The Description of Zapatella davisae, New Species, (: Cynipidae) a Pest Gallwasp of Black Oak () in New England, USA. Author(s): Matthew L. Buffington , George Melika , Monica Davis , and Joseph S. Elkinton Source: Proceedings of the Entomological Society of Washington, 118(1):14-26. Published By: Entomological Society of Washington DOI: http://dx.doi.org/10.4289/0013-8797.118.1.14 URL: http://www.bioone.org/doi/full/10.4289/0013-8797.118.1.14

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THE DESCRIPTION OF ZAPATELLA DAVISAE, NEW SPECIES, (HYMENOPTERA: CYNIPIDAE) A PEST OF BLACK OAK (QUERCUS VELUTINA) IN NEW ENGLAND, USA.

MATTHEW L. BUFFINGTON,GEORGE MELIKA,MONICA DAVIS, AND JOSEPH S. ELKINTON

urn:lsid:zoobank.org:pub:C3906382-DC1D-4D30-B527-F072108FA204

(MLB) Systematic Entomology Laboratory, ARS-USDA, c/o National Museum of Natural History, Smithsonian Institution,10th & Constitution Ave NW, MRC 168, Washington, DC 20560 urn:lsid:zoobank.org:author:603275DE-9AE3-40C6-8AD7- 6A2AF7485F35; (GM) Plant Health and Molecular Biology Laboratory, National Food Chain Safety Office, Directorate of Plant Protection, Soil Conservation and Agrienvironment, Budao¨rsi str. 141-145, Budapest 1118, Hungary. E-mail: [email protected] urn:lsid:zoobank.org:author:A6FEF774-3A77-4058- 94E5-E670D880ED69; (MD & JSE) Department of Environmental Conservation, University of Massachusetts, Amherst, MA 01003 MD: urn:lsid:zoobank.org: author:2AA1D888-3287-409B-B8FF-C185FF987E86 JE: urn:lsid:zoobank.org: author:F9C03B49-E783-4C8B-A44F-F167E03D01CF

Abstract.— Many species of gall wasp (Cynipidae) essentially co-exist with their host oak tree species. Occasionally, the association becomes destructive to the tree, as is the case with Zapatella davisae, new species. This species is a twig galler, and as such, in the cases of heavy infestation, cause flagging, leaf clumping, and dieback of branches and twigs. Historical records of other species of Zapatella suggest that members of this genus have a checkered record with respect to damaging their host plants in North America, and these data are summarized here. Key Words: Cape Cod, Long Island, Martha’s Vineyard, oak pest DOI: 10.4289/0013-8797.118.1.14

INTRODUCTION Significant damage was not reported for several years and, in some cases, it In the early summer of 2013, MLB was was incorrectly attributed to the invasive contacted by JE regarding the identifica- winter moth, Operophtera brumata L. tion of a gall wasp species (Hymenoptera: (Lepidoptera: Geometridae) and fall can- Cynipidae) that was inducing gall damage kerworm, Alsophila pometaria (Harris) on black oak (Quercus velutina Lam.) on (Lepidoptera: Geometridae). Over the last Cape Cod and Martha’s Vineyard, MA. three years, damage has become widespread The first record of what is here described on Cape Cod and Martha’s Vineyard, war- as a new species (previously identified as ranting research to understand this species. ceropteroides Bassett) was Through correspondence between MLB recorded on Martha’s Vineyard in 2007. and GM, molecular sequencing data, VOLUME 118, NUMBER 1 15 comparison with additional specimens sculpture; the pronotum, laterally, is in the National Collection (NIC; delicately reticulate; the metascutellum National Museum of Natural History, is rugoso-reticulate; the metanotal trough Smithsonian Institution), consultation and the lateral area of the propodeum has of the Hopkin’s plant-insect catalogue dense white setae. In Callirhytis, a dis- (NIC) and additional collections from tinct malar sulcus is present; the meso- MA, the wasps submitted appeared to soma is less arched, and is always at least represent a new species of Zapatella slightly longer than high in lateral view; Pujade-Villar & Melika, 2012; herein we the transversely orientated rugae on the describe Zapatella davisae Buffington mesoscutum are much stronger, with and Melika, new species. Furthermore, much fewer