A new species of Elasmosoma ruthe (Hymenoptera: Braconidae: Neoneurinae) from the northwestern United States associated with the western thatching ants, Formica obscuripes forel and Formica obscuriventris clivia creighton (Hymenoptera: Formicidae)

2007·4.1 MB·

by  Scott R Shaw

PROC. ENTOMOL. SOC. WASH.
109(1), 2007, pp. 1-8
A NEW SPECIES OF ELASMOSOMA RUTHE (HYMENOPTERA:
BRACONIDAE: NEONEURINAE) FROM THE NORTHWESTERN UNITED
STATES ASSOCIATED WITH THE WESTERN THATCHING ANTS,
FORMICA OBSCURIPES FOREL AND FORMICA OBSCURIVENTRIS
CLIVIA CREIGHTON (HYMENOPTERA: FORMICIDAE)
Scott R. Shaw
U.W. Insect Museum, Department of Renewable Resources, University of
WY
Wyoming, 1000 East University Avenue, Laramie, 82071-3354, U.S.A. (e-mail:
[email protected])
—
Abstract. A new species of neoneurine braconid, Elasmosoma michaeli., is
described from Oregon, Washington, and Wyoming (Yellowstone National Park).
The new species is an imagobiont endoparasitoid of adult workers of Formica
obscuripes Forel in Washington and Wyoming. This species was recorded as
a parasitoid ofFormica obscuriventris clivia Creighton workers in Oregon. This is the
first record of Elasmosoma attacking Formica obscuripes.
Key Words: new species, Neoneurinae, Elasmosoma, Oregon, Washington,
Wyoming, Yellowstone National Park, ant parasites, Formica
obscuripes Forel
The braconid subfamily Neoneurinae (1979). Shaw (1985) examined Elasmo-
comprises a small group ofrelatively rare soma petukms using scanning electron
parasitic wasps that are associated with microscopy and illustrated diagnostic
worker ants of the Formicafusca L. and features of the head, leg, and metasoma.
F. rufa L. species groups (Shenefelt 1969, The legs of Elasmosoma are unusual in
Shaw 1997). Little is known about their having vestigial tarsal claws and greatly
biology but neoneurine species have long enlarged pulvilh (Shaw 1985: fig. 6). Egg
been assumed to be endoparasitoids of deposition by neoneurine females into
adult worker Formica ants (Muesebeck the abdomens of adult worker ants has
1941, Marsh et al. 1987, Quicke and van been observed by numerous authors
Achterberg 1990, Shaw and Huddleston (Olivier 1893; Pierre 1893; Wasmann
1991, Poinar and Miller 2002, Poinar 1897; Donisthorpe 1927; Kariya 1932;
2004, Shaw 2004). The North American Shaw 1992, 1993) but the details oflarval
species of the neoneurine genus Elasmo- development have only recently been
soma Ruthe were studied by Muesebeck described (Poinar 2004). The process of
(1922, 1941) and later revised by Hud- oviposition is extremely rapid, requiring
dleston (1976), who recognized five only a fraction of a second. Wasmann
species in the continental United States (1897) reported rearing an adult Elasmo-
and Canada. The North American spe- soma from a cocoon found attached to
cies, their distributions, and known host the abdomen of a dead worker Formica
associations were catalogued by Marsh ant, but only recently has any neoneurine
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
been raised from egg to adult in a labo- specimens appeared to be the same as the
ratory setting (Poinar 2004). Neoneur- putative new species from YNP.
ines fly close to the ground and stay very During the summer of 2002, I had the
near nests (Poinar 2004) and raiding opportunity to return to south YNP for
trails (Muesebeck 1941) of^Formica ants. braconid sampling, now armed with the
They are usually found hovering over, or knowledge that F. obscuripes is the likely
near, ant nests and trails, where they can host ant for this wasp species. Despite
be sampled by aspirator. Although For- expectations that this wasp species would
inica ant mounds are quite common in be rather rare and difficult to locate, it
southern Yellowstone National Park was instead found to be quite common.
