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Alate aphid (Hemiptera: Aphididae) species composition and richness in Northeastern USA snap beans and an update to historical lists.

Aphids are a small, but diverse group of insects with an origin in the Jurassic and a total of 4800 species world-wide (Grimaldi & Engel 2005; Dixon 1985a; Dixon 1985b). They are primarily phloem feeders and when present in high densities can damage their host plant. Aphids excrete excess carbohydrates from their diet of phloem sap, providing a nutrient-rich substrate for sooty mold fungi to grow. Sooty mold can be a major problem in a number of agricultural crops because the mold can either render produce unmarketable or reduce plant quality of the commodity. Aphids are also important vectors of viruses that can kill their host plant or substantially reduce crop yield and quality (Agrios 2005). Some viruses are transmitted by aphids in a non-persistent, stylet-borne manner. They are obtained quickly by their aphid vector during short tasting probes, adhere to the stylet lining by binding to helper component proteins or directly to the stylet, and remain there until they are flushed out during another tasting probe (Ng & Falk 2006). Nonpersistently transmitted viruses can be vectored by alates, sometimes by multiple aphid species (Gildow et al. 2008) regardless of whether or not there is reproduction on the plant host, and the epidemiology of these viruses can be influenced heavily by the alate aphid community.

Several viruses of this type have been introduced recently, or increased in frequency, in the northeastern U.S. One, plum pox virus (PPV), threatened the stone fruit industry following its arrival in the U.S. This virus causes sharka disease in parts of Europe and South America where it is endemic (Roy and Smith 1984; Rosales et al. 1998). Type D isolates were detected in the U.S. in Pennsylvania in 1999 (Damsteegt et al. 2001), and surveillance and eradication efforts of this invasive species included destruction of approximately 23% of the non-cherry stone fruit orchards of Pennsylvania (Wallis et al. 2005). As part of these efforts, studies were conducted to determine the potential aphid species that might serve as reservoir or route of transmission in the region where this virus was first detected (Wallis et al. 2005). Soon thereafter, in the early 2000s, Northeastern and Midwestern U.S., snap bean crops (Phaseolus vulgaris L; Fabales: Fabaceae.) had viruslike symptoms (leaf mosaic and blistering, deformed pods) and experienced dramatic yield loss (Larsen et al. 2002). Among the viruses detected were alfalfa mosaic virus, bean common mosaic virus, bean pod mottle virus, bean yellow mosaic virus, clover yellow mosaic virus, clover yellow vein virus (ClYVV), cucumber mosaic virus (CMV), tobacco streak virus and white clover mosaic virus (Grau et al. 2002; Larsen et al. 2002; Shah et al. 2006). CMV was the most prevalent virus detected in these snap bean fields (Larsen et al. 2002; Shah et al. 2006). As is the case with PPV, CMV is transmitted by aphids in a non-persistent, stylet-borne manner (Nault 1997). CMV-infected plants were often found in clumps in snap bean fields, which were consistent with aphid-initiated virus epidemics (Shah et al. 2005). CMV epidemics also occurred more frequently in New York than in Pennsylvania. The CMV epidemics coincided with the appearance of a newly invasive aphid, Aphis glycines Matusmura (Nault et al. 2009), and as was the case with stone fruit, the threat of viral epidemics led to extensive surveys of the alate aphids species composition in the affected crop.

These recent surveys of aphid collected from snap bean fields in Pennsylvania and New York, and peach orchards in Pennsylvania, were quite extensive. Also from Pennsylvania, J. O. Pepper specialized in aphid identification and actively collected them for most of the 20th century. His collections centered at his home in central Pennsylvania (State College) and included much of the surrounding forest and farmland. The bulk of his collection is housed in the Frost Entomological Museum (University Park, Pennsylvania), and he also contributed slides to the United States National Collection (Beltsville, Maryland). Pepper (1965) reported 345 species in a published list of the aphids of Pennsylvania and their host plants. To date, this is the most comprehensive published list of aphids for the state. However, since taxonomy and systematics are in flux, the names that Pepper published are currently out of date and in need of revision.

The purpose of this study was to identify the species composition and estimate aphid species richness in snap bean agroecosystems in the northeastern states from field surveys, and generate a current list of aphid species in this region using field survey data, literature, and an examination of the J. O. Pepper aphid collection.

MATERIALS AND METHODS

Detailed methods for alate aphid collection in snap bean fields in Pennsylvania and New York were published in Nault et al. (2009). To summarize, we used water pan traps baited with a green ceramic tile (Webb et al. 1994) and filled with a 20% propylene glycol solution in snap bean fields in both states from 2002-2006 in NY and 2004-2006 in PA. Traps were installed in a total of 56 fields in western NY (12 each yr, except for 2004 which had 8 fields) and 18 fields in Centre county PA (6 each yr). The traps in Centre County formed an approximately 40 mile transect in the southern portion of the county roughly following state routes 45 and 192. The traps were checked weekly for aphids from the early trifoliate stage (early to mid Jul) until field harvest. Collection methods in the peach (Prunus persica (L.) Stokes; Rosales: Rosaceae) orchard are documented in Wallis et al (2005), and also used the water pan traps baited with a green tile. Trapping occurred during 2 yr in 2 orchards in central Pennsylvania.

For both studies, aphids were removed from pan traps and then stored in 70% ethanol (EtOH), then transferred to potassium hydroxide and heated for 1 h or until clear. Cleared aphids were rinsed for 10 min each in a sequence of 95% EtOH, absolute EtOH, and clove oil. Once rinsed, each aphid was placed on a drop of Canada balsam on a glass slide and positioned to expose diagnostic features before a coverslip was placed on top. Aphids collected in New York were identified by R. Eckel (RVWE Consulting, Frenchtown, New Jersey), whereas those from Pennsylvania were identified by W. Sackett and A. Bachmann using keys by Smith et al. (1992) and Blackman & Eastop (2000). Voucher specimens are located at the New York State Agricultural Experiment Station in Geneva, New York, and the Department of Entomology, Pennsylvania State University, University Park, Pennsylvania.

Species rarefaction curves were calculated for the Pennsylvania and New York collections individually and for both states combined using EstimateS (Colwell 2005).

A complete list of aphids from Pennsylvania was compiled using the J. O. Pepper Aphid Slide Collection, which is housed at the Frost Entomological Museum (University Park, Pennsylvania), as well as species recorded in Pepper (1965). We searched the slide collection in addition to using Pepper (1965) because Pepper continued to collect aphids and make slides into the late 1980s, but did not publish any updates to his original 1965 paper. Because the collection and Pepper (1965) contained aphid species names from the early 20th century, we consulted 2 online aphid databases to ensure that the final list used the most current nomenclature (Aphid Species File - http://aphidspeciesfile. org, accessed 22-IV-2012 and Aphids on the World's Plants - http://www.aphidsonworldsplants.info/, accessed l-XI-2013). We combined our findings from the Pepper collection material with the results of our pan trapping study and Wallis et al. (2005) to create a more current list of the aphids of Pennsylvania (Tables 3-11).

