WIN“... I I I -_I THE SEA$€§NAL HESTCRY AND GEOGRAPHIC DESTRJBUTéCiN 0F APHEDS WFESTlNG RASPBERRY, BLUEBERRY AND STEJAWBERRY 3N THE LQWER PENINSULA OF MICHIGAN Thesis. for the Degree of Ph. D. MJCHIGAN $TATE UNIVERSJTY Francis Edward Giles 1966 THESIS “Ml L LIBRARY Michigan Stan University This is to certify that the thesis entitled The Seasonal History and Geographic Distribution of Aphids Infesting Raspberry, Blueberry and Strawberry in the Lower Peninsula of Michigan presented by Francis Edward Giles has been accepted towards fulfillment of the requirements for Ph.D. degree in Entomology Date May 9. 1966 0-169 ROOM USE ONLY W 74‘ l£12a~ k3,: m“ 4;, -, k; i: midigmm FEB m: @020 ADSTRACT THE EASONAL HISTORY AND GEOGRAPHIC DTSTRdeTlQN 0? A?HIDS INFESTLNG RASPBERRY, BLUEBERRY AND STRAWBERRY IN THE LUK-IER PEN NS'L'LA OF MICE-ilGAN by Francis Edward Giles The seasonal his:ories of Aphids infesting rsspberry, blueberry and strawberry are described. With the exception of two species, all of the aphids were found to be abundspt and not res ricted in distri— bution. Only two S’YJYbCIIy aphids were found to be migratory. imp'30ropl’vora rubi (Rilf .) was commonly found on. raspberry but not on strawberry. The preferred feedirg location was found to be the terminal portions of the canes and the le ves. Specimens were col- lected more frequently rom red raspberry than from pluck ruspberry except in qualitative ssmples from commerci 1 fields where the species was found to be abou: equally ibundsnt on buth hos s. This species appeared to prefer cultivated blsck raspberry to wild black raspberry. Ldrge populations of ibis aphid were no: found uALil :he first or second weeks of August. Amphoropbora sevsori31a Huson was commonly found on :be capes of raspberry. A defini:e preference for both wild and cultivnted blfick raspberry over red raspberry was shown. Large populations were 10: found until the lasf week of July or :he firs: week of August. The sexual forms of :his species are described for the first time. Apnis rubifolii (Thomas) was found [0 be one of :ne mos: common aphids infestirg szpberry and blackberry. Grea:er preference was shown for red raspberry t1an for bluck raspberry. There was some indi— cation that~wild plants of all types are preferred to culzivated stock. This species was found to be almost exclusively n leaf-feeder. The fundatrices appeared early in the spring and were present on the plants for over a month. Sexusles appeared lgLe in the fall and remained on the plants for long periods. Egg production was usually nigh. L rge «4' populations of :nis species were encouctered in 1318 " may and June and were present for lore unbroken periods. \r -_nson;iphis rubicoln (()est.) appears to be in the souzberly limit of i;s range in Michigan. It was not found in quantitative samples from commercial fields and only rarely in other collections. The preferred feeding loca:ion appears :0 be the terminal lQJVCS of rsspberry plifiCS. Because spLerue were found throughout :He season, and alate females were no: particullrly abundant in late fall, this species is no: :xouget to be migrd:ory. The most commox blueberry aphid collected was Masons his pep eri hscGill. This species was found in only Lbe vestern coun1ies of zne Lower Peninsula. The preferred feeding locgiio: nppedrs L0 be the leaves at the base of the plant. s L. Hyzus scammelli Mason was 1 relstively rore blueberry aphid abut was found in bO'b the esszern end western counties. The preferred I I feeding locn:ion appears to be the leaves 4: the bsse of file pl 3‘. No blueberry spbids were found on wild blueberry. The possi— bility of direct or indirect compezition between tlese two species is discussed. Francis Eduard Giles Colonies of Apbis spiraecole Patch were found on blueberry planis on [NO occasions. Rearing teszs showed that it will maintain ifself on blueberry plants for up to 20 days. Two of i:s naiural hos:s, Indian hemp and spires plants, commonly gr u in blueberry plantations. Since this aphid readily moves from :bese two plants to blueberry, further SLudies of g, spiraecola as a possible virus vecLor are indicated. gcyrthosipbon pisum (Harris), a migraiory species, was found in- frequen11y on strawberry. Sexual forms were rarely collec:ed. In- feszations on strawberry plants appear Lo be a resul; of chance sligblings during migrations between preferred host planfs. Acyr'losipbon porosum (Sand.) was found L0 be the mos; common aphid infesting Siruwberr}. Preference was siown for cultiveied plants over wild plents. This species also coloniees rose plants in Hicbigen. Aplis forbesi Weed was :le only sm4ll fruit aphid found on :be roo:s of the hosi plan:. It was no: collected in abundance in Lhe Lower Peninsula and showed a preference for vild scruwberry and home gr vn planzs to commercially grwwn SLOCk. @pbis gossypii Glov. is frequensly mlSLdefl for :. forbe§i_wben found on strawberry. This species was colleceed on.cul:iva:ed striv- berry grown in greenhouses as well as in :be field. if was never found on wild plants. Its scarcity on szrawberry, and :“e fact thin mos: colleCLions consisLed of nymphs, indica c that mos: colonies are be resul: of chance visits by ulate females. A record of en ovi- 9 b parous female on strawberry, :Loueb: to be previously unrecorded, is giVen, C ae‘osipbon fragaefolii (Cock.) was found to be one of :be Sruncis Ldvard Giles ‘ures; of the s rswberry aphids. l: w.s colleczed only in home g rdens, nurseries and imporced experimenrul hybrid plaq:s. It is possible :hut Kris species is present in *he Lower Peninsula only Vien i: is intro- duced on imporced plants. C’aetosiphon minor (forbes), the mos; imported; vector of srr~v~ ..- H berry viruses iv the Lover Peninsula, preferred wild strawberry and home grown plants over commercial stock. Sexuxl forms have u long time span on the plants and egg production was no:ed L0 be moderugely ' o A Wigs on some occasions. fiscrosiphum eupVorbise (Thomas) is a migrutory species that uses SZrawberry as at less: one of i:s secondary hosis in the Lower Peniu— » sula. Large infes*atiors were fOUld in rbe spring and fall but never in midsummer. Prefererce was s“own for s?ruwberries grown in home gurdens and commercial fields. Xacrosiphum rosae (L.) was rarely fourd on s;rswberry; ihe pri- mary “Gs: in the Lower Peninsula is probably rose. The firs; appear aces of :he unducrices of various aphid species are correl;:ed wit“ growirg degree duys. Fvidence is presenied To show thee there is liftle difference in the :ime of :5e species' appearance in :“e weszeru counzies and in the midland, or a: most, this difference amounts :0 orly four d ys. The distribuaiox of LPG aphid species and the parasifies, pred4;ors and lots ussocjgzed vi:h ’bem ere lis:ed. u. The possibilities of aphid populations being influenced by cheeses is Los; pl a; physiology are discussed. THE SEASONAL HISTORY AND GEOGRAPHIC DISTREBUTION U7 APHIDS Y T?)F‘"\f\"’ lifi'ESTING RASPBEBRV", BLUESEE‘L‘TY AND STR Homer: IN THE- LCN‘."ER PENTNSULA J‘s" MICHIGAN if)“. Francis Edward Giles A THESIS Submiited to Micvigun State Ufiiversiry in partiil fulfillment of {he requiremen:s for the degree of DOCTOR or PHILOSOPHY Deparlment of Extomology 1906 ACKNOWLEDGEMENTS I would like LO thank :he members of my commiLtee, composed of Dr. Gordon Guyer, Chairman of the Depirtmen: of Entomology, and Drs. J. Rat“, J. Hoffman, A. Howitt and J. Moulzon for their lelp in the preparation of this thesis. -« To Drs. James bath and James bucher I extend my apprecistio: for their advice, guidance and encouragement. I would also like Lo thank Dr. H. E. HacGillivray, of Lhe Cansda Department of Agriculture, I . . . . and Messrs. Thomas Hlsvuc and Roneld Willson for Lveir nelp in col- lecting and identifying specimens. Finally I would like to ex:end my sincere nppreciazion fo Miss Louise Russell, of tKe United Stiles Depfirlmen: of Agriculture, withou; rlose pnzien: help and consider (ion :His :besis would no: Have been written. ii TAbLE UP CONTTNTS 0:) dc FD LlST OF TABLES . . . . . . . . . . . . . . . . . . . . . . . . . . vii LIST OF FIGURES . . . . . . . . . . . . . . . . . . . . . . . . . ix INTRODUCTION . . . . . . . . . . . . . . . . . . . REVIEW OF LITERATURE . . . . . . . . . . . . . . . . . . . . . . . b Amphoropfora rubi (haltenbscb) . . . . . . . . . . . . . . . . . Amp*oropeora sersoriaza Meson . . . . . . . . . . . . . . . . . Ap is rubifolii (Thomas) . . . . . . . . . . . . . . . . . Husonsp is rubicoln (Oestlund) . . . . . . . . . . . . . . . . . Hasonspbis pepperi FecCillivruy . . . . . . . . . . . . . . . égzus scummelli Meson . . . . . . . . . . . . . . . . . . . . . Acvrrnosipbon pisun (Kultenbdcb) . . . . . . . . . . . . . . \c mmmb C‘ . iggrirrnsipbon porrunnn (Sanderson). . . . . . . . . . . . . . . . épfis forbesi Need . . . . . . . . . . . . . . . . . . . . . . A2 is eossvpii Glover . . . . . . . Cheetosipbon frngnefolii (Cockerell) . . . . . . . . . . . . Ciseiosfplon minor (forbes) . . . . . . . . . . . . . . . . . . liicrosiifmnn eup*mnd3i1e (T mmnas) . . . . . . . . . . . . . . . N4cnuosipfwwn rosgc> L. . . . . . . . . . . . . . . . . . . . . . . CluCLudiio s of Aphid Populstions . . . . . . . . . . . . . . . CDCDN P—‘QOKOCD l—‘P—‘l—JH [\J {4) i I SL1\"‘E;LE}:VI‘ O? :llldrfi{l)DS . . . g o o c o o o 0 a c o o o a o o o o o a 14 General Colleczing . . . . . . . . . . . . . . . . . . . . . . . 