcr dna and gel band purification kit ( ge healthcare ) .
to evaluate any effects that uncertainty in root - branch placement might have on apparent levels of support within the ingroup , we compared ingroup backbone mlbs values from the analysis of our complete data ( full outgroup ) with those from each of six different variations in outgroup composition : ( i ) ingroup only ; ( ii ) ingroup + dryopteris ; ( iii ) ingroup + dryopteris and didymochlaena ; ( iv ) ingroup + dryopteris and notholaena ; ( v ) ingroup + dryopteris and notholaena and pteridium ; and ( vi ) ingroup + our full eupolypod i sample ( fig . 5 ) . this outgroup sampling regime was selected to successively bisect the longest outgroup branches , with a particular emphasis on breaking the proximate root branch ( the branch connecting the ingroup to the first outgroup node ) .
our approach to resolving the eupolypod ii phylogeny couples a considerably expanded taxon sample with moderate character sampling . our objectives include identifying well - supported major ( approximately “family - level” ) clades and determining the backbone relationships among these clades . given the anticipated phylogenetic challenges and potential for artifacts in our data , we explicitly evaluate our phylogenetic hypothesis against these analytical pitfalls , placing strong emphasis on the use of the reduced consensus technique ( wilkinson 1996 ) to isolate the effects of signal weakness from those of signal conflict ( e . g . , wiens 2003 ; cobbett et al . 2007 ) . our study aims for a comprehensive and well - supported phylogeny of this important group of ferns , and for novel inferences about the behavior of our choice of methods , gleaned from their performance on this data set .
discover a faster , simpler path to publishing in a high - quality journal . plos one promises fair , rigorous peer review , broad scope , and wide readership – a perfect fit for your research every time .
in the past two decades , molecular systematic studies have revolutionized our understanding of the evolutionary history of ferns . the availability of large molecular data sets together with efficient computer algorithms , now enables us to reconstruct evolutionary histories with previously unseen completeness . here , the most comprehensive fern phylogeny to date , representing over one - fifth of the extant global fern diversity , is inferred based on four plastid genes . parsimony and maximum - likelihood analyses provided a mostly congruent results and in general supported the prevailing view on the higher - level fern systematics . at a deep phylogenetic level , the position of horsetails depended on the optimality criteria chosen , with horsetails positioned as the sister group either of marattiopsida - polypodiopsida clade or of the polypodiopsida . the analyses demonstrate the power of using a ‘supermatrix’ approach to resolve large - scale phylogenies and reveal questionable taxonomies . these results provide a valuable background for future research on fern systematics , ecology , biogeography and other evolutionary studies .
citation : lehtonen s ( 2011 ) towards resolving the complete fern tree of life . plos one 6 ( 10 ) : e24851 . urltoken
copyright : © 2011 samuli lehtonen . this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
funding : this study received funding from the kone foundation and the academy of finland grants to sl . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
the combined four - gene ( atpa , atpb , rbcl , rps4 ) data set included a total of 5 , 166 sequences ( dataset s1 ) , hence the matrix of all 2 , 957 taxa by four genes had 6662 missing gene sequence entries ( c . 56 % missing data ) . most taxa ( 91 % ) were represented by the rbcl gene , but the least sampled gene ( atpa ) was available for only approximately 18 % of the taxa . less than 10 % of the sampled taxa were represented by all four genes , and 54 % were represented only by a single gene . in most of the fern families at least some taxa were sampled for all markers , with two small families ( diplaziopsidaceae and rhachidosoraceae ) represented by the rbcl gene only and four other families ( psilotaceae , schizeaceae , cystodiaceae and lomariopsidaceae ) lacking one of the studied genes . the parsimony analysis of these data retained 124 equally parsimonious trees of 74 , 910 steps ( dataset s2 ) . the final ml optimization likelihood score was −391724 . 512141 ( figure 1 , dataset s3 ) .
