Description of the female of Acrophotopsis (Hymenoptera: Mutillidae) with synonymy of Sphaeropthalma dirce

2009·4.3 MB·

by  James P Pitts

HYM. RES.
J.
Vol. 18(2), 2009, pp. 205-211
Description of the Female of Acrophotopsis (Hymenoptera: Mutillidae)
with Synonymy of Sphaeropthalma dirce
James P. Pitts and Joseph S. Wilson
Department of Biology, Utah State University, 5305 Old Main Hill, Logan, UT 84322-5305, USA;
[email protected]
—
Abstract. The female of Acrophotopsis campylognatha Schuster is described. Sphaeropthalma dirce
(Fox), known from females only, is transferred to Acrophotopsis and is the senior synonym of A.
eurygnatha Schuster. This represents the first description of females for Acrophotopsis. Most
importantly, the females ofAcrophotopsis canbe diagnosedby the followingunique combination of
characters:havingadistinctbasaltoothonventralmarginofmandibleandatooth-likeprojectionat
the anterior termination of the dorsal mandibular carina; having the mesosoma and second
metasomal tergite moderately punctate to reticulately sculptured and having rasp-like tubercles
situated between the reticulations that are more apparent anteriorly; having the first metasomal
segmentpetiolatewiththesecond;havingthepygidiumlaterallydefinedbycarinaewithgranulate
sculpturing;andhavingthepropodeumandfringesoftergitestwothroughfourwithdistinctwhite
plumose setae.
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Key words. velvet ant, Sphaeropthalminae, Dilophotopsis, ITSl and ITS2
Acrophotopsis Schuster (Hymenoptera: topsis campylognatha occurs in Baja Califor-
Mutilhdae) is an easily recognized genus nia andinthewestern SonoranandMojave
of nocturnal mutillid possessing deeply Deserts of Southern California, while A.
excised mandibles, a flattened hypopygi- eurygnatha occurs in eastern Sonoran Des-
dium with lateral, basal carinae, and ert of Arizona and Mexico, and into the
genitalic parameres that overlap in situ, Mojave Desert as far west as Nevada
while lacking mesostemal processes. The (Ferguson 1967). A third species, A. bergi
genus currently contains four species re- Casal, was added to the genus later and
centlyrevisedbyPitts and McHugh (2002). occurs in central Mexico in the states of
All species ofAcrophotopsis arefound inthe Jalisco, Morelos, and Puebla,Mexico (Casal
southwestern U.S. and Mexico, and are 1967). The last species to be added to the
known only from males (Manley and Pitts genus was A. mickeli Pitts and McHugh,
2002). Although females of Acrophotopsis described from Baja California Sur (Pitts
are unknown, they are presumed to be and McHugh 2002).
active at night similar to males. Nothing The genus is known from a single sex, in
more is known about the biology of part, due to the extreme sexual dimor-
Acrophotopsis. phism that occurs in mutillids (Brothers
R.M. Schuster (1958) described the genus 1995). Nocturnalvelvetantmales are easily
Acrophotopsisbased onmales oftwo species collected in light traps, while females are
of previously undescribed nocturnal rarely collected. Sex associations are fur-
Sphaeropthalmini from the Nearctic re- ther complicated by great morphological
gion, A. campylognatha Schuster and A. similarity among species. This makes as-
eurygnatha Schuster. These two species are sociating sexes nearly impossible based on
sympatric in the Mojave Desert. Acropho- examination of museum specimens and
206 Journalof H^-mexopter-a Research: Festschrift Honoring Roy Sn-elling
sex associations made by catching pairs in (Thompson, et al. 1994) and intraspecfic
copula are rare. More advanced molecular and interspecific genetic distances were
techniques, however, can be used to make calculated from these alignments. DNA
sex associations using species-specific ge- sequences were deposited in GenBank
netic loci (Pilgrim and Pitts 2006; Pitts et al. (Accession Nos. GQ223230-GQ223237).
