Journal of Hymenoptera research 04 1995

Mesosoma very short medially, anterior edge with twoor threepairs oflateralsetae transverse ridge; propodeum with width of median panels about 1.6X their length Figs 17-19,plicaeand me

BANKS, D Male nest defense in the digger wasp Cerceris binodis (Hymenoptera:

CARVER, M Euryischomyia Girault (Hymenoptera:Chalcidoidea: Aphelinidae: Eriaporinae:

Apoidea:Sphecidae: Pemphredoninae),part 1 204

GESS, F. W. Descriptionsof themaleof Riekia nocatunga Richards,themale and two

strik-ingly distinctsympatric colourformsof Riekia confluens (Snelling)and themaleof

Rolandia angulata (Richards) (Hymenoptera: Vespidae: Masarinae) from

GESS,F. W.,S K. GESSand R.W. GESS AnAustralian masarine, Rolandia angulata

(Rich-ards) (Hymenoptera: Vespidae): nestingand evaluation of association with

GOKHMAN,V E. and D L. J. QUICKE. The last twenty yearsof parasitic Hymenoptera

HEYDON,S L. Areviewof theNorthAmericanspecies of ThinodytesGrahamand Mauleus

Graham (Hymenoptera: Pteromalidae) 1

LATTKE,J E. Revisionof the ant genus Gnamptogenys in the New World (Hymenoptera:

PULAWSKI, W J. The wasp genusTachytella Brauns, 1906 (Hymenoptera:Sphecidae) . 121

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Vol 4, 1995,pp 1-24

A Review of the North American Species of Thinodytes Graham and

Mauleus Graham (Hymenoptera: Pteromalidae)

Steven L. Heydon

BohartMuseum, Department ofEntomology, Universityof California,

Davis,CA 95616-8584 USA

Abstract.—The

Halticoptera-group isdefinedascontaining thosepteromaline pteromalid genera

with a reticulate body, acarinate pronotal collar, weakly developed notauli, weakly delimitedfrenum, propodeum with the median carina and plicae connected posteriorly by a W-shaped

carina, petiolewithabasalflange,andthehind marginof the first gastral tergitesinuouslaterallyand usually emarginate medially. Genera included in this group are Halticoptera Spinola, Halti- copterina Erdos, AndersenaBoucek, Thinodytes Graham,Syntomopiis Walker,MauleusGraham, and

Ploskana Boucek Thinodytes and Mauleus are revised for the Nearctic region and keys to the

world's describedspecies are given. New species include T caroticus n. sp., T cyzicopsis n sp.,

T petiolatus n sp.,M cultratus n. sp.,M iligneus n.sp., andM venetus n. sp. Polycystus nigritusHoward is transferred toMauleus as M nigritus n. comb, and Gastrancistrus cephalon Walker is

transferred to Thinodytes as T cephalon n. comb BubekiafallaxGahann syn issynonymizedwith

T.

cephalon Walker

INTRODUCTION are Notoglyptus Masi, Sphegigaster

Spino-la, and Schimitschekia Boucek

de-^ ^^^ Thinodytes and Mauleus, thefined as containing those pteromaline North American species ofall the genera

sculptured, pronotalcollar acarmate(Figs

Nearctic region were recently reviewed

21, 22), notauli weakly developed (Figs ^^ ^^e currently being studied. Andersena

21, 22),frenum weakly delimited(Figs 21, includes

only one species (Boucek 1993)

peti-i^gHalticoptera The worldspecies of

Si/n-rected lateral andventral flange (Figs

20), and hind margin of first gastral ter- by Heydon (1988),'Heydon (1993), and

emar-Heydon and LaBerge (1988), respectively,

ginate medially (Figs 21, 22). Genera of This

this group include Halticopitera Spinola, of Thinodytes and Mauleus [Ploskana was

by Boucek (1976).] Research by

Thinodytes Graham, Syntomopus Walker, the author has revealed the presence of

Ploskana Boucek, and Mauleus Graham, numerous species ofboth thesegenera in

characteristics of the Halticoptera-group

Thinodytes previously contained two and which probably are more or less described species, the Palearctic species

to the Halticoptera-^roup T. and the

Carib-Journal of

Figs 1-10 1, Thinodytescaroticus n.sp., female hind leg;2, Thinodytes cephalon (Walker), 2, female hindleg;

3, 4, Thinodytes cyzicopsisn. sp., femaleheaddorsal view, 4,maleantenna; 5, 6, Thinodytes petiolatusn. sp., 5,

femalehead dorsal view, 6, maleantenna; 7, Thinodytes cyzicus n. sp., femalehead dorsal view; 8, Mauleus

cultratus n.

sp., female habitus; 9, Mauleusiligneus n.

sp., female antenna; 10, Mauleusvenetus n. sp.,female antenna.

bean species T. clypeatus (Girault) 1918

To these I add the New World species T.

cephalon (Walker) 1843, n. comb., and

de-scribe four new Nearctic species—T

car-oticus n. sp., T.

cyzicopsis n. sp., T.

petiol-atus n. and T. santerna n sp Mauleus

was created for the species M. maderensisGraham, 1981 from Madeira However,this species is associated with plants na-tiveto Mexico and is suspected to be na-

Figs 11-16. 11,Thinodytes

petiolatus n sp., female clypeus; 12, Thinodyies cephahn (Walker), femaleclypeus;

13, Maiik'iis iligueus n sp.,male clypeus; 14, Synlomopiisamerkanus Ashmead, femalehead dorsal view; 15,

Maidens iligneiisn.

sp., male headposterior aspect; 16, Halticoptera sp.,male head posterior aspect.

species is given more credence by the new Nearctic species— M. cultratus n. sp.,

/('HSspecies,M. ///^^r/fj/s (Howard), 1897n. These three new species are describedcomb, andthe author's of three herein

Figs. 17-22. 17, Tliinodijtes ceplmlon (Walker), femalepropodeumandpetiole; 18, Tlunodi/tes ci/zicopsis n.sp.,male

propodeumandpetiole; 19, Tlii)iodi/tes petiolatiisn.sp., femalepropodeum andpetiole; 20, 21, Maidensdigneusn.

sp., 20,malepropodeumandpetiole, 21, female habitus; 22, Syntomopus arpedesHeydon,female habitus.

METHODS ^\^ jg used instead of clava In addition,

pet-used instead of hollow and iole. The are used:

the median ocellar diameter is MOD, the

ocellar-ocular distance is OOL, the

poste-rior ocellar distance is POL, the lateral

plate sensilla are MPP sensilla, the lower

ocular line is LOcL, and the antennal

measurements given in the descriptions

can be converted to millimeters by

multi-plying by 0.02. The acronyms for the

mu-seums from whichmaterial was borrowed

are listed in the acknowledgments section.

TJiinodytes Graham

Dicydus Thomson, 1876:221, 253. Typespecies:

Miscogaster cyzicus Walker, 1839:200; by

because ofDicydusWalker, 1833:371, 455.

Thinodytes Graham, 1956:261. Type species:

MiscogastercyzicusWalker, 1839:200;by

orig-inal designation (examined) Peck, Boucek,

and Hoffer, 1964:41. Graham, 1969:150, 167.

Hedqvist, 1975:167. Dzhanokmen, 1978:82.

BoucekandRasplus, 1991:32.

Description.— Body color

sculp-turing sometimes smooth on frenum (T.

cyzicopsis, T. petiolatiis, and T. santenm)

and median panels ofpropodeum(T.

san-terna); petiole alveolate in species with

smooth in species with transverse petiole

(Fig 17); gastral tergites nearly smooth

Clypeus variable, with either three

asym-metrically arranged denticles (Fig. 11) (T.

cyzicopsis, T. cyzicus, T. petiolatiis), three

san-terna and T.

clypeatus)orwithsinglebroad

(T. cephalon and T caroticus). Head with

short genal concavity often present;

anten-naltorulusabove LOcL Antennawith

for-mula 1:1:2:6:3; scape cylindrical, slender;

length of flagellum plus pedicelof female

less than head width (subequal in length

in T.

cyzicopsis),equaltoorslightlygreater

than head width in male; funicular

mentscylindrical; MPP sensilla usually in

length of funicularsegment (Figs 4, 6);

terminal segment, except T. cephalon with

terminal spine and large patch of

micro-pilosity. Male maxilla with stipites larged; palps slender (Fig 15). Mesosoma

very short medially, anterior edge

with twoor threepairs oflateralsetae

transverse ridge; propodeum with width

of median panels about 1.6X their length

(Figs 17-19),plicaeand mediancarina tinct and connected posteriorly by W-

dis-shaped carina (Figs 17-19), basal foveasometimes bordered mesally by short

straightcarina (Fig. 17) (T. clypeatus,T.

zicopsis, and T. cephalon), or by long sinu-ous carina (Figs 18, 19) (T. cyzicopsis, T cyzicus, and T. petiolatiis); spiracles ovate

vein sometimes distinctly shorter (T. oticus and T. cephalon); stigma small, but

car-itsheighthalfthedistancebetweenstigmaand anterior wing margin in T. clypeatus;

three partial distal setal rows; basal cell

bareexceptsometimesa fewsetae distally

(T clypK^atus and T.

cyzicus);basal vein

se-tose except in T. cephalon; speculum

an-teriorlydirected lamellate flange(Figs

trans-verse, unsclerotized ventrally, and out lateral setae (Fig. 17) (T caroticus, T.

with-cephalon, and T. santcrna), or quadrate to

elongate and sclerotized ventrally (7

cly-peatus, T. cyzicopsis, T. cyzicus, and T

pe-lateral setae sometimes

Journal of

(Fig. 19) (T cyzicopsis and T. petiolatus);

weak median carina present in T.

clypea-tiis. Gaster offemaleovate, acuminate

api-cally, 1.4-1.8X as long as wide;

hind margin of Tl sinuous laterally and

Discussion.—

separate Thinodytes from the other genera

of the Halticoptera-group, particularly

Hal-ticoptera, Maiileus, and Si/ntomopiis, is

im-possible because Thinodytes is what is left

when the more distinct genera of the

Hal-ticoptera-group are characterized.

Apomor-phiccharactersamongrelatedgenera

com-mon to all Thinodytes species such as the

reticulatebody, weakly developed notauli,

poorly delimited frenum,propodeumwith

with a basal bracingconsisting of an

ante-riorly directed lateral and ventral flange,

and thehindmarginofthefirstgastral

charac-ters defining the Hnlticoptera-group itself.

American species, some described and

some not, that fit within the present

defi-nition of Thinodytes. Once these are

inves-tigated, it may be possible to divide

Thi-nodytes intomonophyletic generic units.