parsimony and ml trees were largely consistent with each other and with the prevailing view of the fern familial relationships [ 1 ] , [ 18 ] – [ 20 ] . the parsimony analysis positioned horsetails ( equisetopsida ) as a sister group to the marattiopsida - polypodiopsida clade , whereas ml placed them as sister to polypodiopsida . these controversial groupings received low support values . within the tree fern clade , ml and parsimony largely disagreed at the family level . metaxyaceae was positioned as a sister to other tree ferns in the parsimony analysis , whereas in the ml tree the family was placed as sister to dicksoniaceae . the clade composed of thyrsopteridaceae and associated families in the ml tree also included cibotiaceae and dicksoniaceae in the parsimony analysis . similarly , the two methods disagreed in the exact phylogenetic position of dennstaedtiaceae and many small families , including saccolomataceae , cystodiaceae , hypodematiaceae , cystopteridaceae and woodsiaceae . however , most of the incongruent groupings received less than 50 % bootstrap support in both analyses , consistently with the observation that their relationships were also uncertain in previous studies [ 18 ] , [ 23 ] .
a recently published linear fern classification [ 8 ] was largely supported at the family level . at the generic level , however , improved sampling revealed several patterns that were inconsistent with previously published results and current fern taxonomy . some of the most relevant results are shortly described here , otherwise readers are directed to trees available as supplementary information ( dataset s2 , s3 ) and at treebase ( urltoken ) . in ophioglossaceae , the results contradicted those published by hauk et al . [ 21 ] notably regarding the position of cheiroglossa , which is here nested within ophioglossum . in addition , o . lusitanicum l . was here grouped together with helmintostachys zeylanica ( l . ) hook . in the present study , the genus odontosoria ( lindsaeaceae ) was polyphyletic , and sphenomeris was grouped with the tapeinidium - osmolindsaea - nesolindsaea clade , thus contradicting the results of a recent study on lindsaeaceae phylogenetics [ 22 ] .
aspleniaceae was divided into three well - supported lineages , corresponding to hemidictyum and two broadly - defined genera : hymenasplenium and asplenium [ 8 ] , [ 18 ] . several well - supported clades were also present within thelypteridaceae , although not exactly matching the current generic classification . similarly , previous studies have suggested that the current classification of blechnaceae is unnatural [ 8 ] . in this study , the family was divided into three well - supported clades that did not correspond to the currently accepted generic limits ( woodwardia ; salpichlaena - stenochlaena - blechnum p . p . ; blechnum p . p . - brainea - sadleria - pteridoblechnum ) . within athyriaceae , diplazium was strongly supported as monophyletic , but cornopteris was nested within athyrium with a high level of support .
the two subfamilies of dryopteridaceae [ 8 ] were monophyletic ( with the exception of dryopteris inaequalis ( schlecht . ) kuntze , which was placed in elaphoglossoideae ) in the ml analysis , albeit with a very poor support . in the parsimony analysis , on the other hand , the subfamily elaphoglossoideae was divided into two groups with unresolved relationships with dryopteridoideae . the subfamily elaphoglossoideae included pleocnemia winitii holttum , the only member of its genus included in this study . previous studies have considered pleocnemia as a member of tectariaceae [ 8 ] , [ 23 ] , [ 30 ] . at a generic level , polystichum included cyrtomium and cyrtogonellum , arachnioides included leptorumohra and lithostegia , and acrorumohra was nested in dryopteris ( excluding d . inaequalis ) .
the observed uncertainty might , to some extent , reflect the large number of missing entries in the supermatrix . more than half of the taxa were represented by a single gene , rbcl being clearly the best - sampled marker . those markers not as thoroughly sampled were , however , sampled rather evenly across the different fern lineages , so that very few families completely lacked data of one or more genes . furthermore , smith et al . [ 16 ] were able to resolve several difficult problems in the phylogeny of green plants by sampling the rbcl gene only , and it has been shown that supermatrix approach can handle even 90 % missing data without loss of accuracy if the data available contain enough informative characters [ 11 ] , [ 17 ] , [ 32 ] – [ 35 ] .