14 16 IdeniifiCJtiop . . . . . . . . . . . . .~. . . . . . . . . . . . 16 Quantitative Collecting . . . . . . . . . . . . . . . . . . . . 18 Wild Red Raspberry, 1961 :o 1963 . . . . . . . . . . . . . . . 18 Culcivated Raspberry, 1963 . . . . . . . . . . . . . . . . . . 20 Cultivated Blueberry, 1964 . . . . . . . . . . . . . . . . . . 20 CulLivufed Strawberry, 1964 . . . . . . . . . . . . . . .‘lO‘lefll: 111g . . . . . g g a g o O 0 I o I o a o I o a o o o O o o o {ASPBSRRY APMIDS . . . . . . . . . . . . . . . . . . . . . . . {XMmBl OLSOP‘OYa rlei (Ll/Lil‘er‘bilc!‘> - o n u o o I o o n u o a o 0 ' ' ”I" QuanLi;u:ive Studies . . . . . . . . . . . . . . . . . . . . . Wild Red Raspberry. 1961 . . . . . . . . . . . . . . Wild Red Raspberry. 1962 . . . . . . . . . . . . . . . . 1.7' d D.,-. b ‘7 19r/3 Mild Re “asp err“. 3 . . . . Black Raspberry. Hon:e Farm. 1963 . . . . . . . . . . . . iii . Black Raspberry. Jobnson Farm. 1963 Black Raspberry. Maxwell Farm. 1963 Black and Red Raspberry. Valle Farm. Black Raspberry. Vaughn Farm. 1963 . Red Raspberry. Vaugnn Farm. 1963 . . Ampberopbora sensoriata fiason . . . . . . QuanLiLaLive Studies . . . . . . . . . . Wild Red Raspberry. 1961 . . . . . . Wild Red Raspberry. 1962 . . . . . . Wild Red Raspberry. 1963 . . . . . . Black Raspberry. Monte Farm. 1963 . Black Raspberry. Johnson Farm. 1963 Black Raspberry. Haxwell Farm. 1963 Black Raspberry. Kalle Farm. 1963 . Red Raspberry. Kalle Harm. 1963 . . Black Raspberry. VaugVn Farm. 1963 . Red Raspberry. Vaughn Farm. 1963 . . Ap is reoifolii (Thomas) . . . . . . . . Qumn11iigiLixue Stimiies . . . . . . . . . . Wild Red Raspberry. 1961 . . . . . . Wild Red Raspberry. 1962 . . . . . . Wild Red Raspberry. 1963 . . . . black Raspberry. Jonte farm. 1963 . Black Raspberry. Johnson Farm. 1963 Black Raspberry. Haxvell Farm. 1963 Black Raspberry. Talle farm. 1963 . Red Raspberry. Kalle Farm. 1963 . . Black Raspberry. Vang n Farm. 1963 . Red Raspberry. Vang n Farm. 1963 . . Yasonxp'is rebicola (Oesflund) . . . . . . Quantizative Studies . . . . . . . . . . Wild Red {aspberry. 1961 . . . . . . Hild Red Raspberry. 1963 . . . . . Wild Red Raspberry and Commercial Liosomapbis sp. . . . . . . . . . . . . . p—-.— BLULBEi-{fi‘i ZXPELEDS o o o o o o o o o t I o o o Yasonfipbis pepperi MacGilliyray . . . . . Quan_i:acive Studies . . . . . . . . . . Iiichigyui State [hiivers‘CWJ Horiixu11ture Cbickaming Planiacion . . . . . . . . Hutchinson Plantation . . . . . . . . Hartmann Planzation . . . . . . . . Vakeman Planfiation . . . . . . . . . . Double A Blueberry Firm . . . . . . . Wadswor: b Plant aL ion . . . . . . . . DePree Plantation . . . . . . . . Boodoot Plarrarion . . . . . . . . . . Pauls Plantation . . . . . . . . . . 'r F“ C. l—I . Cl. :3 o 963 . . . 1963 arm . . Page CD\1\J\1\J\I\I l0 [\3 l3 [‘0 h.) Ix.) [*3 PJ IQ 0000 30 45 47 47 47 50 50 50 50 5O 51 51 51 A _.. —_——fi__—_ Derkse Plantation . . . Tyfius scammelli Jason . . . . Quanti:arive Studies . . . . Hicligan Stace Universi:y Cbickaming and Hutchinson Hartmana Plan1ation . . . Wakeman Plantation . . . . Double A Blueberry Farm . {{adéuaortii IqxlniciLiCNI . . . DePree Plantation . . . . Booboo: Plantation . . . . ERulls Inainfaltion . . . . . Derkse Plantation . . . . Apjis spiriecola Patch . . . STRAWBERRY APHIDS . . . . . . Acyrfbosip601 pisnm (Harris) . Quan;i:ative S udies . . . . Tidey Farm . . . . . . . . Ric igan SLate Univ-rsify Hendzel Farm . . . . . . §ELY“U‘osipfiuxi perosrm1 (Sand.) QuanLiLative Studies . . . . Tidey Farm . . . . . . . . UicVigan Szafe Universi1y Wendzel Farm . . . . . . . Ap:is forbesi Heed . . . . Quantirative tudies . . . . gp“is gossypii Clov. . . . . . QLuinLj.;ar ivc: Stimiies; . . . . C aetosipbon fr gleffllll (Cock Quantitative Studies . . . . Cbae*osip:on minor ( orbes) . Quantizafiive SLudies . . . . Tidey Parm . . . . . . . . Hicfligan Stare University Wendzel farm . . . . . . . Hacrosip*um eup’orbi e (Tlomas Quan:i:a:ive Studies . . . . Tidey Farm . . . . . . . . liichigpui Straw: Uniaarrsi }’ / Nendzel Farm . . . . . . . Hacrosiplum ros1e (L.) . . . . Quan;ica;ive Studies . . . DISCUSSION AND CONCLUSIONS . . . Raspberry Ap’ids . . . . . . . Blueberry Aphids . . . . . . . dor. '1 \' -zic » ,. -. L; 0 ul ".lll‘C. tiOnS I O 0 C O O O O O O I I L11” (3 O I O O O O O O I C O O O I I C C I O O D O O I O O 0 v 0 I O D l O O O O O O 0 Ci \I\I\I\J\J wwwuismuraxo \] \J \F \1 CD CD\1 Ulw Scravberry ApVids Early Appearance of Apb‘ Conclusions LITERATURE CITED A P P :le D I X I APPENDIX II APPENDIX III APPENDIX IV APPENDIX y . APPENDIX VI APPCNDIX VII APPENDIX VIII APPENDIX IX vi Pct ('40 9O 97 99 100 105 141 146 [—4 -1 m *‘1 )‘r ,_,. T M3 L E S Table l. Tbe Re ative Abundance of Raspberry Aphids Taken in Quali:ative Collections from 1961 Lbrough 1964 . . . . 1. Relative Abundance of :Le Apbid Species Collecred and the Preferred Feeding Loca ion. Commercial P1. :‘i-CIdS, 1963 I I C I O O O O O C O C O I 0 O O . O . U . 35 Relative Abundance of Lhe Apbid Species Collec:ed and :He Preferred ieedinu Location. Mild Red Ras berry D .1 3 1901-1E163 I o o I o o I O I o O O O O I O 0 O a o n o O L\ Time of Occurrence and {be Dura:ion of :be Horpbological Forms of Raspberry Aphids Compiled from Records Obzaified During tie Years 1961 to 1964 . . . . . . . . . . . . . . 5. Relafive Abundance of Blueberry Aphids Taken in Qua1i1aLive Collections from 1961 :brougb 1964 . . . . . 6. Occurrence of Two Apbid Species in Blueberry P1«1“CdZIOTlS. 19(93 0 o o o 0 o O o o o o o o O c O I I I 7. RelaLive Abundance of Two Aphid Species Collec:ed in Quantitative Samples in Blueberry PlanLations. 1964 . . C0 . HYTUS scammelli Mason. Preferred Feeding Location and frequency DiSLribu'ion on Five Varieiies of Blueberry Crown on the HiCLigan SiaLe Universifiy Rorziculture Til 1.7111 g . . g . g . o . a o o o o o o o o o c o o o a o n o-' .ne Morphological ‘O . Time of Occurrence and Duration of Forms of Blueber;y Aphids Compiled from Records Obtained During :be Years 1961 to 1964 . . . . . . . . . . . . . . 10. Relazive Abundance of Sfiravberry ApVids Taken in Qualiz Live ColleCIions from 1961 :o 1964 . . . . . . 11. Rela'ive Abundance of Apbid Species Taken from [no Vari£>ties (3f S*ranfl3err}n 'Tidey’i’arni . . . . . . . . . . _ . ' (Thomas) (T? Strawberry . . Ampaoroplori seesoriz'a fl son. Oviperous Female . . . Ampboropbora sensoriaia HmSOn. (Top) Male. (301.) Hind Tibia of Male, Slide 3—206 . . C I U ' O . . O I I I ix 48 49 7F) 113 INTRODdCTION Aphids are small, delicate insects that are found on most plants throughout the world. Their ability to reproduce parthenogenetically and viviparously, and the short period required for maturation, enables them to reach epidemic numbers very rapidly. Most species can produce winged forms under certain conditions. These can fly into aphid—free fields and start heavy infestations. Aphids are one of the most serious agricultural pests. Their common name, ”plant lice'l connotes the distaste with which they are re- garded by commercial food growers, agricultural workers and consumers. Aphids live entirely on plant sap. Continued feedi g by large colonies can cause plants to become unthrifty and can lower the sugar content of a crop. Toxic substances injected by a feeding aphid may result in stunting, deformation of leaves or fruit, or the formation of galls on leaves, stems or roots. The excess sugars present in the sap are excreted by these insects as honeydew which may disfigure personal property and plants. Frequently large deposits of honeydew support the growth of mold on the plants. These molds may lower the market value of a crop and interfere with photosynthesis and normal pollination. The ability of some aphid Species to transmit viruses results in diseases.which reduce crop