(YNP), Wyoming, U.S.A., until recently Within minutes of first arriving in YNP
no neoneurines were known to occur we successfully located and observed
there. Elasmosoma hovering by a F. obscuripes
The 1988 fires in YNP burned mound near the South Gate Ranger
400,000 ha or 45% of the park and Station. Over the next two weeks ob-
attracted international attention Chris- servations were made and samples were
tensen et al. (1989). Following that event obtained for descriptive purposes. The
there was a renewed interest in YNP purpose of this paper is to describe and
biotic communities, and new studies of name this new neoneurine species as
insects were conducted. During the sum- a contribution to continuing studies of
mer of 1990, we operated four Malaise the insect fauna of YNP, and to stimu-
traps in burned and unburned areas of late and facilitate more detailed biolog-
south YNP (Lockwood et al. 1996). Our ical studies of neoneurine wasps.
study found a total of 603 Hymenoptera
Methods and Terminology
species, representing 36 families (from
a total of2,331 hymenopteran specimens Specimens of Elasmosoma were sam-
sampled). Of these, the parasitic wasp pled using a wet-aspirator and preserved
family Braconidae was well represented, directly into 2 dram vials of90% ethanol.
with 109 species found in south YNP. The wasps are intent on chasing ants and
That study also yielded the first record of not too difficult to observe or aspirate.
the ant-associated subfamily Neoneuri- The main difficulty with this sampling
nae in YNP: a single (damaged) speci- method is that they flyclose to themound
men of Elasmosoma, apparently a new so sand and nest debris is easily aspirated
species. Although intriguing, the dam- as well. The other hazard is simply
aged condition of this single specimen working in close proximity to such active
prevented further taxonomic study. and aggressive ants. The samples were
In 1999, I was contacted by Jack moved to clean 90% ethanol, then later
Longino who kindly sent a nice series into 100% ethanol for 24 hours. Dry
of 13 Elasmosoma. These were collected specimens were prepared by placing them
on the campus of Evergreen State in fine wire baskets, soaking in chloro-
College (Thurston County, Washington) form for 20 minutes, and air-drying un-
in association with the western thatching der a fume hood, prior to point-mount-
ant, Formica obscuripes Forel. He later ing. This method approximates theresults
sent two specimens from Long Beach ofcritical-point-drying at less expense.
(Pacific County, Washington), also Species covered in this paper can be
found attacking workers ofF. obscuripes. identified as members of the subfamily
Finally, in 2001, George Poinar sent Neoneurinae using the subfamily key by
a series of Elasmosoma collected in Sharkey (1997). The definition oi^Elasmo-
Lincoln County, Oregon. All of these soma follows that of Muesebeck (1922,
VOLUME NUMBER
109, 1
1941), Huddleston (1976); Marsh et al. suslength;hypopygiumoffemaledensely
(1987) and Shaw (1997). Specimens can be setose, apical margin of hypopygium
narrowly and deeply U-shaped; clypeus
determined as Elasniosoina by using the
paleyellow orwhite .... E. michaeli, n.sp.
keys of Marsh et al. (1987) and Shaw - Hind tibial spur relatively longer, longer
(1997). Morphological terminology fol- spur just slightly shorter than entire
lows that of Shaw (1992, 1997) and basitarsus length; hypopygium offemale
Sharkey and Wharton (1997). Wing vein sparsely setose, apical margin of hypo-
pygium broadly and shallowly V-shaped;
terminology agrees with the system pro-
clypeus black
posed by Sharkey and Wharton (1997) for E. pergandei Ashmead and E. bakeri
theManualoftheNew WorldGeneraofthe Ashmead {E. bakeri Ashmead is known
A
Family Braconidae. labeled diagram of only from male specimens. It is not
wing veins for Elasmosonia was given by certain if it can be maintained as a distinct
(Sharkey and Wharton 1997: 37, fig. 18). species from E. pergandei Ashmead.]
The following key to species is modified Elasmosoma michaeli Shaw, new species
after Huddleston (1976) and based on
(Figs. 1-6)
examination of the holotypes of all in- —
cluded species. Because of sexual varia- Critical diagnosis. Clypeus pale yel-
tion, Elasmosoma species are most reliably low or white; distance between lateral
identified based on female specimens. ocellus and eye greater than twice the
diameter of lateral ocellus; hind tibial
Key to Females ofthe Described
spur relatively short, longer spur just
North AmericanSpeciesofElasmosoma slightly longer than V2 basitarsus length;
1 Metasoma elongate, conspicuously longer first metasomal tergite about as wide as
(1.5-2.0 times) than head and mesosoma long; hypopygium of female densely
combined,stronglycompressed;tergite3at setose, apical margin of hypopygium
leastaslongaswide . . E. schwarziAshmead narrowly and deeply U-shaped; hypopy-
- Metasoma at most equal in length to
head and mesosoma combined, not com- gium of female flat, not folded along
pressed; tergite 3 conspicuously wider midline or—compressed.
