Misidentification of Vespula alascensis as V. vulgaris in North America (Hymenoptera: Vespidae; Vespinae)

2010·0.2 MB·English

by  James M. Carpenter

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3690, 7 pp., 5 figures, 1 table June 25, 2010
Misidentification of Vespula alascensis as V. vulgaris
in North America (Hymenoptera: Vespidae; Vespinae)
JAMES M. CARPENTER1 AND TRAVIS R. GLARE2
ABSTRACT
Morphological and molecular-based comparisons of the North American yellowjackets
identified as Vespula vulgaris (Linnaeus, 1758) with samples from other parts of the world
demonstratetheyarenotthatspecies.ThenameVespulaalascensis(Packard,1870)isapplicableto
the North American species, new status. Vespa communis de Saussure, 1857, non von Schrank,
1785,is a synonymof Vespula maculifrons (duBuysson, 1905),REVISED SYNONYMY.
INTRODUCTION vespine fauna, these have been treated as
species of Vespula, with Vespula maculifrons
Vespa vulgaris was described by Linnaeus (du Buysson) used as the name for Vespa
(1758) from ‘‘Europa,’’ and recorded from communis.
North America by Kirby (1837). De Saussure Vespula vulgaris was recorded from New
(1853–1858) included North America in the Zealand by Thomson (1923), although it
distribution of this taxon in his worldwide apparently did not become established for
monograph, but he later (de Saussure, 1857) 60 years (Donovan, 1984). Donovan (1984)
decided that the North American specimens noted differences in worker length and nest
were a distinct species, describing them as size between New Zealand specimens and a
Vespa communis. Du Buysson (1905) recog- sample from the western United States, and
nizedV.communisinhismonograph,butalso stated (p. 426) that this disparity ‘‘makes it
recorded V. vulgaris from North America doubtful if the New Zealand population
again, and in this he has been followed by originated in the western United States.’’
subsequent investigators. Since Bequaert’s One of us (T.R.G.) later compared molecular
(1932) synopsis of the North American sequence data for samples of V. vulgaris from
1DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory ([email protected]).
2Bio-Protection Research Centre, PO Box 84, Lincoln University, Lincoln 7647, New Zealand (travis.glare@lincoln.
ac.nz).
CopyrightEAmericanMuseumofNaturalHistory2010 ISSN0003-0082
2 AMERICAN MUSEUMNOVITATES NO. 3690
the United Kingdom, United States and New POLYMERASECHAINREACTION AMPLIFICATION
Zealand, and came to the conclusion (Glare,
2004: 2): ‘‘The UK and New Zealand wasps Polymerase chain reaction (PCR) amplifi-
were identical, but the ‘V. vulgaris’ from the cations were performed in 25 ml volumes
USA was not the same species.’’ This finding containing 0.4 mM of each primer (Invitro-
led the other of us (J.M.C.) to investigate gen Life Technologies), 2 ml DNA, 0.2 mg/ml
BSA (NEB, Biolab), and the following re-
whether diagnostic differences could be found
agents from ABgene, Innovative Sciences:
inthemalegenitalia.Thisturnedouttobethe
case. In the present paper we document both 200mMdNTPs,13ReddyMixTMPCRbuffer,
1.5 mM MgCl , and 0.6 U Thermoprime Plus
the molecular and morphological differences. 2
DNA polymerase. PCR primers which target
the mitochondrial region of part of the CO1
MATERIALS AND METHODS
gene, using primers mtd6-GGAGGATTT-
DISSECTIONS GGAAATTGATTAGTTCC (CI-J-1718 in
Simon et al., 1994) and mtd9- CCCGGTA-
Male specimens in the collection of the AAATTAAAATATAAACTTC (CI-N-2191
American Museum of Natural History were in Simon et al., 1994) were used. PCR prod-
dissected, the genital capsuleextracted, cleared ucts destined for sequencing were amplified in
slightly in warm lactophenol, and examined in 50mlvolumescontaining0.4mMofeachprimer
glycerin. Some previouslydissectedmales were (Invitrogen Life Technologies), 4 ml DNA,
also available. North American specimens 0.2 mg/ml BSA (NEB, Biolab), 200 mM
identified as V. vulgaris were from Alaska, dNTPs (ABgene, Innovative Sciences), 13
Alberta, California, New York, Oregon, and buffer containing 1.5 mM MgCl (Roche),
2
‘‘Hudson Bay Territory.’’ Eurasian specimens and1.75UExpandHighFidelityPCRsystem
werefromChina,theCzechRepublic,England, (Roche).Amplificationswerecarriedoutinan
and‘‘Europe,’’aswellasfromNewZealand. EppendorfMastercyclergradientthermocycler
using30cyclesof30secat94uC,45secat54uC,
DNA EXTRACTION and 1 min at 72uC. Positive (previously
successfully amplified DNA) and negative
Extraction of DNA was successfully per- (sterile dH O) controls were included in each
2
formedonanumberofwaspsamplescollected PCRrun.