Halticopteraisdistinguishedfrom

states including a bidentate clypeus, the

antennal torulus located at or below the

LOcL, the scape usually nonmetallic, the

male maxilla with lamellately expanded

palps andusually withanother lobe onthe

stipites, and a median longitudinal carina

on the petiole. Thinodytes has the clypeus

of clypeal denticles, butno known

Thinod-ytes species has a bidentate clypeus Theantennal torulus in Thinodytes is located

distinctly abovethe LOcL (except in T.

tiolatus),and thescape usuallyhasmetalliccoloration The male maxilla of Thinodytes

lobes on the stipites. Thinodytes clypeatus is

the only species of Thinodytes that has a

median carina on the petiole. Halticoptera

species are commonly bright metallicgreen; those of Thinodytes areusually dark

by thebidentate clypeus (Fig 13), the

(Figs 8, 21), the median panels of the

pro-podeum short (2X as wide as long) (Fig 20), and the lateral flanges of the petiole

enlarged and thickened (Fig. 20). In

meso-somaisdistinctlylower thanthe vertex, the

median panels of thepropodeum are

lon-ger (Figs. 17-19), and the basal flanges of

the petiole are lamellate (Figs. 17-19)

(lengthabout V3 its width) (Fig 22), threebroad symmetrically arranged clypealdenticles (Fig 14), and usuallya flattened

Thi-nodytes is much shorter, those Thinodytesspecies having symmetrically arrangeddenticles have them fingerlike rather than

nev-er so flattened as it usually is in

Syntomo-pus species

Biology.— The knownhosts of Thinodytes

species are all small Diptera living in

plants as leaf or stem miners

KEYTO HOLARCTICSPECIESOF THINODYTES GRAHAM

- Petiole as

2. Scape andlegsbeyondcoxaepale,nonmetallic.Frenumandmedianpanelsofpropodeumsmooth Plicae rounded and smoothly convergent santema

- Hind margin of Tl entire medially Dark bands on tibiae with diffuse borders (Fig. 2).

Female clubwithterminal spineatapex cephalon (Walker)

4. Clypeal denticles symmetrically arranged Petiole with weak median carina. Ovipositor

-Clypealdenticles asymmetrically arranged, the mediandenticle displaced to the left

(Fig.

11) Petiolewithout a median carina (Figs 18-19). Ovipositor sheaths hardly exserted 5

5. Costalcell withthreerowsof setaedistally. Eye length<4X as longas the templelength

(Fig 5). Body moreor less alldark cyzicus (Walker)

- Costalcell with two rowsof setaedistally.Eye length >4X aslongas the temple (Figs 3, 7). Body dark with diffuse metallic patcheson head and mesosoma 6

6. Male with terminal segment of funicle appearing as wide or wider than long (Fig 4).

Petiole usually less than 1.7X as longas wide (Fig. 18). Female withpetiole usually less

than 1.5X aslong aswide Bothsexes withhind margins ofTl and T2 as long medially

aslaterally, (eastern United Statesand Canada) cyzicopsis n. sp.

- Males withterminal

segmentof funicleappearinglonger thanwide(Fig.6).Petioleusuallymore than 1.7x as long as wide Female with petiole usually more than 1.5x as long as

wide (Fig. 19). Both sexes with hind margins ofTl and T2 usually longer laterally than

Thinodytes caroticus Heydon, new species

(Fig 1)Holotype, female.—Color:

wing veins brown; pretarsi black; knees,

apical K; of tibiae, middle and hind tarsi

white, border between the light ends and

dark median band oftibiae sharp (Fig 1).

alveo-late; median panels of propodeum

ob-scurely alveolate.

10);anteriormargin ofclypeus with single

genal concavity; eye height 1.2X length

(10:8); 2.0X malar distance (10:5), eye

length 4.0X temple length (8:2); ratio of

MOD, OOL, POL, LOL as 2.0:3.0:6.5:3.0;

vertexroundingregularly into occiput;

to-rulus Vi own diameter above LOcL

An-tenna with of

lengths of scape, pedicel, annelli, Fl-6,club as 6.5:2.5:0.5:2.0:2.0:2.0:2.0:2.0:2.0:5.0;

segment simple apically, with

length 1.3X width(25:19);dorsellum shortsmooth band; propodeum with basal fo-

an-teriormarginofmedianpanels,withshort

weak longitudinal carina crossing groove

with spiracles on anterior margin of

car-inate anteriorly. Fore wing with ratio of

lengths of submarginal, marginal,

post-marginal, stigmal veins as19.0:11.5:9.0:5.0;

with row of 4 setae. Petiole conical, verse, smooth, with mediancarina Gasterfusiform, length 1.8X width (30.0:16.5);hind margin of Tl emarginate medially;ovipositor sheaths hardly exserted; hypo-

Journal of

Allotype, male.—Color

pattern similar to

blue; fore tarsi palebrown; pale portions

of legs pale yellow-brown instead of

white Body length1.2 mm Antennawith

length of pedicel plus flagellum 0.98X

head width (21.0:21.5); relative lengths of

scape, pedicel, annelli, Fl-6, and club as

6.0:2.5:0.5:2.0:2.0:2.0:2.0:2.0:2.0:6.0; widths

of Fl, F6,clubas2.5:3.0:3.0;funicularsetae

sparse, reclinate.Gasterovate,length 1.5X

width (23:15).

Variation.— The body length of females

examined varied between 1.2and 1.8 mm

color ofthe dorsumof themesosoma

darkbands onthelegsarealwaysdistinct,

but the intensity of their metallic

colora-tionis variable

Discussion.—Thinodi/tes caroticus most

closelyresemblesT. cephalon becauseboth

species are dark in color; have a single,

broad,asymmetricallyplaced clypeal

den-ticle; and have smooth, transverse

peti-oles.Thinodytescaroticus differsfromT

ce-phalon in the following: 1. The hind

straight in T. cephalon 2. Thebasal vein of

the fore wing is setose in T caroticus, but

bare in T. cephalon 3. The median panels

in T caroticus, but distinctly alveolate in

T.

car-oticus,but about%the gastrallength in T.

cephalon 5. The dark bands on the tibiae

are distinct with sharp borders in T

caro-ticus,butare lessdistinctand havediffuse

tibialcolorbandsareuniquetoT.caroticus

and will identify the species at a glance

(Fig 1).

Etymology.— The species name comes

from the Greek karotikos, meaning

stupe-fying orsoporific,and refers tothe general

nondescript appearanceof this species.

Type Material.— The holotype, allotype

(bothUCDC) and one male were

all reared by the author from the leaf

mines ofCalycomyzapromissa (Frick)

(Dip-tera: Agromyzidae) collected 30 June 1985

Illinois, near Champaign, Illinois.

Fifty-nine additional paratypes seen were

UCDC, USNM): Bermuda DEVONSHIRE

male FAGOT PARISH: Berry Hill Road,29.VI.1988, 2 females, 3 males; Botanical

Marsh, 29.VI.1988, 1 female SMITHS

fe-male Canada ONTARIO: Chatham, 1952

male United States. CALIFORNIA:

Haw-thorne, IX.1940 (ex Aster blotch), 3 males, 1 male;Jepson Prairie Preserve (13

fe-km s. Dixon), 20.V.1983, 1 female; Lake

(bred fromdipterousleafminer),1 female;

pus-ilia{prob.= Liriomyzapusilla)],1 female;

So-quel, 26.VIII.1948 (ex Agromyza sp.), 2 males, 1 male; 11 km e St. Helena (Lake

fe-male, 7.IX.1991 (on Heraculeum), 1 male; 6

West-wood Hills (Los Angeles County),

5.XI.1940 (ex serpentineleafminer in

Zin-nia),4 females, 1 male.FLORIDA:

{proh.= Liriomyzapusilla)],1 male; ville, 2 females GEORGIA: Savannah,

Jackson-5.VI.1943 (parasite of goldenrod

leafmin-er), 3 females ILLINOIS: South Farms ofthe University of Illinois, 19.V.1985, 1 fe-

male; White Heath, 24.IX.1939, 1 female

INDIANA:4 miless.New Harmony

(Har-mony State Park), 28.VI 1983, 2 females; 2miles s. New Lisbon, 14 VII.1981, 1 male

IOWA: SiouxCity (reared from leafminer

(rearedfrom mineofleafon sunflower), 1female KANSAS: Lawrence, 14.V.1955, 1

26.VII.1960, 1 female NEW MEXICO:

19 IX.1905 (on Grindelia squarrosa (Pursk)

27.IX.1905 [ex dipterousleafminer of

(Com-positae)], 2 females; Roma, 26 III.1948 (ex

pupa of dipterous leaf miner), 3 females,

3 males

Biology.—This species has been reared

from leaf-mining Agromyzidae, mostly

Ca-lycomyza promissa (Prick) and Liriomyza

pusilla (Meigen) BecauseL pusilla is a

Pa-learctic agromyzid species (Spencer 1976)

and isnotknown from theNearctic region

(Spencer and Steyskal 1986), it is likely

that this host record is in error. There is

one record from Chatham, Ontario from

also reared from leaf miners on

Machaer-antheraannua, Zinnia,goldenrod,

sunflow-er, and "Aster" It has been reared from

both linear-mining and blotch-making

leaf miners Other plant associations of a

more uncertain nature include the

com-posites Baccharis sp., Heracleum sp., and

Grindelia squarrosa

Thinodytesclypeatus (Girault)

Polycystus clypeatus Girault, 1918:128.Holotype,

female (USNM); Hym. TypeNo. 20682;

(ex-amined)

Thinodytes clypeatus (Girault): Heydon, 1989:

193.

Redescription.—

Mesosoma, petiole black, with blue tints

dark brown; legs with basal % of femora

brown, remainderof legs white

Sculpture: Dorsum of mesosoma

panels of propodeum weakly alveolate;

petiole alveolate

Structure: Head withanterior marginof

LOcL Antenna with and widths

19.5X3.6: pedicel 6.0x4.5: annelli 2.6x3.6:

P54.6X6.8: F6missing: club 13.4X6.8; club

rather flat, length 1.4X width (33:24);

2), sides converging posteriorly; notauli

pairs oflateral setae, frenum almost

indis-tinguishablefrom remainderofscutellum;

0.45X width,withrowoffoveae separated

by carinae along anterior margin, spiracle

a raised smooth crescent Fore wing with

relative lengths ofsubmarginal, marginal,postmarginal, stigmal veins as 21:12:11:6;

cell with one seta; basal vein with three

setae. Petiolelength 1.2X width (6:5);withweak median carina; lacking lateral setae.Gaster fusiform, length 1.7X width (31.0:18.5); Tl emarginate medially; hypopy-

ovi-positor sheaths exserted for length equal

to thatof T7

Discussion.— The

removed and crushed on a slide.

Thinod-ytes clypeatus differs from all other scribed Thinodytes species because it has

sheaths, and a median carina on the

peti-ole. Thinodytes clypeatus has three metrically arranged denticles like Synto-mopusspecies,but T. clypeatus differsfrom

denticles ofT. clypeatusarefingerlikelobes

like those of the other Thinodytes species

withthree denticles,whereasthedenticles

triangular (Fig 14). The pronotal collarof

Thinodytes clypeatus is short, eleven times

aswideas long;the pronotalcollar in

as long (Fig 22). Most species oftera have a median carina on the petiole,but there are no other characters to indi-

Halticop-10 Journal of

cate a particularly close relationship

be-tween T. clypeatus and Halticoptera.