anastomoses between them; records in the NIC indicate that other the pronotum has distinctly strong rugae species of Zapatella may be considered laterally; the metascutellum is rugose, to be pests of oak-dominated ecosystems never reticulate; and, the metanotal in the northeastern USA. trough and the lateral area of the propo- Zapatella was established for two deum either lack or have very few setae. newly described Neotropical species, Z. The most striking characters that dif- nievesaldreyi Melika & Pujade-Villar ferentiate Zapatella from Callirhytis, (from Colombia) and Z. grahami Pujade- and Plagiotrochus (the three Villar & Melika (from Costa Rica); genera most morphologically similar to both species are known to induce stem Zapatella) are the long, prominent part of swelling galls (Pujade-Villar et al. 2012). the ventral spine of the hypopygium, be- PartoftheresearchonwhichZapatella ing 6.0x–8.5x longer than broad; and the was originally described included the ex- hind coxae with dense white setae on the amination of additional Nearctic species of dorso-posterior surface. In the other three gallwasps, particularly those in Callirhytis mentioned genera, the prominent part of Foerster; this research indicated that five the ventral spine of the hypopygium is Nearctic species of Callirhytis belong to very short, at most 2–3 times longer than Zapatella: Z. cryptica (Weld), Z. herberti broad, and hind coxae lack dense setae. (Weld), Z. oblata (Weld), Z. quercusmedullae Aside from the new species described (Ashmead), and Z. quercusphellos (Osten here, several other species of oak gall- Sacken). Phylogenetically, Zapatella is wasps have been reported as harmful to a core member of , or the oak the oak trees on which they develop. gall wasps, the most speciose lineage of Plagiotrochus amenti Kieffer, 1901 can Cynipidae with a host preference for be a serious pest of Q. suber L. in the Quercus spp. (M.L. Buffington and Mediterranean area (Benia et al. 2009, G. Melika, pers. obsv.). Garbin et al. 2005), as well as in North Members of Zapatella most closely and South America, where Q. suber has resemble those in Callirhytis. However, been introduced to provide cork for the in Zapatella, the malar sulcus is absent; wine industry (Bailey & Stange 1966, the mesosoma is strongly arched, short, Zuparko 1996, Dı´az 1973). Disholcaspis and as long as high in lateral view; the cinerosa (Bassett, 1881) induces stem mesoscutum has numerous fine, short, galls in Texas, sometimes severely dam- interrupted transverse striae with several aging urban ornamental live oak trees longitudinal anastomosis connecting (Frankie et al. 1992); D. quercusmamma transverse striae, and together, forming has been reported as a street pest of native a net-like, delicately reticulate, irregular Quercus spp. planted as ornamentals in 16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Colorado (Eckberg and Cranshaw, 1994). prunings. This may be an effective Severe outbreaks of Andricus quercus- method of control at low densities, how- laurinus Melika & Pujade-Villar, 2009 ever on Cape Cod and Martha’s Vineyard, cause the death of thousands of trees in gall wasp populations are much too ex- natural stands of Q. laurina Humb. & tensive and densities are too high to Bonpl. Tree decline and mortality are eradicate using a pruning method. associated with twig and stem swellings caused by asexual galls (Melika et al. 2009). Andricus breviramuli Pujade-Villar, MATERIALS AND METHODS 2014. was reported as a serious pest of Repositories.—USNM: National Museum Quercus laeta Liebm. (section Quercus); of Natural History, Smithsonian Insti- in this case, the gall wasps kill and distort tution, Washington DC. branches of individual stressed trees. PHMB: Plant Health and Molecular After emergence of the adult wasp, the Laboratory, National Food Chain Safety galled tissues (twigs and branches) die, Office, Budapest, Hungary. but the basal part of the twig is alive and Description.