than long 2 Female. Habitus of adult female as
2(1) Distance between lateral ocellus and eye illustrated by (Poinar 2004: fig. 6c). Body
abouttwicediameteroflateralocellus, or length 2.0 mm.
slightly less; hypopygium of female fol-
ded along midline and compressed Color: Body mostly black, except
E. petulans Mue. s.ebeck metasomal tergum 2 sometimes with
- Distance between lateral ocellus and eye yellowish brown along anterior and
greater than twice diameter of lateral posterior margins. Clypeus and labrum
ocellus; hypopygium of female flat, not pale yellow to white. Mandible white
3(2) Hfiolnddedtiablioalngspmuirdlilnoengoerrctohmapnrehsisneddbas.i.-. 3 except tip reddish brown. Compound eye
tarsus; first metasomal tergite wider than silvery gray. Front legs mostly white,
long; hypopygium offemale arising near front femur and tibia infused with pale
middle of metasoma, hypopygium con- yellow, apical tarsomere reddish brown.
cealed below metasoma Middle and hind legs with similar color
E. vigilans Cockerell
- Hind tibial spur at least slightly shorter pattern except darker, more extensively
than hind basitarsus; first metasomal infused with pale yellow to yellowish
tergite about as wide as long; hypopy- brown. Apex of hypopygium pale yel-
gium of female arising in distal 1/3 of lowish brown to white.
metasoma, hypopygium wide and flat, Head: Compound eyes, extremely
with lateral lobes projectingbeyond sides
ofmetasoma 4 large and strongly converging anteriorly
4(3) Hind tibial spur relatively short, longer and ventrally (Fig. 2). Malar space ex-
spurjust slightly longer than 'A basitar- tremely narrow. Width of malar space
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-2. Elanwsonui niichaeli. L Metasoma, lateral view showing ovipositor and densely setose
hypopygium. 2, Head and antenna, lateral view.
near anterior condyle ofmandible slight- Mesosoina: Mesosoma very compact.
ly less than width of mandible near tip. Mesosoma length in profile about 1.6X
Face narrow, width of face slightly less head length. Mesonotum finely coria-
than clypeus width. Antenna short ceous and densely covered with micro-
(Fig. 2), flagellum length equal to meso- scopic setae, no trace of notauli. Prono-
soma length (in lateral profde). Antennal tum finely coriaceous. Dorsoposterior
scape short, slightly shorter than first lobe of mesopleuron (Figs. 3^) smooth
flagellomere. Pedicel Vi length of scape. and shining, devoid of sculpture and
Flagellum with 10 compact flagello- setae. Remainder of mesopleuron with
meres, each of similar size and just dense, fine, foveolate sculpture (Fig. 4).
slightly longer than wide, except apical Sternaulus not clearly differentiated
(IQth) flagellomere about 2X the length of from broad patch of foveolate meso-
first flagellomere. Apical flagellomere pleural sculpture. Propodeum coarsely
tapers to pointed apex. Apical flagello- rugulose. Hind tibial spur relatively
mere partly divided dorsally, thus ap- short, longer spur just slightly longer
pearing to have 1 1 flagellomeres in dor- than '/2 basitarsus length (Fig. 5). Hind
sal view but only 10 flagellomeres in tarsus (Fig. 6) with pulvillus greatly
lateral or ventral view. Ocellar triangle enlarged, tarsal claws greatly reduced
equilateral. Distance between lateral and nearly vestigial.
ocellus and eye slightly greater than Wings: Venation extremely reduced, as
twice the diameter of lateral ocellus. illustrated in (Shaw 1997: fig. 2). Pattern
Surface sculpture of head very finely of venation highly distinctive for genus
coriaceous. but with little discernable variation be-
VOLUME NUMBER
109, 1
^5^"??ws^v-%
Figs. 3-6. Elamosoma michaeli. 3, Mesosoma, lateral view. 4, Mesopleuron, lateral view showing
detail ofsurface sculpture. 5, Hind leg, lateral view showing apex ofhind tibia and length oftibial spurs
relative to basitarsus. 6, Apex ofhind tarsus showing vestigial tarsal claws and modified pulvillus.
tween species. Forewing length 1.0 mm. black. Antenna longer, thicker, and more
Costa and pterostigma dark brown to densely setose than in female, with 12
black. Remaining veins light brown. flagellomeres. Each flagellomere about
Metasoma: Metasoma about equal in 2x longer than wide, and densely setose.