DistributionandBiology.— Thetype

spec-imen wasrearedfroma leafminer oncorn

West Indies by F. Watts De Santis (1979)

reports this species from Barbados also,

andgives thenameofitshostasAgromyza

parvicornis Loew (Diptera: Agromyzidae).

Thinodytes cephnlon (Walker),

new combination

Figs 1, 17

?Pteromnlus Rhseo Walker, Walker 1839b:88-89

Lectotype, female (BMNH); Hym. Type No.

5.772 (examined).

Gastmncistrus cephalon Walker 1843:30

Lecto-type, male (BMNH); Hym. Type No. 5.661

(examined)

Bubekiafallax Gahan, 1933:114-116 Holotype,

female(USNM); TypeNo 44841 (examined)

Allen and Painter, 1937:225. Nikol'skaya,

1937:25. Peck, 1951:538.Thompson,1958:587.

Peck, 1963:610-611 Morrill and Kieckhefer,

1971:1130. Allen and Pienkowski, 1973:616,

617. Burks, 1979:789. Hendrickson, 1979:300,

302 n syn.

Discussion.—Thinodytes cephalon iseasily

distinguished by its single broad clypeal

denticle, apical spine on the female club,

notauli traceable tothehind marginofthe

mesoscutum asimpressed lines,bare

dor-sal vein, smooth and short petiole (Fig.

17), and first gastral tergite having the

hind margin entire.

A possible seniorsynonym of T.

cephal-on isPteromalus rhaeo describedfrom

(designated herein) ofP. rhaeo is

unfortu-nately missingthehead. Itappearssimilar

which many specimens in my collection

Island, but it is

significantly larger than

any other specimen examined The

medi-anpanelsofthepropodeuminP. rhaeoare

entirely rugose and the plicae are

poste-riorly divergent In other T. cephalon

ru-gae which are located mostly posteriorly,

and the plicae are parallel. Thehind tibia

in P. rhaeo is almost uniform in color; in

T. cephalon, the hind tibia are distinctly

Pter-omalus rhaeo may be within the range of

variation of what is recognized as T

ce-phalon,but I am reluctant to formally

syn-onymize the name until more is known

Distribution.—Thinodytes cephalon is one

ex-tends throughout most of North and

Biology.—Thinodytes cephalon is a

para-sitoid of dipterous larvaethat mine leaves

or grass stems Thinodytes cephalon was

has also been reared from the wheat stem

(Dip-tera: Chloropidae), ina numberofstudies(Gahan 1933; Allen and Painter 1937; andMorrill and Kieckhefer 1971). Allan andPainter report that T. cephalon probably

ovipositsinto the larval stage ofthis host.

One additional host added herein is

Lir-iomyza trifoliearumSpencer(Diptera:

Thinodytescyzicopsis Heydon,

new speciesFigs 3, 4, 18

Holotype, female.—Color: Head,

mesoso-ma black with metallic green patches as

follows: entire dorsellum, pairs of spotsalongsidemedianocellus,lateralhind cor-

ners of pronotum and upper epimeron;metallic blue patches as follows: on inner

orbits, anterior part of lateral lobe of

scu-tum and propodeum Antennawithscape,

pedicel dark blue;flagellumblack Petioleblack Casterbrownishblackwithmetallicblue reflections dorsally, green ventrally

black with weak blue

re-flections; trochanters, femora brownish

black with weak green reflections, knees

yellow, tibiae with diffuse-edged dark

tarsi brown; middle and hind tarsi with

pretarsi black, remainder yellow.

Sculpture: Head, mesosoma, regularly

propo-deumshallowlyandindistinctly alveolate;

petiole finely alveolate

width 1.2X height(27:22),2.2X length(27:

12); anterior marginofclypeus with three

minute asymmetrically arranged apical

denticles; genal concavity absent; eye

height 1.4X length (13:9), 1.9X malar

dis-tance (13:7), length4.5X temple length (9:

2) (Fig 3); ratio ofMOD, OOL, POL, LOL

as 2.5:4.0:7.0:3.0; vertex rounding

above LOcL Antennawith length of

ped-icel plus flagellum l.Ox head width (27:

27); relative lengths of scape, pedicel,

an-nelli, Fl-6, club as 11.0:3.0:1.0:2.5:3.0:2.75:

2.75:2.5:2.5:7.0; widths of Fl, F6, club as

3.0:3.5:3.5, Fl^ appearing quadrate, F5-6

slightly transverse; club simple apically,

with minute patchof micropilosityon

dorsal-ly, length 1.6X width (36:22); pronotum

with humeral angles squared;

bandlike, short, weakly alveolate;

propo-deum with basal fovea bordered mesally

weakly carinate anteriorly; spiracle <

of propodeum. Fore wing with ratio of

lengths of submarginal, marginal,

post-marginal, stigmal veins as 25:13:14:8;

more than stigmal vein width; basal cell

bare;basalveinsetose Petiole length 1.2X

width (6:5); without median carina; with

one pair of lateral setae. Gaster length

1.4X width (31:22); hypopygium

sheaths hardly exserted.

male.—Color similar to

holo-type except frons blue, mesoscutum with

a pair of large diffuse green spots; axilla

green Bodylength 1.3 mm Antenna(Fig.4) with length of pedicel plus flagelluml.Ox head width (24:23); relative lengths

ofscape,pedicel, annelli, Fl-6, clubas 7.0: 3.0:1.0:2.0:2.5:2.5:2.5:2.5:2:6.5; widthsof Fl,F6, club as 2.0:2.5:2.5; setae fine, reclinate.Petiole length 1.2X width (5:4)(Fig. 18).Gaster ovate, length 1.2X width (20.0: 16.5).

Variation.— The body color varies from

in-tensity, size, number, and hue of the tallic spots on the body are highly vari-

me-able.Thecolor of the metallicpatches

blue-green The body length ranges between

mm in males The length of the marginal

veinvaries from0.9 to 1.2times the length

about equal its length [x=1.02±

the petiole are sometimes difficult to see

flanges of the petiole.

Discussion.-Thinodytes cyzicopsis

resem-bles the Palearctic species T. cyzicus

the following: 1. Thinodytes cyzicopsis has

a series of metallic patches on the body,

whichare lackingin T cyzicus 2.Theratio

of the eye length to temple length ages 4.25±0.12 (n=6, range 3.8^.5) in fe-male T. cyzicofisis (Fig. 3) but measured

cell of the fore wing of T.

cyzicopsis has

two rows of setae distally, whereas there

arethree rows distally in the costal cellofT.

cyzicus. 4. The petiole has a pair of

lat-eral setae in T.

cyzicopsis, which are

Journal of

Thinodytescyzicopsisand T.

petiolatus are

verysimilarspeciesand specimens cannot

twospecies aredistinctfromother

Thinod-ytes in their commonpossession of a

sim-ilarpatternof metallicpatchesonthehead

and mesosoma. Individuals ofT.cyzicopsis

differfromthoseofT petiolatus in the

fol-lowing: 1. Theratio ofeye heighttomalar

1.74±0.035 (n=8, range 1.5-1.8) infemales

males;inT petiolatus theratiois1.48±0.23

1.77±0.038 (n=10, range 1.6 to 2.0) in

males 2. All funicular segments of the

T cyzicopsis; all funicular segments are

usually longer than wide in male T

pe-tiolatus 3. The ratio of petiole length to

gas-tral tergite in T. cyzicopsis isabout as long

shortermedially than laterallyin T.

petiol-atus 5. The hind margin of T2 is straight

to just noticeably concave in T.

cyzicopsis,whereas it is usually distinctlyconcave in

T. petiolatus. 6. The terminal segment of

slightlytransversein T.

cyzicopsis,whereas

it

ad-dition to these structural characters, T.

from Kouchibouquac National Park, New

1977 by S. J. Miller The allotype (USNM)

col-lectedon vernalalfalfa on 22June 1968by

A G.Wheeler.Sixteenparatypeswere

Canada ALBERTA: Elkwater Lake,

fe-male NEW BRUNSWICK:

Phy-tobia {Calycomyza)solidaginison Solidago], 1male; Crosby, 31.VII.1952 (on apple), 1

(swept from Salix hlanda), 1 male,

29.VI.1972 (swept from Salix blanda), 3

males, 24 VII.1972 (swept from Salix

21.VII.1947, 1 female, 25.VII.1947 (swept

from Rosa rugosa), 1 female, 9.VIII.1945, 1female United States. ILLINOIS: 2 miles

Farms of the University of Illinois, nr.Champaign, 23.VI.1981, 1 male MASSA- CHUSETTS: Hopkinton, 9.VIII.1951 (ex

Ilex leaf miner), 1 female NEW MEXICO:

VIR-GINIA: Winchester, 16.VI.1964, 1 male,

be-tween T. cyzicopsis and T cyzicus.

Biology.— Known hosts of T. cyzicopsis

include Phytobia {Calycomyza) solidaginis

(Composi-tae)] and anIlexleafminer.Thespecieshas

(Salica-ceae), Rosa rugosa (Rosaceae), and alfalfa Salixand Rosa are probablyprimarily nec-

tarsourcesbecauseneitherhasmany

leaf-mining agromyzids, but these plants are

Thinodytes cyzicus (Walker)

Miscogaster cyzicus Walker, 1839a:200.

Lecto-type,female(BMNH); Hym.TypeNo.5.2570(examined)

Syntomopus cyzicus (Walker): Walker, 1846:28.

Schmiedeknecht,1909:376.

Dicyclus circulusThomson, 1876:253. Lectotype,

Thinodytes cyzicus (Walker): Graham, 1956:261.

Hedqv-ist, 1975:180. Boucek, 1977:56. Kamijo, 1978:

Hedqvist, 1983:167. Boucek and Rasplus,

1991:32.

Discussion.— The synonymy of Dicyclus

circiilus with Thinodytescyzicus isaccepted

on the authority of Graham (1969).

Thi-nodytes cyzicus resembles T. cyzicopsis and

T. petiolatus,becauseall three specieshave

three small, sharp, asymmetrically

long, reticulate petiole. Besides the

geo-graphic separation of their ranges, T.

Nearctic species by its body color.

Thino-dytes cyzicus is uniformly dark and lacks

the distinctive diffuse metallic patches on

the head and mesosoma present in T

cy-zicopsis and T. petiolatus. Thinodytescyzicus

alsohas threerows ofsetae distally in the

costal cell,whereas T cyzicopsis and T.

pe-tiolatus have only two rows The eye

lengthin T. cyzicus is4X or lessthe length

and T.

petiolatus havethe eye lengthmore

than 4X the length of the temple (Figs 3,

7).