—Specimens were examined produces new sprouts. However, repeated using a Leica 205cÒ™ microscope attacks may kill branches and distort and with fluorescent lighting. Photographs reduce growth of an entire tree (Pujade- were captured with a LeicaÒ™ DMRB Villar et al. 2014). compound microscope with a GT-VisionÒ™ Specimens housed in the NIC suggest Lw11057C-SCI digital camera attached. species of Zapatella have been either Lighting was achieved using techniques a nuisance or a pest of Quercus species summarized in Buffington et al. (2005), for decades. In particular, specimens of Z. Kerr et al. (2009) and Buffington and quercusmedullae (Ashmead) (identified Gates (2009). Single montage images were by Middleton as Callirhytis cryptus) produced from image stacks with the pro- taken in Atlanta, GA (Hopkin’s Number gram CombineZPÒ™. Scanning electron 13659d) indicate that this species was micrographs were generated using a causing serious damage to Quercus nigra HitachiÒ™ TM3000 desktop scanning L. Other specimens of Z. quercusme- electron microscope; specimens were dullae associated with Hopkin’s number coated in 25–30 nm gold-palladium alloy 10194 tell a more interesting story. (CressingtonÒ™ 108 auto sputtercoater), Mr. Kessler notes in areas where Q. mar- using ‘analysis’ voltage, running in ‘compo’ ilandica, Q. velutina,andQ. rubra grow mode. All images are available for down- together (around Birmingham, AL), load from www.morphbank.com. All spec- a scale insect infestation can weaken imens are housed at either the USNM trees, and a gall wasp (in this case, Z. (M.L. Buffington) or PHMB (Plant Health quercusmedullae) can become a problem; and Molecular Biology Laboratory, Na- identification of the wasps by Rohwer. tional Food Chain Safety Office, Budapest) Lastly, Hopkins number 13604f records (G. Melika). Morphological terminology Z. quercusmedullae (as C. cryptus,de- follows that of Ronquist and Nordlander termined by Middleton) as a pest on Q. (1989), Fontal-Cazalla et al. (2002) and catesbaei Michx. around Atlanta, GA, Buffington et al. (2007); cuticular surface that has ‘injured shade trees on two terminology follows that of Harris (1979). premises severely’; the suggested remedy Collection and Rearing methods.— was to selectively prune galled twigs in Specimens used for morphological analysis the winter and early spring, then burn the were reared from branch samples of VOLUME 118, NUMBER 1 17 infested trees at a site on Martha’s morphology; will key to this species Vineyard, Massachusetts (N41.3975414, in Pujade-Villar et al. (2012). Zapatella W -70.6771814) and at four sites on Cape davisae differs from Z. quercusmedullae Cod (in the towns of Dennis, Barnstable in the morphology of the mesopleuron: in and West Harwich (N 41.7370257, W Zapatella davisae there exists a relatively -70.193331; N41.7285832,W -70.1869863; smooth area just ventral to the meso- N41.6605355, W-70.496351; N 41.672801, pleural triangle, running posteroventrally W -70.119433, respectively). In March to the posterior margin of the meso- 2014, two samples per tree were removed pleuron; in Z. quercusmedullae (and all from each site, including new and last other Zapatella examined here), this area year’s growth, and placed in individual is evenly scultptured throughout. In Z. 1-gallon ZiplocÒ bags. The bags were davisae POL longer than OOL, F5-F11 storedina4°C PercivalÒ for 6 weeks forming slight club, each segment as until adult gall wasps emerged. After wide as long, while in Z. quercusme- emergence a total of 50 adult gall wasps dullae POL equal OOL, F5-F11 not were collected from the four Cape Cod forming slight club all segments are lon- sites and 2 from Martha’s Vineyard. , ger than broad. Each wasp was placed in an individual Description.