yields and may bring about the death of the host plant. The presence of only a few virus vectors, feeding for [\J only a short time, is all that is needed to inoculate a stand of healthy Since very little work has been done on aphids in Michigan, it is difficult to estimate how many species are present in the State. Leonard (1963) estimated that between 303 and 450 species occur in New York, and it is possible that as many can be found in Michigan. It is unfortunate that so little is known about the aphid fauna of Michigan as it is one of the leading agricultural states in the union. It ranks first in the production of black raspberry, fourth in red raspberry, second in blueberry and, third in strawberry. When this research was started, only four aphid species were definitely known to infest small fruits in Michigan. Nothing was known of their distribution, relative abundance and biology. Since small fruit farming is so important to the Michigan economy, and promises to become even more important in the near future, an investigation of these pests appeared to be a worthwhile project. A brief outline of the generalized aphid life cycle follows. This will serve to acquaint the reader with terminology used by the author . Nymphs hatch from OVerwintering eggs in the early spring; these develop into stem mothers or fundatrices. The fundatrices reproduce parthenogenetically and viviparously to give progeny which mature into the next morphs, or forms. The summer viviparae consist of apterous and alate females which also reproduce parthehogenetically and viviparously. The wingless summer viviparae are commonly spoken of as apterous females or viviparous females. The term apterac includes both the nymphs and sexually mature individuals, or imagines, of the npterous vivipnrous femele. The winged summer vivipnrae are called slates, or slate females. Since the nymphs that will develop into winged forms have external wing pads, they are called alatoid nymphs. The summer vivipnrae are the most abundant forms and are present during the greatest portion of the season. Occasionally, anomalous specimens are produced which have abortive wing pads and show characteristics of both the slate and apterous females; these are known as intermediate forms. In the fall of the year, the summer vivipnrae produce nymphs which mature into scxunles, or true male and female forms. After these morphs mate, the females produce eggs which overwinter and complete the life cycle. The ovipnrous females are usually apterous and are termed ovipnrne; males are usually winged. Since the summer vivipnrne are most often encountered in the field, all taxonomic keys are based on these morphs, and especially upon the slate females. The identification of the other forms can be very difficult, if not impossible. REVIEW OF LITERATURE Amphorophora rubi ( altenbnch) Mason (1925) and Ninter (1929) stated that this is a cosmopolitan species found mainly on the plants of the genus Rubus. Winter reported it to be the most active aphid on raspberry. He stated that it was a leaf-feeder and that it was most abundant on red raspberry. Populations fluctuated greatly and specimens were hard to find after hot, dry weather. Alates were uncommon in Minnesota. Eggs were laid on the leaves but were hard to find. In testing seven varieties of red raspberry for resistance to this species, Winter found that the variety Latham ranked second in resistance. In Canada, Stace-Smith (1960) reported that Latham was susceptible to this species, and Daubeny and Stace-Smith (1963) reported large colonies of this aphid on Latham. é. ruhi_is the principle vector of raspberry mosaic (Cooley 1936, Daubeny and Stace—Smith 1963) and has been shown to transmit black raspberry necrosis virus (Stace Smith 1960). It is also a vector of gubus yellow net (Lennedy, Day and Eastop, 1962). Cooley (1936) found that practically all wild red raspberry and some wild black raspberry were infected with mosaic in New York. There are races of g, rub} that differ in ability to colonize different hosts and to transmit diseuses (Russell 1962, Dauben and / cl Stace-Smith 1963). In western United States, Canada and, to some extent, Nova Scotia, it is a pest of strawberry (Craig and Stultz 1964, Frazier 1951). Plakidas (1955) reported that it transmits some or all of the components of the strawberry yellows and crinkle virus complex. Mellor and Forbes (1960) cited Frazier as being successful in transmitting strawberry mottle virus with this species. Kennedy §£.ilx (1962) listed this aphid as a vector of strawberry leaf banding. Amphorophora sensoriata Mason Mason (1923, 1925) erected this species but did not describe the sexual forms. He found it to be exclusively a cane~feeder and thought that it might be migratory as few specimens were found in the summer. Winter (1929) did not consider it migratory and reported that black raspberry was the preferred host. Both authors reported it widely distributed in North America with a range as far south as Virginia. Patch (1938) did not list it from any hosts except 3223: spp. Cooley (1936) stated that it is a minor vector of raspberry mosaic. Kennedy t al. (1962) listed it as a vector of black raspberry necrosis. -—.——_— Aphis rubifolii (Thomas) O This species was reported only from the leaves of raspberry and blackberry plants (Patch 1938, Palmer 1952, Leonard 1963). Winter (1929) reported it common throughout North America and stated that it preferred the undersurfaces of wild red raspberry leaves. In early SGPtember he found an average of 54.7 specimens on a leaf but reported counts as high as 219 per leaf. He also observed that the eggs hatched pl when the buds first turned green. Cooley (1936) reported this species an inefficient cerrier of raspberry mosaic and the sole vector of leaf curl in raspberry and blackberry. Kennedy E£.il' (1962) listed it as a possible vector of black raspberry necrosis. Uasonaphis rubicola (Oestlund) This aphid was reported only from the plants of the genus Bubus (Patch 1938, Palmer 1952, Leonard 1963). MacCillivray (1958) recorded it from New Brunswick, Nova Scotia, Quebec, ntario and British Columbia. Winter (1929) reported it from California, Minnesota and Maine. Cooley (1936) and Palmer (1952) found it rare in Colorado and New York. In Michigan, Benrett (1932) had to import colonies for experimental work. MacGillivray (1958) stated that the life cycle was not clear; 5111 although specimens were found on raspberry all summer, it could have alternate host. She recorded epterous and alate females from June 16 to October 22 and sexuales on October 4. In Colorado and Utah, Palmer (1952) recorded apterous and alate females from August 28 to October 23 and slate males 0? Scp1ember 25' Cooley (1936) did not consider it an important vector of rasp- A (1962) listed it capable of transmitting berry mosaic. Kennedy et a1. -—-.—- black raspberry necrosis. Mnsonaphis pepperi MacCillivray MacGillivray (1958) reported this aphid from Maine and Pennsylvania on Vaccinium augustifolium Ait., i. cor mbosum L., \_ y. stamineum L. and Vaccinium_sp. Leonard (1963) reported it from New York on Vaccinium sp. In Pennsylvania, MacGillivray (1958) recorded fundatrices on day 25 and June 1, apterous and slate females from May 25 to September 1 and oviparous females on October 16. In Michigan, Tuttle (1947) found only a few aphids. He reported that they appeared to be of one species and were causing no noticeable damage. fiyzus scammelli Mason Mason (1940) recorded this nphid on cranberry in New Jersey. MacGillivray (1965) stated that it occurred on the low sweet blueberry (V. augustifolium) in Nova Scotia. Marucci (1964) found that it pre~ ferred the soft, succulent growth at the base of the cranberry plant. He considered it a rare aphid but stated that in recent years it has become more abundant. The greatest numbers were fOUhd in May ard after June it became scarce. His attempts to transfer this species from cranberry to blueberry were not successful. He stated that the toughness of the leaves may have been responsible for the failure of this aphid to colonize blueberry. No literature was found to indicate that N. pepperi or g. scammelli are known to transmit virus diseases. As shoestring disease is becoming more common in Michigan plantations it is interesting to note that Lockhart and Hall (1962) have found this disease to be present in all lowbush plants (V. eugustifolium) tested in Nova Scotia. This blue— berry species also occurs in Michigan. Agyrthosiphon pisum (Inltenbsch) This is a cosmopolitan species that