Distribution.—Thinodytes cyzicus occurs

1970; Kamijo 1978)

Biology.— The hosts of T. cyzicus are all

Agromyzidae Askew(1970) recorded this

species from an agromyzid, probably

Phy-tomyza atricornis Meigen, on Senecio

jaco-baea Linnaeus Kamijo (1978) recorded T.

cyzicus from Chromatomyia horticola

(Gou-reau)(Diptera: Agromyzidae) on pea and

from an agromyzid on Lathyrusmaritimus.

Takada and Kamijo (1979) recordedT.

T. cyzicus may parasitize the larval stage

of its host

Thinodytespetiolatus Heydon,

new species

Figs 6, 7, 11, 19

Holotype, female.—Color: Bodyblackbut

ver-tex, gena, lateral region of pronotum,

lat-eral lobe of mesoscutum, middle lobe of

mesoscutumwithpairoflarge diffusetallic patches, anteriorlateralcorner ofax-

me-illa dark metallicgreen; gasterwithish reflections. Antenna with scape dark

green-green; remainder brown, pedicel with

black with greenish reflections; femora

Sculpture: Clypeus weakly alveolate;face alveolate, cells elongate in radiatingfashion from clypeus; remainder of head

deli-cately and regularly alveolate except

propo-deum alveolate; petiole finely alveolate.

width 1.4X height (28:21), 2.6X length(28.5:11.0); clypeus with threesmall asym-metricallyarranged clypeal denticles (Fig 11); weak genal concavity extending Vs

length4.8X temple length (9.5:2.0)(Fig 7)

vertex rounding regularly into occiputantennal torulusjust above LOcL. Anten-

0.91X head width (26:28.5); ratio oflengths of scape, pedicel, annelli, Fl-6,clubas 11.5:3.5:1.0:2.5:2.5:2.5:2.5:2.5:2.5:6.0;

apically, with small patchofmicropilosity

notauli shallow posteriorly; propodeum

(Fig. 19) withbasalfovea margined

sculptured band, carinate anteriorly; spi- arate these two species are given in the

marginofpropodeum.Forewingwithrel- hind margin of T2 in many specimens ofative lengths of submarginal, marginal, T petiolatus is distinctly concave, but thepostmarginal, stigmal veins as 27.0:14.5: visibilityofthis characterdepends on how15.0:9.0; stigma small;basal veinwith row the specimen has dried. This character is

fe-length 2.0X width (8:4); without median males

carina;withtwopairsoflateral setae.Gas- Type Material:— The

(33:24); hind margin of Tl strongly sinu- June 1931, on Salsola pestiferby D. E. Fox

ous laterally, emarginate mesally; ovipos- and the allotype (USNM) was collected at itor sheaths hardlyexserted; hypopygium TwinFalls, Idaho,on 5August 1920by R

Elk-blue-green; dark bands on tibiae very water, 9.VI.1956, 1 male; Elkwater Lake,

6) with length of pedicel plus flagellum (swept from barley), 1 female, 5.VIII.1956

1.1X head width (28:25); ratio of lengths (swept from barley), 1 male; nr.

Leth-of scape, pedicel, annelli,Fl-6, club as9.0: bridge, 1924.1925, 1 female BRITISH

CO-3.5:1.0:2.5:3.0:3.0:3.0:3.0:3.0:8.0; widths of LUMBIA: Bowser, 28.V.1955, 1 female;

fine, reclinate. Petiole length 2.0X width 18.VI.1973, 1 male United States. (8:4). Gaster length 1.2X width (20:16); ZONA:Mesa,3.VI.1958 (swept fromalfal-

ARI-truncate apically. fa), 1 male CALIFORNIA: Albany,Variation.— The body color varies from 29.IV 1958, 1 male;

sosoma reduced and obscure, the frenum male; Boca, 22.VII.1970, 1 female; Bolinas,

metallicgreen The bodylength of females Phytomyza aqiiilegiana), 10 females; Cerro

examined variedbetween 1.3 and 1.8 mm Noroeste (sw. corner of Kern Co.),

Dar-sometimes extends only Va the length of win Falls (nr. Panamint Springs),

the propodeum The row of setae on the 29 III.1984(onEncelia), 1 male; Emeryville,basal vein sometimes curls proximally, 28.V.1958, 1 male; Eureka Dunes (Inyo

fe-Disciission.—

Thinodytespetiolatus and T. male; Lake Tahoe, 29.VI 1927, 1 female;

cyzicopsis are very similar species and are Lily Pond (alpine lake), VI 1971, 1 female;

7.V.1961, 2 males; PlacerCo., Vlll, 1 male; (5), 1 male; Moscow, 6.VIII.1926, 1 male;Sagehen Creek (near Hobart Mills), 24. Murtaugh, 29.V.1930 (3 & 5), 1 male; Oak-

VI.1970, 1 female, 21-25.VI.1982 (sweep- ley, 7.VIII.1929(1,3 &5), 1 female; Rupert,ing Primus), 1 male, 12.VII.1972, 1 male, 29.V.1930, 1 female; Tuttle, 22.V.1931 (4),

11-15.VII.1982, 1 female, 23.VII.1968, 1 fe- 1 female; Twin Falls, 7.VI.1930, 1 male;male, 1.VIII.1970, 1 female; San Bernardi- Wendell, 22.V.1931 (5), 3 males MON-

OR-19.VII.1982, 1 male; Santa Cruz, EGON: Corvallis, 15.VI.1981, 1 male,22.V11I.1948 (ex PJnjtonn/zasp B), 1 female; 26.VI.1985, 1 female, 3 males; Near Cor-

Santa Rosa, 25.V.1990, 2 males; Sheppard vallis(St. Mary'sPeak), 15.VIII.1984

6-Sphenosciadiumcapitellatum),! male; Shive- 12.VI.1984 (Malaise trap), 1 female, 11

Soquel, 26.VIII.1960 (ex Agromijza sp.), 2 20.VIII.1984, 5 females, 2 males; 1 mile w

females; Titus Canyon (Death Valley Na- McKinzie Pass, 1 female, 1 male UTAH:

mexicana), 1 male; Tomales Bay State Park MonteCristo,6.VII.1976, 1 male; Myton, 3

WASHING-(1500'), 29.III.1955, 1 male; 6 miles w Bas- TON: San Juan Island (Barney's Place),

setts (Yuba Pass), 9.VII.1970, 2 females, 2 23.VII.1944, 1 male Vancouver, 15.VI

males COLORADO: Chambers Lake(Lar- 1911, 1 female, 1 male, 16.VI.1911, 1 male,

Collins, 20 VIII.1895 (on boxelder foliage), 13.VII.1985 [Artemesia spinosa

{?=spines-1 female, 1 male; Glacier Basin, Rocky cens D.C Eaton], 1 male; Snowy Range,

Mountain National Park, 24.VII.1977, 1 23.VIII.1951, 2 females, 1 male,

male; Echo Lake (Mt. Evans, 10,500'), Etymologxj.— The

species name refers to

4.VIII.1961, 1 female IDAHO: Boise, the long petiole, characteristic of this 28.V.1984 (Malaise

spe-trap), 2 males; Buhl, cies.

petiolatus is

14.VI.1930 (3), 1 female, 9 VII.1931, 1 fe- unknown but it has been taken in

associ-male; Eden, 11 VIII.1930 (2), 1 female; ation witha number ofplants, suchas

1 male; Hollister, 16.V.1931 (3), 1 male, Salsola pestifer, and Sisymbrium altissimum20.V.1931, 2 males, 2.VI.1931, 1 female, 1 in Idaho and Encelia in California It has

male, 5.VI.1931 (3), 1 female, 7.VI.1931 (3 also been collected in association with

&5), 1 female, 13.VI.1931,1 female;Hubbs crop plants, such as beets in Idaho and

Butte,22.V.1931, 1 male, 6.VI.1931, 1 male; barley in Alberta

Holotype, female.—Color: Body black

'

Thehostplantsfor thespecimens fromIdahoare

j^^^^g ^f mesoscutum, SCUtellum, gaster;

numbered as follows: 1= A.? rosae. 2= Beta viiharis n <- i i i

L. 3= Soplua sophia {=Descuraima sophia (L.)V.B. coppery reflectionson frenum and Webb).4= SalsolapestiferA Nelson.5= N.(ortn ) al- lum; yellow-green reflechons on middle

dorsel-tissiiniim or = lobe ofSCUtellumand propodeum.

Anten-Journal of

dorsally, brownish white ventrally. Legs

Sculpture: Clypeus alveolate; frenum,

with T5-7coriaceous, remainder smooth

width 1.4X height(28:20), 2.3X length(28:

genal concavity extending Va malar

dis-tance; eye height 1.3X length(11:10), 2.2X

length (10:2); ratio of MOD, OOL, POL,

LOL as 2:3:6:3; vertex rounding regularly

into occiput; antennal torulus

just above LOcL Antennawithlength of pedicelplus

flagellum 0.86X head width (24:28); ratio

of lengths ofscape, pedicel, annelli, Fl-6,

club as 10:3:1:2:2:2:2:2:2:6; widths of Fl,

F6,clubas2:2:2;clubsimpleapically,with

small patch of micropilosity ventrally on

dorsal-ly,length 1.5X width(32:22);notauli

shal-lowposteriorly;propodeumwithbasal

fo-vea obscure, nucha lunate strip and

cari-nate anteriorly, spiracles on anterior

relative lengths ofsubmarginal, marginal,

postmarginal, stigmal veins as 24:12:12:6;

width (36:23); hind margin of Tl nearly

straight, slightly convex mesally;

Discussion.—This species is distinct

from all other Thinodytes species by the

characterslistedin the key:scapeand legs

beyond coxae pale, nonmetallic; frenum

and medianpanelsofpropodeumsmooth;

posteriorly Inthesecharacters,T. santerna

is phenetically similar to species of

Noto-glyptiis.

Type Material—The holotype (USNM)

Monument, Inyo County, California(USA) by E E. Grissell on Distichlis in a

brackish marsh

Etymology.— The

specific epithet of this

species is from the Latin noun santerna,

meaning borax, and refers to the localitywhere the type specimen was collected.Biology.— Nothingis knowofthehost(s)

of T. santerna

Gra-ham, 1981 (examined); original

designa-tion.

Description.— Body very dark green or

blue; scapebrownish yellow, nonmetallic.Head, pronotum, mesoscutum, scutellum

(including frenum), dorsellum, medianpanels of propodeum, petiole alveolate;gastral tergitesnearly smooth Head with

(Fig 13), lateral part of mouth marginwith shortshallowgenal concavity; anten-

nal torulus IX owndiameterabove LOcL Antenna with scape cylindrical, >6X as

head width in females, about equal tohead width in males; funicular segmentscylindrical; MPPsensilla in single row; fe-

Male maxilla with palps slender, stipites

dorsally; pronotum withcollar short(Figs

23-25), nearly level with vertex dorsally,

notauli shallow, impressed lines at most;

median groove, frenum indistinguishable

ridge across anterior margin of

propo-deum; propodeum (Fig. 20) with medianpanels short (width about 2X median

length), plicaeand median carina well

W-shaped carina; spiracles strongly ovate

ofveins asfollows: >

Figs 23-25. 23, Maidensmaderensis Graham,femalepronotumand mesonotum;24, Maidens iligneus n sp.,femalepronotumand mesonotum;25, Maidens venetiis n.

sp., femalepronotum and mesonotum

stig-mal vein; costalcell with complete row of

setaeand sometimesa partialsecond row;

basal cell bare; basal vein setose;

specu-lumpresent,openposteriorly Petiole(Fig.