RESULTS

In snap bean fields in New York and Pennsylvania, a total of 8,821 aphids were identified, with 7,484 from New York and 1,337 from Pennsylvania. A total of 97 species were caught; 71 from New York and 41 from Pennsylvania. We were unable to identify only 254 (2.8%) of the aphids. Of the aphids captured, those species representing 1% or greater of the total number caught in either state are listed in Table 1 (originally published in Nault et al. 2009) with their abundances. A comprehensive list of all aphid species found in Pennsylvania and New York snap bean fields is shown in Table 2 along with their host associations based on Blackman & Eastop (1994, 2000, and 2006). From this host information we estimated that 61 percent of the species dispersing through snap bean fields in both states were most likely coming in from the surrounding forests as their hosts are woody, not herbaceous, species (Fig. 1).

Species accumulations followed asymptotic patterns (Fig. 2) suggesting reasonably adequate sampling of the aphid species present as alates in commercial snap bean fields. Overall, there were fewer aphids collected in Pennsylvania, but based on the rarefaction curve there were a similar number of total species represented in a sample of the same number of individuals (Figs. 2 and 3, at 1,250 individuals there would be 45 species sampled in Pennsylvania and 50 in New York). Based on the historical collections reported by Pepper, there are approximately 350 aphid species in Pennsylvania. Historical reports in Leonard (1963) suggest that there are approximately 430 aphid species in New York.

Combining the list of aphids collected from snap bean fields, peach orchards and those published by J. O. Pepper in 1965, we developed a new, more comprehensive list of the aphids present in Pennsylvania. We found 7 species present in our collections that were not present in the slide collection housed in the Frost Entomological Museum (University Park, Pennsylvania) or published in Pepper (1965) (Table 3). One of these aphids, Aphis glycines Matsumura, was introduced to the US around the turn of the 21st century and is now widespread throughout the Midwest, Northeast and southeastern Canada (Ragsdale et al. 2011).

DISCUSSION

Our passive trapping in snap bean fields alone yielded a surprisingly high percentage of the species present throughout Pennsylvania and New York (~14% and ~18% respectively). Our sampling method concentrated on only one habitat (commercial snap bean fields), but did intercept aphids moving from the surrounding forests and hedgerows. The high degree of landscape heterogeneity and crop diversity in the trapping areas includes plant species that serve as hosts for many of the aphid species that represented less than 1% of the total capture (Pfleeger et al. 2006). These aphids were captured in very small numbers (mostly singletons), and are not important contributors to the plant virus epidemics reported by Wallis et al. (2005) and Nault et al. (2009).

Of the aphids we captured, 2 species were especially notable; Therioaphis trifolii Monell, which comprised 31.8% of the identified aphids, and A. glycines which represented 18.2 % of the identified aphids. Both of these aphids were introduced to North America (A. glycines from Asia and T. trifolii from Europe) and were quite destructive to crops immediately after their introduction (in soybean and alfalfa, respectively). Aphis glycines continues to cause significant economic damage in soybean (Ragsdale et al. 2011). While not known to colonize Phaseolus spp., both species are competent vectors of the legume strain of CMV (Gildow et al. 2008).

The intermittent appearance of CMV in central Pennsylvania snap bean crops could be influenced by a unique agricultural landscape. Agricultural fields are located in valleys bordered by the low, but steep, forested ridges of the Appalachian Mountains. The ridge and valley system might be acting like a barrier, keeping CMV out for most of the season. We did not search for a CMV reservoir outside of testing a few alfalfa fields, which were also negative for CMV. It is possible, that much like our A. glycines population, legume strains of CMV may be transient. If this is the case, migrating aphids may be scrubbed of virions when they land in one of the many bordering forests containing many non-host plants.

The Pepper (1965) aphid list in addition to the Pepper slide collection allowed us to compile a comprehensive list of the aphids present in Pennsylvania, but the nomenclature was in need of updating. Our efforts to update the nomenclature, and incorporate our more recent sampling efforts, resulted in a modern list of aphids of Pennsylvania that includes recently introduced species.

Caption: Fig. 2. Individual-based rarefaction curves showing aphid species accumulation in Pennsylvania and New York.

Caption: Fig. 3. Individual-based rarefaction curve showing aphid species accumulation from the combining of samples from PA and NY (solid line) and the 95% confidence intervals for the curve (dashed lines).

ACKNOWLEDGEMENTS

The authors would like to thank Fred Gildow, William Sackett and Dana Roberts. Funding for this work was provided by the Pennsylvania Vegetable Growers Association.

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AMANDA C. BACHMANN (1), BRIAN A. NAULT (2) AND SHELBY J. FLEISCHER (3) *

(1) South Dakota State University Department of Plant Science, 412 W Missouri Ave, Pierre, SD 57501, USA

(2) Cornell University Department of Entomology, New York State Agricultural Experiment Station, 630 W. North Street, Geneva, NY 14456, USA

(3) Penn State Department of Entomology, 501 ASI, University Park, PA 16802, USA

* Corresponding author; E-mail: shelby.fleischer@gmail.com

TABLE 1. ALATE APHID SPECIES REPRESENTING > 1 % OF THE CAPTURE FROM
WATER PAN TRAPS IN COMMERCIAL SNAP BEAN FIELDS IN PENNSYLVANA
(2004-2006) AND NEW YORK (2002-2006). DERIVED FROM TABLE 1 IN NAULT
ET AL. (2009).

                       New York            Pennsylvania

Species                Total    Percent    Total   Percent
                                of Total           of Total

Therioaphis            2,274      30.4      535      40.0
  trifolii (Monell)
Aphis glycines         1,475      19.7      131      9.8
  Matsumura
Acyrthosiphon pisum    1,106      14.8      28       2.1
  (Harris)
Rhopalosiphum           685       9.2       75       5.6
  maidis (Fitch)
Pemphigus               239       3.2        0       0.0
  populieaulis Fitch
Aphis eraeeivora        179       2.4       123      9.2
  Koch
Aphis gossypii          130       1.7       201      15.0
  Glover
Hayhurstia              128       1.7        1       0.1
  atriplieis (L.)
Lipaphis                128       1.7        0       0.0
  pseudobrassieae
  (Davis) *
Myzus persicae           97       1.3       26       1.9
  (Sulzer)
Capitophorus             79       1.1        7       0.5
  eleagni (Del
  Guerica)
Aphis sp.                77       1.0        0       0.0
Rhopalosiphum padi       77       1.0       45       3.4
  (L.)
Aphis fabae Scopoli      15       0.2       14       1.0
Anoecia sp.              1        <0.1      14       1.0
Braehyeaudus             2        <0.1      15       1.1
  persicae
  (Passerini)
Unknown                 216       2.9       38       2.8
Others                  576       7.7       84       6.3
Total                  7,484     100.0     1,337    100.0

                       Overall

Species                Total   Percent
                               of Total

Therioaphis            2809      31.8
  trifolii (Monell)
Aphis glycines         1606      18.2
  Matsumura
Acyrthosiphon pisum    1134      12.9
  (Harris)
Rhopalosiphum           760      8.6
  maidis (Fitch)
Pemphigus               239      2.7
  populieaulis Fitch
Aphis eraeeivora        302      3.4
  Koch
Aphis gossypii          331      3.8
  Glover
Hayhurstia              129      1.5
  atriplieis (L.)
Lipaphis                128      1.5
  pseudobrassieae
  (Davis) *
Myzus persicae          123      1.4
  (Sulzer)
Capitophorus            86       1.0
  eleagni (Del
  Guerica)
Aphis sp.               77       0.9
Rhopalosiphum padi      122      1.4
  (L.)
Aphis fabae Scopoli     29       0.3
Anoecia sp.             15       0.2
Braehyeaudus            17       0.2
  persicae
  (Passerini)
Unknown                 254      2.9
Others                  660      7.5
Total                  8,821    100.0

* Published in Nault et al (2009) as Lipaphis erysimi (Kaltenbach).