is found mainly on plants of the Leguminosae (Hille Ris Lambers 1947). Evans and Gyrisco (1956) stated that this aphid is generally considered to be nonmigratory although it flies from perennial to annual legumes. No records of this species transmitting strawberry viruses were found. Acyrthosiphon porosum (Sanderson) . This species is found throughout North America on Ross spp. and strawberries (Mason 1940). Pnlmer (1952) reported it only from rose in Colorado and Utah; Mason (1940) found that it was more common on rose thnn stranberry although the eggs were 111d on both plants. Craig and Stultz (1964) found it to be the mos: abundant strnyberry aphid in Nova Scotia where it represented 53.11 and 73.31 of the aphids collected in a two year period. Demdree and lecus (1951) stressed the importance of knowing the exact status of this species as e strawberry pest in view of its potential as a virus disease vector. it has been shown to be capable of transmitting strawberry mettle and some, or all, of the components of the yellow‘ and crinkle virus complex (Plakidos 1955, Kennedy et al. 1962, Craig nnd Stultz 1964). gphis forbesi Weed This species is apparently restricted to strawberry Patch 1938, Palmer 1952). It is distributed throughout North America but nppenrs to be more abundant in the eastern section of the United States (Palmer 1952, Allen 1959). In Nova Scotia it comprised only 17.11 8rd 2.31 of the collections made in a two year period (Craig and Stultz 1964). Many authors, including Hottes and Prison (1931) and Cutright (1925), reported that this aphid is carried to the plant's roots by ants. Marcovitch (1925) stated that Lasius alienus (Foerster) is an important ant in this respect but that Pheidola vinelandica is the most common ant doing this in Tennessee. Wheeler (1910) listed ants of the genera Myrmica and Crematogaster among those thnt commonly attend aphids but stated that the genera Lasius and Prenolepis are among those most per- fectly developed in this respect. Attempts to transmit strawberry diseases with this aphid have never been successful (Kennedy, Day and Eastop 1962). Aphis gossypii Glover This is a cosmopolitan species (Patch 1925b) consisting of many races with,different life cycles (Bodenheimer and Swirski 1957). Leonard (1963) listed 60 food plants for this sphis in New York. Patch (1925b) reared it on strawberry experimentally but reported that its primary host in Maine was orpine (Sedum purpureum Teusch). firing (1959) found the primary host in Connec:icut to be Catalpa bignonioides Walt. and showed that this species is a faculative migrant. Batchelder (1927), Walle (1933) and firing (1955), described the many color and morphological variations of this aphid. Kennedy et nl. (1962) listed this species as capable of transmittirg strawberry mottle. Chaetosiphon fragaefolii Cockerell) This is a cosmopolitan aphid (Schaefers 1960) that occurs on 10 rose, strawberry and Potentilla spp. (hottes and Frison 1931, Palmer 1952); Schaefers (1960) stated that Potentilla spp. is not a preferred host. Patch (1938) and Plakidas (1955) reported it from rose and straw- berry, and Leonard (1963) reported it only from strawberry in New York. Although Palmer (1952) observed migration of alates from rose to straw- berry, this species has no alternate host and only makes dispersal flights (Dicker 1952, Schaefers and Allen 1962). This species is a serious pest in California but is relatively scarce east of the Mississippi (Demaree and Marcus 1951, Plakidas 1955, Schaefers and Allen 1962). Craig and Stultz (1964) did not collect it in Nova Scotia during a two year study, and Fulton (1954) did not find any aphids of this genus in Michigan during a four year study. C. fragaefolii is the most important vector of strawberry viruses. Plakidas (1955) stated that it is the principle vector of xanthosis and crinkle, and that it also transmits stun' and witches broom. Craig and Stultz (1964) reported that it transmits mottle and vein banding and Kennedy 55.51: (1962) listed it as transmitting strawberry lesion, mild yellow edge and vein chlorosis. Plakidas (1955) stated that yellow edge is now present in wild Strawberries east of the Rocky Mountains, but that wild roses in Washington and Oregon which harbor this aphid did not appear to be hosts of any strawberry viruses. Cheetosiphon minor (Forbes) This aphid is found throughout the United States but is more abundant east of the Mississippi (Demaree and Marcus 1951, Palmer 1952, Plakidas 1955). it appears to be restricted to s:rewberry (Mottes ard ! 11 Frison 1931, Patch 1938, Palmer 1952, Leonard 1963), but Knowlton (1954) reported it from Potentilla Sp. in Washington. Craig and Stultz (1964) cited Rorie as saying it was the dominant vector in Arkansas. However, in Nova Scotia these two authors did not find it in 1961, and in 1962 it accounted for only 0.57 of the collections. Fulton (1954) did not collect any aphids of this genus during a four year study in Michigan. 9. minor has been shown to transmit virus type 1 and 2 of Demaree and Marcus (Plakidas 1955) and strawberry mottle (Kennedy et 21,, 1962). ~— Demaree and Marcus (1951) reported that Potentilla simplex Michx. can serve as a symptomless host of type 1 and 2 viruses. In Michigan, Fulton (1954) found that P. anserina L., P. argentea L. and P. recta L. acted as latent hosts of type 2 virus. Macrosiphum euphorbine (Thomas) This is a polyphagous and cosmopolitan species (Castop 1958). Patch 0925a) and Leonard (1963) stated that its primary host is Rosa sp. Patch observed some oviposition on strawberry in Maine but stated that the fundatrices rarely hatched. She also stated that spring colonies had been reported in New Jersey. Demaree and M rcus (1951), however, reported it was a frequent h hitant of strawberry and stressed the importance of knowing this species' ex ct status in view of its potential as a virus vector. Craig and Stultz (1964) found that it account for 22.7% of their collections in 1961 and 15.52 in 1962 and Steted that it is a possible vector of latent C virus. Macrosiphum rosae L. This is a cosmopolitan species that is apparently restricted to Rosa spp. (Patch 1914, Hottes and Prison 1931, Palmer 1952, Leonard 1963). Mellor and Forbes (1960) stated that this aphid is capable of trans- mitting mild yellow edge and vein banding viruses. Fluctuations of Aphid Populations Many investigations have been instigated to explain the fluctua- tions of aphid populations. In recent years, attention has been focused on the reactions of aphid populations to the physiological activities of the host plant. Bodenheimer and Swirski (1957) gave several examples in which aphid species apparently respond to physiological Changes of the host plant. They also stated that indirect competition can occur when any sucking insect feeding on a leaf, changes the physiological condition of the leaf, and renders it unsuitable for future aphid feeding. Kennedy, lbbotson and Booth (1950) showed that Myzus persicae (Sulz.) and Aphis fabae Scop. preferred young, growing leaves and senescing leaves to mature leaves. lbbotson and Kennedy (1950) showed that this preference was modified by the rate with which the leaf was growing or senescing. The apparent reason for these reactions is the young leaves are areas in which nitrogenous substances are being mobilized for protein synthesis. The senescing leaves are areas where proteins are being hydrolyzed and translocated. Both leaf types by concentrating nutrients, apparently provide a more suitable site for aphid feeding and reproduction than do mature leaves. Dicker (1952) working with g. fragaefolii on strawberry in 13 Great Britain, found that in first year plants the population of this aphid increased steadily thrOUghout the season. in older plants, the PM 'rdo "r, aphid population peaked in early spring and then declined rapidly. l hypothesis, although admittedly not the complete explanation, was that the first year plants produced leaves steadily all season and the physiological activities of the plant, not being disrupted by fruit formation, yielded a steady supply of nutrients that permitted a steady build up of aphids. Older plants exhibited a spring flush of growth and this increased physiological activity enabled the aphid population to increase around the time of early fruit ripening. After the fruit had ripened, leaf production and development decreased, and the aphid population declined due to lack of suitable feeding sites. Dicker also suggested that at the time of fruit ripening, physiological changes occur in the leaves