the advances in fern systematics over the past decades have provided a rather good taxonomic understanding at the family level , and the recently proposed fern classification [ 8 ] was largely supported by the current study . generic delimitation , however , has remained ambiguous in a number of fern families [ 8 ] , [ 23 ] . the analyses presented here shed new light on several unresolved issues , and can be used as a starting point to a more robust classification at this taxonomic level . a good example was that of blechnaceae , a family composed of three well supported clades that ( apart from woodwardia ) do not correspond well with the currently accepted generic classification .
until recently , most of the molecular systematic studies of ferns were based on classical fern taxonomy . the most convenient way of overcoming the impact of outdated taxonomies , as well as detecting contaminated or misidentified sequences [ 11 ] , [ 40 ] , [ 41 ] , is through the use of supermatrix analysis of all available data . the results presented here corroborated most recent findings in molecular fern systematics , but also provided a much wider view for future studies in fern evolution , taxonomy , and beyond . instead of relying on the classical fern taxonomy , pteridologists can now select proper outgroups and delimit their ingroups in an appropriate way from an evolutionarily perspective . as yet , only about one - fifth of the extant fern diversity is currently covered by genbank , but the road is open for a fully sampled fern tree of life , and ultimately , for a natural fern classification .
sequence data was retrieved from genbank release 176 ( feb . 23 , 2010 ) using phylota browser ( urltoken ) . phylota assembles blast clustering for all sequences in the genbank release file [ 42 ] . clusters corresponding to four protein coding plastid genes , rbcl , rps4 , atpa , and atpb , were downloaded for root node “moniliformopses” . this data set was further supplemented by downloading rbcl data of japanese ferns [ 43 ] and adding several fern sequences produced with standard methods and primers [ 3 ] , [ 19 ] , [ 44 ] – [ 49 ] in our laboratory and submitted to genbank , but not yet available on the queried release ( genbank accession numbers hq157300–hq157307 , hq157324–hq157330 , hq157332–hq157334 , hq245099–hq245103 , hq680978 ) . when multiple sequences were available for one taxon , the most complete one was retained and the other sequences excluded . a few sequences in the preliminary test analyses were positioned into highly questionable taxonomic groups , and these apparently misidentified or contaminated sequences were also excluded from the final analyses . the finally accepted fern sequences ( 2 , 656 taxa ) were further supplemented with 301 outgroup taxa representing lycophytes ( 205 taxa ) , angiosperms ( 61 taxa ) and gymnosperms ( 35 taxa ) .
multiple sequence alignments were produced for each data set with muscle [ 50 ] using default settings followed by one round of refinement . due to variable sequence completeness all the alignments had high amounts of missing data at the 5′ and 3′ ends . these ambiguous regions were eliminated from the final data sets after visual inspection , as well as ambiguously aligned segment within the rps4 gene . however , possible errors in the sequences ( such as stop - codons ) were not investigated . indels inserted during the sequence alignment were treated as missing data in the corresponding phylogenetic analyses . because all the markers included were plastid genes they were expected to share a common evolutionary history and were analyzed simultaneously . aligned sequence matrices were concatenated with sequencematrix software [ 51 ] . in total , the data set consisted of 2 , 957 taxa ( rbcl 2 , 681 ; rps4 1 , 134 ; atpb 825 ; atpa 526 taxa ) and 4 , 406 aligned base pairs of molecular data ( rbcl 1 , 332 ; rps4 379 ; atpb 1 , 188 ; atpa 1 , 507 bp ) . the aligned data matrices and resulting trees are available at treebase ( urltoken , [ 52 ] ) .