TABLE 2. SPECIES OF ALATE APHIDS WITH HOST ASSOCIATIONS, COLLECTED
FROM WATER PAN TRAPS IN COMMERCIAL SNAP BEAN FIELDS IN PENNSYLVANIA
(2004-2006) AND NY (2002-2006), AND FROM SIMILAR TRAPS IN PEACH
ORCHARDS IN CENTRAL PENNSYLVANIA (2003-2004, WALLIS ET AL. 2005).
PRIMARY AND SECONDARY HOST PLANT ASSOCIATIONS FOR NORTH AMERICA TAKEN
FROM BLACKMAN & EASTOP (1994 [AWT], 2000 [AWC], AND 2006 [HPS]).

                                Crop Sampled

                                Snap bean  Peach

Aphid Species                   PA    NY    PA

Acyrthosiphon kondoi Shinji      *    *
Acyrthosiphon pisum Harris       *    *      *
Amphorophora rubi Kaltenbach     *    *
Anoecia corni (Fabricius)        *    *      *
Anoecia cornicola (Walsh)        *
Anoecia oenotherae Wilson                    *
Aphis carduella Walsh            *    *
Aphis cephalanthi Thomas              *
Aphis craccivora Koch            *    *
Aphis fabae Scopoli              *    *      *
Aphis forbesi Weed                    *
Aphis glycines Matsumura         *    *
Aphis gossypii Glover            *    *      *
Aphis hederae Kaltenbach              *
Aphis lugentis Williams                      *
Aphis (Protaphis)                *           *
  middletonii Thomas
Aphis nasturtii Kaltenbach                   *
Aphis oestlundi Gillette         *
Aphis pomi DeGeer                     *
Aphis pulchella Hottes                *      *
  & Frison
Aphis rubifolii (Thomas)              *
Aphis rumieis L.                      *
Aphis spiraecola Patch                *      *
Aphis viburniphila Patch         *    *
Aulacorthum solani               *    *
  (Kaltenbach)
Brachycaudus (Scrophulaphis)          *      *
  persicae group
Brevieoryne brassicae (L.)            *      *
Capitophorus elaeagni            *    *      *
  (del Guercio)
Capitophorus hippophaes          *    *
  (Walker)
Carolinaia (Glabromyzus)                     *
  rhois (Monell)
Chaitophorus populifolii         *           *
  Essig
Cinara atlantica (Wilson)        *
Drepanaphis aeerifoliae               *
  (Thomas)
Drepanaphis carolinensis              *
  Smith
Drepanaphis nigricans Smith           *      *
Drepanaphis sabrinae Miller                  *
Drepanosiphum platanoidis             *
  (Schrank)
Dysaphis (Pomaphis)                   *
  plantaginea (Passerini)
Dysaphis tulipae (Boyer                      *
  de Fonscolombe)
Eriosoma lanigerum                           *
  (Hausmann)
Essigella pini (Wilson)          *    *
Eulachnus rileyi (Williams)           *      *
Geoica squamosa Hart                  *
Hayhurstia atriplicis (L.)       *    *
Hyadaphis foeniculi              *
  (Passerini)
Hyalopterus pruni (Geoffroy)          *
Hyperomyzus lactucae (L.)             *      *
Hysteroneura                          *
  setariae (Thomas)
Illinoia liriodendri                  *
  (Monell)
Kaltenbachiella ulmifusa         *
  (Walsh & Riley)
Lipaphis pseudobrassicae              *      *
  (Davis)
Macrosiphoniella                 *
  ludovicianae (Oestlund)
Macrosiphoniella sanborni        *
  (Gillette)
Macrosiphum euphorbiae           *    *      *
  (Thomas)
Macrosiphum pallidum             *    *
  (Oestlund)
Macrosiphum (Neocorylobium)           *
  pseudocoryli (Patch)
Macrosiphum rosae (L.)                *
Melaphis rhois (Fitch)           *
Monellia caryella (Fitch)        *    *      *
Myzus (Nectarosiphon)            *    *      *
  persicae (Sulzer)
Myzocallis sp.                        *
Nearctaphis bakeri (Cowen)       *    *      *
Nearctaphis clydesmithi               *      *
  Hille Ris Lambers
Nearctaphis crataegifoliae       *    *      *
  (Fitch)
Ovatus crataegarius (Walker)          *
Pemphigus populieaulis Fitch          *      *
Pemphigus populitransversus           *      *
  Riley
Pemphigus populivenae Fitch           *      *
Periphyllus americanus Baker                 *
Periphyllus testudinaceus             *
  (Ferni)
Phorodon humuli (Schrank)             *
Prociphilus (Meliarhizophagus)               *
  fraxinifolii (Riley)
Pterocomma bicolor (Oestlund)                *
Pterocomma smithiae (Monell)                 *
Rhodobium porosum (Sanderson)    *    *
Rhopalomyzus poae (Gillette)          *      *
Rhopalosiphoninus latysiphon          *
  (Davidson)
Rhopalosiphum maidis (Fitch)     *    *      *
Rhopalosiphum nymphaeae (L.)                 *
Rhopalosiphum oxyacanthae        *    *
  (Davidson)
Rhopalosiphum padi (L.)          *    *      *
Rhopalosiphum rufiabdominale     *    *
  (Sasaki)
Schizaphis graminum (Rondani)    *    *
Sipha flava (Forbes)                  *      *
Sitobion avenae (Fabricius)      *    *
Tetraneura nigriabdominalis           *      *
  (Sasaki)
Therioaphis (Rhizoberlesia)           *
  riehmi (Borner)
Therioaphis (Pterocallidium)     *    *      *
  trifolii (Monell)
Uroleucon (Lambersius)           *
  anomalae (Hottes & Frison)
Uroleucon pseudambrosiae              *
  (Olive)
Utamphorophora crataegi               *
  (Monell)
Vesiculaphis carieis                  *
  (Fullaway)