andsuppliedbyNewZealand,Australian,U.S.
and U.K. collaborators (table 1). Both adults SEQUENCING
andlarvaewereextractedusingamodification
ofthemethodofHenryetal.(1990).Wholeor Expand amplified PCR products were
parts of wasp larvae or adults were ground in cleaned using a High Pure PCR Product
liquid nitrogen and 1 ml of TENT (10 mM Purification kit (Roche) and sent to the
Tris-HCl [pH 7.4], 25 mM EDTA, 10 mM Waikato DNA Sequencing Facility (Hamilton,
NaCl, and 0.5% Triton X-100) buffer added. New Zealand) for sequencing. Genbank num-
After5minutesofcentrifugation,thepelletwas bersforsequencesobtainedaregivenintable 1.
resuspended in 600 ml TEN (10 mM Tris-HCl
[pH7.4],25mMEDTA,10mMNaCl)buffer, ALIGNMENT
0.2 mg/ml of proteinase K (Invitrogen Life
Technologies),andafinalconcentrationof1% Sequences were aligned using DNAMAN
SDS.Following4hincubationat37uC,60mlof (Lynnon BioSoft, Quebec, Canada) and
5 M NaCl was added and the DNA twice ClustalX (Thompson et al., 1997).
extracted with an equal volume of phenol-
chloroform. The DNA was precipitated with
CLUSTERING
isopropanol, resuspended in 50 ml sterile
distilled water, and stored at 4uC or frozen Ropalidia romandi (Le Guillou) [Polistinae],
for long-term storage. The optimal concentra- Genbank accession number AF146677, was
tion of DNA for use in subsequent PCRs was used as an outgroup. Symmetric resampling
determinedempirically. (Goloboffetal.,2003)ofthealignedsequenc-
2010 CARPENTER AND GLARE: MISIDENTIFICATION OFVESPULA ALASCENSIS 3
Collector JoReesetal.,Landcare,NewZealand NicolaRichards,AgResearch,NewZealandAndyAustin,WaiteInstitute,UniversityofAdelaide DerekDaly,UniversityofLiverpoolHalC.Reed,OralRobertsUniversity
p
bleedtrapRobertsaitedtrabledtrap.bledtrap.bledtrap.
E1bankaccessionnumbers Locality Pelorus,SouthIsland,NZPelorus,SouthIsland,NZPelorus,SouthIsland,NZChristchurch,Canterbury,NZ Adelaide?,AustraliaBrukunga,AustraliaBotanicGardens,AustraliaUK AntrimCo.,Michigan,USA,AuSaInstitute,Aug.152001AAIB-baitTulsaCo.,Oklamhoma,USA.OralU.campus,Aug.272001AAIB-bAntrimCo.,Michigan,USA,AuSaInstitute,Jul.252001AAIB-baiteAntrimCo.,Michigan,USA,AuSaInstitute,Jul.252001AAIB-baiteAntrimCo.,Michigan,USA,AuSaInstitute,Jul.252001AAIB-baite
Ln
TABndGe us) DNA dult dult
DNAextractionsa Specimen VespulavulgarisadultVespulavulgarisadultVespulavulgarisadultV.germanica(FabricifoundressqueenV.germanicaadultV.germanicaadultV.germanicalarvaeSuppliedasextractedV.vulgarisfromVespulamaculifronsa Vespulamaculifronsa V.flavopilosa V.vulgaris‘‘’’ V.vulgaris‘‘’’
kno. 53555451 485052495756 59 58 60 61
nbansion 2078207820782078 207820782078207820782078 2078 2078 2078 2078
Geces GUGUGUGU GUGUGUGUGUGU GU GU GU GU
c
a
Code v1v2v3FQ Aw_W1conBK2AustGRLUkvul13Ukvul16RC_mac USAmac2 USAVflav USAvul1 USAvul2
d
e
DatereceivedDNAextract 24/03/0324/03/0324/03/0318/11/98 18/12/0122/07/0222/07/0208/04/0308/04/0327/01/03 27/01/03 27/01/03 27/01/03 27/01/03
r
o
4 AMERICAN MUSEUMNOVITATES NO. 3690
Fig.1. SymmetricresamplingtreegeneratedwithTNT,basedonapproximately400bpsequencesofthe
CO1gene ofmitochondrial DNA. Valuesdisplayed are frequency differences.
es was carried out with the program TNT paramere through several views to appreciate
(Goloboffetal.,2008),using10,000replicates. the difference).