—
2007; Pilgrim et al. 2008). Taxonomic methods. The following acro-
The purpose of this study is to associate nyms are for institutions or collections
the females with the two species of housing the material discussed in the
Acrophotopsis found in the United States, current study: Department of Entomology,
A. campylogjiatha and A. eurygnatha. Academy of Natural Sciences, Philadel-
phia, Pennsylvania, U.S.A. (ANSP); De-
MATERIALS AND TERMINOLOGY
partment of Entomology, California Acad-
—
Trapping methods. Field studies were emy of Sciences, San Francisco, California
conducted throughout the Southwestern (CASC); and Entomological Museum, De-
U.S. during the summers of 2005-2008 to partment ofBiologv, Utah State University,
collect fresh specimens of both sexes of Logan, Utah, U.S.A. (EMUS).
nocturnalvelvet ants to attemptassociating We adopt the following notation after
the sexes using molecular techniques. Male Ferguson (1967) for punctures in the order
and female nocturnal mutillids were col- of decreasing coarseness: reticulate, coarse,
lected at 60 field sites across the South- moderate, small, fine and micropunctate.
western U.S. Micropunctate refers to punctures that are
Specimens were collected using black extremely shallow and do nothave vertical
light and fluorescent lantern traps, and by waUs or sharp margins. Small refers to
hand. Specimens collected with light traps punctures that do have slight vertical walls
were captured in soapy water and trans- and are separated by at least 5X their
ferred into 95° ethanol, while all hand- diameter. We use the term "simple setae"
collected specimens were placed directly for setae that are smooth and do not have
into 95% ethanol. barbed surfaces. "Brach\q5lumose setae"
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Molecular methods. The two internal refers to setae ^vithbarbs that are less than,
transcribed spacers (ITSl and ITS2) were or equal to, the diameter of the shaft at the
sequenced for representatives of each attachmentofthebarb. The term "plumose
available species and sex, sequences were setae" is used for setae that have longer
aligned, and females were associated with barbs. The term "tibial spurs" is used
malesbased onidentical or nearly identical instead of "calcaria." The term"paramere"
DNA
sequences for those loci (i.e., ver\^ is used instead of "gonoforceps" to remain
small genetic distances). The methods consistent with previous mutilHd litera-
proposed by Pilgrim and Pitts (2006) were ture. The acronyms T2, T3, etc., denote the
followed for performing sex associations. second, third, etc., metasomal tergites,
ITSl and ITS2 were sequenced for at least respectively. Similarly, S2, S3, etc., signifies
one female of each morphospecies and the second, third, etc., metasomal stemites,
several male specimens of each described respectively.
species. PCR was used to amplifv' the ITSl
and ITS2 regions of the nuclear genome Acrophotopsis Schuster
using the molecular protocols described in
DNA AcrophotopsisSchuster1958. Ent. Amer. (n. s.) 37:
Pilgrim and Pitts (2006). samples 4(inkey),61,male,ly^pespecies:Acrophotopsis
were sequenced in both directions and eurygnathus Schuster, orig. desig.
combined using Sequencher 4.0 (Gene
—
Code Corp., Ann Arbor, MI). DNA se- Diagnosis of females. The females of
W
quences were aligned using Clustal Acrophotopsis can be diagnosed by the
x^.cEER2. :XV 207
—
rial view. 2. Dorsal \ieT.v c: e5-:5:r-_a. 3. Dcrsa^ '.~e-
secondmetasomal rer^Te ^. J:
following unique combination of charac- multitude of longitudinal ridges (Hg. 3);
ters: they are nocturnal with reddish the pygidimn is granulate and defined
brown to brown integument; the com- laterally by carinae (Fig. 4); and the pro-
pound eyes axe only slightly ovate podeum and fringes of tergites t^vo
(Fig. 1); the mandible has a distinct basal through four consists of distinct white
tooth on ventral margin and a tooth-like plumose setae ^Figs 2 and 3"^.
projectionattheanteriortermination ofthe
Acrophotopsis campvlognatha Schuster
dorsal carina; the mesosoma is longer than
broad and only slightly wider at the
mesonotal spiracle than elsewhere
(Hg. 2); the first metasomal segment is
petiolate ^\-ifh the second (Bgs 1 and 2);
the mesosoma and second metasomal <C\SC . —
tergite are moderately punctate to reticu- Diagnosis cf female. ^The female of A.
lately sculptured and have rasp-like tuber- :-'.:: r t:/i5 can be separated from that
clessituatedbet^seenthereticulations^\"ith :: - - .: by the mesosoma and second
the tubercles being more apparent anteri- metasomal tergite being reticulately sculp-
orly than posteriori}^ (Figs 2 and 3); the tured and the setae on the dorsum of the
punctures, at least on the anterior half of mesosoma and centrally on the second
the second tergite, have lateral margins tergite being whitish and onl\- sUgJitly
that extend posterioriy apj>earing as a tinned reddish-bro^NTi.