—Adult female (type se- cell of a well plate with 2 µl of ethanol. ries). Color (alive or freshly killed): two The plate was sealed and sent to MLB forms: form I, head and mesosoma or- and GM for morphological identification. ange to dark brown-orange, metasoma Molecular diagnostics.—Freshly emerged anteriorly bright orange, darker posteri- specimenswereplacedinPCRtubes,con- orly, legs and antenna medium orange; taining 95% ethanol, and sent to the Bar- form II, whole insect bright yellow- code of Life project in Guelph, Canada for orange. DNA extraction and sequencing of the CO1 Head: Oval in anterior view (Fig. 3A); barcoding gene; extractions and sequencing frons shagreen, with slight longitudi- protocols follow Hebert et al. (2003) and nal microsculpture, moderately setose; Gwiazdowski et al. (2015). CO1 sequences clypeus impressed, pinched laterally by were retrieved for each specimen and were clypeo-pleuralstomal line; strong striae uploaded onto the BOLD database (http:// radiating from clypeo-pleuralstomal line www.boldsystems.org/index.php/Public_ dorsally towards ventral margin of eye, BINSearch?searchtype=records) under the malar sulcus absent; gena and vertex Project ID ‘MOND’ along with images of gently shagreen, nearly glabrous; gena each voucher; original sequences can be broadly rounded (Fig. 2A, B). Toruli located on Genbank under accession num- ring-like; antenna with 11 flagellomeres, bers KU567119 - KU567202. semi-clavate, F1 longest, 1.3x longer than F2, F2-4 sub-equal in length, F5-F11 forming slight club, each segment as Zapatella davisae Buffington & wide as long, darker than proceeding Melika, n. sp. flagellomeres (Fig. 1B). urn:lsid:zoobank.org:act:C9ACDAD4- Mesosoma: Lateral surface of prono- 6D76-4D0F-AD12-68A297E942C4 tum slightly rugulose, sparsely covered in short appressed setae (Fig. 1B); lateral Figs. 1-4 pronotal carina lacking, pronotal plate Diagnosis.—Most similar to Z. narrow, dorsally shiny-smooth, sub- quercusmedullae, both in biology and medial pronotal depressions deep, open 18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Light microscope images of Zapatella davisae, new species, female. A and B, habitus, lateral view. VOLUME 118, NUMBER 1 19

Fig. 2. Light microscope images of Zapatella davisae, new species, female. A, head and mesosoma, dorsal view. B, head and mesosoma, lateral view. 20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Scanning electron micrographs of Zapatella davisae, new species, female. A, head, anterior view; B, clypeus and mandibles, anterior view; C, pronotal plate, anterior view; D, mesosoma, lateral view; E, scutellum, postero-lateral view; F, scutellum and propodeum, postero-lateral view. laterally (Fig. 3C). Mesopleuron heavily (Figs 1B and 3D). Mesoscutum heavily shagreen save for dorsal-ventral patch shagreen to gently rugulose, generally below mesopleural triangle, running to glabrous, with sparse setae randomly metapleuron (Figs 1B and 3D); meso- distributed; anteroadmedian signum pleural triangle deeply impressed, se- distinct; median mesoscutal line present, tose, clearly defined along all edges short, extending from posterior margin VOLUME 118, NUMBER 1 21

Fig. 4. Scanning electron micrographs and light microscope images of Zapatella davisae,new species, female. A, metasoma, lateral view; B, anterior base of metasoma and petiole, lateral view; C, hypopygium, lateral view; D, marginal cell of forewing (detail); E, fore and hind wings. 