RESULTS TAXONOMY
Figure 1 is the tree resulting from symmet- TheNorthAmericanspeciesidentifiedasV.
ric resampling. The North American speci- vulgaris is clearly not that species. The
mens identified as V. vulgaris group with question now is whether there is a name
Vespula flavopilosa Jacobson rather than with available for the North American species.
European and New Zealand V. vulgaris. Bequaert (1932) treated Vespa communis de
North American males are readily distin- Saussure,describedfrom‘‘Americaseptentr.,’’
guished from Eurasian and New Zealand asasynonymofVespulavulgaris,althoughhe
males by characters of the genitalia. The had earlier (Bequaert, 1930) treated it as a
digitus of V. vulgaris (fig. 2) is smoothly synonym of Vespula maculifrons. He did not
rounded and symmetrical toward the apex, state reasons for either synonymy, although
whereas in the North American form (fig. 3) presumablythelatterfollowedRohwer(1926).
thedigitusisasymmetricaltowardtheapex.In The synonymy of communis with vulgaris was
V. vulgaris (Fig. 4) the paramere is somewhat followed in the checklist of Vespinae by
more emarginate beside the dorsal terminal Carpenter and Kojima (1997). However,
process, so that the process appears slightly J.M.C. has seen type material of communis in
longer, than in the North American form the Muse´um National d’Histoire Naturelle in
(fig. 5; it may be necessary to rotate the Paris, and the taxon should be treated as a
2010 CARPENTER AND GLARE: MISIDENTIFICATION OFVESPULA ALASCENSIS 5
Figs. 2–5. Male genitalia. Figs. 2–3, digitus in ventral view: 2, V. vulgaris. 3, V. alascensis. Figs. 4–5,
paramerewithdorsalterminal processin dorsalview: 4,V.vulgaris. 5, V.alascensis.
synonymofmaculifrons,revisedsynonymy.In ed.’’Comparing these featureswith thekey in
any case, Vespa communis de Saussure, 1857, Akre et al. (1981) leads to the couplet for V.
is a junior primary homonym of Vespa vulgaris. Other details of the coloration
communis von Schrank, 1785, as was pointed mentionedinthedescriptionmatchV.vulgaris
out by Rohwer (1926). as well, viz. ‘‘the indentation in the eyes
Bequaert (1932) also listed Vespa alascensis yellow’’ [evidently referring to the clypeus],
Packard, 1870, described from ‘‘Lower ‘‘with a central broad black band, dilated
Yukon,’’ as a synonym of Vespula vulgaris. below into a rounded, larger termination,’’
Packard (1870) is an obscure publication ‘‘Mandibles yellow, cutting edge and teeth
indeed, because most of the copies of the black’’ and mesonotum ‘‘with the usual
publication were destroyed in the Great oblique line in front of the wings, and the
Chicago Fire of 1871. The publication was triangular spot beneath; two yellow spots on
overlookeduntilBanks(1920)calledattention the scutellum, and two lunate yellow spots
to a reprint in the Museum of Comparative behind.’’ The illustrations of the variation in
Zoology. Presumably Bequaert consulted this V. vulgaris in Jacobson et al. (1978) and Eck
copy, as the whereabouts of the type of V. (1999)showallthesedetails.AccordingtoEck
alascensis are unknown. The description and (1999)NorthAmericanspecimensasaruleare
figure do allow recognition of this taxon, a darker than Eurasian V. vulgaris, and in
conclusion also reached by Jacobson et al. particular never have yellow markings on the
(1978). Packard (1870: 27) mentioned eyes propodeum, and none are mentioned in the
reaching to the base of the mandibles, yellow description of alascensis. However, this fea-
markings, no markings on the scutum, black ture may have little diagnostic value. Eck
antennae, an upper and lower yellow spot on (1999) illustrated Palearctic vulgaris lacking
the gena the only markings on the orbits, and such spots, and Miller (1961: 10) mentioned
metasomal apical bands on terga II–IV with propodeal spots present in a small fraction of
‘‘the middle sinus triangular, the outer round- colonypopulations(,5%),althoughJacobson
6 AMERICAN MUSEUMNOVITATES NO. 3690
et al. (1978) and Eck (1999) observed only only the characters of the male genitalia
black propodea in North American vulgaris. reliably diagnose Vespula vulgaris and V.