X
208 Journalof Hymenoptera Research: Festschrift Honoring RoySnelling
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Description of female. Coloration and tubercles posteriorly. Propleuron punctate
Setal Pattern: Body reddish-brown to anteriorly. Humeral angle dentate. Epaulet
brown. Mandibular apicesblack. Flagellum prominent. Scutellar scale absent. Meso-
and legs yellow to dark yellow. Setae pleuron punctate medially. Mesostemum
sparse in general, not concealing sculpture. with low transverse tubercle present medi-
Head, pleurae, and vertical face of propo- allyjust anterior to mesocoxa. Metasternum
deum with decumbent and erect white tridentate, median tooth ~4X as long as
brachyplumose setae. Dorsum of meso- lateralteeth.Extremeventralregionoflateral
soma with decumbent and erect brachy- margin of propodeum punctate. Mid- and
plumose setae; setae white, but slightly hind-tibiaewithtworowsofspinesonouter
tinged reddish-brown. Propodeal dorsum margin and eachwithpairoftibial spurs.
—
and vertical face with distinct, sparsely- Metasoma. Segment 1 distinctly petio-
spaced, white plumose setae. Tl covered late with segment 2. Tl with small sparse
with both decumbent white plumose setae punctures. T2 with large reticulations on
and erect white brachyplumose setae. T2 anterior half with tubercles situated be-
with erect white brachyplumose setae, tween reticulations, becoming more
tinged light brown centrally; sparse short sparselypunctate posteriorly. Reticulations
white plumose setae present on posterior and punctureswithlateral margins extend-
third. T2-4 and S2-S5 with fringe of white ing posteriorly forming longitudinal car-
plumose setae; fringe becoming sparser on inules, evenin sparselypunctate region. T2
more apical tergites. Fringe of T5 medially withfeltline;length 0.20X lengthoftergite.
with light golden brachyplumose setae, T3-T5 shagreened. T6 with distinct pygid-
laterally with white plumose setae. Legs ial area defined laterally by weak carinae;
with white brachyplumose setae. surface granulate. S2 with slight anterome-
—
Head. Head rounded posteriorly, not as dian tumid region. S2-S5 with punctation
wide as mesosoma, densely punctate. Eye similar to tergites.
slightly ovate, distance fromposteriorman- Length: —5.6 mm.
—
dibular articulation —2.5X length of pedi- Dna voucher specimen data. California, San
cel. Clypeus protruding anteriorly, poster- BernardinoCo.:5 mi SBarstow, 19,30.May.2005,
omedially produced into low triangular E.E. & K.A. Williams, KW14; 1 S, 30.May.2005,
swelling with central tubercle. Antennal E.E. & K.A. Willi—ams, JP324 (EMUS).
scrobe without dorsal carina. Antennal Distribution. Acrophotopsis campylog-
tubercle glabrous, except with carinate natha is present in the southern regions of
apical margin. Flagellomere I ~1.2X length the Mojave Desert of California and into
of pedicel. Flagellomeres II-III ~1.0-1.2 the Sonora—n Desert of Baja California.
length of pedicel. Flagellomeres I-III sub- Remarks. This sex association is based on
A
equal in length. Flagellomeres II-X pro- molecular data. total of 1,432 base pairs
duced apically on ventral side; appearing (504 bpforrrSland928 bpfornS2)wasused
crenulate.Mandiblebidentateapically.Dor- toassociatethemaleandfemaleofthisspecies.
sal mandibular carina with tooth-Uke pro- Both the rrSl and ITS2 loci are identical
jection at anterior termination of carina. betweenthemaleandfemaleandthisdistance
Ventralmandibularmarginwithlargebasal is much smaller than the interspecific genetic
tooth; lacking excision apical to ventral distancebetweenA.campylognathaandA.dirce
tooth. Genal carina absent. (8% forrrSl; 11% forITS2).
—
Mesosoma. Mesosoma wider anteriorly
Acrophotopsis dirce (Fox)
than posteriorly, longer than broad. Meso-
soma reticulate on dorsum, some reticula- Mutilla dirce Fox, 1899. Amer. Ent. Soc, Trans.
tions with margin appearing tuberculate; 25: 257, female. Holotype: Arizona, Tucson,
punctures becoming larger and without coll. Wickham, type no. 4651 (ANSP).