length to head and mesosoma combined, Flagellum tapering gradually from base
not compressed. Tergite 3 conspicuously to apex, apical flagellomere (12th) only
wider than long. Metasomal terga finely about Vi as thick as first flagellomere.
coriaceous, sparsely setose, setae not Compound eye smaller than in females,
obscuring surface sculpture. Hypopy- and malar space broader. Width of
gium of female densely setose, flat, not malar space about equal to length of
folded along mid-line or compressed. antennal scape. Male genitalia mostly
Apical margin of hypopygium narrowly retracted into metasoma and not visible,
and deeply U-shaped. Lateral lobes of except for small narrow parameres,
hypopygium extensively desclerotized about equal in size to apical hind
and thin, nearly transparent. Ovipositor tarsomere. —
(Fig. 1) about as long as apical tergite, Holotype. Female, United States of
thin, sha—rply pointed, and flexible. America: Washington, Thurston Coun-
Male. Similar to female except more ty, Evergreen State College, 20 Septem-
extensively black, lacking light colors on ber 1999, Jack Longino, #4191, at
clypeus and metasoma, and hind coxa Formica obscuripes nest. Deposited in
—
,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
University of Wyoming Insect Museum, in the vicinity. It is possible that some
Laramie, Wyoming. workers of F. propinqua wandered near
—
Paratypes. Oregon: 16 2, Lincoln the F. obscuripes mound. Another possi-
County, between Newport and Wald- bility is that F. propinqua were temporary
port, 8 September 2001, aspirated near social parasites in nests of F. obscuripes
ant nest, George Poinar, coll.; 2 $, 1 S and that YNP mounds were mixed nests.
Waldport, 7 August 2001, George Poi- In all three cases the ants usually make
nar, coll. Washington: 8 $, 5 S, similar, very large, thatched dome nests.
Thurston County, Evergreen State Col- It seems likely that the parasitoid attacks
lege, 20 September 1999, Jack Longino, any of these host ants with similar
#4191, at Formica obscuripes nests; 2 2, behavior. The behavior of adult E.
Pacific County, Long Beach, #4160, 46° michaeli in YNP was the same as de-
20.8'N, 124° 03.7'W, attacking Formica scribed by Poinar (2004) in Oregon.
obscuripes; 9 2, Yakima County, Pleas- Elasmosoma michaeli adults were quite
ant Valley Campground, Highway 410, active on sunny mornings between 9am
46° 57'N, 121° 20'W, 1,000 m, 30 and 1 1am, with activity tapering off at
August 2003, J. Longino, #5136. Wyo- midday.
ming: 21 2,1 S, Yellowstone National Discussion. Elasmosoma michaeli is
Park, near South Entrance, 1 August most similar to E. pergandei. Females of
2002, Michael J. Shaw and Scott R. both species have a broad, flat hypopy-
Shaw coll., aspirated by ant nest, USDL gium that projects beyond the sides and
NPS YNP Scientific Research Permit apex of the metasoma. The lateral lobes
Number YELL-2002-SCL5310; 20 2, of the hypopygium are extensively de-
same data except collected 30 July sclerotized and thin. When viewed from
2002. Deposited in University ofWyom- above these hypopygial lobes are rather
ing Insect Museum, Laramie, Wyoming transparent and windowlike. Elasmo-
and National Museum of Natural His- soma michaeli can be distinguished by
tory, Smithsonian Institution, Washing- the hypopygium of the female being
ton, D.C. — more densely setose (Fig. 1), and the
Etymology. This species is named for apical margin of the hypopygium being
my son, Michael Joseph Shaw, for his narrowly and deeply U-shaped. The
assistance during the 2002 field season in hypopygium of E. pergcmdei is less
Yellowstone and Grand Teton. densely setose, with the apical margin
—
Biology. The egg-laying behavior, more broadly V-shaped, and having
embryonic development, larval develop- a distinct sharp notch at the midline
ment, and cocoon-forming behavior of where the hypopygium margin meets
this species was described by Poinar the ovipositor base. Elasmosoma michaeli
(2004) as "an undescribed species of is also similar to E. bakeri (a species
Elasmosoma near pergcmdei Ashmead." known only from male specimens). Elas-
Poinar reported this species as attacking mosoma michaeli can be distinguished
workers of Forf7iica obscuriventris clivia from both E. pergandei and E. bakeri by
Creighton workers in Oregon. In Wash- the shorter tibial spurs (Fig. 5). The
ington and Wyoming it was found tibial spurs are much longer in E. per-
attacking workers ofFormica obscuripes. gandei and E. bakeri (nearly as long as
According to Jack Longino, the majority the entire basitarsus). Elasmosoma mi-
YNP
of ants in the samples are F. chaeli females can be distinguished from
obscuripes but a few individuals from those of E. pergandei by the pale yellow
the same mound resembled F. propinqua or white clypeus. The clypeus is black
Wheeler. There were other ant mounds in females of E. pergandei. The cly-
VOLUME NUMBER
109, 1 7
peus color of E. bakeri females is not Donisthorpe, H. St. J. K. 1927. The Guests of
known. British Ants, Their Habits and Life Histories.