TheonlypointinPackard’sdescriptionthat alascensis. Females of V. alascensis are gener-
is difficult to evaluate is the characterization ally said to be darker than V. vulgaris (Eck,
of the clypeus as ‘‘deeply cleft,’’ however the 1999), but there is overlap. Even the propo-
figureshowsnothingunusual.Thedescription deal spots, often present in V. vulgaris and
compared alascensis to the new species Vespa absent in V. alascensis, can be absent in V.
tripunctata, now considered a synonym of vulgaris (see figures in Eck, 1999). But the
Vespula austriaca (Panzer). The description of matter should be further investigated: the
tripunctata mentioned ‘‘Clypeus with two character differentiating D. norvegicoides and
prominent sharp black teeth on the front D. pacifica is a subtle one, and a similar
edge’’ (Packard, 1870: 26), and sharply pro- feature may yet be found to diagnose V.
duced apical angles of the clypeus are indeed vulgaris and V. alascensis.
characteristic of vespine social parasites, such
asV.austriaca.Packard’sstatementaboutthe ACKNOWLEDGMENTS
clypeus of alascensis may have meant to
contrast it with that of tripunctata, in lacking We thank Michael Archer and Libor
sharp apical angles. Dvorˇa´k for sending males of V. vulgaris for
The identification of alascensis as North dissection. Adrien Perrard kindly confirmed
American V. vulgaris is therefore accepted the identification of Vespa communis with
here, and that name is available for the Vespula maculifrons. Hal Reed, Oral Roberts
distinct species: Vespula alascensis (Packard), University, Tulsa, OK; Derek Daly, Univer-
NEW STATUS. sity of Liverpool; Andrew Austin, Waite
Institute, University of Adelaide; Jo Rees,
LandcareResearch,NewZealand;andNicola
DISCUSSION
Richard, AgResearch, New Zealand, supplied
Five Holarctic species were recognized in wasp material or DNA from wasps. Nicola
the checklist of Vespinae by Carpenter and Richards and Tracey Nelson provided techni-
Kojima (1997); now there are four. In each cal assistance. Steve Thurston rendered the
illustrations in figures 2–5. Matthias Buck,
case, names are available for both the pale-
Leopoldo Castro, Libor Dvorˇa´k, and Alison
arctic and nearctic forms, as these were
Stewart commented on a draft of the manu-
described as separate species. These are
script, and we thank Lynn Kimsey and an
Vespula rufa (Linnaeus) with V. intermedia
anonymous referee for their reviews.
(du Buysson) its Nearctic synonym, V. aus-
triaca with V. infernalis (de Saussure) in the
Nearctic,Dolichovespulanorwegica(Fabricius) REFERENCES
withD.albida(Sladen)intheNearctic,andD.
Akre, R.D., A. Greene, J.F. MacDonald, P.J.
adulterina (du Buysson) with D. arctica
Landolt, and H.G. Davis. 1981. The yellow-
(Rohwer) in the Nearctic. In view of our
jackets of America North of Mexico. United
present findings it would seem that reinvesti-
States Department of Agriculture Handbook
gation of the status of all these forms is 552: 1–102.
warranted. Archer, M.E. 1989. A key to the world species of
Another Holarctic species was recognized the Vespinae (Hymenoptera). ResearchMono-
by Matsuura and Yamane (1990) and graphs of the College of Ripon & York St.
Pekkarinen (1995), who treated the nearctic John 2.York.
Dolichovespula norvegicoides (Sladen) and the Banks, N. 1920. A rare pamphlet (Hym., Lep.,
Neur.).Entomological News 31: 176.
palearctic D. pacfica (Birula) as subspecies of
Bequaert, J. 1930. On the generic and subgeneric
D. norvegicoides. In this case, a diagnostic
divisions of the Vespinae (Hymenoptera).
difference between D. norvegicoides and D.