20) longer than wide, with basal flange

thickened

laterally, without median

lanceolate,length1.6ormoretimes width;

sinuous laterally, emarginate medially

(Figs8, 21).

Discussion.—This genus is placed in the

Halticoptera-group as definedinthispaper

by: the rounded pronotum, shallow

propo-deum with a median carina and plicae

reticu-late petiole with a complete basal flange,

emar-ginate medially Maidens and Halticopitera

exhibit considerable phenetic

similarity

bi-dentate(Fig 13);thelefthandclypeal

den-ticle is divided by a sulcus formed as a

den-ticles are shown in their plesiomorphic

separated state in Thinodytes (Fig 11).The

den-ticle is found in Halticoptera Spinola

the unrelated genus Sphegigaster Spinola,the left tooth in the bidentate clypeus is a

struc-ture and the nonmetallic scape are

rela-tionshipbetweenHalticopteraandMauleus

Halticoptera is well-defined cladisticallyrelative to Mauleus by the very low inser-

LOcL) and by themalemaxilla,which hasthe terminal two segments of the palps

expand-ed Two apomorphic characters readilydefineMauleus relative to Halticoptera and

relatedgenera: 1.The propodeumis

exam-ined) 2. The basal flanges of the petioleare exceptionally large and thick, givingthepetiole a connateappearance.Thebas-

al flanges in related genera are generallyfree-standing lamella and the petiole is

more or less cylindrical.

reared from pupae ofthe native hollyleaf

miner, Phytouiyza ilicicola Loew (Diptera:

18 Journal of

- Pronotalcollarwithsideswidestnearanteriormargin and converging

posteriorlyindorsal

2. Vertexandmesoscutumwith conspicuouspalesetae. Propodeumwithreticulationsmuchlesscoarse thanonscutellum,medianpanel eachwithbroadshallowgroovealonganterior

- Vertexandmesoscutumwithindistinctdarksetae.Propodeumwithreticulations ascoarse

ason scutellum, median panel each with a pair ofelongate sublateral depressions along

3. Propodeum withanteriordepressionbetweenbasal foveae;median panelswithextensivearea ofweak, almost smooth sculpture. Pronotumwith humeral angles acute, coming to

blunt pointsin dorsalview (Fig 25). MPPsensilla in twoor more rows onfunicular

-Propodeum without distinct anteriordepression betweenbasal foveae;medianpanels

al-mostentirely alveolate,withatmostasmallcentral patchofweaksculpturing. Pronotum

withhumeralangles eithersquared orslightly andsmoothly convergentposteriorly (Fig.

24). MPP sensilla in singlerow onfunicularsegments(male ofM cultratus unknown) 4

4. Eyeheight2.5-2.7X genaldistance. Female with combined lengthofheadand mesosoma

longer thangaster; gaster lessthan twice aslongaswide; hypopygiumextendingaround

% gastral length (UnitedStates) iligneusHeydon-

Eyeheight2 1-2.4X genaldistance. Female with combinedlengthofhead and mesosomaless than or equal to length of gaster (Fig 8); gaster more than twice as long as wide;

hypopygium extending to near tip ofgaster (southern Mexicoto Argentina)

cultratusHeydon

ovi-Fig. 8 positor sheaths) 1.6mm Head width1.2X

height (26:21), 2.2X length (26:12); genal

Holotype female.-Color: Head, pleural

concavityextendingVs malar distance;eye

j^^-g^t 1.4X length (13.5:9.5), 2.2X malar

bluishblack; collar, dorsum ofmesosoma

distance (13.5:6.0), length 4.8X temple

greenish black; gaster dark brown, Tl

with dark blue reflections. Antenna with

2:4:6:3; torulus IX own diameter abovescapebrownishyellow withweakmetallic lqcL Antennawithlength of pedicelplus

reflections;pedicel, flagellumbrown. Legs

flagellum 0.81X head width (21:26); ratiowithcoxae, trochanters,femora darkblue; of

lengths of scape, pedicel, annelli, Fl-6,

tibiae brown except basal and apical tips dub as 9.0:3.5:1.0:1.5:2.0:2.0:2.0:2.0:2.0:5.0;

brownishyellow;foretarsibrown, middle widthsof Fl, F6, clubas2:3:3;MPPsparse,

andhindtarsiyellow-brownwithpretarsi only one or two visible per segment from

Sculpture: Clypeus, median portion of (33:23); pronotum with sides

converging

propo-mesonotum, scutellum, frenum, median deum with width of median panels 2.2Xpanels of propodeum alveolate; petiole length (11:5); basal fovea a shallow, tri-

finely alveolate; gastersmooth exceptT5- angular depression;

plicae fading out in

7 coriaceous anterior 0.5X own diameter

from anterior margin of propodeum.

submargin-al, marginal, postmarginal, stigmal veins

as 27:16:14:7; costal cell with single

com-plete row of setae; basal vein setose with

one seta posteriorly on cubital vein

Peti-ole length 1.5X width (9:6); sides

narrow-ing posteriorly Gaster lanceolate (Fig. 8),

length2.6X width(42:16),length l.OX that

of head and gaster (42:42); hypopygium

reaching to apex of T7; ovipositor sheaths

exserted for distance equal to half length

ofhind tibia.

Variation.— The color of the head,

pleu-ral regions, and petiole varies from dark

blue, as in the holotype,to dark greenand

nearlyconcolorous withthedorsumofthe

mesosoma The body length ofspecimens

whereas the length of the specimen from

Ixtapan is 2.3 mm, the one from Morelia

areexserted for a distance equal to ¥ito Vi

the lengthof the hind tibia.

Discussion.—Maidens cultratus can be

distinguishedfrom M. iligneusby the

char-acters given in the discussion section for

that species

Eti/mologx/.— The species name comes

from the Latin word cultratus, meaning

knife-shaped, and refers to the shape to

(CNCI) andone paratype femalewere

para-type females were collected as follows

(CASC, CDAE, CNCI, SEMC, USNM):

4-8.XI.1983 (Malaise trap). Mexico

CHIA-PAS: San Cristobal de las Casas,

1-12.V.1969; MICHOACAN: Morelia,

TA-MAULIPAS: 6 miles n. Ciudad Victoria

Panama. Chiriqui, XII.1946

Biology.— The host(s) of this species are

Sculpture Clypeus and immediate

vi-cinity finely alveolate (Fig 13); remainder

of head, mesoscutum, scutellum, frenum,

median panels of propodeum alveolate;petiole finely alveolate; gaster smooth ex-

ceptT6 and T7coriaceous

width1.3X height(29:23), 2.2X length(29:

13); eye height 1.4X length (15.0:10.5),

OOL, POL, LOL as 2.5:4.0:7.0:3.0; toruluslocatedIX owndiameterabove LOcL An-

flagel-lum 0.88X head width (25.5:29.0); ratio of

lengths of scape, pedicel, annelli, Fl-6,clubas11.0:3.0:1.0:2.5:2.5:2.5:2.5:2.5:2.5:7.0;

Me-sosoma length 1.5X width (36.0:24.5);

sides convergent posteriorly (Fig 24);

no-tauli extending to hind margin of

(Fig. 20) with plicaefading outin anterior

half,basal foveaextending halfway down median panels and bordered mesally by

marginofpropodeum. Forewingwith

postmarginal, stigmal veins as 25:15:11:6;

and one partial row distally; basal vein

with row of three setae. Petiole (Fig. 20)

length 1.3X width (8:6); narrowing riorly; basal flanges large. Gaster fusiform

poste-(Fig 21), length 0.87X length ofhead and

gaster; ovipc^sitor sheath exserted for a

distance equal to/ length ofhind tibia.

Allotype.—Male Color similar to

paler ventrally; tibiae brownish yellow, allotype (USNM), and an additional nine

Bodylength 1.8 mm Head with ocelli rel- female and seventeen male paratypes

Phijto-LOL as 3.0:3.0:7.5:3.5 Antenna with mi/za ilicicolaon IlexopacaSolanderatlengths of pedicel plus flagellum l.Ox ington, Kentucky byD A Pottercollected

Lex-head width (31:31); relative lengths of 16 May 1984 An additional 43 paratypesscape, pedicel, annelli, Fl-6, club as 10.0: were collected as follows (CNCI, UCDC):

3.5:1.0:3.0:3.5:3.0:3.0:3.0:3.0:8.0; widths of ILLINOIS: Cave-in-Rock State Park, near

Fl, F6, club as 2.5:3.0:3.0; setae reclinate. Cave-in-Rock, 4.VI.1981, 1 female

KAN-Gaster ovate, length 1.8X width (42: SAS:Oswego, 17.V.1976, 1 female

in air-dried specimenwill be shorter) miner) 1 male, VII-VIII.1937(exPhytomyza

Variation.—Length of female specimens ilicis), 4 females, 8 males; Laurel,

between 1.1 and 1.8 mm. Other than the Brunswick, 26.V.1947 (ex Phytomyza

ilici-ratherlargevariationin size, thisisamor- cola), 1 female TEXAS: Houston,

8.XII.-phologicallyuniform species. 1929 (ex leafminer on IlexvomitoriaDiscussion.— Mauleus

9.V.-from M. cultratus in the following: 1. The 1912, 1 female VIRGINIA: Norfolk,

12); range 2.5-2.7] females, 4 males

than inM. cultratus [x=2.24±0.038

(n=

Un-M. cultratus 3. The ratio of the length of derhill 1943 (as Sphegigastrinae, new

the gaster is

relatively greater in M. ilig- 1936 (as Sphegigaster sp.)]. It may also be

(n=10); range 1.2- the species called Halticoptera sp. by Kulp

8) 4. Theratio of the gastral length divid- 1968 are questionable because the Phy

to-ed by its width is less for M. iligneus myza complex onhollywasnotstudied in

2.2-3.1)] 5. The hypopygium extends to Mauleus iligneus was a primary parasite

/^news(Fig 21),but isnearlyevenwiththe leafminer, Phytomyza ilicicola, was

unpar-tip of T7 in M. cultratus (Fig. 8). 6. The asitized, but was a facultative

hyperpar-ovipositor sheaths are exserted for a asite on Opius striativentris Gahan

(Hy-length equal to Va the length of the hind menoptera: Braconidae)whenthatspecies

tibia in M. iligneus (Fig 21), but for about had already parasitized the agromyzidV3 to Vi the length of the hind tibia in M. maggot. Potter and Gordon reared no

Mau-theLatin word ilex, meaningholly leus iligneus parasitizes the fly late in its

Mauleus maderensis Graham

Fig. 23

No 5.3454 (examined) Boucek and

Rasplus, 1991:41.