Aphid Species                   Primary Host for Aphid

Acyrthosiphon kondoi Shinji     Leguminosae, Trifoleae, Loteae
Acyrthosiphon pisum Harris      Leguminosae, Genistae,
                                  Trifoleae, Fabae, Hedysareae
Amphorophora rubi Kaltenbach    Rubus spp.
Anoecia corni (Fabricius)       Cornus sanguinea
Anoecia cornicola (Walsh)       Comus spp.
Anoecia oenotherae Wilson       Cornus spp.
Aphis carduella Walsh           Cornus stolonifera
Aphis cephalanthi Thomas        Cephalanthus occidentalis
Aphis craccivora Koch           polyphagous, Leguminosae
Aphis fabae Scopoli             Euonymous europaeus,
                                  Viburnum opulus
Aphis forbesi Weed              Fragaria spp.
Aphis glycines Matsumura        Rhamnus spp.
Aphis gossypii Glover           Catalpa, Hibiscus, Celastrus,
                                  Rhamnus, Punica
Aphis hederae Kaltenbach        Hedera helix, Araliaceae,
                                  Cuscuta
Aphis lugentis Williams         Senecio spp., Erigeron sp.
Aphis (Protaphis)               Compositae, Cruciferae,
  middletonii Thomas              Umbelliferae, Grami-nae
Aphis nasturtii Kaltenbach      Rhamnus cathartica, R. alnifolia
Aphis oestlundi Gillette        Oenothera biennis
Aphis pomi DeGeer               Pyroidea
Aphis pulchella Hottes          Euphorbia
  & Frison
Aphis rubifolii (Thomas)        Rubus spp.
Aphis rumieis L.                Rumex spp., Rheum spp.
Aphis spiraecola Patch          Citrus, Spiraea spp., polyphagous
Aphis viburniphila Patch        Viburnum spp.
Aulacorthum solani              Polyphagous
  (Kaltenbach)
Brachycaudus (Scrophulaphis)    Prunus persica, P. armeniaca
  persicae group
Brevieoryne brassicae (L.)      Cruciferae
Capitophorus elaeagni           Elaeagnus spp.
  (del Guercio)
Capitophorus hippophaes         Elaeagnaceae
  (Walker)
Carolinaia (Glabromyzus)        Rhus glabra, R. typhina
  rhois (Monell)
Chaitophorus populifolii        Populas spp.
  Essig
Cinara atlantica (Wilson)       Pinus spp.
Drepanaphis aeerifoliae         Acer saccharinum, A. rubrum,
  (Thomas)                        A. saccharum
Drepanaphis carolinensis        Acer saccharum, A. rubrum
  Smith
Drepanaphis nigricans Smith     Acer rubrum
Drepanaphis sabrinae Miller     Acer saccharum
Drepanosiphum platanoidis       Acer pseudoplatanus, Acer
  (Schrank)                       spp., sycamore
Dysaphis (Pomaphis)             Malus spp., Pyrus
  plantaginea (Passerini)
Dysaphis tulipae (Boyer         many monocots
  de Fonscolombe)
Eriosoma lanigerum              Pyroidea, apple, Crataegus,
  (Hausmann)                      Coloneaster
Essigella pini (Wilson)         Pinus spp.
Eulachnus rileyi (Williams)     Pinus spp.
Geoica squamosa Hart
Hayhurstia atriplicis (L.)      Chenopodiaceae, Atriplex,
                                  Chenopodium spp.
Hyadaphis foeniculi             Lonicera spp.
  (Passerini)
Hyalopterus pruni (Geoffroy)    Prunus domestica, P. armeniaca
Hyperomyzus lactucae (L.)       Ribes spp.
Hysteroneura                    Prunus domestica
  setariae (Thomas)
Illinoia liriodendri            Liriodendron tulipifera
  (Monell)
Kaltenbachiella ulmifusa        Ulmus rubra
  (Walsh & Riley)
Lipaphis pseudobrassicae        Cruciferae
  (Davis)
Macrosiphoniella                Artemisia ludoviciana,
  ludovicianae (Oestlund)         A. vulgaris
Macrosiphoniella sanborni       Dendranthema indicum, morifolium,
  (Gillette)                      frutes-cens, Compositae
Macrosiphum euphorbiae          Rosa spp
  (Thomas)
Macrosiphum pallidum            Rosaceae, Rosa spp.
  (Oestlund)
Macrosiphum (Neocorylobium)     Ostrya virginiana, Corylus spp.
  pseudocoryli (Patch)
Macrosiphum rosae (L.)          Rosa spp.
Melaphis rhois (Fitch)          Rhus spp. (glabra, typhina)
Monellia caryella (Fitch)       Carya spp.
Myzus (Nectarosiphon)           Prunus persica, Prunus spp.
  persicae (Sulzer)
Myzocallis sp.                  Fagaceae
Nearctaphis bakeri (Cowen)      Crataegus, Cydonia, Malus, Pyrus
Nearctaphis clydesmithi         Crataegus
  Hille Ris Lambers
Nearctaphis crataegifoliae      Crataegus spp
  (Fitch)
Ovatus crataegarius (Walker)    Crateagus spp.
Pemphigus populieaulis Fitch    Populus deltoides, P. tremuloides
Pemphigus populitransversus     Populus spp.
  Riley
Pemphigus populivenae Fitch     Populus spp.
Periphyllus americanus Baker    Acer spp.
Periphyllus testudinaceus       Acer spp., Aesculus spp.
  (Ferni)
Phorodon humuli (Schrank)       Prunus spp.
Prociphilus (Meliarhizophagus)  Fraxinus spp.
  fraxinifolii (Riley)
Pterocomma bicolor (Oestlund)   Populus spp., Salix spp.
Pterocomma smithiae (Monell)    Populus spp., Salix spp.
Rhodobium porosum (Sanderson)
Rhopalomyzus poae (Gillette)    Lonicera alpigena
Rhopalosiphoninus latysiphon    bulbs (Tulipa, Gladiolus),
  (Davidson)                      runners
Rhopalosiphum maidis (Fitch)    Gramineae
Rhopalosiphum nymphaeae (L.)    Prunus spp.
Rhopalosiphum oxyacanthae       Alus, Pyrus, Cotoneaster,
  (Davidson)                      Crataegus, Sorbus
Rhopalosiphum padi (L.)         Prunus virginiana
Rhopalosiphum rufiabdominale    Prunus spp.
  (Sasaki)
Schizaphis graminum (Rondani)   Gramineae
Sipha flava (Forbes)            Gramineae
Sitobion avenae (Fabricius)     Gramineae
Tetraneura nigriabdominalis     Ulmus spp.
  (Sasaki)
Therioaphis (Rhizoberlesia)     Melilotas spp.
  riehmi (Borner)
Therioaphis (Pterocallidium)    Leguminoseae
  trifolii (Monell)
Uroleucon (Lambersius)          Aster novaeangliae
  anomalae (Hottes & Frison)
Uroleucon pseudambrosiae        Compositae, Lactuca spp.
  (Olive)
Utamphorophora crataegi         Crataegus spp.
  (Monell)
Vesiculaphis carieis            Rhododendron spp.
  (Fullaway)