Volume 18, Number1, 2009 209
Acrophotopsis eurygnathus Schuster 1958. Ent. length of pedicel. FlageUomeres I-III sub-
Amer. (n. s.) 37: 11 (in key), 65, male. equal in length. FlageUomeres II-X pro-
Holotype: USA, Arizona, Gila Co., Globe, duced apicaUy on ventral side; appearing
8.Vn.l949, coll. Werner & Nutting (CASC). crenulate.MandiblebidentateapicaUy.Dor-
NEW SYNONYM.
sal mandibular carina with tooth-like pro-
Diagnosis of female.—The female of A. jection at anterior termination of carina.
Ventralmandibularmarginwithlargebasal
dirce can be separated from that of A.
campylognathabythemesosoma and second tooth; lacking excision apical to ventral
tooth. Genal carina absent.
metasomaltergitebeingonlydenselypunc- —
tate (Fig. 3) and the setae on the dorsum of Mesosoma. Mesosoma wider anteriorly
the mesosoma and centrally on the second than posteriorly, longer thanbroad (Fig. 2).
Mesosomaconfluentlypunctateondorsum,
tergite being distinctly re—ddish-brown.
Redescription offemale. Coloration and some reticulations with margin appearing
Setal Pattern: Body reddish-brown to tuberculate;puncturesbecomingsomewhat
reticulate posteriorly, butwithout tubercles
brown. Mandibular apices black. Flagel-
lum, scape and legs yellow to darkyellow. (Fig. 2). Propleuron anteriorly, meso-
pleuron medially, and extreme ventral
Setae sparse in general, not concealing
region of lateral margin of propodeum
sculpture (Figs 1 and 2). Head, pleurae,
and vertical face of propodeum with punctate. Humeral angle dentate. Epaulet
prominent. Scutellarscale absent. Mesoster-
decumbentand erectwhitebrachyplumose
num with low transverse tubercle present
setae (Fig. 2). Dorsum of mesosoma with
mediallyjustanteriortomesocoxa.Metaster-
decumbent and erect brachyplumose setae
numtridentate,mediantooth~4X aslongas
(Fig. 2); setae reddish-brown. Propodeal
dorsum and vertical face with distinct lateral teeth. Mid- and hind-tibiae with two
rows of spines on outer margin and each
sparsely spaced white plumose setae
(Fig. 2). Tl covered with both decumbent withpair of—tibial spurs.
white plumose setae and erect white bra- Metasoma. Segment 1 distinctly petio-
chyplumose setae (Fig. 2). T2 with erect late with segment 2 (Fig. 1 and 2). Tl with
small sparse punctures. T2 confluently
whitebrachyplumose setae, reddishbrown
punctate on anterior half with tubercles
centrally; sparse short white plumose setae
situated between reticulations, becoming
present on posterior third (Fig. 3). T2-4
(Figs 3 and 4) and S2-S5 with fringe of sparselypunctate posteriorly (Fig. 3). Retic-
whiteplumose setae; fringebecomingmore ulations andpunctureswithlateralmargins
extending posteriorly forming longitudinal
sparse on apical tergites. Fringe of T5
carinules, but not in sparsely punctate
medially with light golden brachyplumose
setae, laterally with white plumose setae. region. T2withfeltline;length0.20X length
of tergite. T3-T5 shagreened. T6 with
Legs with white brachyplumose setae.
Head.—Head rounded posteriorly, not as distinct pygidial area defined laterally by
wide as mesosoma, densely punctate. Eye weak carinae; surface granulate (Fig. 4). S2
with slightanteromediantumid region. S2-
slightlyovate, distance fromposteriorman-
S5 with punctation similar to tergites.
dibulararticulation —2.5X lengthofpedicel Length: ~7 mm.
(Fig. 1). Clypeus protruding anteriorly, —
posteromedially produced into low trian- SanDtnaaCnviozucChoe.r: 5spkecmimWenPdeantaa.BlUanScAa:LaAkreizaotnaR,t
gularswellingwithcentraltubercle. Anten-
39;Atascosa Mts, 1 ^,3/7.May.2004,M.E. Irwin
nal scrobe without dorsal carina. Antennal & F.D. Parker, JP84 (EMUS). MEXICO: Sonora,
tubercle glabrous, except with carinate Rancho Palo Injerto, 20 km E Alamos: 1 o,
apical margin. Flagellomere I ~1.2x length JP680, Jun.2006, 1 ^^ 28/31.Jun.2007,JP686 M.E.
of pedicel. Flagellomeres Il-m ~1.2-1.3x InAdn & F.D. Parker (EMUS).