Diagnosis.—Mauleus maderensis differs

from the threenewly described species in

dorsal view as for most other

Pteromali-dae— more or less parallel, but weakly

convex, with the broadest point being

posterior edge (Fig 23). The three new

speciesofMauleus described in this paper

have the pronotum in dorsal view

broad-est near its anterior margin and distinctly

issimilar inpropodeal structuretothe

Ca-ribbean speciesM. niritus. However, these

twospeciesareeasilydistinguishedby the

characters given in the key

Distribution.— Madeira (Pico das

Arru-das, near Sao Martinho), and possibly

Mexico (Boucek and Rasplus, 1991)

ofMexican origin (Boucek, pers. comm.).

Biology.— The insect host(s) of M.

mad-erensis remainunknown.

new combination

No 5.876 (examined) Heydon, 1989:193

Diagnosis.— ThetypeofMauleus nigritus

gone and the gaster and petiole mounted

on thecard separatelyfrom theremainder

how-ever, to confirm that this species belongs

in Mauleus Mauleus nigritus differs from

the three newly described species in that

ithas the sidesof the pronotum in dorsal

view moreorless parallel,withthe

broad-est pointbeing about halfwaybetween its

anteriorandposterioredge.Thethreenew

have the pronotum in dorsal viewest near its anterior margin and distinctly

similar in pronotal structure to M.

mader-ensis; however, these two species are

eas-ily distinguished by the characters given

in the key.

Distribution.—Mauleus nigitus is known

Biology.— The insect hosts ofM. nigritus

Fig. 25

Holotype, female.—Color: Body black

ex-cept anterior aspect of head, collar, soscutumsteelblueand lateralportionsofmetanotum, propodeum, Tl blue Anten-

me-na with scape, ventral side ofpedicel and

Fl brownishyellow, remainderof pediceland flagellum brown. Legs with coxae

blue reflections, except basal and apicaltipsbrownish yellow; tibiaebrownish yel-

Sculpture.—

Clypeus and immediate

vi-cinity finely alveolate; remainder ofhead,mesoscutum, scutellum, frenum, medianpanels of propodeum alveolate; petiolefinely alveolate; gaster smooth except T6 and T7 coriaceous

width 1.4X height (42:30), 2.5X length(42.0:16.5); eye height 1.5X length (20:13),

POL, LOL as 3.0:6.5:9.0:4.0. Antenna with

length of pedicel plus flagellum 0.90X

scape, pedicel, annelli, Fl-6, club as 15.0:

4.0:1.0:4.5:4.5:4.5:4.0:4.0:3.5:8.0; relative

sensilla in two rows on each

22 Journal of

ventral patch of micropilosity Mesosoma

length 1.5X width (26.5:18.0); pronotum

with humeral angles acute (Fig. 25);

parallel; propodeumwith regionbetween

basal foveae depressed (this depression

shortermesally than laterally and

bound-edposteriorlyby weakcarina), plicae

fad-ing out before reachfad-inganteriormarginof

propodeum, nuchal region raised,

al-most on anterior margin of propodeum.

sub-marginal, sub-marginal, postsub-marginal, stigmal

veins as 37:23:15:8; costal cell with 1

com-pleteand 1 distal partialrowofsetae;

bas-al cell bare; basal vein with row of setae;

speculum open posteriorly Petiole length

1.1X width(9:8).Gaster length 1.4X width

(43:31), 0.66X combined length of head

gastral length;ovipositorsheaths hardly extending beyond hind

marginof T7

Allotype, male.—Similar to female

ex-cept:bodylength1.8 mm Head withratio

ofMOD, OOL, POL, LOLas3.5:4.0:7.0:3.0.

Antenna with length of pedicel plus

lengths of scape, pedicel, annelli, Fl-6,

club as11.0:3.0:1.0:4.5:4.5:4.0:4.0:4.0:4.0:9.0;

relative widths of Fl, F6, club as 3.0:3.5:

distrib-uted over funicular segments in many

rows; setaereclinate,nearlyabsent.Gaster

length 1.1X width (29:26).

Diagnosis.—Mauleus venetus is

Mau-leus species by the acute humeral angles

of the pronotum, the broad depression

propo-deum, andthe lack ofsculptureover most

of the median panels of the propodeum.

iligneus—thecoloration ofthe two species

isbrighterblue inM. venetus Inaddition,

fu-nicular segment in both sexes, whereasthey are arranged in only one row in M.iligneus.

Etymology.— The species name comes from the Latin word venetus, meaning

blueor sea-blue, and refers to the

Type Material.— The holotype (IRCW) is

a female, collected in Grant Co (T6N,

26.V.1985 Six paratypeswerecollected as

follows (CNCI, IRCW, UCDC, USNM):

1 female United States. ILLINOIS:

Uni-versity of Illinois South Farms, near

Champaign, 26.V.1985, 1 male

MICHI-GAN: Midland Co., 2.VII.1943, 1 male

male WISCONSIN: Grant, T6N, R6W,

S17, 3-8.VI.1976 (gypsy moth Malaise

trap), 1 female, 14-21 VI.1976(gypsymoth

Biology.— The host(s) of M. venetus are

unknown.

ACKNOWLEDGMENTS

I thank Melissa Bennett and two anonymous

re-viewsfor their careful reading of this manuscript I

also thank thefollowing peopleforthe loan of

ma-terialusedin this study: Dr J S.Noyes, TheNatural

HistoryMuseum,London, ENGLAND(BMNH);Dr.

W.J Pulawski, CaliforniaAcademy of Sciences,San

Francisco,CA(CASC);Dr F G.Andrews,California State Collection of Arthropods, Sacramento, CA(CDAE);Dr G A P Gibson,CanadianNational Col-

lection, Ottawa,ON(CNCI);Dr. W E LaBerge,

Illi-nois Natural History Survey,Champaign,IL (INHS);

S Krauth, University of Wisconsin, Madison, WI(IRCW); Dr R. W Brooks, Snow Entomological

(SEMC);Dr E E Grissell, United States Nationalseum, Washington, D.C. (USNM) The acronym for

Mu-the collection of Mu-the BohartMuseumat theUniversity

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J RES Vol 4, 1995, pp 25-32

(Hymenoptera: Vespidae): Nesting and Evaluation of

Abstract.— An account is given ofsome aspects of the nesting of Rolandia angulata (Richards)(Masarinae) This wasp nests in multicellularburrows in

compacted sandy soil. The sloping trance tothe burrow, notsurmountedby a turret, isconcealed beneatha

en-pebble, a plantletor a suitableitemof debris.The mainthrust of the shaft is verticallydownwards,however,at intervals

it curves outwards to end in a horizontal cell so that each cell, except the last excavated and

thereforedeepest one, appears tobe accessed bya lateral shaft. Thecells areunlined The tecture of the nest and the method of its construction are discussed The association between

archi-Rolandia angulata andGoodenia (Goodeniaceae)flowers, the source ofnectar and

pollen, is

evalu-ated It isshown that the associationwith Goodeniapinnatifida Schldl., at least, is mutually

bene-ficial. Indeed it is suggested that in some areas, at some times, R angulata may be the mostimportantpotential pollinator of thisplant.

The genus Rolandia Richards, 1962 is (28.03S, 148.30E), and 85 km E of St

listed by van der Vecht and Carpenter George (28.03S, 148.30E) (27-29.X.1993,

Meade-Waldo, 1911 As the analysis on Thereisonlyone publishedobservationwhich this isbased has not yet beenpub- on the nesting of Rolandia—that of R. ma-

in the present paper Rolandia is restricted oblique, blindly ending burrow in sandy

Hous-R maculata (Meade-Waldo) and R. hous- ^^^ is currently investigating further the

third, R borreriae Snelling, from Northern ^^^^i^^ available for comparison

angii-(Richards), the subject of this paper, from ^"^" published m the present paper were Queensland and New South Wales undertaken by the authors during the

r, ^ J- 1 I u u J J course of a fieldtrip to Australia in Rolandia angulata has been recorded , ^^^^ w ,

from southwestern Queensland and , , , »„

n u noo/ic 1/ir /.niri /o- u j gulata, the forage plants, and thebee Cunnamulla [28.04S, 145.40E (Richards ; « i.u r i x x ^u a i-

visi-.^^„ T^ ,

, , ^ , . tors to the forage plants), the Australian

, ,

'^ t> / National Insect Collection, Canberra [R.

m thenorthto90 km WofCobar, Barnato

,,,^,,,,^,,1 and the Australian NationalTanks [31.38S, 144.59E] (about 400 km

the south, and from three sites to the east NESTING

of Cunnamulla, 80 km E of Cunnamulla Description of the Nesting Areas.— Two

catedbetweenStGeorge and Cunnamulla

of St George on 28 October and the other

80 km east ofCunnamulla on 29 October,

in dry open scrub dominated by Acacia

(Fig 2). Both areas had recently received

rain intheformoflocalized

thundershow-erswhich had resulted in agrowth of

an-nuals which were in flower on the road

vergesand in thelow lyingareas Thesoil

inboth areas was but

fri-able It was increasingly moist at least to

Flozvers Visited.— The

rec-ord offlowervisitingbyR angulataisthat

Goodenia cycloptera R.Br, inC.Sturt

(Good-eniaceae) atBarnatoTanks In the present

of aprostrateherb also identified as G

cy-cloptera (Fig. 11) and an erect herb

Good-enia pinnatifida Schldl. (Fig. 6) in the twoareas where nests were discovered and

also in the area 85 km east of St George

Atthemosteasterlysite,thatis85km east

Cunnamulla it was observed that G

whereasG puinntifidabecameincreasingly

common andatthesite80kmeastof

fol-lowed a similar pattern to that of G

pin-natifida.

Visits to the flowers by female R

angii-lata were abundant by lOhOO and

afternoon Visits became fewer in the late

afternoon and ceased after IThOO Males

only laterin thedayto visit them for

nec-tar.