22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Z. davisae damage on Quercus velutina ranging from moderate (left) to severe (middle). Damage can include flagging, leaf clumping, and dieback of branches and twigs. of mesoscutum about 1/15th length of widened ventrally, setose, bordered pos- mesoscutum, notch-like in some speci- teriorly by distinct metapleural sulcus mens (Fig. 2A); notaulus incomplete, (Fig. 3D); upper episternum deeply ex- originating at posterior end of meso- cavated by spiracular groove, bounded scutum, gradually obliterated about half- posteriorly by calyptra, with longitudinal way to anterior margin of mesoscutum sculpturing; lower episternum dorsally (Fig. 2A). Disk of mesoscutellum deeply glabrous, smooth, ventrally setose; pit rugulose laterally, more gently so medi- at anteroventral margin of metapleuron ally, with few long setae laterally (Figs indistinct, setose; propodeum short, in- 2A and 3E); scutellar ridge separating visible laterally; posterioly impressed scutellar fovea narrow, short; scutellar beneath mesoscutellum (Fig. 3F); fovea oval, obliquely angled relative to propodeal carinae thin, complete, widely midline, posterior rim present, bottom spaced, parallel; lateral propodeal cari- smooth, glabrous (Figs 1B and 3E). nae absent; area between propodeal ca- Metapleural-propodeal complex: rinae smooth, dorsally setose (Fig. 3F); Metapleuron glabrous anterodorsally, nucha short, glabrous, deeply rugulose gradually more setose posteroventrally longitudinally. (Figs 1B and 3D); anterodorsal depression Wings: Entire wing glabrous, with of metepisternum elongate, triangular, scattered scars presumably representing VOLUME 118, NUMBER 1 23 setal sockets; marginal cell incomplete, in early June. In the fall of 2013 and elongate, open along anterior margin 2014, tree tips on Cape Cod were bagged (Fig. 4D); 2r distinct; R1 absent or to evaluate a possible fall emergence. No hardly visible, Rs very inconspicuous, gall wasps were found, which suggests nearly straight; Rs+M indistinct, reach- that Z. davisae does not have an au- inges basalis at half of its height; areolet tumnal generation. Gall cavity develop- indistinct, represented by glabrous area; ment starts in late July to early August. marginal, radial sector and cubital veins Larval development begins August, with represented by trace veins; wing margin pupal and adult development following without cilia; distal margin complete, suit. Z. davisae overwinters in multiple evenly rounded (Fig. 4E). life stages, including larval, pupal, pha- Legs: Coxae with even, dense patches rate adult and adult stages. From Octo- of setae, especially prominent on hind ber to April, multiple life stages are coxa; sparse, appressed setae present on simultaneously present in twigs. Thus all femorae and tibiae; femora with far, there is no evidence of a sexual shagreen sculpture, all tibiae and tarso- generation or alternate galls on black oak meres gently reticulate. Tarsomeres or other host species. Current research is evenly covered in short, appressed setae; ongoing to investigate a possible alter- length of tarsomere 1 equal to combined nate summer generation. length of tarsomeres 2-5; claw simple Etymology.—Named in honor of the (without basal tooth). discoverer of this species, Monica Davis. Metasoma: Petiole obscured by ante- Material examined.—Holotype (fe- rior margin of tergum 3 (T3); all post- male): USA: NY, Suffolk Co. Long petiolar terga free; T3 6x length of T4, Island, Oakdale, Bayard Cutting Abor- dominating metasoma in lateral view etum, N 41.7370257, W 70.1933310, (Fig. 4A); posterior margins of all met- 8.IV.2014, Coll. Monica Davis, UMASS asomal terga parallel, gradually angled LIA. USNMENT01119081. away from midline posteriorly; T4-T9 Paratypes: USA: NY:SuffolkCo. with extremely minute micropores Long Island, Oakdale, Bayard Cutting (Fig. 4C); sparse setae present on T8 Aboretum, N 41.7370257, W 70.1933310, (Fig. 4C); hypopygium (sternite 7) 8.IV.2014, Coll. Monica Davis, UMASS not extended ventrally; sparse, short LIA45 (USNMENT01119082- USNMENT setae presently along extreme ventral 01119083); Long Island A, N 40.960373, margin. W -72.714072, 4.IV.2014, D. Gilrean, Description of gall and plant damage.— coll. (USNMENT01119710-USNMENT Galled twigs can be slight to obvious. 