Bulletin of the Brooklyn Entomological
pacifica, the elevation between gena and Society25: 59–70.
clypeus, was adduced by Archer (1989) (and Bequaert, J. 1932. A tentative synopsis of the
confirmed by Pekkarinen, 1995). At present hornets and yellow-jackets (Vespinae; Hyme-
2010 CARPENTER AND GLARE: MISIDENTIFICATION OFVESPULA ALASCENSIS 7
noptera)ofAmerica.EntomologicaAmericana Matsuura,M.,andS.Yamane.1990.Biologyofthe
(newseries) 12(2):71–138. vespine wasps. Berlin: Springer-Verlag, [trans-
Buysson, R.du. 1905 (1904). Monographie des lationofthe1984book‘‘Comparativeethology
gueˆpes ou Vespa. Annales de la Socie´te´ Ento- of the vespinewasps,’’ published byHokkaido
mologiquedeFrance73(3):485–556,565–634. University Press, Sapporo,in Japanese]
Carpenter,J.M.,andJ.Kojima.1997.Checklistof Miller,C.D.F.1961.Taxonomyanddistributionof
the species in the subfamily Vespinae (Insecta: Nearctic Vespula. Canadian Entomologist 93
Hymenoptera: Vespidae). Natural History Supplement 22:1–52.
Bulletin ofIbaraki University1: 51–92. Packard, A.S. 1870. Notice of Hymenoptera and
Donovan, B.J. 1984. Occurrence of the common nocturnal Lepidoptera collected in Alaska by
wasp, Vespula vulgaris (L.) (Hymenoptera: W.H. Dall, Director of the Scientific Corps of
Vespidae) in New Zealand. New Zealand the Western Union Telegraph Company.
Journalof Zoology11: 417–427. Transactions of the Chicago Academy of
Sciences 2: 25–32.
Eck, R. 1999. Zur Populationsgliederung und
Pekarrinen, A. 1995. Geographic variation and
Zoogeographie von Vespula vulgaris Faktoren
taxonomy of the species of Dolichovespula in
der intraspezifschen Merkmalsa¨nderungen
the boreal zone of the Holarctic region
(Insecta: Hymenoptera: Vespidae). Entomolo-
(Hymenoptera, Vespinae). Acta Zoologica
gische Abhandlungen (Dresden) 58 (17):
Fennica 199: 61–70.
309–342.
Rohwer,S.A.1926.Remarksonthenameofoneof
Glare, T. 2004. All wasps are not the same.
our common yellowjackets. Proceedings of the
Stowaways4: 2.
Entomological Society of Washington 28:
Goloboff, et al. (2003). Improvements to resam-
93–94.
plingmeasuresofgroupsupport.Cladistics19:
Saussure, H.de. 1853–58. Monographie des gueˆpes
324–332.
sociales ou de la tribu des vespiens. Paris:
Goloboff, P., J.S. Farris, and K.C. Nixon. 2008.
Masson.
TNT,afreeprogramforphylogeneticanalysis.
Saussure, H.d.e. 1857. Bemerkungen u¨ber die
Cladistics 24:1–13.
Gattung Vespa, besonders u¨ber die amerika-
Henry,J.M.,A.K.Raina,andR.L.Ridgway.1990.
nischen Arten. Stettiner Entomologische
Isolation of high molecular-weight DNA from
Zeitung 18: 114–117.
insects.Analytical Biochemistry 185: 147–150.
Simon, C., et al. (1994). Evolution, weighting, and
Jacobson, R.S., R.W. Matthews, and J. phylogenetic utility of mitochondrial gene
MacDonald. 1978. A systematic study of the sequences and a compilation of conserved
Vespula vulgaris group with a description of a polymerase chain reaction primers. Annals of
new yellowjacket species in Eastern North the Entomological Society of America 87:
America (Hymenoptera: Vespidae). Annals of 651–701.
the Entomological Society of America 71: Thomson, G.M. 1923. Naturalized animals and
299–312. plants. New Zealand Journal of Science and
Kirby, W. 1837. Fauna Boreali-Americana. Part Technology 6 (4):223–231.
IV.TheInsects.Norwich,CT:JosiahFletcher. Thompson, J.D., T.J. Gibson, F. Plewniak, F.
Linnaeus, C. 1758. Systema naturae per regna tria Jeanmougin, and D.G. Higgins. 1997. The
naturae, secundum classes, ordines, genera, ClustalX windows interface: flexible strategies
species, cum characteribus, differentiis, syno- for multiple sequence alignment aided by
nymis, locis. 10th ed. Holmiae [Stockholm]: quality analysis tools. Nucleic Acids Research
Laurentii Salvii. 24: 4876–4882.
Complete lists of all issues of the Novitates and the Bulletin are available at World Wide
Websitehttp://library.amnh.org/pubs.Inquireaboutorderingprintedcopiesviae-mailfrom
[email protected] or via standard mail from: American Museum of Natural History,
Library—ScientificPublications,CentralParkWestat79thSt.,NewYork,NY10024.TEL:
[phone]. FAX: [phone].
ThispapermeetstherequirementsofANSI/NISOZ39.48-1992(PermanenceofPaper).