that are unfavorable to aphid reproduction. Schaefers and Allen (1962) found evidence that Q. fragaefolii populations also tended to follow fluctuations in Strawberry physiolOgy; however, under California conditions the population peaks were not as closely tied to fruit production as reported by Dicker. Schaefers and Allen pointed out that the lower leaves of the strawberry provided a cooler and more humid microclimate that is more favorable to aphid survival and increase. They found, as did Dicker, that high aphid mortality occurred due to splashings of rain and mud, especially after fields had been topped. STATEMENT OF METHODS General Collecting Most of the aphids collected were taken by means of a #1 w ter color brush, the ends of which were trimmed to a fine point. This in— strument allowed the delicate specimens to be removed from the pltnt without damage and provided the collector with a positive record of aphid-host plant association. Aphids were collected and stored in 957 ethanol. Storage in 702 alcohol, as recommended by Essig (1948) and Palmer (1952), resulted in poorly preserved specimens. Nymphs found in the field were frequently ceged and allowed to mature and reproduce; the entire colony was then removed for identifica- tion. Clip cages used by Mc Clanahan (1961) were found to be too small. New ones were fashioned from microscope cover slip boxes (Tig. 1). These boxes had plastic screen windows and were tied to the plant stem. Loose string or cotton packing was used to make the cages "aphid tight”. Although adequate, the cages were awkward to handle and were replaced by cages made of plastic centrifuge tubes as described by Peschke (1959). To collect alnte forms, sticky traps and water traps were placed in test plots. Sticky traps were made from the bottoms of commercial one pint ice cream containers (Fig. 1). These were painted with Canary Yellow Effecto Enamel (Pratt and Lambert Co.) and coated with 14 Fig. 1.--Aphid cages and sticky traps used by the author. lb Hepco—Stikem (Michel d Pelton 00., Oakland, Calif.). These traps worked well but becsme covered with earth and debris. Trapped aphids were in such poor condition that identification was impossible and the use of sticky traps was discontinued. Water traps were made from miscellaneous metal and plastic pans measuring approximdtely 18" X 12" X 8”. Aphids collected in these con- tainers were in fdirly good condition except for mold and detritus which adhered to them. In addition to the problems described by McClanahan it was found that the traps had to be cleaned and refilled about every three days. This limited their vglue and it was decided to use the presence of slntoid nymphs to indicate the appearance of slates (Dicker, 1952). Mounting Freshly killed mnterial was heated in the storage alcohol at just below boiling point for five minutes to eliminate internal air bubbles. Specimens preserved for more than 24 hours did not require this treatment. Initially, aphids were cleared by the techniques recommended by Essig and mounted in Turtox CMC - 10 medium (General Biological Supply House, 8200 So. Hoyne Ave., Chicxgo). Both the techniques and mounting medium proved unsatisfactory and the clearing and mounting procedures developed by Hille Ris anbers (1950) were followed. Identificgtion Most specimens were identified by means of a compound microscope l7 equipped with a phase contrast optical system and a measuring eyepiece. Large samples of apterae were frequently determined with a Wild Model V dissecting microscope equipped with both substage and conventional illumination and having a magnification of up to 200 X; when this micro- scope was used, the aphids were first cleared and then examined while still in the clearing solution. Vibrations and convection currents made focusing difficult and questionable specimens had to be mounted for inspection with a compound microscope. In retrospect, it is doubtful if the use of a dissecting microscope saved any time over conventional methods of identification. The two ”standard” references, ”The Plant Lice, or Aphididae, of Illinois” (Hottes and Frison, 1931) and ”Aphids of the Rocky Mountain Region" (Palmer, 1952) were foun to be inadequate. Publications proving helpful were those of MacGillivray (1958), Mason (1925, 1940), Schaefers (1960), Richards (1963), Bachelder (1927) and Hille Ris Lambers (1953). Sampson's keys to nymphs (Sampson, 1946) were of limited value and most nymphs were determined through the aid of Miss Louise Russell of the U.S. Department of Agriculture. Through the cooperation of Miss Russell and Dr. H. E. HacGillivray of the Canada Department of Agriculture the author was able to accumulate enough determined materigl to enable him to make his own identifications. Parasitized aphids were collected along with the leaf or stem to which they were attached and placed in screw cap vials. To avoid the growth of molds, the vials were left uncapped for 24 hours to allow the moisture from the plant material to evaporate. The parasites that 18 emerged were sent to the U.S. Department of Agriculture, Insect Identi- fication and Parasite Introduction Research B anch for identification. Heavy clearing of the parasitized aphid was necessary due to dis- coloration of the integument and the presence of exuviae and internal frass pellets from the parasite. This was accomplished by placing the plant material and the aphid on a slide in a drop of KOH and adding a cover slip. The slide was heated until the specimen was clear and‘ pliable. The aphid was then washed, separated from the plant material, and mounted. Identification was made with a compound microscope. Although many specimens could not be identified due to parasite— induced distortions, this method of identification is more positive than the common one of identifying the parasitized specimen by means of a normal specimen collected from the same plant. Quantitative Collecting The location and description of each test plot with the dates of collections and the numbers of aphids taken are presented in Appendices VI through IX. Wild Red Raspberry, 1961 to 1963 (Fig. 2) This stand was too small for true random sampling. To reduce personal bias, sampling was begun at different ends of the stand every week. At every five paces a plant was selected. Three compound leaves were picked from each plan:. These leaves-~one from the tip of the Plant, one from the base of the. plant and one midway between top and 19 Fig. 2.--Stand of wild red raspberry on the Michigan State University campus. bottom~-were placed in separate jars of alcohol. Any aphids found on the canes were placed in vials of alcohol with a fine brusw. Specimens were removed from the alcohol with the aid of a dissecting microscope and identified. In 1963 an attempt was made to sample these raspberry plants by treating the leaves with heat and Methyl isobutyl ketone in Berlese funnels. It was found that Aphis ruhifolii (Thomas) was too sedentary to pass through the apparatus and died among the leaves. As a result, leaf sample data from September 4, 11 and 18 are not available. Cultivated Rispberry, 1963 The rows of plants were numbered consecutively. The plants to be sampled were selected by pacing off along the rows. The numbers used in selecting the samples were ‘nken from J table of random numbers (Dixon and Massey, 1957). Three plants in eech plot were sampled at weekly intervals. The collections were made and the samples were treated as described for wild red raspberry. Sampling was without replacement. / Cultivated Blueberry, 19‘4 Blueberries growing on the Michigan State University Horticulture Farm were sampled wi2hou: replacement until October 11. On this djte all the plants in CJCh varie:sl row had been Ssmpled. Subsequent ssmples had to be made from plants which had been sampled earlier in the season. Each plant was numbered and the plant to be sempled was chosen 3 at random. Two plants of each variety were sampled at approximately two week intervals. Collections were made with the D-Vac model 12 Sampler (D-Vac Co., 1462 Callens Rd., Ventura, Calif.). An evaluation of the efficiency of this machine was made by Naki (1965). In taking the sample, the running D-Vac machine was positioned in the foliage so that a branch was in the net. The machine was then pushed deeply into the plan: until the branch stopped its forward motion. This operation was repeated five times around the top and around the bottom of the plant so that the plant was completely encircled. The collections from the crown and :he base of the plant were treated