01119725); Long Island B, N 40.7354614, Pin-size exit holes made by emerging W -73.1629878, 4.IV.2014, D. Gilrean, wasps are often present on galled twigs. coll. (USNMENT01119726-USNMENT Removing bark and outer wood from 01119739, USNMENT01119707- infested twigs exposes oval cells produced USNMENT01119708; MA: Barnstable by the wasp, often containing adults, pupae, Co., Cape Cod, Dennis, N 41.7370257, or larvae. W 70.1933310, 12.VI.2014,Coll. Monica Biology.—Only the asexual twig- Davis, DC16, USNMENT01119700; galling stage on black oak (Quercus ve- Barnstable, N41.6605355, W-70.496351, lutina) is known for Z. davisae. Wasps 31.III.2014, Coll. Monica Davis emerge in May, following leaf expansion (USNMENT 01119755-USNMENT and their respective parasitoids emerge 01119768, USNMENT 01119709); Dennis, 24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

N41.7285832,W -70.1869863, 31.III.2014, of this species were located in the USNM Coll. Monica Davis (USNMENT (including one paralectotype) and com- 01119769-USNMENT 01119777); Dennis, pared with Zapatella davisae.While N41.7285832,W -70.1869863, 7.III.2014, clearly not the same species as Z. davisae, Coll. Monica Davis (USNMENT this species was determined to conform 01119778-USNMENT 01119780); Dennis quite clearly to Zapatella, and may indeed Cemetery, N 41.7370257, W -70.193331, be a synonym of Z. quercusmedullae.As 7/III/2014 (USNMENT 01119781- this latter question is beyond the scope of USNMENT01119791); Dennis Ceme- this paper, we only provide here the new tery, N 41.7370257, W -70.193331, 31/III/ combination (below), as the species itself 2014 (USNMENT 01119792-USNMENT requires redescription. 01119794); Martha’s Vineyard, N41.3975414, W -70.6771814, 13. XI.2013, Coll. Monica Davis (USNMENT DISCUSSION 01119795); Martha’s Vineyard, N Impact.—Symptoms of tree in- 41.3975523, W -70.6771685, 13.XI.2013, festation by Zapatella davisae include Coll. Monica Davis (USNMENT 01119796); dieback, sparse growth in spring, epi- West Harwich, N 41.672801, W -70.119433, cormic growth, flagging, and swollen 31.III.2014, Coll. Monica Davis (galled) twigs. Galled twigs can be slight (USNMENT 01119797- USNMENT to obvious. Damage caused by the cells 01119805); West Harwich, N 41.672801, and emerging wasps significantly dis- W -70.119433, 7.III.2014, Coll. Monica rupts the vascular system, and can cause Davis (USNMENT 01119806- USNMENT twig mortality, leading to flagging. In- 01119810 n); Yarmouth Port, Ancient festation typically starts in the upper Cemetery, ex black oak, 29.III.2013, coll. portion of crown and moves downward. Russell Norton (USNMENT01119701- Trees often do not show dieback until USNMENT01119706); Cape Cod Coop.; the second or third year of infestation. ex twigs, not swollen, on Quercus velutina Heavily infested trees can have near leg. Russell Norton, early summer (12 100% dieback and typically produce females in PHMB). epicormic growth. Outbreaks of this Comments.—Zapatella davisae was species on black oak have occurred previously identified as Callirhytis cer- along the east coast including Cape Cod, opteroides and it has been implicated in areas in South Shore, MA, Martha’s damaging Quercus velutina in the past. Vineyard, Nantucket, parts of Rhode Is- As mentioned in the Introduction, Za- land, and coastal Connecticut; the cir- patella most closely resembles Cal- cumstances of these outbreaks is not lirhytis. Nearctic Callirhytis has already known. been shown to be polyphyletic (Nylander Part of the story surrounding this 2004, Liljeblad et al. 2008) and many particular interaction may include the North American species assigned to fungal pathogen Botryosphaeria sp., Callirhytis (Weld 1952, Burks 1979), which has caused a great deal of damage in fact, are not Callirhytis ‘sensu stricto’ to Q. velutina in the past (1990’s) (Pike (Melika & Abrahamson 2002, Melika et al. 2001). It is often an opportunitic et al. 2009). Callirhytis ceropteroides, pathogen that acts as a secondary stress which was transferred from Bassettia agent. It could be that the widespread Ashmead to Callirhytis ‘sensu stricto’ mortality of Q. velutina recently ob- (Melika & Abrahamson 2002). Specimens served on Cape Cod and Martha’s VOLUME 118, NUMBER 1 25