G. Routledge and Sons, Ltd., London. 244pp.
Acknowledgments Huddleston, T. 1976. A revision of Elasmosoma
Ruthe (Hymenoptera: Braconidae) with two
Thank you to Christie Hendrix and new species from Mongolia. Annales Histor-
Nicki Tippets for their assistance with ico-Naturales Musei Nationalis Hungarici 68:
scientific research and collecting permits 215-225.
in Yellowstone National Park and Grand Kariya, M. 1932. Observations on a braconid
species parasitic on ants. Konchu Sekai 36:
Teton National Park, respectively. This 298-301 (in Japanese).
work was supported, in part, by a 2002 Lockwood, J. A., S. R. Shaw, and J. M.
research grant from the UW-NPS Re- Struttmann. 1996. Biodiversity ofwasp species
search Station, and by Mclntire-Stennis (Insecta: Hymenoptera) in burned and un-
project number WYO-362-02, braconid burnedhabitatsofYellowstoneNational Park,
Wyoming, U.S.A. Journal of Hymenoptera
wasps in forest ecosystems. Park science
Research 5: 1-15.
ranger Rick Mossman was of great Marsh,P.M. 1979. FamilyBraconidae,pp. 144-313.
assistance by pointing out the location In Krombein, K. V., P. D. Hurd Jr., D. R.
of several ant mounds in southern Yel- Smith, and B. D. Burks, eds. Catalog of
lowstone NP. Special thanks to my son Hymenoptera in America North of Mexico,
fMiiecldhaewlitJh. Sshetatwinfgoruhpis aanssdistmaonncietionritnhge MarsSDh.m,Ci.tP,h.sVMoo.nl,iauSnm.eRI.1n,sSt1hi1ta9uw8t,ipopan.ndPrRe.ssW,haWratsohni.ng1t9o8n7,.
Malaise traps in Grand Teton National An identification manual for the North Amer-
Park and with aspirator sampling in ican genera of the family Braconidae (Hyme-
southern Yellowstone. His keen eyesight noptera). Memoirs ofthe Entomological Soci-
wAdadsitciroucniaall ttohtahnekssucacreess doufethitsoproHjaecntk. Mueseetbyeocfk,WaCs.hiF.ngWt.on1,92N2o..A1r3,ev9i8siopnp.oftheNorth
American ichneumon-flies belonging to the
Harlow for assistance with local arrange- subfamilies Neoneurinae and Microgastrinae.
ments at the UW-NPS Research Station. Proceedings of the United States National
Thank you to Jack Longino and George Museum 61: 1-76.
Poinar for providing wasp specimens, ant . 1941. A new ant parasite (Hymenoptera,
associations, data, and observations from Braconidae). Bulletin of the Brooklyn Ento-
mological Society 36: 200-201.
Washington and Oregon, respectively.
Olivier, E. 1893. [no title]. Annales Societe En-
Stefan Cover (Harvard University) and tomologique France 1893: 71.
Jack Longino (Evergreen College) pro- Pierre, A. 1893. Un parasite des fourmis Elasmo-
vided identifications of host ants from soma berolinense Ruthe. Revue Scientifique
Yellowstone. David Smith arranged the Bourbon Centre France 6: 112-114.
loan of type specimens of Elasmosoma Poinar, G. Jr. 2004. Behavior and development of
Elasmosoma sp. (Neoneurinae: Braconidae:
from the collection of the National Hymenoptera), an endoparasite of Formica
Museum of Natural History, Smithso- ants (Formicidae: Hymenoptera). Parasitology
nian Institution. Zhaojie Zhang, director 128: 521-531.
ofthe University ofWyoming Microsco- Poinar, G. Jr. and J. C. Miller. 2002. First fossil
record of endoparasitism of adult ants (For-
py Facility, assisted with the production
micidae: Hymenoptera) by Braconidae (Hy-
ofscanning electron microscope images.
menoptera). Annals of the Entomological
Society ofAmerica 95(1): 41-43.
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