All plants flowering together with the

twoGoodeniaspecies,mostnotablyseveral

Asteraceae and a Wahlenbergia species

flow-er visitors. None was being visited by R.

angulata

Provision.—Provision in the form of a

firm, white pollen loafwas obtainedfrom

each ofthree cells. Pollen from the loaves

was examined microscopically and found

to match that obtained from the Goodenia

flowers

the flowers but rested on a neighbouring

plant, forexample on a grass stem, or on

copu-lationsatflowerswereobservedanda

nest-exca-vatingfemalewasnoted Several instances

ground next to plants and of "hot

pur-suit" were noted

Description oftheNest.— Thenest (n =

8)

consists ofa subterranean burrow (Fig 5)

clearing(Fig 3).Theentranceisconcealed

item of debris (Fig. 4). It is a

3-4 mm in diameter, not surmounted by

a superstructure For approximately the

first10 mm the shaft slopes gently

verti-cally downwards, however, at intervals itcurvesoutwardstoendinahorizontalcell

so that each cell, except the lastexcavated

and therefore deepest one, appears to be

accessed by a lateral shaft. These "lateral

shafts" radiate out through 360° each

continues constant The first cell in the

be-tween 180 and 370 mm, the "lateral

shafts" were 30 mm in length, and the

cells 13 mm long and 4 mm in diameter

packed with sand afterthe cell which

cells are unlined

Provisioning.— Water is not required forthe excavation of the nest as the sandysoil, though compact, is friable. The sand

extracted from the burrow is carried out

oftheshaftheldbetweenthehead andthe

ammochaetae. Whilst excavation is in

progress, the female, when leaving the

nest,backsout Duringtheinitialstagesofburrow excavation the extracted sand isdropped in flight in a more or less con-

stant area to one side of theentrance and

dropped further from the nest in a

con-stant arc about250 mm from theentrance

develops

in a sealed, fully provisioned cell

indicat-ed that mass provisioning is practised

with-28 Journal of

shaft leading to thenext cell.

Thefullyfedlarva spinsa whitecocoon

which completely fills the cell. Like the

sealed cell it is therefore rounded at the

inner end and truncate at the outer end

Discussion of Nesting.— Thenest ofR

an-gulata is essentially similar to that of R.

(pers. comm.) as a vertical burrow in

sandy soil, about 300 mm deep and

un-lined, with unlinedcells atthe lowerend

Thusthebasicnesttypeforthesetwo

spe-cies of Rolnndia can be defined as a

mul-ticellularsub-verticalburrowinhorizontal

ground excavated by the nester, without

an entrance turret and with excavated

When comparedwith the sevenbasic nest

types recognized for the Masarinae as a

whole by Gess and Gess (1992) this

ap-pears to fitNest type1 except for the lack

ofan entranceturret. Whenthe methodof

construction is compared with that of the

specieslistedforthisnesttype, abasic

dif-ference is apparent Water is used in the

excavation and construction of Nests of

type1, likenestsoftypes 2and 3,whereas

the nests of the two Rolandia species are

is possibledue to the friablenature of the

soil in which they are sited. As noted in

GessandGess(1992)nestinginfriable soil

than primitive as in the Pompilidae and

Sphecidae.Thenest typeofthetwo

Rolan-dia species is therefore seen as a sub-type

which canbe derived from Nest type 1.

Both species of Rolandia carry sand,

ex-tracted from the shaft, out of the burrow

held between the head and the

am-Fig 5 Plan of vertical section of a nest of Rola7tdia

attgulata Cells 2—5 followed cell I, radiating out in

sequence through 360° and at successively greater

50mm

Figs 6-10 6, Goodeniapinnatifida, an erect herb; 7, a flower of Goodeniapiiiiiatifida,indusium concealedby

the bases of the adaxialpetals;8, Rolandia angulata entering a flower of Goodeniapinnatifida, indusiumof the flower exposed (X 2.8); 9, Rolandia angulata in nectar drinking position in a flower of Goodenia pinnatifida,

indusium of the flower fittingsnugly overthe wasp'smesosoma (X 2.8); 10, Rolandia angulatawithdrawingfroma flower of Goodeniapinnatifida,showingdustingofpollen onthe head,dorsumof theprothorax, and

anterior part of themesoscutum(X 2.8).

ammochae-tae fringing the genae is a generic

char-acter (Snelling 1986) and it is therefore

excavate their nests in a similar manner

The only other vespid genus recorded as

the spoils of excavation is Pterochihis

in vertical burrows in friable soil by two

specieshasbeendescribed

(Isely 1914and Evans 1956).

EVALUATION OF ASSOCIATION

WITH FLOWERS VISITED

Both R angulata males and females

ob-tain nectar and for their

nec-tarand pollen for provisioning their

nest-cells apparentlysolelyfromGoodenia

flow-ers. They are therefore probably ant on Goodenia flowers To determine

depend-whether or not the association between

thewasp and theflowers is mutually

ben-eficial or not, that is whether or not thewasp in addition pollinates the flowers,

requires a consideration of the functionalmorphology ofthe flowers, thebehaviour

of the wasps in the flowers, and wasp/flowerfit.

ad-axial petalsare differentiatedin their

Journal of

Figs 11-13 11, Goodeniacycloptera, a prostrate herb; 12, three flowers of Goodenia cycloptera, indusium cealedbythe bases of the adaxialpetals;13,Rolandia angulata in nectar drinking position in aflowerofGoodenia

con-cycloptera,indusiumof the flower presseddownonthewasp'sfoldedwings(X 3.3).

indusium in the flower (Figs 7and 12) so

thatit is

seeks the nectar at the base of the flower

(Figs 8 and 13). The indusium is a cup at

the top of the style and the surrounding

stigmatic initial that collects and retains

pollenfromthe stamens,whichdehisce in

thebud, and presents it thus tothepollen

1992) Later the stigmatic initials mature

and grow outofthe indusium andcollect

R. angulatawhen visitingtheflowersfor

nectaralways alights on thelowerlip and

indu-sium so that, if the flower is in the pollenpresenting phase, thewaspreceives pollen

onitshead When it isin thenectar ing position in a flower of G. pinnatifida,

the indusium but, when it is in a flower

of G

cycloptera, which is deeper, it is the

folded wings which are pressed beneaththeindusium (Fig 13).On emerging from

a flower of G canbe

seen tohave beenwell dusted withpollen

thus laden with pollen to a flower with

receptive stigmas the wasp would be

ide-ally suited to pollinate it It is not clear,

effec-tively pollinate the deeper flowers of G

cycloptera. It ispossiblethat pollenmay be

transferred from its head to a receptive

transferred

distribution, R. angulata is a potential

pol-linator, at least,of the widespread species

G pinnatifidawithwhichit thereforehas a

atnoneofthesiteswasR.angulatathe sole

visitor to the Goodenia flowers

fascicu-laris F.Muell & Tate at Kondar to the

southeast, was a

relatively abundant itor to G

vis-pinnatifida. Furthermore the

flowers of both species of Goodenia,

par-ticularly those of G cycloptera, were

com-mon species was Leioproctus

{Chri/socolle-tes) moretonianus (Cockerell) (Colletidae)

which was also recorded from

Michener (1965). Less common visitors

were a second, but slightly smaller,

spe-cies of Leioproctus {Chrysocolletes) and a

species of Megachile (Megachilidae)

fur-ther species of Megachile, an additional

coUetid, a halictid, a few anthophorids

the bees were uncommon at the site 27

km west of St George and at the site 80

species of Leioproctus (Chrysocolletes) were

absent At the latter site a third much

smaller species of Leioproctuswas also

re-corded However, R. was the

only abundant visitor suggesting that in

pinnatifida.

ACKNOWLEDGMENTS

Thefollowingarethankedwithappreciation:Terry

Houstonof theWesternAustralianMuseumfor mission to quote hisunpublished observations con- cerning thenesting of Rolandia maciilata; Jo Cardale

per-of the Australian NationalInsect Collection,Canberra

forfacilitatingthe identification offorageplants; sten Cowley, Australian National Herbarium, Can-

Kir-berra for the identifications of theforageplants;Alan

Weavingof theAlbanyMuseumforproducingblack

and white negatives from the authors' colour parencies for Figs1-4 and 6-13;and theSouth Afri- canFoundationfor ResearchDevelopmentfor a roll-

trans-ing support grant to F.W.Gess,whichmadepossiblethe fieldwork in Australia. The manuscriptwas re-

viewed byKarl V. Krombein and ananonymous

re-viewer,whoarethankedfor their constructive

com-ments.

LITERATURE CITED

Bohart, R.M 1940. Arevision of theNorth American

species of Pterocheilusandnoteson relatedera (Hymenoptera, Vespidae) Annals of the En- tomological Society of America33(1): 162-208 Carolin, R.C.,M.T.M.Rajput,andD Morrison 1992 Goodeniaceae In:

gen-George, A.S ed., Flora of

Aus-tralia,volume35: Brunoniaceae, Goodeniaceae.

Can-berra: Australian Government Publishing

Ser-vice, pp 4-300.

Evans, H.E 1956. Notes onthebiologyof four cies of ground-nestingVespidae(Hymenoptera) Proceedings of the Entomological Society of Washing- ton58(5):265-270.

spe-Gess, F.W. and S.K Gess 1992 Ethology of threesouthern Africangroundnesting Masarinae,two

Celonites species and a silk-spinning Quartinia species, with a discussion ofnestingbythe sub- family as a whole (Hymenoptera: Vespidae).

Journal of Hymenoptera Research 1(1): 145-155.

Houston,T.F 1984. Bionomicsof a pollen-collecting

wasp, Paragia tricolor (Hymenoptera: Vespidae: Masarinae), in Western Australia Records of the

Western AustralianMuseum11(2): 141-151.

Isely, D 1914. The biologyof someKansasidae Kansas University Scietice Bulletin (2)8(7):

Eumen-233-309.

Meade-Waldo G 1911. Notesonthefamily

Masar-idae (Hymenoptera), with descriptions of a newgenus and three newspecies. Annals andMaga-zine of NaturalHistory(8)8: 747-750.

Michener,CD 1965. A classification of the bees of AustralianandSouth

Journal of

oftheAmericanMuseumofNaturalHistory 130:

1-362.

Richards,O.W 1962. Arevisional study of the Masarid

wasps {Hymenoptera, Vespoidea). London: British

Museum(natural History).

Richards,O.W 1968. Newrecordsand newspecies

of Australian Masaridae (Hymenoptera:

Vespo-idea) Journal of the Australian Entomological

Soci-ety 7: 101-104.'

Snelling,R.R. 1986. Thetaxonomyand nomenclature

ofsome Australian paragiinewasps

(Hymenop-tera: Masaridae) Contributions in Science 378:

Vol 4, 1995, pp 33-10

Two Strikingly Distinct Sympatric Colour Forms of

Rolandia angiilata (Richards) (Hymenoptera:

Abstract.—The maleof Riekianocatunga Richards fromsouthern Queensland,the maleand two

strikingly distinctsympatric colourformsofRiekia confluens (Snelling),comb,nov., fromWestern

Australia,andthe maleof Rolandia angnlata (Richards) from southernQueensland are described.

The charactersdistinguishingR. nocatungaand R. confluens are discussed

TheAustralian Masarinaeand their

nat-ural history are relatively poorly known.

Duringa recentcollectingtrip toAustralia

by the author, S.K.Gess and R.W.Gess,

material collected included males ofthree

species, Riekia nocatunga Richards, Riekia

confluens (Snelling),comb, nov.,and

Rolan-dia angnlata (Richards), hitherto described

distinct sympatric colour forms of Riekia

thesewas previously unknown and helps

to elucidate the identity of a single male

to place it in a species

Riekia Richards, 1962 and Rolandia

Meade-Waldo, 1911 by van der Vecht and

Car-penter (1990) on thebasis ofanas yet

pub-lished and maybestudied, thepresent

au-thor considersitbest tocontinuetoaccept

Riekia and Rolandia as genera in theirown

right.