210 TouRXALOF H>:mexopter-a Research: FestschriftHoxorixg Ro^. Sxellixg
—
Materinl examined. USA: Arizona, Cochise There are other nocturnal females that
Co.: Leslie Canyon MX'R, 1 o, 19.May.2000, have a subset of these characters that could
W.R. Radke (EMUS);Nevada,\'yeCo.: Mercur\-: be confused \vith Acrophotopsis, but all lack
1 9, 12.Aug.l964, 1 9, 14.Aug.l964, 1 c, the scattered tubercles on the mesosoma
13.Jim.l961 (B\U'C); New Mexico, Hidalgo Co.: and metasoma. Specifically, Sphaeropthalnia
Stone Cabin, U-Ranch, 1 9, 15.Jul.l977, Muma & laodamia (Fox) and Stethophotopsis maculata
Packard (EMUS); Socorro Co.: SeviUeta MN'R, 1
Pitts both have longitudinal carinae on the
9, 26.Oct.1992 (E—MUS).
second tergite, but Sp. laodamia has a
Distribution. Acrophotopsis dirce has
distinct dorsal carina on the scrobe, \vhile
been collected from the Mojave Desert of
Xevada to the Sonoran Desert of Arizona St. maculata lacks this carina (Pitts and
Manley 2002). Additionally, Sp. laodamia
and Mexico.
— and St. maculata have neither a large
Remarks. The sex association is based
\'entral tooth, nor a dorsal tooth on the
on the similarities of the female described
mandible. They also lack tubercles on the
here %vith the female associated with A.
mesosoma and second tergite, and lack a
campylogiiatJm and the knowTi distribution
laterally defined pygidium. The male of Sp.
of A. eiin/giiatha. The t\~pe specimen of A.
laodamia is unkno^vn, but this species
dirce ^vas collected in Tucson, Arizona, and
seems to be placed in the correct genus.
does not differ from other specimens from
Lastly, Sp. unicolor (Cresson) has a dorsal
farther east in Arizona and Xe^v Mexico.
tubercle on mandible, but lacks a large
These specimens are found in the same
ventral tooth and has a sessile attachment
areas as the A. euri^giiatha male. WTiile no metasomal segment 1 to metasomal seg-
females ^vere available for molecular com- ment
2.
parisons, the available intraspecific genetic
It is rather difficult to differentiate the
distances bet^veen males was lo^s' (0.0- species of Acrophotopsis based on females.
0.3%forTrSl).
This is not surprising given the difficult^' of
DISCUSSIOX separating the females of other related taxa
(e.g. Pitts et al. 2004; Pitts 2006). The t\vo
These are the first females to be associ- Acrophotopsis species apparently differ
ated \vith this genus. Only three nocturnal only in the coarseness of the sculpturing
genera in the Xearctic region, Aca?ithopho- onthe dorsumofthemesosomaandsecond
topsis Schuster, Laminatilla Pitts, and Schus- tergite ofthe metasoma, as^vell as in subtle
terphotopsis Pitts remain kno^vn only from a setal coloration differences in these same
single sex. Ferguson supposedly associated areas. The two species do not overlap
a female ^vith A. euri/gnatha during his greatlv in range, and, therefore, locality
study at the Xe\"ada Test Site, ^vhich ^vas data can also be a good indicator for
cr\-ptically listed in .Allred (1973), but he identifying the females. The males of these
apparently never described the female and species, on the other hand, are not difficult
we have been unable to find the specimens to distinguish and differ in several charac-
referred to in Allred's manuscript. ters, such as shape of the cuspis of the
The females of Acrophotopsis are easy to genitalia (Pitts and McHugh 2002).
recognize as belonging to the genus. Wilson and Pitts (2008) recently con-
Disregarding setal color, thev will kev out curred along\vith Pitts and McHugh (2002)
to Dilophotopsis Schuster in Manley and and Pitts (2003) in suggesting that Dilopho-
Pitts (2002), from which they can be topsis and Schusterphotopsis Pitts are closely
immediately separated by the presence of related to Acrophotopsis. The females of D.
the anterior tooth at the termination of the concolor and D. steiiognatha (Cresson) have
dorsal mandibular carina and the presence been described (Mickel 1963; Pitts et al.
of scattered tubercles on the metasoma. 2007) and can be compared to the females
Volume 18, Number1, 2009 211
ofAcrophotopsis. The female ofDilophotopsis Casal, O. H. 1967. Comentarios sobre Acrophotopsis
pawn (Cameron) remains unknown. The Schuster, con la descripcion de una nueva
especies de Mexico (Hymenoptera: Mutillidae).
females of these two genera are morpho-
Physis 74: 1-4.