separately with Methyl isobutyl ketone in J portable Berlese funnel designed by Niemczyk (1963). In 1963 it was decided tint the author would cooperate with Hr. T. L. Burger who was also doing graduate research on blueberry in- sects. Aphids collected by Hr. Burger from plantations in 1963 and 1964 were submitted to the JULhOf for identification and the results are presented in this thesis. For a description of the collecting and sorting techniques used in this pert of the research see Burger (1960). letivated StrawberryJ 1964 The IWNMS of plants VKMNB consecutivelyriunnbered and the areas to be sampled were selected by pacing off along the rows. The numbers used in this operation were drawn from a table of random numbers. Sampling, which was done at approximately two week intervals, was without repldcement. Five areas were sampled in each field on each collecting date. TO IQ When the area of the strawberry row had been selected, the running D-Vac machine was lowered to the ground over the plants and then raised. This cycle was repeated ten times, allowing about one second for each motion. The contents of the net were then transferred to a portable Berlese funnel. As the amount of debris picked up in taking five samples was too great for the capacity of the Berlese canister, the samples were treated as follows. After placing the first sample in the canister, the second sample was taken. This was added to the canister and the two samples allowed to remain for fifteen minutes. The Berlese funnel was then sharply bounced on the ground five times to dislodge any aphids that were clinging to the vegetation. As the upper chamber was being cleared of debris, a search was made for any remaining aphids. This procedure was repeated for the remaining three samples. When sampling stravberries on the Michig n State University Horticulture Farm, it was HCCQSSdrY to modify these methods slightly as the plants were plaited by varieties in short rows. To select the sample area, a forked stick was constructed having a spread equal to the net diameter of the D—Vac machine. This forked stick was 'walket” up the rows, as with a pair of iavigational dividers, until the chosen area was reached. Two samples from each varietal row were placed. in the Berlese funnel and treated in the manner previously described. Sampling without replacement was practiced until September 13 when the entire length of the varietal rows had been sampled. After this date sub— sequent samples were mede from areas that had been sampled earlier in the season. RASPBERRY APHIDS Wild and cultivated raspberry plants were qualitatively examined from July 1, 1961 through 1964.‘ Data summarizing the relative abun- dance of the aphid species in various collecting situations are pre- sented in Table 1. .No aphids were ever found on the roots of the plants or in the leaf litter from the raspberry beds. TABLE 1.-—The relative abundance of raspberry aphids taken in qualitative collections from 1961 through 1964 Number of Collections Amphorophora Amphorophora Aphis Masonaphis Made in Various Sites rubi sensoriata rubifolii rubicola Commercial fields Red raspberry 12 33.3% 16.7% 33.3% 16.71 Black raspberry 40 35.02 42.5Z 12.5f 10.07 Home gardens ,_ A Red raspberry 54 44.42 18.5% 40.71 7.4? Black raspberry l 0.0% 100.0% 0.02 0.07 Nursery stock ' m . Red raspberry 4 25.0% 0.0% 25.0} 25.0; Black raspberry 3 0.07 66.0% 0.0: 0.0; Wild plants ~ m Red raspberry 18 38.9% 11.12 55.0} 5.5: Black raspberry 18 11.11 38.9% 38.9L 16./a Blackberry n C m wild plants 3 0.0? 0.0; 100.0: 0.0; Cultivated plants 7 0.0? 0.01 42.9. 0.0: Amphorophora rubi (Kaltenbach) . o . , ‘ 91 ‘ l . {4" 3. (ITS - This was a common speCies that was found on cultiv.ted rtd r1 p berry (Rubus idaeus L. X Rubus strigosus (Hichx.)), Wlld red raspberry 23 [\J L\ (Rubus strigosus L.) and on cultivated and wild black raspberry (Bubus occidentalis L.) throughout most of the Lower Peninsula. In 1961, large infestations were encountered in Clare Co. on August 27, in Emmet Co. on September 19 and in Monroe Co. on September 30. in 1962 this aphid was not collected too frequently in May and June but appeared to become more abundant during the summer. A. rubi was the mosc active aphid studied. Eben disturbed, this large, rapidly moving species would frequently release its hold on the plant and drop to the ground. Observations indicate that this species occurred most often on the terminal portions of the canes and on the upper leaves. There appeared to be no preference for either the top or bottom surfaces of the leaves. Copulation and oviposition were not observed. Quantitative Studies (fig. 3) Wild Red Raspberry. 1961 The first samples were taken on July 4. This species was not found until August 9, when apterous females and nymphs were taken. The population of this species increased rapidly and reached its peak on August 14. The population maintained itself at this peak until August 21 and then began to decline. No specimens were collected from September 7 until September 27 when a number of nymphs were found. After that date, only a few scattered nymphs were taken. Wild Red Raspberry. 1962 The first specimen, an apterous female, was collected on June 29. No Specimens were found the following week, but on July 13 the numbers I‘) U! Wild red raspberry, 1961 Johnson. Black raspberry ()0 .. 60 - 30_ 30 q 1 I 7 T T I“ _-_T r *fi Wild red raspberry, 1962 VaUghn. Black raspberry 60- 6O _ 3O . 30 - T r_4.7 u fl" T— fir} T— j I fi—l Wild red raspberry, 1963 Vaughn. Red raspberry 60. 60 . 30 u 30 . Jun Jul Aug Sep Oct Nov Jun Jul Aug Sep Oct Nov fig. 3.--Seasonal population trends of Amptorophora rubi (Kalt.) on raspberry. of apterae increased. This species was not found again until August 10, when its numbers again began to increase. The population reached its peak on August 17, and then began a steady decline. No specimens were taken from August 2’ until Occober 12; on October 19 the numbers in- creased slightly, but this was the last time this species was found. hild Red Raspberry. 1963 This species was taken iknf the first time on July 3. No specimens were found again until July 24. On this day the population began to build up and reached a minor peak on August 14. After a decline on August 21, the population reached another minor peak on August 28. The numbers declined after this date, and this species main— tained iiself at a low level during mid~September until it began to recover on September 25. In the first part of October the numbers declined again; on the following week the population began to build up 9 strongly and the peak occurred on October 23. The population fell off rapidly after this, and 10 specimens were found on the lest day of. collecting. Leaf sample data for September 4, 11 and 18 are not evailsble. The information presented for this time interval is for the numbers of It is assumed that the \— this species taken only in the cane samples. population decreased during these three weeks, for on September 25, very few specimens were collected. Black Raspberry. Monte Farm. 1963 A.rubi was not found in this field. [\J \l 13‘ Black Raspberry. Johnson .arm. 1963 A single nymph was taken on September 21. Black Raspberry. Maxwell Farm. 1963 A. rubi was not found in this field. t Black and Red Raspberry. Valle Farm. 1963 A. rubi was no: found in these fields. Black Raspberry. Vaughn Farm. 1963 A single apterous female was collected on August 10. Red Raspberry. Vaughn Farm. 1963 The first specimens of A. rubi were collected on June 15. The numbers increased on the following week but then began to decline a: the end of June. No specimens were taken from July 6 to July 20. On July 27 the numbers of this small population increased but a steady decline then set in. From August 10 to August 29 no specimens were The taken. On August 29 the large5t number of specimens was taken. population then began a steady decline thnt l 810d for the rest of the season. Amphorophora sensoriafa Mason This was a common species that was collected from both wild and cultivated red and black raspberry throughout mos: of tee Lower Peninsula. In 1961, large infestations were encountered in Berrien Co. on July 26, in Kent Co. on July 30, in Hillsdale Co. on August 11 and in Newaygo Co. on September 17. Since Mason did nor describe the sexusl forms of this aphid, descriptions of these morphs are presented in Appendix I. In 1962 this aphid appeared to become more abundant during July and August. On August 18, an intermediate morph was collected in herrien Co. was found most often on the canes of the host plant. A A.SCHSOFlFta It was easily noticed because of its habit of forming linear colonies on the canes. This species was nored to be feirly active; in any colony, there were always a few individuuls, usually imagines, moving among the colony. The nymphs were usually fairly 'ndctive. When disturbed, the individual aphids of the colony would move quite rapidly, but were no: Copulation and oviposition particularly prone to drop from the plant. were not observed. Quantitative Studies (Fig. 4) I Uild Red Raspberry. 