Institutions in which the material

stud-ied is are: Albany Museum,

Grahamstown, South Africa (AMG); tralian National Insect Collection, Canber-

Perth (WAM).

Riekia nocatunga Richards

Riekia nocatunga Richards, 1962: 55-57, female.

This specieswasdescribedfrom 7 miles

NofNocatunga [on mapsas Nockatunga]

paratype)

his original description with regard to the

de-tails. Two of these females, in the tionofthe Natural History Museum, Lon-don, have been examined by the present

collec-author Theyareconfirmedasbeing R. catunga They are less melanistic than the

areas are less extensive and some, in

Journal of

ticularthoseonthepronotum and tergites each side of mesoscutum where its

fur-1 and 2, are brownish rather than black, rows meet pronotum (usually but not The light areas are more extensive and ways present in females) absent in boththeircolour is a strongyellow ratherthan males examined,

Snelling (1986) figured and briefly dis- slightly longer than interantennal

dor Homestead (25.08S, 116.54E) in West- as long as scape (with radicle), one and aernAustralia Whereas it agreedgenerally third times as long as wide at its distal

with the originaldescriptionofR. nocatun- end, and one and one fifth as long as the

ga, Snelling stated that there was no cer- second flagellomere. Last three

be-species and that he suspected that it was neath; ultimate flagellomere narrowing

not He listed some discrepancies, allow- apically and distinctly curved to form a

tobe availablebefore the specific status of Genitalia(Figs3and4);parameralspine

Recentlycollected Riekia material, ofrel- length and not hook-like apically; ventral

onomicquestion, consistsofassociated fe- and short, subtriangular, without a

and Western Australia The specimens Length 7.5-7.8 mm; length of forewing

from Queensland are certainly R. nocatun- 5.8 mm, hamuli 14.

ga, as established by the comparison by New Material Examined.—Queensland:

from WesternAustralia on theotherhand Goodenia fascicularis F.Muell & Tate,represent two strikingly distinct colour Goodeniaceae); SouthwoodRoad, western

cially similar looking to Riekia nocatunga (27.56S, 149.30E), 26.x.1993 (F.W.,S.K.&

this form show the characters noted by of St George (28.03S, 148.30E), 27.X.1993Snelling for hismale. (F.W.,S.K.& R.W.Gess) 8 females, 1 maleMale.—

verysimilarto thatoffemalebutdiffering Schldl., Goodeniaceae) Two females and

inthat the following parts are palelemon- 1 male in ANIC; 2 females inWAM; rest

yellow: small streakonscapesdistally,en- of material in AMG.

spotbetween and above antennal sockets,

^ Riekia sp. Snelling, 1986: 6, 8 and Figs 10,

19-sinus(not risingabove upperlimitof

fron-21 maletal spot and separated from it by about

width of antennal socket), uninterrupted Snelling (1986)

erroneously described

Figs. 1-4. Riekia nocatunga 1, dorsofrontal view of vertex and dorsal view of anterior third of thorax of female (X17.6); 2, dorsal view of posterior third of thorax of female (X17.6); 3-4, ventral and ventrolateral

views of genitalia ofmale(X57).

Figs 5-8. Riekiaconfluens 5, dorsofrontalviewof vertexanddorsalviewof anterior third of thorax of female (X17.6); 6, dorsalviewof posterior third of thorax of female(X17.6); 7-8, ventraland ventrolateralviewsof genitaliaofmale(X63);[all lemon-yellowcolour form]

generic key, the species with its

Riekia Richards, 1962 Further, with the

se-tae attheapexof the mid- andhindtibiae

(a specificcharacter) the charactersgiven

by him in his ofthe species are

all common to both coiiflnciii^ and

nocn-tungn and may therefore be considered

to be generic characters pertaining to

Riekia. Similarly the confluence of the

(leading to the name confluens), or

ca-36 Journal of

rina, is a character shared with

confluens (and nocatiinga) generically

apart from Paragia.

Material collected at a single site near

distanc-esof 165km and260kmin awesterlyand

west-south-westerlydirection respectively

from the typelocalitiesofconfluens (16km

WSW Lyons River Homestead, 24.38S,

115.20E—paratypefemale, and36km ESE

Minnie Creek Homestead, 24.00S,

115.42E—holotype female) consists of 53

females and 4 males The material is

di-visible into two very distinct groups, one

black and reddish-brown and the other

black and lemon-yellow. Whereas

intra-groupvariability isnegligible, inter-group

differences with respect to colour

gener-ally and to colour pattern on the

metaso-ma in particular are striking.

in-termediate forms initially led to thebelief

that two species were represented.

be found that supported this view and it

inoverall facies is a product ofthe

differ-ences in colour and colour pattern. The

reddish-brown colour form, consistent

with the description of Snelling's two

para-type), is represented in the present

mate-rialby femalesonly,thelemon-yellow

col-our form bybothsexes Tofacilitate

intra-specific comparison, descriptions limited

to colour pattern are givenofboth female

inter-specificcomparisonwithnocatiingathe

de-scription of the male is

detail.

description it isincorrectly stated that the

probasitarsus is

inad-vertent error as it is in fact slightly less

than halfas wide as long

Reddish-brown (RB) Colour FormFemale.—Black The following reddish-brown: mandiblesotherthanforteethand

fused spots) between and above antennalsockets and narrowly separated ventrally

from clypeal marking, variously sizedelongate spot in upperhalfofeach ocularsinus (rising to level of anterior ocellus

and lateral spotssometimesbroadlyfusedabove], a large oval spotbehind eyes dor-

sally, entire upper surface of prothorax

other than for narrow streak bordering

posterior margin, tegulae except for cleartestaceous central spot and narrow mar-gin, scutellar disc other than for its ante-

me-dially, a large spot on angles of deum, large spot on mesopleuron below

propo-tegula, distal end of mid- and hindcoxae,

part of foretrochanter and whole of andhindtrochanters, femur, tibia and tar-

than for anterior declivity, tergite 2 other

than for a variously developed anterior

black band which may be triangularly

produced in the middle and for a pair of

on posterior margin, an anteriorly ing triangular spoton posterior marginoftergite 3, tergite 4other than for a narrow

transverse posterior band on gastral

ster-nites 2-5 (those of sternites 2 and 3 and sometimes 4 strongly and widely anteri-orly produced in the middle)

Lemon-yellow (LY) Colour Form

Female.—

(Figs 5 and 6). Black The

neath, trochanters of mid- and hindlegs, eral and anterior margins, single large

hind-heavily black suffused) and inner and coxae, entire outer aspect of midfemora,

pect and distal end oftibiae, all tarsomer- distalends),narrowlongitudinalstreakon

es, posterolateral portions of dark mark- all basitarsi, transverse posterior band onings on sternites2-4 and sternite 6 apical- gastral sternites 2-5 (all with median lobe

ly (allblack-suffused) Thefollowing lem- strongly and widely anteriorly produced,on-yellow: proximal half to two-thirds of together giving the effect of a wide me-

dis-arcuateblack lines ondistal half, a pair of tance; firstflagellomere slightlymorethanirregularly shaped frontal spots between half (0.53) as long as scape (with radicle),and above antennae, a narrow streak on one and three quarter times as long as

of pronotum, tegulae except for clear tes- Tergite 7 truncate and narrowly

trans-taceous centralspotand narrowmargin, a verse

apically

large transverse suboval shield-like spot Genitalia (Figs 7and 8); similartothose

on scutellum (not quite reaching anterior of R nocatungabut differing in detail

par-margin), a small spoton raised partof

ax-ticularly with respect to the form of the

illae,a largespotonanglesofpropodeum, inwardly directed lobe of the volsella

a small spot on mesosternum anterior to Length 7.8-8.3 mm; length of forewing

coxalcavities, a small lateral spoton mid- 5.5-5.7 mm; hamuli 12-13

and hindcoxae, distal half of outer aspect Material Examined.— Western Australia:

of forefemora, distal spot on outer aspect 8 km NE of Carnarvon (24.51S, 113.45E)

all tibiae (except distal ends), wide ante- (F.W.,S.K.& R.W.Gess) 16 RB females, 29

tergites 1-5, tergite6 (other than for vari- flowers ofLechenaiiltia sp.,Goodeniaceae),

ously developed posteriorly pointing V- 4 RB females, 1 LYfemale (atwater), 2 RB

shapedblackmarkleaving lateraland me- females, 1 LY female (without biological

dian spots or almost eliminating these), data). Two RB females, 2 LY females andtransverse posterior band on gastral ster- 1 LY male in both ANIC and WAM; rest

lobed with median lobe strongly and Discussion.—R confliieiis may be

Male.—

indi-very similar to that offemalebutdiffering cated morphological characters,

in that the followingparts are palelemon- InR. confluens theratioofPOL(distance

yellow: entire anterior aspect of scapes, between posterior ocelli):OOL (distance

Journal of

catunga it is 1:1.0-1.1. relative proportionsofthe firsttwo

flagel-In R confliiens the pilosity of the pro- lomeres, in the form of gastral tergite 7

dividual hairs being slender and only Present data would indicate that the

slightly curved apically whereas in R no- distributionsof the species arewidelycatunga the pilosity is much denser and arated, R nocatungci occurring in New

sep-the individual hairs are more robust and South Wales and Queensland and R

con-markedly and evenly curved. fluens in Western Australia

In R confluens thescutellum (Fig 6)has

its disc posteriorly widely and evenly to Rolandia angulata (Richards)bluntly rounded and abruptly declivitous

^.^^^.^^,^^^^^^^^j^.^j^^^^^^ ^^^^. ^q^_^q2, female,

over-^^,^„^^,.^ ^^^^^^^^^ (Richards), Snelling, 1986: 8.

hangs; theposterolateral freeedgeofeach

lateral wing is almost straight. The meta- This species was described from

ible from above In R. nocatunga the scu- Tanks [31.38S, 144.59E] (about 400 km

tellum (Fig 2) has itsdisc posteriorly nar- south of Cunnamulla), New South Wales

thereforeof even depth; it is sloping, vis- Insects, Canberra, and found to be

identi-ible from above cal.

it is two and a quarter times wider than lighter in hue and brighter and contrast

whereas in R. nocatunga it is 2-4, most marking, and the distinct median and

recogni-theapexofthemetatibia is1-2,most com- tion of the male even in the field,

The species differ in the female sex in than interantennal distance; first

flagello-the form of the pronotum as seen from mere three times as long as broad at itsabove In R. confluens (Fig 5) the anterior distal end, about two-thirds as long as

and lateral margins describe a semicircle scape (withradicle) [three-quarters aslong

whereas in R. nocatunga (Fig 1) the as scape withoutradicle],anda littlemore

so that the anteriorand lateralmarginsdo flagellomeres 3-9 subequal in length andnot describe a semicircle but the anterior with the exception of 3 wider than long;

margin appears subtransverse. 3-7 increasing in width; 7 and 8 of same