Aphid Species                   Secondary Host for Aphid   Source

Acyrthosiphon kondoi Shinji                                 AWC
Acyrthosiphon pisum Harris                                  AWC
Amphorophora rubi Kaltenbach                                HPS
Anoecia corni (Fabricius)       Gramme ae                   AWT
Anoecia cornicola (Walsh)       Gramme ae                   AWT
Anoecia oenotherae Wilson       Oenothera biennis           AWT
Aphis carduella Walsh           Umbelliferae                AWT
Aphis cephalanthi Thomas                                    HPS
Aphis craccivora Koch                                       AWC
Aphis fabae Scopoli             polyphagous                 AWC
Aphis forbesi Weed                                          HPS
Aphis glycines Matsumura        Glycine max                 AWC
Aphis gossypii Glover           polyphagous, cotton,        all
                                  cucurbits
Aphis hederae Kaltenbach                                    HPS
Aphis lugentis Williams                                     HPS
Aphis (Protaphis)                                           AWC
  middletonii Thomas
Aphis nasturtii Kaltenbach      wide range                  HPS
Aphis oestlundi Gillette                                    HPS
Aphis pomi DeGeer                                           AWC
Aphis pulchella Hottes                                      HPS
  & Frison
Aphis rubifolii (Thomas)                                    HPS
Aphis rumieis L.                                            HPS
Aphis spiraecola Patch                                      AWC
Aphis viburniphila Patch                                    HPS
Aulacorthum solani                                          HPS
  (Kaltenbach)
Brachycaudus (Scrophulaphis)    Scrophulariaceae            AWT
  persicae group
Brevieoryne brassicae (L.)                                  AWC
Capitophorus elaeagni           tubuliferous Compositae     AWT
  (del Guercio)
Capitophorus hippophaes         Polygonum spp.,             AWT
  (Walker)                        Persicaria spp.
Carolinaia (Glabromyzus)        Gramineae                   AWC
  rhois (Monell)
Chaitophorus populifolii                                    AWT
  Essig
Cinara atlantica (Wilson)                                   AWT
Drepanaphis aeerifoliae                                     AWT
  (Thomas)
Drepanaphis carolinensis                                    AWT
  Smith
Drepanaphis nigricans Smith                                 AWT
Drepanaphis sabrinae Miller                                 AWT
Drepanosiphum platanoidis                                   AWT
  (Schrank)
Dysaphis (Pomaphis)             Plantago spp.               AWT
  plantaginea (Passerini)
Dysaphis tulipae (Boyer                                     AWC
  de Fonscolombe)
Eriosoma lanigerum                                          AWC
  (Hausmann)
Essigella pini (Wilson)                                     AWT
Eulachnus rileyi (Williams)                                 AWT
Geoica squamosa Hart
Hayhurstia atriplicis (L.)                                  HPS
Hyadaphis foeniculi             Umbelliferae spp.           AWC
  (Passerini)
Hyalopterus pruni (Geoffroy)    Phragmites communis,        AWC
                                  Arundo donax
Hyperomyzus lactucae (L.)       Sonchus spp.                AWC
Hysteroneura                    Graminieae                  AWC
  setariae (Thomas)
Illinoia liriodendri                                        AWT
  (Monell)
Kaltenbachiella ulmifusa        Labiatae                    AWT
  (Walsh & Riley)
Lipaphis pseudobrassicae                                    AWC
  (Davis)
Macrosiphoniella                                            HPS
  ludovicianae (Oestlund)
Macrosiphoniella sanborni                                   AWC
  (Gillette)
Macrosiphum euphorbiae          highly polyphagous,         AWC
  (Thomas)                        Solanaceae
Macrosiphum pallidum                                        AWC
  (Oestlund)
Macrosiphum (Neocorylobium)                                 AWT
  pseudocoryli (Patch)
Macrosiphum rosae (L.)          Dipsacaceae                 AWC
Melaphis rhois (Fitch)          mosses                      AWT
Monellia caryella (Fitch)                                   AWC
Myzus (Nectarosiphon)           polyphagous, over           AWC
  persicae (Sulzer)               40 families
Myzocallis sp.                                              AWT
Nearctaphis bakeri (Cowen)      Leguminosae                 AWC
Nearctaphis clydesmithi         unknown                     AWT
  Hille Ris Lambers
Nearctaphis crataegifoliae      Trifolium spp.              AWC
  (Fitch)
Ovatus crataegarius (Walker)    Labiatae esp Mentha         AWT
Pemphigus populieaulis Fitch    unknown                     AWT
Pemphigus populitransversus     Cruciferae                  AWC
  Riley
Pemphigus populivenae Fitch     Chenopodiaceae              AWT
Periphyllus americanus Baker                                AWT
Periphyllus testudinaceus                                   AWT
  (Ferni)
Phorodon humuli (Schrank)       Humulus lupulus (hops)      AWC
Prociphilus (Meliarhizophagus)                              AWT
  fraxinifolii (Riley)
Pterocomma bicolor (Oestlund)                               AWT
Pterocomma smithiae (Monell)                                AWT
Rhodobium porosum (Sanderson)
Rhopalomyzus poae (Gillette)    grasses                     AWC
Rhopalosiphoninus latysiphon                                AWC
  (Davidson)
Rhopalosiphum maidis (Fitch)                                AWC
Rhopalosiphum nymphaeae (L.)    water plants                AWC
Rhopalosiphum oxyacanthae       grasses                     AWC
  (Davidson)
Rhopalosiphum padi (L.)         Gramineae                   AWC
Rhopalosiphum rufiabdominale    Gramineae, Cyperaceae,      AWC
  (Sasaki)                        Solanaceae
Schizaphis graminum (Rondani)                               AWC
Sipha flava (Forbes)                                        AWC
Sitobion avenae (Fabricius)                                 AWC
Tetraneura nigriabdominalis     Gramineae                   AWC
  (Sasaki)
Therioaphis (Rhizoberlesia)                                 HPS
  riehmi (Borner)
Therioaphis (Pterocallidium)                                AWC
  trifolii (Monell)
Uroleucon (Lambersius)                                      HPS
  anomalae (Hottes & Frison)
Uroleucon pseudambrosiae                                    HPS
  (Olive)
Utamphorophora crataegi                                     AWT
  (Monell)
Vesiculaphis carieis            Cyperus spp.                HPS
  (Fullaway)

TABLE 3. NEW APHID RECORDS FROM PENNSYLVANIA REPORTED
IN NAULT ET AL. (2009) AND/OR WALLIS
ET AL. (2005), BUT NOT FOUND IN PEPPER
(1965).

Species              Nault    Wallis
                     et al.   et al.

Acyrthosiphon          *
  kondoi
Aphis glycines         *
Aphis lugentis                  *
Aphis (Protaphis)      *        *
  middletonii
Aphis pulchella                 *
Nearctaphis                     *
  clydesmithi
Tetraneura                      *
  nigriabdominalis

TABLE 4. SPECIES IN SIX SUBFAMILIES OF THE FAMILY APHIDIDAE
OCCURRING IN PENNSYLVANIA.