1961 Nymphs of this species were taken on August 9. The population increased the following veck and reached its peck on August 14. The numbers of this species then begun to decline, until on August 30, no specimens were found. A single numph tgken on September 7 is the lust record of this species in this plat. Wild Red Raspberry. 1962 Only nymphs of this species were found. These were collected on July 13 end August 17. [J \D Wild red raspberry, 1961 Maxwell. Black raspberry 50- 50- ] 1 l 5—] I l +1 I lfi 50. Wild red raspberry, 1962 50. Walle. Black raspberry 25¢ 25~ H_l—-—'—"I‘I l [—1 'fi—LI'L'IRI (fit 50. Wild red raspberry, 1963 50- Vaughn. Black raspberry 25- 25- Monte. Black raspberry 50. Vaughn. Red raspberry 200 | 25 ‘““'“-'"""_"" 25- , l T I ‘1 ‘ I *4 I —[ , Jun Jul Aug Sep Oct Nov . Jehnson. Black raspberry 000 400 200 __,____l_|_l___....--._.._. 50 25 T Jun Jul Aug Sep Oct Nov Fig. 4.--Seasonnl population trends of Amphorophora sersorieta Mason on raspberry. 30 wild Red Raspberry. 1963 é. sensoriata was not collected until August 7. The smell population maintnired itself et a fairly steady level until August 21 when it diminished. During the next two weeks, it began to recover and the largest number of specimens were taken on September 4. The numbers then diminished but begun to build up again during the last of September. On October 2 no specimens were found. On Cetober 9 the numbers again rose, but this was the lest time the species was found in this plot. Leaf sample data for September 4, 11 and 18 are not available. The information presented for this time interval is for the numbers of this species taken only in the cane samples, and it is possible that this species cas present in larger numbers during this time. Black Raspberry. Monte Farm. 1963 This species was not found until July 13 when the population abruptly reached its peak. The numbers declined rapidly in the fol- lowing week, and after July 20 this aphid was not found again. However, a visual inspection of the plants showed that small numbers of aphids, believed to be é. sensorinte, were present throughout the collecting 888.8017. Black Raspberry. Johnson Farm. l9o3 On June 8 a single specimen of A. sensorinta was found. This i Species was not taken again until July 20 when a rapid bUild-up began. After a slight decline on July 27 the population began a steady increase that terminated in a population peak on August 17. During the next two weeks the population declined steadily but attained a minor peak on 31 September 7. From this date uxtil the end of September, the population declined steadily, and none of this species was present in the samples of the last day. Black Raspberry. Maxwell Farm. 1963 The only time that this species was taken was on August 10, when two apterous females were collected. Black Raspberry. Walle Farm. 1963 This species was not abundant in this plot. The first specimens of apterae were taken on July 20. The following week no specimens were found. On August 10 the population reached its peak, but declined sharply the following week. No specimens were taken on August 17 and 24. On August 29 the numbers increased slightly, but this was the last time that this species was found in this field. Red Raspberry. Walle Farm. 1963 A. sensoriate was not found in this field. Black Raspberry. Vaughn Farm. 1963 This species was not found until July 20. On this date the POpulation rose abruptly but immediately declined, and no spec1mens were collected on the following week. On August 10 the population had A Deoun a steady increase that terminated in a population peak on A“gust 17. The iumbers declined in the following weeks and by 3 r) 1‘ ;‘.~ - september 7 no specimens were collected. On September -8 t»e popula tion had begun to rise again. Red Raspberry. Vaughn Farm. 1963 This species was not found until August 10. After this date the species was not present in the samples until August 29. On this date the population reached its peak and then declined abruptly. No specimens were found on September 7. The last specimen collected, a viviparous female, was found on September 14. éphis rubifolii (Thomas) This was a common aphid that was found on both wild and cultivated black and red raspberry throughout most of the Lower Peninsula. It was also collected from wild and cultivated blackberry (Rubus allegheniensis Porter). Ants were frequently found in colonies of this aphid. A list of these ant species is presented in Appendix V. g. rubifolii was found to be an extremely sedentary species that was not prone to move when disturbed. The preferred feeding location appeared to be close to the veins on the under—surface of the leaf. On a few occasions it was found on the canes or the leaf petioles. ‘ Trouble was frequently experienced in removing this small aphid from the plan: because of the tenacity it showed in maintaining its feeding position. Heat and chemicals were not successful in driving this species from the leaves. Intraspecific copulation was not observed. However, on October 25, 1964, in Ingham Co., an oviparous female was taken E§.EQBElj. l . .. . ‘ with an alate male of the genus Nearctaphis. In 1963 ov1posi-ion has Determination by Dr. M. E. MacGillivray. 33 observed in Ingham Cb. on wild red raspberry. Oviposition was first observed on October 9 and continued until October 25. The eggs were yellow when first laid but turned black with age. Some of these eggs were stored in 95? alcohol and later mounted in glycerine. The average measurement of seventeen newly laid eggs was .54 x .25 mm. The average measurement of twelve black eggs was .52 x .22 mm. The preferred oviposition site seemed to be the axils of the leaf petioles at the tips of the plant. As oviposition activity increased, the eggs were deposited over a greater area of the canes until the entire cane and its branches were black with eggs (Fig. 5). This was an extremely large infestation, and a leaf measuring l-l/Z inches wide and 3 inches long contained 370 aphids. Quantitative Studies (Fig. 6) Wild Red Raspberry. 1961 The numbers of this species varied erratically during July ahd early August. On August 9 the population began to build up and reached its peak on Augus: 14. The numbers of this species then began a fairly steady decline that lasted until the first week of September. On September 14 the population increased again and maintained itself at this level until the following week. On September 27 the population had begun to decrease. This decrease lasted until October 18 when the POPUIation recovered slightly. After October 24 the numbers began to ,- T r) .. .,. dwindle. No aphids were collected on November 7 cfld only 4 mere Luke“ the following week. 34 Fig. 5.--A2his rubifolii (Thomas). Oviparous females and eggs on wild red raspberry. 35 Wild red raspberry, 1961 Maxwell. Black raspberry 6O - lOO 30‘l ‘ 1‘ 601 Wild red raspberry, 1962 160 Walle. Black raspberry 30‘ 1500. Wild red raspberry l9 3 60“1 Walle. Red raspberry 900- 300. 50 'l 30-1 251 I ' I I 7 60‘ Monte. Black raspberry 607 Vaughn. Black raspberry 30- 30- 100 Johnson. Black raspberry 60‘ Vaughn Red raspberry 30- Jun Jul Aug Sep Oct Nov Jun Jul Aug Sep Oct Nov F’s. 6.-—Seasonal population trends of Aphis rubifolii (Thomas) on raspberry. 36 Wild Red Raspberry. 1962 On May 25 a few apterous females and iymphs were taken. The population increased the following week, but no specimens were found on June 12. On June 22 the population abruptly reached its peak and then began to decline. This species maintained itself at low levels for the rest of the summer. No specimens were found in the samples of July 6 and 27 or those of August 3, 10 and 31. The population increased sharply to a minor peak on September 7. During the following week the numbers declined, then began a slow build up that lasted until the end of October. In early November the population began to dwindle. Only two specimens were found on the last day of collecting. Wild Red Raspberry. 