phylogenetic analyses were performed for the concatenated supermatrix under equally weighted parsimony criteria using tnt [ 53 ] and maximum likelihood criteria using raxml [ 54 ] . in the parsimony analyses 500 ‘new technology’ [ 55 ] , [ 56 ] search replications were used as a starting point for each hit . these replications saved no more than 10 trees per replication , and were run until the best score was hit 10 times , using tbr - swapping , random and constraint sectorial searches , five ratchet iterations , and five rounds of tree fusing ( xmult = repl 500 hits 10 css rss ratchet 5 fuse 5 hold 10 ) . the memory was set to hold 80 , 000 trees . branch support was evaluated by running 500 bootstrap replicates . tbr - swapping , sectorial search , and five rounds of tree fusing were employed in each replicate ( resample = boot replications 500 savetrees [ xmult = rss css fuse 5 ] ) . maximum likelihood ( ml ) analyses were performed using the parallel pthreads - version of the computer program raxml 7 . 2 . 8 [ 54 ] , [ 57 ] running in 2×2 . 26 ghz quad - core intel xeon macintosh with 8 gb of ram . the search was initiated with 500 rapid bootstrap replications followed by a thorough ml search on the original alignment ( - t 16 - f a - x 12345 - p 12345 - # 500 - m gtrgamma ) . free model parameters were estimated by raxml under the gtr + γ model . this is the most commonly used model for real data sets , and provides good performance for large data sets [ 58 ] .
congruence among the data sets was examined by running parsimony bootstrap analyses for each gene separately [ 59 ] . visual inspection of the family - level nodes did not reveal well - supported ( > 70 % support ) conflict at this phylogenetic level , with the exception of nested position of lonchitidaceae within lindsaeaceae in the atpa analysis ( data not shown ) . at lower phylogenetic levels the highly variable taxon sampling made the assessment of phylogenetic conflict highly problematic , and simultaneous analysis of all data sets was considered appropriate based on family - level congruence .
concatenated supermatrix in nexus - format ( file can be opened after unzipping for example with mesquite [ 60 ] ) .
the strict consensus tree of parsimony analysis with bootstrap support values in nexus - format ( file can be opened for example with figtree [ 61 ] ) .
the ml tree with bootstrap support values in nexus - format ( file can be opened for example with figtree [ 61 ] ) .
i thank all the researchers who have produced and shared dna sequences via genbank . the willi hennig society is acknowledged for making tnt publicly available . two anonymous reviewers are acknowledged for their constructive comments on earlier draft version of this manuscript .
conceived and designed the experiments : sl . performed the experiments : sl . analyzed the data : sl . contributed reagents / materials / analysis tools : sl . wrote the paper : sl .
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institute of ecology and evolutionary biology , national taiwan university , no . 1 , section 4 , roosevelt road , taipei 10617 , taiwan
broad phylogeny of ferns . approximate number of species per clade is given in parentheses . modified from smith et al . ( 2006 ) .
five plastid loci were selected for analysis : atpa , atpb , matk , rbcl , and the trng - trnr intergenic spacer ( henceforth “ trng - r ” ) . all loci , except for matk , were amplified according to either the “standard” or “difficult” reaction protocols ( below ) depending on the source of the material ( standard for most extractions ; difficult for those from herbarium specimens greater than 10 years old ) , using the primers listed in table 1 . the standard amplification reaction used standard taq polymerase with the following cycle : a 3 min initial denaturation at 95 о c , followed by 35 cycles of 30 s denaturation at 95 о c , 1 min annealing at 54 о c , and 2 min elongation at 72 о c , followed by a final elongation of 10 min at 72 о c . the difficult amplification reaction , using phusion high fidelity dna polymerase ( finnzymes ) , was 1 min initial denaturation at 98 о c , followed by 35 cycles of 10 s denaturation at 98 о c , 30 s annealing at 58 о c , and 1 min elongation at 72 о c , followed by a final elongation of 8 min at 72 о c . amplification of all matk sequences followed the protocol of kuo et al . ( 2011 ) .
notes : f = forward ; r = reverse ; a = used in amplifications ; s = used in sequencing reactions . while most primers were applied across the phylogeny , superscripts following primer names indicate lineage specificity : a aspleniaceae ; b athyriids ; c blechnaceae ; d cystopteris / gymnocarpium ; e depariids ; f diplaziids ; g diplaziopsis / homalosorus ; h rhachidosorus ; i onocleaceae ; j pteridaceae ; k thelypteridaceae ; l woodsia and allies .