Subfamily          Tribe            Species

Anoeciinae                          Anoecia corni
                                    Anoecia cornicola
                                    Anoecia oenotherae
                                    Anoecia setariae
                                      Gillette & Palmer
Hormaphidinae      Cerataphidini    Cerataphis lataniae (Boisduval)
                   Hormaphidini     Hamamelistes spinosus Shimer
                                    Hormaphis hamamelidis Fitch
Mindarinae                          Mindarus abietinus Koch
Phyllaphidinae                      Phyllaphis fagi (L.)
                                    Stegophylla quercicola
                                      (Monell)
                                    Stegophylla quercifoliae
                                      (Gillette)
                                    Stegophylla quercina
                                      Quednau
Pterocommatinae                     Fullawaya terricola
                                      (Hottes & Frison)
                                    Plocamaphis flocculosa
                                      Weed
                                    Pterocomma bicolor
                                    Pterocomma medium Baker
                                    Pterocomma populifoliae
                                      (Fitch)
                                    Pterocomma smithiae
Saltusaphidinae    Saltusaphidini   Iziphya flabella (Sanborn)
                                    Iziphya vittata Richards
                                    Strenaphis elongate (Baker)
                   Thripsaphidini   Allaphis verrucosa (Gillette)
                                    Subsaltusaphis virginica (Baker)
                                    Thripsaphis ballii (Gillette)

TABLE 5. SPECIES IN THE SUBFAMILY APHIDINAE, TRIBE
MACROSIPHINI OCCURRING IN PENNSYLVANIA.

Abstrusomyzus phloxae (Sampson)
Acuticauda solidaginifoliae (Williams)
Acyrthosiphon kondoi
Acyrthosiphon lactucae (Passerini)
Acyrthosiphon malvae (Mosley)
Acyrthosiphon pisum
Acyrthosiphon pseudodirhodum (Patch)
Amphorophora agathonica Hottes
Amphorophora ampullata Buckton
Amphorophora rossi Hottes & Frison
Amphorophora rubi
Amphorophora sensoriata Mason
Aulacorthum solani
Brachycaudus (Prunaphis) cardui (L.)
Brachycaudus /ieZic/irysi(Kaltenbach)
Brachycaudus (Scrophulaphis) persicae group
Brachycaudus (Brachycaudina) rociadae (Cockerell)
Brachycaudus (Thuleaphis) rumexicolens (Patch)
Brachycaudus (Appelia) schwartzi (Borner)
Brachycorynella asparagi (Mordvilko)
Brevicoiyne brassicae
Cachryphora canadensis Hille Ris Lambers
Cachryphora serotinae (Oestlund)
Capitophorus carduinus (Walker)
Capitophorus elaeagni
Capitophorus hippophaes
Carolinaia carieis Wilson
Carolinaia (Galbromyzus) howardii (Wilson)
Carolinaia (Glabromyzus) rhois
Catamergus Kickapoo (Hottes & Frison)
Cavariella aegopodii (Scopoli)
Cavariella cicutae (Koch)
Cavariella hendersoni Knowlton & Smith
Cavariella pastinacae (L.)
Cavariella salicis (Monell)
Cavariella theobaldi (Gillette & Bragg)
Ceruraphis eriophori (Walker)
Ceruraphis viburnicola (Gillette)
Chaetosiphon (Pentatrichopus) fragaefolii (Cockerell)
Chaetosiphon (Pentatrichopus) minor (Forbes)
Chaetosiphon (Pentatrichopus) tetrarhodum (Walker)
Coloradoa rufomaculata (Wilson)
Cryptomyzus ribis (L.)
Decorosiphon corynothrix Borner
Diuraphis (Holcaphis) hold (Hille Ris Lambers) Dysaphis
(Pompahis) plantaginea
Dysaphis tulipae
Ericaphis scammelli (Mason)
Ericaphis wakibae (Hottes)
Hayhurstia atriplicis
Hyadaphis foeniculi
Hyalomyzus collinsoniae (Pepper)
Hyalomyzus eriobotryae (Tissot)
Hyalomyzus mitchellensis Smith
Hyalomyzus sensoriatus (Mason)
Hyalopteroides humilis (Walker)
Hyperomyzus lactucae
Hyperomyzus (Neonasonovia) nabali (Oestlund)
Hyperomyzus (Neonasonovia) picridis (Borner & Blunck)
Idiopterus nephrelepidis Davis
Illinoia azalea (Mason)
Illinoia borealis (Mason)
Illinoia canadensis (MacGillivray)
Illinoia goldmaiyae (Knowlton)
Illinoia liriodendri
Illinoia pepperi (MacGillivray)
Illinoia (Masonaphis) rhokalaza (Tissot & Pepper)
Illinoia richardsi (MacGillivray)
Illinoia (Oestlundia) rubicola (Oestlund)
Illinoia spiraecola (Patch)
Linosiphon sanguinarium (Hottes & Frison)
Liosomaphis berberidis (Kaltenbach)
Lipaphis pseudobrassicae
Longicaudus trirhodus (Walker)
Macrosiphoniella abrotani (Walker)
Macrosiphoniella frigidicola Gillette & Palmer
Macrosiphoniella leucanthemi (Ferrari)
Macrosiphoniella ludovicianae
Macrosiphoniella millefolii (De Geer)
Macrosiphoniella (Phalangomyzus) pennsylvanica (Pepper)
Macrosiphoniella sanborni
Macrosiphoniella subterranean (Koch)
Macrosiphoniella tanacetaria (Kaltenbach)
Macrosiphoniella tapuskae (Hottes & Frison)
Macrosiphum adianti (Oestlund)
Macrosiphum californicum (Clarke)
Macrosiphum (Neocorylobium) carpinicolens Patch
Macrosiphum (Neocorylobium) coryli Davis
Macrosiphum cystopteris Robinson
Macrosiphum euphorbiae
Macrosiphum gaurae (Williams)
Macrosiphum gei (Koch)
Macrosiphum geranii (Oestlund)
Macrosiphum lilii (Monell)
Macrosiphum paZZidwra(Oestlund)
Macrosiphum (Neocorylobium) pseudocoryli
Macrosiphum ptericolens Patch
Macrosiphum rosae
Macrosiphum tiliae (Monell)
Mastopoda pteridis Oestlund
Metopolophium dirhodum (Walker)
Microlophium sibiricum (Mordvilko)
Microparsus desmodiorum Smith & Tuatay
Microparsus olivei Smith & Tuatay
Microparsus singularis (Hottes & Frison)
Muscaphis music Borner
Myzaphis rosarum (Kaltenbach)
Myzodium modestum (Hottes)
Myzus cerasi (Fabricius)
Myzus formosanus Takahashi
Myzus lythri (Schrank)
Myzus ornatus Liang
Myzus (Nectarosiphon) persicae
Nasonovia (Kakima) aquilegiae (Essig)
Nasonovia compositellae (Theobald)
Nasonovia (Kakima) cynosbati (Oestlund)
Nasonovia (Kakima) heucherae (Thomas)
Nasonovia (Ranakimia) purpurascens (Oestlund)
Nasonovia ribisnigri (Mosley)
Nearctaphis bakeri
Nearctaphis clydesmithi
Nearctaphis crataegifoliae
Neomyzus circumflexus (Buckton)
Neotoxoptera formosana (Takahashi)
Neotoxoptera violae (Pergande)
Ovatus crataegarius
Papulaphis sleesmani (Pepper)
Phorodon humuli
Pleotrichophorus ambrosiae Hille Ris Lambers
Pleotrichophorus asterifoliae (Strom)
Pleotrichophorus glandulosus (Kaltenbach)
Pleotrichophorus patonkus (Hottes & Frison)
Pleotrichophorus wasatchii (Knowlton)
Pseudacaudella rubida Borner
Rhodobium porosum
Rhopalosiphoninus latysiphon
Rhopalosiphoninus (Myzosiphon) solani (Thomas)
Rhopalosiphoninus staphyleae (Koch)
Rhopalomyzus (Judenkoa) lonicerae (Siebold)
Rhopalomyzus poae (Gillette)
Sitobion avenae
Uroleucon ambrosiae (Thomas)
Uroleucon anomalae
Uroleucon (Lambersius) caligatum (Richards)
Uroleucon chrysanthemi (Oestlund)
Uroleucon chrysopsidicola (Olive)
Uroleucon (Lambersius) erigeronense (Thomas)
Uroleucon eupatoricolens (Patch)
Uroleucon (Uromelan) eupatorifoliae (Tissot)
Uroleucon floricola Robinson
Uroleucon (Lambersius) gravicorne (Patch)
Uroleucon (Uromelan) helianthicola (Olive)
Uroleucon (Uromelan) Mini (Hottes & Frison)
Uroleucon impatiensicolens (Patch)
Uroleucon lanceolatum (Patch)
Uroleucon leonardi (Olive)
Uroleucon (Lambersius) luteolum (Williams)
Uroleucon nigrotibium (Olive)
Uroleucon nigrotuberculatum (Olive)
Uroleucon obscuricaudatum (Olive)
Uroleucon paucosensoriatum (Hille Ris Lambers)
Uroleucon pepperi (Olive)
Uroleucon pieloui (Richards)
Uroleucon pseudambrosiae
Uroleucon rudbeckiae (Fitch)
Uroleucon (Uromelan) rurale (Hottes & Frison)
Uroleucon russellae (Hille Ris Lambers)
Uroleucon sonchellum (Monell)
Uroleucon sonchi (L.)
Uroleucon (Uromelan) taraxaci (Kaltenbach)
Uroleucon (Uromelan) tar doe (Hottes & Frison)
Uroleucon (Uromelan) tuataiae Olive
Utamphorophora crataegi
Utamphorophora humboldti (Essig)