logically quite similar and share several Ferguson,W.E.1967.Malesphaeropthalminemutillid
notable characteristics, such as a large wasps of the Nevada Test Site. Brigham Young
basal tooth on the ventral margin of the UniversityScienceBulletin, BiologicalSeries8: 1-26.
mandible,aswellasthe dorsalcarina ofthe Manley, D. G. andJ. P. Pitts. 2002. Key to the genera
andsubgeneraofMutillidaeofAmericaNorthof
mandible terminating in a semi-erect tooth Mexico with description ofnew genus. Journal of
and the granulate sculpturing ofthepygid- Hymenoptera Research 11: 72-101.
ium. The females ofDilophotopsis,however, Mickel, C. E. 1963. Description of the female of
have a longer first flagellomere, have more Dilophotopsis stenognatha Schuster (Hymenoptera:
sdiosmtai,nctbuptlulmaocskethseetaelrefcrtintguebserocnletsheomnettah-e PilgrmMiuemt,tihllEo.iddaMef.)o.raPaansnds-oPcaJi.caitfPii.ncgPEintttthose.modl2io0gm0i6os.rtphA39i:cm1os8le3ex-ce1us8l4a.orf
dorsumofthe second metasomaltergite. In velvet ants (Hymenoptera: Mutillidae). Journal of
some cases the dorsum ofthe mesosoma of Kansas Entomological Society 79: 222-230.
D. concolor has indistinct tubercles, but Pitts, J. P. 2003. Schusterphotopsis, a new genus of
Sphaeropthalmini (Mutillidae: Sphaeropthalmi-
never to the degree of Acrophotopsis. Al-
nae) from California, with notes on the closely
though not all of the females of Dilophotop- related genera Acrophotopsis Schuster and Dilo-
sisareknown,thesimilaritiesofthefemales photopsis Schuster clade. Zootaxa 333: 1-7.
of these taxa further strengthens the asser- . 2006. Review of the Sphaeropthalma imperialis
tion that Dilophotopsis andAcrophotopsis are species-group (Hymenoptera: Mutillidae), with
descriptions of females and taxonomic notes.
sister groups. In addition, the females of
Zootaxa 1248: 1-20.
these genera share many characteristics and V. McHugh. 2002. Revision ofAcropho-
J.
with females of the Sphaeropthalma orestes topsis (Mutillidae: Sphaeropthalminae), with a
species-group, more so than with other new species from Baja California. Journal of
Sphaeropthalmafemales,andsuggestingthat Hymenoptera Research 11: 363-374.
and D. G. Manley. 2002. Description of the
Sphaeropthalma may be a paraphyletic as- females of Stethophotopsis Pitts and Photopsioides
semblage. Schuster (Hymenoptera: Mutillidae). Proceedings
of the Entomological Society of Washington 104:
ACKNOWLEDGMENTS
672-679.
We would first like to thank Carol von Dohlen at sex,T.asJ.soBcoiuadt,ioanndofE.tMhr.eePilsgpreicmi.es200o7f. Mnoolcetcuurlnaalr
Utah State University for the use of laboratory space velvet ant (Hymenoptera: Mutillidae). Journal of
andequipment.WealsothankErikPilgrimandCarrie theKansas Entomological Society80: 136-145.
Drake for help with DNA extraction, PCR, and DNA -, F. D. Parker, and T. L. Pitts-Singer. 2004. A
sequencing and Sarah Clark for label data entry and review of the Sphaeropthalma uro species-group
voucher specimen curation. We thank David Tanner (Hymenoptera: Mutillidae), with taxonomic
(Utah State University, Logan, UT) and Kevin Wil- changes.JournaloftheKansasEntomologicalSociety
liams (UtahStateUniversity,Logan,UT) forcritically 77: 223-234.
reviewing drafts of this paper. This research was Schuster, R. M. 1958. A revision ofthe sphaeropthal-
supported by the Utah Agricultural Experiment mine Mutillidae of America north of Mexico. II.
Station, Utah State University, Logan, UT and was Entomologica Americana 37: 1-130.
approved asjournalpaperno. 8063. Thompson,J.D.,D.G.Higgins,andT.J.Gibson. 1994.
CLUSTAL W: improving the sensitivity of pro-
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Brothers, D. J. 1995. Mutillidae. Pp. 541-548 in genusDilophotopsis(Hymenoptera:Mutillidae)by
Hanson,P. E. andI. D. Gauld,eds. TheHymenop- using molecular and morphological data with
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