pcr products from carlson - yoon extractions were purified using multiscreen plates in a vacuum manifold ( millipore ) and sequenced by macrogen inc . ( south korea ) . for material extracted under the protocol of pryer et al . ( 2004 ) , each pcr product was cleaned using 0 . 5 μ l of exonuclease i and 1 μ l of shrimp alkaline phosphatase ( usb , cleveland , oh ) ; reaction tubes were incubated at 37 о c for 15 min and then heated to 80 о c for 15 min to inactivate the enzymes , prior to sequencing on a applied biosystems 3730 xl at the duke igsp sequencing facility ( duke university ) . material extracted under the protocol of kuo et al . ( 2011 ) was sequenced at genomics ( taipei , taiwan ) . we completed our sampling with an additional 100 previously published sequences from genbank ( appendix 1 ) .
sequences were manually aligned in mesquite 2 . 72 ( maddison and maddison 2009 ) . indels ( limited to matk , trng - r , and the ends of the atpa alignment ) were assessed by eye , and ambiguously aligned areas were excluded prior to phylogenetic analysis . any gaps associated with unambiguous indel regions were treated as missing data . in one rapidly evolving region of the trng - r alignment , we were unable to confidently align the pteridaceae sequences to those of the other taxa . in order to retain this otherwise unambiguous region , we excised those portions of the pteridaceae sequences , replacing them with question marks .
notes : missing data include both uncertain bases ( e . g . , ? , r , y ) and gaps ( - ) . support values are listed as mlbs or bayesian pps .
to obtain a point estimate of the phylogeny , we performed 10 tree bisection - and - reconnection heuristic searches of the concatenated ( unpartitioned ) data , each from a different random - addition - sequence starting tree , under ml using a gtr + i + g model as implemented in paup * 4 . 0b10 for unix ( swofford 2002 ) . the values for the exchangeability parameters , base frequencies , gamma shape parameter , and proportion of invariant sites were fixed at their ml values as optimized under a garli 0 . 951 ( zwickl 2006 ) tree search , using default genetic algorithm and termination settings .
our approach to phylogeny evaluation involved permutations of both the models and the implementation of those models ( i . e . , programs ) . the models were deliberately selected according to their varying degrees of susceptibility for each of the risk factors in question , in an attempt to isolate potential model - based biases . the choice to additionally vary the programs used was in part due to constraints of implementation—no single program offered all the models we wished to compare . this approach has the added benefit of demonstrating a further level of robustness : if our phylogenetic results are insensitive to both the differing models and the myriad of incompletely quantified differences among programs , we can be all the more confident in our conclusions . additionally , varying both the models and their implementation more closely matches the options available to empirical phylogeneticists seeking to resolve ancient rapid radiations . this approach suffers a clear liability , however , in that the effects of model differences and implementation differences are conflated . in the event of differing results , we may not be able to isolate the effects of one from the other ; therefore , the added value to empirical phylogeneticists comes at the cost of reduced utility of our results to program developers and theorists .
the specific evaluations performed are described more thoroughly in the results section . computation - intensive analyses were run on either the duke shared cluster resource ( urltoken ) or the uppsala multidisciplinary center for advanced computational science ( uppmax ) . when appropriate , multiple tree files were summarized onto a target phylogram with sumtrees 2 . 0 . 2 ( sukumaran and holder 2010 ) for subsequent inspection or manipulation in figtree 1 . 3 . 1 ( rambaut 2006 ) .
tree - wide mean ml bootstrap support ( mlbs ) values ( summed bootstrap values on the ml tree divided by the number of internodes in that tree ) for the individual loci ranged from 74 % ( atpb ) to 84 % ( matk ) . the concatenated data have a mean mlbs of 92 % and strongly support ( i . e . , have ≥ 70 % mlbs and ≥ 0 . 95 posterior probability [ pp ] ) 90 % of the partitions ( table 2 ) . across data sets , ml and bayesian inference consistently inferred strong support for a comparable number of bipartitions ( table 2 ) , a result that offers further empirical corroboration for the approximate equivalence of 70 % mlbs and 0 . 95 pp ( hillis and bull 1993 ; alfaro et al . 2003 ) .