TABLE 6. SPECIES IN THE SUBFAMILY APHIDINAE, TRIBE APHIDINI
OCCURRING IN PENNSYLVANIA.

Aphis angelieae Lee & Seo
Aphis asclepiadis Fitch
Aphis caliginosa Hottes & Frison
Aphis carduella
Aphis cephalanthi
Aphis coreopsidis (Thomas)
Aphis cornifoliae Fitch
Aphis craccivora
Aphis debilicornis Gillette & Palmer
Aphis decepta Hottes & Frison
Aphis fabae
Aphis farinosa Gmelin
Aphis feminea Hottes
Aphis folsomii Davis
Aphis forbesi Weed
Aphis frangulae Kaltenbach
Aphis gerardiae (Thomas)
Aphis glycines
Aphis gossypii
Aphis hamamelidis Pepper
Aphis hederae
Aphis illinoisensis Shimer
Aphis impatientis Thomas
Aphis (Protaphis) knowltoni Hottes & Frison
Aphis lugentis
Aphis maculatae Oestlund
Aphis (Protaphis) middletonii
Aphis nasturtii
Aphis neilliae Oestlund
Aphis nerii Boyer de Fonscolombe
Aphis (Bursaphis) oenotherae Oestlund
Aphis oenotherae sanborni
Aphis oestlundi
Aphis pawneepae Hottes
Aphis pomi
Aphis pulchella
Aphis rubicola Oestlund
Aphis rubifolii
Aphis rumieis
Aphis sambuci (L.)
Aphis spiraecola
Aphis spiraephila Patch
Aphis (Burspahis) varians Patch
Aphis vernoniae Thomas
Aphis viburniphila
Hyalopterus pruni
Hysteroneura setariae
Pseudasiphonaphis corni (Tissot)
Rhopalosiphum cerasifoliae (Fitch)
Rhopalosiphum enigma Hottes & Frison
Rhopalosiphum maidis
Rhopalosiphum musae (Schouteden)
Rhopalosiphum nigrum Richards
Rhopalosiphum nymphaeae
Rhopalosiphum oxyacanthae
Rhopalosiphum padi
Rhopalosiphum parvae Hottes & Frison
Rhopalosiphum rufiabdominale
Rhopalosiphum sanguinarium McVicar Baker
Sanbornia juniper Pergande ex Baker
Schizaphis graminum
Schizaphis nigra (Baker)

TABLE 7. SPECIES IN THE SUBFAMILY CALAPHIDINAE OCCURRING IN
PENNSYLVANIA.

Tribe         Species

Calaphidini   Betulaphis quadrituberculata
                (Kaltenbach)
              Calaphis alni Baker
              Calaphis betulaecolens
                (Fitch)
              Calaphis betulella Walsh
              Calaphis leonardi Quednau
Panaphidini   Chromaphis juglandicola
                (Kaltenbach)
              Eucallipterus tiliae (L.)
              Hoplochaitophorus
                heterotrichus Quednau
              Hoplochaitophorus
                quercicola (Monell)
              Lachnochaitophorus
                obscuras (Tissot)
              Melanocallis
                caryaefoliae (Davis)
              Monellia caryella
              Monellia hispida Quednau
              Monellia microsetosa Richards
              Monelliopsis bisselli Quednau
              Monelliopsis caryae (Monell)
              Monelliopsis nigropunctata
                (Granovsky)
              Myzoeallis alhanibra Davidson
              Myzoeallis (Neomyzocallis)
                aselepiadis (Monell)

Tribe         Species

Calaphidini   Calaphis (Cepegillettea)
                myricae (Patch)
              Callipterinella
                calliptera (Hartig)
              Euceraphis betulae Koch
              Euceraphis gillettei Davidson
              Euceraphis lineata Baker
Panaphidini   Myzoeallis (Lineomyzocallis)
                bella (Walsh)
              Myzoeallis (Castaneomyzocallis)
                castaneae (Fitch)
              Myzoeallis (Agrioaphis)
              Myzoeallis coryli (Goetze)
              castanicola Baker
              Myzoeallis (Neomyzocallis)
                discolor (Monell)
              Myzoeallis (Lineomyzocallis)
                exultans Boudreaux & Tissot
              Myzoeallis (Lineomyzocallis)
                frisoni Boudreaux & Tissot
              Myzoeallis (Lineomyzocallis)
               granovskyi Boudreaux & Tissot
              Myzoeallis (Lineomyzocallis)
                longiunguis Boudreaux & Tissot
              Myzoeallis (Lineomyzocallis)
                melanocera Boudreaux & Tissot
              Myzoeallis (Lineomyzocallis)
                multisetis Boudreaux & Tissot
              Myzoeallis (Neomyzocallis)
                punctata (Monell)
              Myzoeallis (Lineomyzocallis)
                spinosa Boudreaux & Tissot
              Myzoeallis (Neomyzocallis)
                tuberculata Richards