1963 A number of apterous females and nymphs were taken on the firs& day. The population of this species declined sharply on June 19, but toward the end of the month and early in July the numbers be an to build up. On July 10 the numbers decreased again but recovered somewhat *he following week. On July 24 the population had again decreased, but in the following week a build up began that resulted in the largest numbers ever observed in this Stand or in any of the te5t plo’s of cultivated raspberry. The first population peak occurred on August 14; this was followed by a sharp drop in numbers, but on August 28 the population began to increase again. In an effort to cope with the large number of specimens collected, the author attempted to berlese the leaf samples with heat and chemicals. It was found that most of the specimens were killed rather than driven from the leaves, and accurate counts could not be made. As a result data from leaf samples collected on September 4, 11 and 18 are no: available. On September 11, five viviparous females and two nymphs were taken from cane samples, but these were not included in the weekly counts. . The following week to V1 The second peak occurred on September the population fell sharply bu; began to build up again on October 9. On October 16 the population peaked for the third time. in the following two weeks it began to decline although it continued to main- tain itself in large numbers. On November 6, the last day of collecting, the population had dropped sharply. Black Raspberry. Monte Farm. 1963 A single viviparous female was found on June 8. This species was no: taken again until July 13 when the population reached its peak. The following week the numbers declined, but on July 27 the population reached another peak. During August the numbers declined steadily; no specimens were found from August 10 until September 28 when viviperous females were collected. Black Raspberry. Johnson Farm. 1963 The first specimens were taken on June 8. This species was not found again until June 29 when the population increased sharply. The following week the numbers of this species declined abruptly but began to increase again on July 13. After that date this species began a steady decline that terminated on August 10. On August 17 there was a sharp increase in numbers, and the population reached its peak. The following week there was a sudden decline in numbers, and no specimens 38 were found. The population began to build up slightly during the last of August and the first week of September; on September 14 no specimens were taken. he following week a few specimens were collected, but on September 28, the last day of collecting, only one nymph was found. Black Raspberry. Maxwell Farm. 1963 The first specimens of A. rubifolii were not found until June 29. The numbers begun to increase but declined affer July 6, and this ! t species m intuined itself at a low level for the rest of the month. No specimens were found on July 27 or August 10; on August 17 the numbers increased but diminished on August 24. In the last of August, this species began an increase that culminuted in a populdtion peak on September 7. On the following week the population declined sharply, bu: on September 21 the populution :gein climbed to s::ain the second peak of the sensor. The numbers dropped to zero on September 28. Black Raspberry. Wulle Farm. 1963 On Jure 8 only nymphs of this species vere taken. The populution remained fairly steady un:il June 29 when it increased abruptly and attained the first peak of the season. The population declined sharply the following week, bu; on July 13 it uinn rose sharply. A second peak occurred on July 20 which was followed by e steady decline thnt con- tinued into the first part of August. On August 17, the populatiom increased sharply to nttain the third und highest peak of :he season. On Augus: 24 the numbers 5nd declired rupidly, and no specimens were found, Following this decline, tfle population increased rapidly, end the fourth peak of the season was realized on August 29. Tris species maintained itself at this level, diminishing only a little on September 7, then it began a decline. On September 21 the population recovered a little and continued more or less at this level for the remainder of the collecting season. Red Raspberry. Walle Farm. 1963 A. rubifolii was not found in high numbers in this field. The first Specimens were collected on July 6 when the small population reached its peak; the numbers decreased sharply in the following week and on July 20 no specimens were found. The next collection of this aphid was made on August 29, when a single apterous female was takei. Black Raspberry. Vaughn Farm. 1963 The first specimens of A. rpbifglii were taken on June 8. The population built up quickly, reached its peak on June 22, and began a decline that lasted for two weeks. On July 6 no specimens were col- lected; then the numbers began a slow increase, but after July 20 the numbers again declined. No specimens were collected on August 10, but on August 17 the population increased sharply and attained the second peak of the year. In the following week the numbers decreased sharply and no specimens were taken on August 24. During the last days of August the population began a steady increase that continued until September 28. On this day the population of this species had reached a third peak that was second in magnitude to the peak that occurred in mid-June. Red Raspberry. Vaughn Farm. 1963 The first specimens were found on June 8; the population rose p (.t rapidly and reached a peak on June 22. In the following week the numbers declined rapidly but recovered slightly on July 6. No specimens were found on July 13, but on the following week the population began to recover. On July 27 the numbers increased and a second population peak occurred; after this date the numbers decreased, and no other specimens were taken from August 10 to August 29. On September 7 the population decreased, then began to build up during the last of September. On September 28, only nymphs were present in the samples. Mas naphis rubicola (Oestlund) This species was never found in abundance. It was collected from both wild and cultivated black and red raspberry. Because fl. Egbicola was encountered infrequently, it is difficult to make any statements concerning its habits. In most cases when apierous forms were found, the author did not recognize the species until the specimens had been mounted on slides. Quite frequently in- dividual specimens were found in collections of A. rubi. This would tend to substantiate the author's opinion that this species also tends to feed on the terminal portion of the host plant. Very few large colonies of this aphid were ever found. Whether this is due to its relative scarcity in Michigan or to some innate characteristic is not known. When this species was recognized on the plant, it was observed to be only moderately active and not prone to drop from the plant when disturbed. _L\ r—u Quantitative Studies Wild Red Raspberry. 1961 A few apterous females were collected on September 27. On October 18 oviparous females and immature males were found. Wild Red Raspberry. 1962 Large numbers of M. rubic l appeared suddenly on September 21. C 1' The population then began a steady decline. The last specimen was col- lected on October 12. Wild Red Raspberry and Commercial Fields. 1963 H. '3 This species was not found in the quantitative samples taken Liosomsphis sp. In October of 1963 a large infestdtion of this aphid occurred in the test plot of wild red raspberry. A number of these aphids were transferred to potted wild red raspberry plants. These plants were placed in a grOch chamber thut was regulated to give e 12 hour day with temperatures ranging from 73 to 800 F. and relative humidity from 55 to 607. These colonies fdiled to maintain themselves, and all specimens were dead within seven dnys. As the only recorded hos: of this genus are members of Gremineae, it is presumed that these aphids crawled to the raspberry plants from the tall grasses that were growing in this 1010?. ‘ 0 Determination by Miss Louise Russell. L\ N Data from the quantitative collecting si:es, showing the rela— tive abundance and the preferred feeding locations of the aphid species, are summarized in Tebles 2 and 3. The time of occurrence and duration of the morphological forms of the raspberry aphids, compiled from data gathered during four years of collecting, are presented in Table 4. No great numbers of predators, parasitized or fungus infected aphids, were observed by the author. 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