there are two well - supported conflicts among the individual - locus ml trees . the first involves a tip relationship ( matk unites deparia pterorachis with d . unifurcata , with 72 % mlbs , whereas rbcl places d . pterorachis as sister to the rest of the genus , with 75 % support ) that is peripheral to the focus of this study . the second is deeper in the tree : matk has 80 % mlbs for a clade uniting thelypteridaceae with the athyrioids , blechnaceae , and onocleaceae , to the exclusion of woodsia and allies , whereas both atpa and atpb place woodsia and its allies within that clade ( 92 % in atpa ; 71 % in atpb ) . given that we confirmed this conflict to not be attributable to laboratory or identification errors , and because the loci involved are linked and the taxa are long - diverged , we do not ascribe this conflict to differences in evolutionary history and proceeded to concatenate all the data for subsequent analyses .
each of our ten ml best - tree searches of the concatenated data in paup * ( from different random - addition - sequence starting trees ) inferred the same tree ( fig . 3a ) , suggesting that tree space is unimodal for our data set . most partitions in this topology point estimate were strongly supported by both ml bootstrapping and bayesian pps ( table 2 ) ; the different ml programs ( paup * , garli , and raxml ) inferred very similar mlbs levels ( data not shown ) .
phylogeny evaluation : rate heterogeneity and the bayesian star - tree paradox artifact . a ) unrooted ml phylogram of the concatenated data , with the backbone internodes highlighted and labeled . b ) accounting for the impact of rate heterogeneity on backbone support values . the four values listed for each backbone internode are : mlbs on full data , mlbs with aspleniaceae ( asplenium and hymenasplenium ) pruned from trees , mlbs with aspleniaceae removed from analysis , and posterior support from beast . c ) controlling for the bayesian star - tree paradox artifact using the polytomy prior in phycas . the four pps listed for each internode are from : mrbayes 3 . 1 . 1 ( susceptible to the star - tree artifact ) , phycas with c = 1 , phycas with c = e , and phycas with c = 10 . d ) controlling for the bayesian star - tree paradox artifact using the yang branch - length prior . the four pps listed for each internode are from : mrbayes 3 . 1 . 1 ( susceptible to the star - tree artifact ) , mrbayes 3 . 1 . 2 with branch - length prior mu1 / mu0 = 0 . 01 , mrbayes 3 . 1 . 2 with branch - length prior mu1 / mu0 = 0 . 001 , and mrbayes 3 . 1 . 2 with branch - length prior mu1 / mu0 = 0 . 0001 .
for certain branches , we observed very high bayesian pps from the mrbayes analysis , but much lower levels of support from the ml bootstrapping ( fig . 4a ) ; these support discrepancies are disproportionately represented among short branches ( fig . 4b ) . this pattern is consistent with artificially high bayesian support due to the star - tree paradox artifact—most implementations of bayesian phylogenetic inference do not consider polytomies among the option set for the mcmc sampler and thus can return high pps for branches that are unsupported by the data ( lewis et al . 2005 ; yang and rannala 2005 ; yang 2008 ; see early hints in cummings et al . 2003 ) .
discrepancies between ml and bayesian support values . a ) ml phylogram of the concatenated data , internal branches only ( all tip branches have been deleted ) . branches are colored according to the magnitude of the difference between their pp ( from mrbayes 3 . 1 . 1 ) and their percent mlbs ( from 5000 pseudoreplicates in paup * ) . b ) internal branches rotated to be vertical and sorted by length . colors follow figure 4a .
to ensure that this “star - tree paradox” artifact was not influencing our assessment of support , we compared the results from our original mrbayes 3 . 1 . 1 analysis ( ronquist and huelsenbeck 2003 ; mrbayes 3 . 1 . 1 is potentially vulnerable to the star - tree paradox artifact ) with those of a non - bayesian analysis ( ml bootstrapping in paup * from our initial assessment of support ) , as well as with two implementations of bayesian inference that use different approaches to reduce their vulnerability to the star - tree paradox artifact .