Vineyard arises from the combined Benia, F., Khelil M-A. & Pujade-Villar, J. 2009. impact of general defoliators, Bo- Pre´sence en Alge´rie de Plagiotrochus amenti tryosphaeria spp. and Z. davisae. (Hymenoptera, Cynipidae): une espe`ce gal- licole potentiellement dangereuse pour le Further research needs to be done to cheˆne-lie`ge (Quercus suber L.). Nouvelle understand the mechanism Zapatella Revue d’Entomologie 25: 291–296. davisae uses to induce Q. velutina stress Buffington M.L., Burks R., & McNeil, L. 2005. and mortality. In addition, we need to Advanced techniques for imaging micro- clarify how Z. davisae has spread so hymenoptera. American Entomologist 51: 50–54. rapidly over the past eight years. It is still Buffington M.L., Nylander J.A.A., & Heraty J. unknown whether this is an exotic or 2007. The phylogeny and evolution of Figi- native species and whether its native tidae (Hymenoptera: Cynipoidea). Cladistics predators may help control it’s spread in 23: 1–29. coastal New England. This species de- Buffington M.L., Gates M (2009) Advanced scription has created a platform for fur- imaging techniques II: using a compound ther research and management strategy microscope for photographing point-mount specimens. American Entomologist 54: 222– development that aims to inhibit the 224. dispersal and damaging capabilites of Z. Burks, B.D. (1979) Superfamily Cynipoidea. In: davisae. Finally, future research on the Krombein, K.V.,Hurd, P.D., Jr., Smith, D.R. & morphological and molecular limits of Burks, B.D. (eds), Catalog of Hymenoptera in Zapatella davisae and Z. quercusme- America North of Mexico. Vol. 1. Symphyta dullae and . Smithsonian Institution Press, warrants considerable attention. Washington, DC., pp. 1045–1107. Dı´az, N. B. 1973. Una nueva plaga del alcorno- que en la Repu´blica Argentina. Revista de ACKNOWLEDGEMENTS la Sociedad Entomolo´gica Argentina 34: GM would like to thank Russell 85–88. Norton (Cape Cod Cooperative Exten- Eckberg, T.B, & Cranshaw, W. 1994. Occurrence sion) for sending specimens. MD of the oak rough bulletgall wasp, Dish- olcaspis quercusmamma (Walsh) (Hyme- acknowledges Dan Gilrien for field col- noptera: Cynipidae), as a street tree pest in lections from Long Island sites, and Colorado. Journal of the Kansas Entomo- Arborjet Inc. for funding support. MLB logical Society 67: 290–293. was supported by the Systematic Ento- Fontal-Cazalla, F.M., M. Buffington, G. Nord- mology Laboratory, ARS-USDA, and lander, J. Liljeblad, P. Ros-Farre´,J.L. wishes to thank Paul Goldstein (Sys- Nieves-Aldrey, J. Pujade-Villar and F. Ron- quist. 2002. Phylogeny of the Eucoilinae tematic Entomology Laboratory) for (Hymenoptera: Cynipoidea: Figitidae). Cla- fruitful discussions on this topic. Men- distics 18: 154–199. tion of trade names or commercial Frankie, G.W., Morgan, D.L. & Grissell, E.E. products in this publication is solely for 1992. Effects of urbanization on the distri- the purpose of providing specific in- bution and abundance of the cynipid gall formation and does not imply recom- wasp, Disholcaspis cinerosa, on ornamental live oak in Texas. In: J. D. Shorthouse and O. mendation or endorsement by the Rohfritsch [eds.], Biology of - In- USDA. USDA is an equal opportunity duced Galls. Oxford University Press, New provider and employer. York and Oxford, pp. 258–279. Garbin, L., Durfot, M., Dı´az, N.B. & Pujade- References Villar, J. 2005. Histological modifications on Quercus suber twigs (Fagaceae) caused by Bailey, S.F. & Stange, L.A. 1966. The twig wasp the gall wasps Plagiotrochus suberi (Hyme- of cork oak - its biology and control. Journal noptera, Cynipidae). Entomologia Generalis of Economic Entomology 59: 663–668. 28: 91–102. 26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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