Tribe         Species

Calaphidini   Euceraphis mucida (Fitch)
              Euceraphis punctipennis
               (Zetterstedt)
              Hannabura alnosa (Pepper)
Panaphidini   Myzoeallis (Lineomyzocallis)
                walshii (Monell)
              Neosymydobius albasiphus
                (Davis)
              Patchia virginiana Baker
              Protopterocallis
                fumipennella (Fitch)
              Protopterocallis
                gigantean Bissell
              Protopterocallis
                pergandei Bissell
              Pterocallis
                alnifoliae (Fitch)
              Therioaphis ononidis
                (Kaltenbach)
              Therioaphis riehmi
              Therioaphis trifolii
              Therioaphis trifolii
                forma maculate (Buckton)
              Tinocallis ulmifolii (Monell)
              Tuberculatus punctatella Fitch

TABLE 8. SPECIES IN THE SUBFAMILY CHAITOPHORINAE OCCURRING
IN PENNSYLVANIA.

Tribe           Species

Chaitophorini   Chaitophorus longipesi Tissot
                Chaitophorus nigrae Oestlund
                Chaitophorus nigricentrus Richards
                Chaitophorus nudus Richards
                Chaitophorus populicola Thomas
                Chaitophorus populifoliae Oestlund
Siphini         Sipha (Rungsia) elegans del Guercio

Tribe           Species

Chaitophorini   Chaitophorus populifolii
                Chaitophorus pusillus Hottes & Frison
                Chaitophorus saliniger Shinji
                Chaitophorus stevensis Sanborn
                Chaitophorus viminalis Monell
Siphini         Sipha flava

Tribe           Species

Chaitophorini   Chaitophorus viminicola Hille Ris Lambers
                Periphyllus americanus
                Periphyllus californiensis (Shinji)
                Periphyllus lyropictus (Kessler)
                Periphyllus negundinis (Thomas)
Siphini         Sipha glyceriae

TABLE 9. SPECIES IN THE SUBFAMILY DREPANOSIPHINAE
OCCURRING IN PENNSYLVANIA.

Drepanaphis
acerifoliae
Drepanaphis
carolinensis
Drepanaphis
kanzensis Smith
Drepanaphis
monelli Davis
Drepanaphis
nigricans
Drepanaphis
parva Smith
Drepanaphis
sabrinae Miller
Drepanaphis
simpsoni Smith
Drepanaphis
spicata Smith
Drepanosiphum
platanoidis
Shenahweum
minutum (Davis)

TABLE 10. SPECIES IN THE SUBFAMILY ERIOSOMATINAE OCCURRING IN
PENNSYLVANIA.

Tribe                      Species

Eriosomatini   Colopha graminis (Monell)
               Colopha ulmicola (Fitch)
               Eriosoma americanum (Riley)
               Eriosoma crataegi (Oestlund)
Fordini        Forda marginata Koch
               Geoica ultricularia group
Pemphigini     Grylloprociphilus
                 imbricator (Fitch)
               Mordwilkoja vagabunda (Walsh)
               Neoprociphilus aceris (Monell)
               Pachypappa pseudobyrsa (Walsh)
               Pemphigus bursarius (L.)
               Pemphigus monophagus Maxson
               Pemphigus nortonii Maxson
               Pemphigus populieaulis

Tribe                       Species

Eriosomatini   Eriosoma lanigerum
               Eriosoma (Mimaphidus) lanuginosum
                 (Hartig)
               Eriosoma mimicum Hottes & Frison
               Eriosoma rileyi Thomas
Fordini        Melaphis rhois
               Smynthurodes betae Westwood
Pemphigini     Pemphigus populitransversus
               Pemphigus populivenae
               Prociphilus americanus (Walker)
               Prociphilus caryae caryae (Fitch)
               Prociphilus caryae spp. Fitchii
                 Baker & Davidson
               Prociphilus (Neoparacletus)
                 corrugatans (Sirrine)
               Prociphilus erigeronensis (Thomas)
               Prociphilus (Meliarhyzophagus)
                 fraxinifolii

Tribe                      Species

Eriosomatini   Eriosoma wilsoni Remaudiere
               Kaltenbachiella ulmifusa
                 (Walsh & Riley)
               Tetraneura nigriabdominalis
               Tetraneura ulmi (L.)
Fordini
Pemphigini     Prociphilus longianus Smith
               Prociphilus (Pulvius)
                 probosceus (Sanborn)
               Prociphilus (Paraprociphilus)
                 tessellatus (Fitch)
               Thecabius affinis (Kaltenbach)
               Thecabius (Parathecabius)
                 gravicornis (Patch)
               Thecabius populimonilis (Riley)

TABLE 11. SPECIES IN THE SUBFAMILY LACHNINAE OCCURRING IN
PENNSYLVANIA.

Tribe                    Species

Eulachnini  Cinara atlantica
            Cinara banksiana Pepper & Tissot
            Cinara braggii (Gillette)
            Cinara canatra Hottes & Bradley
            Cinara costata (Zetterstedt)
            Cinara (Cupressobium)
              cupressi (Buckton)
            Cinara fornacula Hottes
            Cinara gracilis (Wilson)
            Cinara harmonia Hottes
            Cinara juniperi De Geer
Lachnini    Lachnus allegheniensis McCook
            Longistigma caryae (Harris)
Tramini     Trama rara Mordvilko

Tribe                 Species

Eulachnini  Cinara juniperivora (Wilson)
            Cinara laricifex (Fitch)
            Cinara laricis (Hartig)
            Cinara pergandei (Wilson)
            Cinara pilicornis (Hartig)
            Cinara pinea (Mordvilko)
            Cinara pinivora (Wilson)
            Cinara pruinosa (Hartig)
            Cinara spiculosa Bradley
            Cinara strobe Fitch
Lachnini    Tuberolachnus
              salignus (Gmelin)
Tramini

Tribe                   Species

Eulachnini  Cinara taedae Tissot
            Cinara (Cupressobium)
              tujafilina (Del Guercio)
            Cinara watsoni Tissot
            Essigella pini
            Eulachnus agilis (Kaltenbach)
            Eulachnus americanus Takahashi
            Eulachnus rileyi
            Schizolachnus parvus (Wilson)
            Schizolachnus piniradiatae
              (Davidson)
Lachnini
Tramini

Fig. 1. Proportion of aphids from the pan trapping
collection in Pennsylvania and New York that use herbaceous
plants, trees, or crops as primary hosts. Host
associations for North America characterized from
Blackman & Eastop (1994, 2000, 2006).

Crop         3%
Herb/grass   29%
Woody        61%
Unknown      7%

Note: Table made from pie chart.


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Article Details
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Author:Bachmann, Amanda C.; Nault, Brian A.; Fleischer, Shelby J.
Publication:Florida Entomologist
Article Type:Report
Geographic Code:1USA
Date:Sep 1, 2014
Words:6270
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